Handbook of the Bruchidae of the United States and Canada Introduction to the Acrobat Pdf Edition

Handbook of the Bruchidae of the United States and Canada Introduction to the Acrobat pdf edition

The Acrobat pdf version of this publication, though identical in content to the print version, differs slightly in format from the print version. Also, in volume 2 the items on the errata list for the print version have been corrected.

[THIS PAGE INTENTIONALLY BLANK] United States Department of Agriculture Handbook of the

Agricultural Research Bruchidae of the United Service Technical States and Canada Bulletin Number 1912 November 2004 (Insecta, Coleoptera)

Volume I

I II United States Department of Agriculture Handbook of the

Agricultural Research Bruchidae of the United Service Technical States and Canada Bulletin Number 1912 November 2004 (Insecta, Coleoptera)

John M. Kingsolver

Volume I

Kingsolver was research entomologist, Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture. He is presently research associate with the Florida State Collection of Arthropods.

III Abstract Hemisphere. It provides the means to identify these insects for taxonomists, students, museum curators, biodiver- Kingsolver, John M. 2004. Handbook of sity workers, port identifiers, and ecolo- the Bruchidae of the United States and gists conducting studies in rangeland, Canada (Insecta, Coleoptera). U.S. Depart- pasture, and forest management in the ment of Agriculture, Technical Bulletin United States and Canada. 1912, 2 vol., 636 pp. Mention of commercial products in this Distinguishing characteristics and diag- publication is solely for the purpose of nostic keys are given for the 5 subfami- providing specific information and does lies, 24 genera, and 156 species of the not imply recommendation or endorse- seed beetle family Bruchidae of the Unit- ment by the U.S. Department of Agricul- ed States and Canada (including Hawaii). ture over others not mentioned. Associated data for each species description include a history of the name, syn- While supplies last, single copies of onymical names, type specimen infor- this publication can be obtained at no mation, geographical distribution, host cost from John M. Kingsolver, Florida plants, and parasitoids. Appendices give State Collection of Arthropods, P.O. Box species, host, fossil, and parasitoid lists 147100, Gainesville, FL 32614–7100. as well as a glossary of terms and a bibli- Copies of this publication may be pur- ography. chased from the National Technical In- Bruchidae are found on every major land formation Service, 5285 Port Royal Road, mass except Antarctica and New Zea- Springfield, VA 22161. ARS has no addi- land. Eggs are usually laid on the seed or tional copies for free distribution. fruit of a plant suitable for development of the larva. Immature stages are spent inside seeds that have been excavated by The U.S. Department of Agriculture (USDA) larval feeding. Adults live free and feed prohibits discrimination in all its programs and on pollen and nectar. activities on the basis of race, color, national Johnson (1970) estimates that approxi- origin, age, disability, and where applicable, sex, mately 84 percent of the known hosts of marital status, familial status, parental status, Bruchidae are in the plant family Legu- religion, sexual orientation, genetic information, minosae. The remaining hosts are scat- political beliefs, reprisal, or because all or part of tered among 31 other families. Sixteen an individual’s income is derived from any public plant families support larval feeding in assistance program. (Not all prohibited bases the United States and Canada. Several apply to all programs.) Persons with disabilities who require alternative means for communication species of Bruchidae, especially those of program information (Braille, large print, with a cosmopolitan distribution, are no- audiotape, etc.) should contact USDA’s TARGET torious pests of stored leguminous seeds. Center at (202) 720-2600 (voice and TDD). To Keywords: Coleoptera, Bruchidae, Am- file a complaint of discrimination, write to USDA, blycerinae, Bruchidiinae, Bruchinae, Ky- Director, Office of Civil Rights, 1400 Independ- torhininae, Megacerinae, United States, ence Avenue, S.W., Washington, D.C. 20250- Canada, Hawaii, taxonomy, insect-plant 9410, or call (800) 795-3272 (voice) or (202) 720- interactions. 6382 (TDD). USDA is an equal opportunity This is part of a series of studies of provider and employer. bruchid genera contributing to a comprehensive database for this important seed-feeding beetle family in the Western

November 2004

IVii 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 ...... 24 Genus Caryedon Schoenherr ...... 24 Caryedon serratus (Olivier) ...... 24 Tribe Pachymerini Bridwell ...... 26 Genus Caryobruchus Bridwell ...... 26 Caryobruchus gleditsiae (Linnaeus)...... 26 Subfamily Kytorhininae Bridwell ...... 28 Genus Kytorhinus Fischer von Waldheim ...... 28 Kytorhinus prolixus (Fall) ...... 28 Subfamily Amblycerinae Bridwell...... 30 Tribe Amblycerini Bridwell ...... 30 Genus Amblycerus Thunberg ...... 30 Amblycerus eustrophoides (Schaeffer) ...... 32 Amblycerus ireriae Romero, Johnson, and Kingsolver ...... 33 Amblycerus nigromarginatus (Motschulsky)...... 34 Amblycerus obscurus (Sharp) ...... 34 Amblycerus robiniae (Fabricius) ...... 35 Amblycerus schwarzi Kingsolver ...... 37 Amblycerus vitis (Schaeffer) ...... 38 Tribe Spermophagini Borowiec ...... 39 Genus Zabrotes Horn ...... 39 Zabrotes amplissimus Kingsolver ...... 42 Zabrotes arenarius (Wolcott)...... 42 Zabrotes bexarensis Kingsolver ...... 43

iiiV Zabrotes chandleri Kingsolver ...... 44 Zabrotes chavesi Kingsolver ...... 44 Zabrotes cruciger Horn...... 45 Zabrotes cynthiae Kingsolver ...... 46 Zabrotes densus Horn...... 47 Zabrotes eldenensis Kingsolver...... 47 Zabrotes humboldtae Kingsolver ...... 48 Zabrotes obliteratus Horn ...... 49 Zabrotes planifrons Horn ...... 50 Zabrotes spectabilis Horn ...... 50 Zabrotes stephani Kingsolver ...... 51 Zabrotes subfasciatus (Boheman) ...... 52 Zabrotes subnitens Horn...... 55 Zabrotes sylvestris Romero and Johnson ...... 55 Zabrotes victoriensis Kingsolver ...... 56 Subfamily Bruchinae Pic ...... 58 Tribe Megacerini Bridwell ...... 58 Genus Megacerus Fahraeus ...... 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) schaefferianus Bridwell ...... 68 Tribe Bruchini Bridwell ...... 69 Genus Bruchus Linnaeus ...... 69 Bruchus brachialis Fahraeus ...... 70 Bruchus pisorum (Linnaeus) ...... 72 Bruchus rufimanus Boheman...... 74 Tribe Bruchidiini Bridwell ...... 75 Genus Borowiecius Anton ...... 75 Borowiecius ademptus (Sharp) ...... 75 Genus Bruchidius Schilsky ...... 76 Bruchidius cisti (Fabricius)...... 77

VIiv Bruchidius villosus (Fabricius) ...... 78 Genus Callosobruchus Pic ...... 79 Callosobruchus chinensis (Linnaeus) ...... 80 Callosobruchus maculatus (Fabricius) ...... 82 Callosobruchus phaseoli (Gyllenhal) ...... 86 Callosobruchus pulcher Pic ...... 87 Tribe Acanthoscelidini Bridwell ...... 88 Genus Abutiloneus Bridwell ...... 88 Abutiloneus idoneus Bridwell ...... 88 Genus Acanthoscelides Schilsky ...... 89 Acanthoscelides aequalis (Sharp) ...... 96 Acanthoscelides alboscutellatus (Horn) ...... 97 Acanthoscelides atomus (Fall) ...... 98 Acanthoscelides aureolus (Horn) complex...... 99 Acanthoscelides baboquivari Johnson ...... 101 Acanthoscelides bisignatus (Horn)...... 102 Acanthoscelides biustulus (Fall) ...... 103 Acanthoscelides calvus (Horn)...... 104 Acanthoscelides chiricahuae (Fall) ...... 105 Acanthoscelides compressicornis (Schaeffer) ...... 106 Acanthoscelides comstocki Johnson ...... 107 Acanthoscelides daleae Johnson ...... 108 Acanthoscelides desmanthi Johnson ...... 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 helianthemum Bottimer ...... 116 Acanthoscelides herissantitus Johnson ...... 117 Acanthoscelides inquisitus (Fall)...... 118 Acanthoscelides kingsolveri Johnson ...... 119 Acanthoscelides lobatus (Fall) ...... 120 Acanthoscelides longistilus (Horn)...... 121 Acanthoscelides macrophthalmus (Schaeffer)...... 122 Acanthoscelides margaretae Johnson...... 123

VIIv Acanthoscelides mixtus (Horn) ...... 124 Acanthoscelides modestus (Sharp) ...... 125 Acanthoscelides mundulus (Sharp)...... 126 Acanthoscelides napensis Johnson ...... 127 Acanthoscelides obrienorum Johnson ...... 128 Acanthoscelides obsoletus (Say) ...... 129 Acanthoscelides obtectus (Say) ...... 130 Acanthoscelides oregonensis Johnson ...... 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 subaequalis Johnson ...... 151 Acanthoscelides submuticus (Sharp) ...... 152 Acanthoscelides tenuis Bottimer ...... 154 Acanthoscelides tridenticulatus Bottimer ...... 155 Genus Algarobius Bridwell ...... 155 Algarobius bottimeri Kingsolver ...... 156 Algarobius prosopis (LeConte) ...... 157 Genus Althaeus Bridwell ...... 158 Althaeus folkertsi Kingsolver ...... 158 Althaeus hibisci (Olivier) ...... 160 Althaeus steineri Kingsolver ...... 161 Genus Caryedes Hummel ...... 163 Caryedes helvinus (Motschulsky) ...... 164 Caryedes incensus (Sharp) ...... 165

VIIIvi Genus Gibbobruchus Pic ...... 166 Gibbobruchus cristicollis (Sharp) ...... 167 Gibbobruchus divaricatae Whitehead and Kingsolver...... 168 Gibbobruchus mimus (Say) ...... 169 Genus Lithraeus Bridwell...... 170 Lithraeus atronotatus (Pic) ...... 171 Genus Meibomeus Bridwell ...... 172 Meibomeus desmoportheus Kingsolver and Whitehead ...... 172 Meibomeus musculus (Say) ...... 173 Meibomeus surrubresus (Pic) ...... 174 Genus Merobruchus Bridwell...... 175 Merobruchus insolitus (Sharp) ...... 176 Merobruchus julianus (Horn) ...... 177 Merobruchus knulli (White)...... 179 Merobruchus lysilomae Kingsolver ...... 180 Merobruchus major (Fall)...... 181 Merobruchus placidus (Horn) ...... 182 Merobruchus terani Kingsolver ...... 183 Merobruchus vacillator (Sharp)...... 184 Genus Mimosestes Bridwell ...... 185 Mimosestes acaciestes Kingsolver and Johnson ...... 186 Mimosestes amicus (Horn) ...... 187 Mimosestes insularis Kingsolver and Johnson...... 188 Mimosestes mimosae (Fabricius) ...... 189 Mimosestes nubigens (Motschulsky)...... 191 Mimosestes protractus (Horn) ...... 192 Mimosestes ulkei (Horn) ...... 193 Genus Neltumius Bridwell ...... 194 Neltumius arizonensis (Schaeffer)...... 195 Neltumius gibbithorax (Schaeffer) ...... 196 Neltumius texanus (Schaeffer) ...... 197 Genus Sennius Bridwell ...... 198 Sennius abbreviatus (Say) ...... 200 Sennius cruentatus (Horn) ...... 201 Sennius discolor (Horn) ...... 202 Sennius fallax (Boheman) ...... 203 Sennius lebasi (Fahraeus) ...... 205

viiIX Sennius leucostauros Johnson and Kingsolver ...... 206 Sennius medialis (Sharp) ...... 206 Sennius morosus (Sharp) ...... 207 Sennius obesulus (Sharp) ...... 208 Sennius simulans (Schaeffer) ...... 209 Sennius whitei Johnson and Kingsolver ...... 210 Genus Stator Bridwell ...... 211 Stator beali Johnson...... 212 Stator bottimeri Kingsolver ...... 213 Stator chihuahua Johnson and Kingsolver ...... 214 Stator coconino Johnson and Kingsolver ...... 214 Stator limbatus (Horn) ...... 215 Stator pruininus (Horn) ...... 216 Stator pygidialis (Schaeffer)...... 218 Stator sordidus (Horn) ...... 219 Stator subaeneus (Schaeffer)...... 219 Stator vachelliae Bottimer ...... 220 Genus Stylantheus Bridwell ...... 221 Stylantheus macrocerus (Horn) ...... 221

Appendix I. Hawaiian Bruchidae ...... 223 Key to Hawaiian Bruchidae ...... 224 Host Associations of Hawaiian Bruchidae ...... 225 Appendix II. Synonymical List of the Bruchidae of the United States and Canada ...... 227 Extant Species ...... 227 Extinct Species ...... 237 Appendix III. Hosts of the Bruchidae of the United States and Canada, by Bruchid ...... 239 Appendix IV. Hosts of the Bruchidae of the United States and Canada, by Host Plant ...... 246 Appendix V. Natural Enemies of Bruchidae of the United States and Canada ...... 260 Glossary of Morphological Terms ...... 264 References ...... 269 Index ...... 319

Volume 2 (Illustrations) Volume 2 contains all figures referenced in Volume 1.

Xviii Acknowledgments

I gratefully acknowledge the assistance and History Survey, Urbana, IL; Michael Ivie, suggestions of the following individuals Bozeman, MT; C.D. Johnson, Flagstaff, AZ; who read or commented on the manu- Los Angeles County Museum, Los Angeles; script; C.D. Johnson, G.S. Pfaffenberger, Michigan State University, East Lansing, D.R. Miller, R.D. Gordon, the late D.R. MI; Museum of Comparative Zoology, Cam- Whitehead, the Systematic Entomology bridge, MA; Museum National d’Histoire Laboratory hymenopterists, and my late Naturelle, Paris; C.W. O’Brien, Tallahas- wife, Cynthia, who assembled the host and see, FL; Naturhistoriska Riksmuseet, parasite lists and the index. Stockholm, Sweden; Ohio State University, Specimens were loaned for this study by Columbus, OH; Purdue University, West the following institutions and individuals: Lafayette, IN; Karl Stephan, Red Oak, OK; American Museum of Natural History, New Texas A&M University, College Station, TX; York; Arizona State University, Tempe, AZ; Robert H. Turnbow, Ft. Rucker, AL; Na- Charles Bellamy, Escondido, CA; Bernice P. tional Museum of Natural History, Wash- Bishop Museum, Honolulu; British Muse- ington, DC; University of Arizona, Tucson, um of Natural History, London; California AZ; University of California, Berkeley, CA; Academy of Sciences, San Francisco; Cali- University of California, Davis, CA; Univer- fornia Department of Agriculture, Sacra- sity of Kansas, Lawrence, KS; University of mento, CA; Canadian National Collections, Michigan, Ann Arbor, MI; University of Ottawa, ON; Carnegie Museum, Pitts- Missouri, Columbia, MO; University Zoo- burgh; Donald S. Chandler, Durham, NH; logical Museum, Copenhagen, Denmark; Cornell University, Ithaca, NY; Field Mu- Zoological Museum, University of Moscow, seum of Natural History, Chicago; Florida Russia; Utah State University, Logan, UT. State Collections, Gainesville, FL; H.F. & A.T. Howden, Ottawa, ON; Illinois Natural

XIix 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 d’Histoire 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, CA USNM: U.S. National Museum of Natural History, Washington, DC UZMC: University Zoological Museum, Copenhagen, Denmark ZMUM: Zoological Museum, University of Moscow, Russia

XIIx State and Province Abbreviations NC North Carolina AB Alberta ND North Dakota AK Alaska NE Nebraska AL Alabama NF Newfoundland AR Arkansas NH New Hampshire AZ Arizona NJ New Jersey BC British Columbia NM New Mexico CA California NS Nova Scotia CO Colorado NT Northwest Territories CT Connecticut NV Nevada DC District of Columbia NY New York DE Delaware OH Ohio FL Florida OK Oklahoma GA Georgia ON Ontario HI Hawaii OR Oregon IA Iowa PA Pennsylvania ID Idaho PE Prince Edward Island IL Illinois PQ Quebec IN Indiana RI Rhode Island KS Kansas SC South Carolina KY Kentucky SD South Dakota LA Louisiana SK Saskatchewan MA Massachusetts TN Tennessee MB Manitoba TX Texas MD Maryland UT Utah ME Maine YT Yukon Territory MI Michigan VA Virginia MN Minnesota VT Vermont MO Missouri WA Washington MS Mississippi WI Wisconsin MT Montana WV West Virginia NB New Brunswick WY Wyoming

XIIIxi II Introduction cosmopolitan stored-product species in the genera Acanthoscelides, Bruchus, Calloso- The beetle family Bruchidae has been bruchus, Caryedon, and Zabrotes to provide recognized for centuries for the ability of as broad a coverage as possible since most certain species to destroy stores of comes- of the biological studies on these species tible leguminous seeds. About 30 species were undertaken outside North America. of bruchids in the world are serious pests, This handbook contains distinguishing and at least 9 are cosmopolitan as the characterizations for 156 species in 24 result of commercial activities. Bruchid lar- genera, plus generic and specific descrip- vae feed entirely within seeds, making their tions, synonymical names, references to detection and control difficult. The most papers on biology, and new or revised successful species are those that attack taxonomic keys for all subfamilies, genera, stored seeds. Because they have adapted and species. New information on hosts and to oviposit and develop in several hosts, distribution was recorded primarily from they are inadvertently assisted by humans’ specimens deposited in the National Col- mobility and storage methods. Although lection of Insects in Washington, DC, and the larva is the destructive life stage, de- from specimens loaned by many other U.S. scriptions of this form are available for (including Hawaiian) and Canadian col- only a few species. For most genera of lections. Included, however, were records bruchids, identification is tedious and dif- from faunal lists such as Kirk (1969, 1970) ficult because many species are closely re- for South Carolina and Kirk and Bals- lated, individuals are small (average length baugh (1975) for South Dakota. Approxi- 1.0–6.0 mm), and dissection of genitalia is mately 30,000 specimens were examined often necessary for positive identification. during 28 years’ study of the Bruchidae. Bruchids are found naturally on all ma- Each species is illustrated with photo- jor land masses except New Zealand and graphs and line drawings to facilitate iden- Antarctica. More intense speciation has tification. A separate key to the Hawaiian occurred in the tropical regions than in the Bruchidae is included in appendix I. Other temperate, and fewer species are found in appendices include a cross-referenced tropical rain forests than in more xeric re- index to host plants (III and IV), a list of gions. Small isolated islands usually have parasitoids (V), and a synonymical check- little or no bruchid fauna because estab- list (II). There is also a glossary of terms lishment of most bruchids depends on the used in this manual. All of the elements previous invasion and establishment of necessary to produce a future annotated suitable food plants for oviposition. catalog of U.S. and Canadian Bruchidae are included. No single information source exists for the Bruchidae of the United States and Can- The geographical scope of the handbook ada. The literature is scattered through a is primarily the 50 United States, Canada, wide range of publications generally un- and the associated continental islands. available to most workers. Because many Beyond these limits, only general distribu- bruchids are critically important pests of tions are provided. stored legumes, their correct identification Although bruchids are difficult to identify, is vital to effective control. The U.S. and information presented in this handbook Canadian literature was combed for data should be useful to taxonomists, biological on classification, host plant associations, control specialists, port identifiers, plant geographical distributions, and parasitoids quarantine officers, museum curators, for this compilation. The world literature ecologists, students and teachers, and was scanned for the several species of Federal and State entomologists.

11 Importance of the Bruchidae country of the plant that might assist in controlling the spread of the pest species. This family has received much attention Since larvae usually destroy the seeds, during the past three decades for several the feeding habits of Bruchidae reduce the reasons. Demands for more food for the potential for reseeding. Other programs (for expanding world population has focused instance, in Argentina) have attempted to on the need for better control of stored- introduce foreign bruchids to help in con- product insects, especially those attacking trolling a native plant that is invading graz- comestible leguminous seeds in larders of ing land, but native parasites destroyed developing countries. Storage losses can the introduced populations (Erb and Frías be ruinous not only from initial damage 1983). Efforts to control the weedy shrub by bruchid larvae but also from ensuing Mimosa pigra Linnaeus using Acanthos- invasions of other organisms (Howe 1973; celides quadridentatus (Schaeffer) and A. Yoshida 1990). puniceus Johnson are under way in Aus- Morallo-Rejesus (1990) estimated that tralia and Thailand (Kassulke et al. 1990; in the Philippines, losses of mung beans Forno et al. 1989; Cock and Evans 1984). infested by Callosobruchus chinensis (Lin- The effectiveness of bruchids as biocontrol naeus) ranged from 7.8 percent to 9.8 agents needs further exploration. In South- percent at harvest, and without protection east Asia and the southwest Pacific, experi- by insecticides or proper storage post- har- mental efforts are underway to control the vest losses rose to 80 to 100 percent within two weedy leguminous plants Cassia tora 3 months. Warthen (1989) summarized Linnaeus and Senna obtusifolia Linnaeus the uses of neem (Azadirachta indica Adr. by the Neotropical bruchid Sennius instabi- Juss.) as a control agent for bruchids in lis (Sharp) (Cock and Evans 1984). stored grains. Recent studies in the use of resistant Laboratories in Colombia, England, varieties of host plants as a means of France, India, Japan, Nigeria, and the biological control of harmful Bruchidae United States are investigating many as- have been conducted by Brewer and Horb- pects of physiology, morphology, plant host er (1984), Dobie (1981), Dobie et al. (1979), resistance, and pheromones in relation to Fernandez and Talekar (1990), Horber the principal cosmopolitan species of Bru- (1978), Kitamura et al. (1990), Minney chidae. A voluminous literature on these et al. (1990), Nwanze and Horber (1976), target species is being generated. South- Osborn et al. (1988), and others. gate (in Aitken 1975) summarized avenues of dispersal of the major economic bruchid species. Medical Importance Another group of bruchids presents a Bruchid adults and larvae feed only on potential threat to the propagation of trees plant tissue and are not associated with targeted for forest plantings in undevel- any disease-causing organisms of warm- oped countries where firewood sources blooded animals. Instances of allergic have been depleted. Seed stocks can be de- reactions have been recorded, however. stroyed by bruchid larvae (Johnson 1983c; Chittenden (1912a) noted that handlers of Southgate 1983). broad beans (Vicia faba Linnaeus) infested by Bruchus rufimanus Boheman developed irritating rashes from adults crawling on Biological Control their skin. Wittich (1940) reported two The accidental introduction of undesirable cases in Minnesota of allergic reaction by plants into certain parts of the world (Aus- women who had been sorting stored beans tralia, Hawaii, Thailand) has stimulated and peas infested with Zabrotes subfas- searches for biological agents in the native

2 ciatus (Boheman). Symptoms included ed 544 names, of which 366 are valid. rhinitis, asthma, skin rashes, and conjunc- tivitis. Relationships With Other Families Bruchidae belongs to a group of plant-feed- Biology of the Bruchidae ing beetle families loosely grouped in older classifications as the Phytophaga. Included Definition in this group are the families Bruchidae, Anthribidae, Cerambycidae, Chrysomeli- Bridwell (1932:101) provided an extended dae, Curculionidae (broad sense), and Sco- definition of this family, but for purposes of lytidae, all of which produce plant-feeding this handbook the following characteristics larvae. A configuration of 5-segmented tar- are sufficient to define the Bruchidae: si with the 4th segment small and nearly Adults.—Body surface setose; head hypog- hidden in the emargination of the 3rd is a nathous or opisthognathous; ocelli absent; characteristic common to all. eye shallowly to deeply emarginate; anten- Contemporary coleopterists generally na 11-segmented with insertion adjacent to agree that the family Bruchidae is closely eye; mandibular apex acute, medial margin related to Chrysomelidae, probably repre- entire, not dentate; gular sutures short, senting a seed-feeding phyletic branch of ending in tentorial pits; frontoclypeal Chrysomeloidea having highly developed suture well marked; elytral striae always characteristics for the spermatophagus present, usually 10 in number; metati- mode of life (Chen 1940; Crowson 1946, bia usually longitudinally carinate; tarsal 1960; Monros 1955; Lawrence and New- claws appendiculate; pygidium exposed ton 1982). Crowson (1960) stated, “Larval beyond elytral apices; male genitalia with characters, and to some extent those of base of median lobe and ventral strut of the adults, clearly establish the affinity of tegmen modified into a pump to evert in- Bruchidae and Sagrinae insomuch that it ternal sac during copulation, lateral lobes becomes difficult to justify familial sepa- (parameres) always present. ration between them. Bruchids appear to Larvae.—Labial sclerome present in instars be the most derivative group.” Kingsolver 2–4; legs reduced or absent (except some (1995b) listed the characters he felt were 1st instar forms); mandible gouge-shaped; sufficient to give bruchids full family sta- pronotal sclerites present in 1st instar. tus. Lawrence and Newton (1982) and Reid (1996) placed the bruchids as a subfamily Phyletics in the Chrysomelidae based on cladistic analysis. Schmitt (1989) and Hawkeswood Number of Genera and Species (1996) argued that Bruchidae should be a Compared with other families of beetles, subfamily within the Chrysomelidae but Bruchidae is relatively small. The esti- that it should retain the “-idae” ending for mated number of species generally cited in stability and unambiguity of scientific com- texts is 1,200. This estimate is based on munication. the 818 species cataloged by Pic (1913a) plus the species described since then. The Evolution world catalog published in 1989 by Udaya- No extensive evolutionary or cladistic giri and Wadhi (which had a cutoff date of studies have been made for the entire 1982) included 1,346 valid species names. family, although Slobodchikoff and John- My working catalog for the Western Hemi- son (1973) analyzed the phylogeny of sphere species lists 2,200 proposed names the American species of Acanthoscelides. up to 1990, of which 895 are valid. John- son and Kingsolver’s (1982) checklist for North America and the West Indies record-

3 Some concurrent evolutionary trends have system (Kasap 1978b); swollen metafemur been observed, however, that allow us to with ventral denticles; metepisternum with speculate. an angular sulcus (also found in some The Bruchidae is apparently a monophy- primitive Cerambycidae); similar wing letic, spermatophagus, chrysomeloid group venation patterns including the presence springing from an ancestor common to the of a wedge cell in some species (Suzuki chrysomelid subfamily Sagrinae (Crowson 1969); larvae as internal feeders in plants 1946; Monros 1955). Similarities in wing (sagrines in stems or crowns, pachymer- venation, male genitalia, form of the meta- ines in seeds). Differences are mostly femur, presence of tibial spurs, internal external: Elytra throughout the family feeding by the larvae, and other characters Bruchidae always striate whereas those of point to a relatively recent divergence of Carpophagus and most other sagrines lack the bruchids followed by development of distinct striae; frons in the pachymer ines spermophagy to a fine degree (Kingsolver with a median carina, sagrines with an 1995b; Verma and Saxena 1996). X-shaped sulcus; larval mandibles gouge- shaped in pachymerines, toothed in those Kingsolver (1995b) listed the characters he sagrines whose larvae are known. Maulik thought were sufficient to give the bru- (1941) presented useful comparative char- chids full family status, and his view was acters for the Sagrinae. supported by Hawkeswood (1996) and Verma and Saxena (1996). Lawrence and Evolutionary Trends Newton (1982) and Reid (1996), however, Certain evolutionary trends are apparent placed the bruchids as a subfamily of the within the Bruchidae. They indicate that Chrysomelidae based on cladistic analy- the most likely primitive genera are in the sis. Schmitt (1989) argued that Bruchidae Pachymerinae and that the Rhaebinae, should properly be a subfamily within the which are found only in central Asia, are Chrysomelidae but that Bruchidae should probably a relict group. The Amblycerinae, be retained with the -idae family ending Eubaptinae (found only in South America), for stability and unambiguity of scientific and Kytorhininae are successively more communication. Monros (1955) treated the specialized, and the Acanthoscelidinae are Bruchidae as a subfamily of the Chrysome- probably the most derived group. lidae. 1. Trochantins are present in the fore The groups seemingly nearest to the Sagri- and middle legs in the Pachymerinae, nae are in the bruchid subfamily Pachym- Amblycerinae, Eubaptinae (South erinae, especially the genera Caryoborus America), Rhaebinae (Asia), and Kyto- and Caryobruchus. Although resemblance rhininae but are modified or lost in the of the sagrine Carpophagus to these pa- Bruchinae except in the mid legs of chymerines is striking, Crowson (1946) did Megacerini. not think that this genus was in any way 2. Median lobe of male genitalia has an related to the Bruchidae. Without know- unmodified apex in Pachymerinae and ing what bruchid species he used in this Rhaebinae (figure 2) but is fractured comparison, I am of the opinion that Car- into a separate ventral valve in all other pophagus has more in common with the subfamilies (figure 3). pachymerine genera than with any other chrysomeloid genus. Similarities are male 3. Ocular sinus is shallow in Pachymeri- genitalia with a simple, curved, tubular nae (figure 1) and inAmblycerus of the median lobe, bases of the lateral lobes Amblycerinae (figure 4) but deep in fused straplike with only the apices ex- more specialized subfamilies. In some panded; absence of a crop in the digestive species of Meibomeus (figure 5), the sinus nearly divides the eye.

4 4. Adult antennal forms range from sim- the egg, through the pod or seed integu- ple serrate or flattened moniliform in ment, through the endocarp, and into the Pachymerinae, Rhaebinae, and Eu- cotyledon, whereas others cut through baptinae to clavate, strongly serrate, or the top of the egg and walk around on the pectinate in males in the Kytorhininae substrate searching for a crack or crevice and in some species in Bruchinae (fig- to help them gain purchase to bore into ures 6–13). the seed. Some species are provided with 5. Mesepimeron is gradually reduced by a suckerlike appendage on the tenth ab- ventral encroachment of mesepister- dominal segment to help them adhere to num (figures 25–27) as the subfamilies the surface during their search for a suit- progress in degree of specialization. able entry site (Pfaffenberger and Johnson 1976). Special adaptations of the neonate 6. Venation of anal area of wing gradually larva include well-developed legs in those lost the “wedge cell” and veins assumed species that emerge from the top of the egg an “h” shape in most of the more and shorter legs in those forms that drill derived families. directly through the floor of the egg. 7. Lateral pronotal carina modified or lost The most remarkable character, however, as specialization progressed. is a toothed dorsal prothoracic plate that assists the larva in escaping from the egg Genetic Studies and gives leverage in penetrating the pod Skaife (1925, 1926) was a pioneer in bru- or seed. All bruchid species for which the chid genetics. Certain species of Bruchidae first larval instar is known possess this are now being used in several laboratories toothed plate (figure 16). Pfaffenberger and throughout the world as test organisms for Johnson (1976) and Kannan (1923) illus- genetic studies because they are readily trated this character for several species. In cultured in the laboratory and have short its first molt, the larva loses these special life cycles. Species used are those that adaptations and assumes a scarabaeiform breed continuously in stored seeds—Ac- aspect with legs much reduced and with anthoscelides obtectus, Callo- sobruchus gouge-shaped mandibles. It feeds and chinensis and C. maculatus, and Zabrotes molts through its remaining three instars subfasciatus, for example. inside an excavation scarcely larger than its own bulk. Life Histories In most cases, the larva, immediately before entering the pupal stage, will bore For a general discussion of bruchid biolo- to the surface of the seed or pod, leav- gies, see Skaife (1926), Southgate (1979), ing a circular cap in the epidermis that and Zacher (1929 and 1930). See also the can easily be cut and pushed off by the Proceedings of the Second International adult during emergence. After excavating Symposium on Bruchids and Legumes (Fu- this escape route, the larva returns to the jii et al. 1990) for other life history studies. feeding chamber and pupates. In some Johnson (1990b) contributed a paper to species, however, pupation occurs partly this symposium. or completely outside individual seeds (for example, Caryedon serratus). Species too Larval Feeding large to develop in individual seeds may A first larval instar is faced not only with feed on several seeds inside an envelope escaping from the egg but also with pen- (the pod or capsule, for example) and spin etrating one or two layers of plant integu- a cocoon inside the envelope (examples ment before it can reach its food source. include some species of Amblycerus and of During eclosion, larvae of most species Sennius). Glands producing the material bore directly through the floor layer of for the cocoon have not been investigated.

5 Adult Nutrition and oviposition. Lago and Mann (1987) recorded several species of bruchids on Although bruchid larvae are well known to inflorescences ofDaucus carota Linnaeus subsist mainly on tissue of the seed coty- (wild carrot). ledon, comparatively little is known about adult food habits for most species. Most Jarry (1987) identified the pollen of 18 observations on adult feeding have been different species of plants in the diges- made on the species affecting crops—the tive tracts of Acanthoscelides obtectus in so-called “economic species.” Adult man- France. Pollen from the principal host, dibles with sharp medial edges and lack of Phaseolus vulgaris, made up only 9 percent teeth are not especially adapted for biting of the total, whereas weedy plants in areas tissue except to complete cutting of the surrounding the plots of P. vulgaris made operculum of the seed prior to emergence. up the bulk of pollen identified. Grass pol- Maxillae in many species are equipped len was well represented. with well developed, feathery setae that as- Adults of Acanthoscelides obtectus can sist in raking pollen into the oral cavity. reproduce and oviposit without first tak- Most so-called “host” records for adults, ing food, but Leroi (1978) determined that usually floristic associations with what- well-fed, newly emerged individuals can ever plant happens to be in bloom at the survive more than 200 days on honey-pol- time the adults are flying about, may have len mixtures. Ovarian production by well- little or no relationship with the larval host fed females was 50 percent more than that plant. Because adults cause little dam- by starved females. Shinoda and Yoshida age by their food intake, it has been pre- (1987a) found that when newly emerged sumed that either they do not feed or they Callosobruchus chinensis were fed on feed sparingly on nectar or pollen. Female powdery mildew (Sphaerotheca fuliginea) or Acanthoscelides obtectus have been ob- rust (Uromyces azukiola) their mean lon- served biting the sides of green pods or the gevity was three times that of control indi- suture of more mature pods of Phaseolus viduals and the numbers of eggs produced to make an opening through which they doubled. Zacher (1929) thought that adults deposit eggs into the interior of the pods. should be able to retain sufficient fat from Whether or not they ingest tissue is not the larval stages to carry them through known (Slingerland 1892). Brindley et al. adult life. (1946, 1952) and Pajni and Sood (1975) Larson and Fisher (1938) included a sum- reported that Bruchus pisorum can only mary of the literature on adult feeding to reproduce after ingesting pollen of its larval that date. Many of the observations were host, the pea plant (Pisum sativum). Adults made on caged bruchids and do not neces- of Bruchus pisorum overwinter in trash or sarily reflect field conditions. Slingerland under bark and emerge from diapause to (1892) claimed that bean weevils (Acan- feed on pea blossoms; then they are ready thoscelides obtectus) confined on live bean to oviposit on green pods. plants apparently fed on the parenchyma De Luca (1966) compiled a list of flower- of the leaves, but he did not actually ob- visiting records from the world literature. serve feeding. No descriptions of actual ingestion were Leroi et al. (1984) found that for Acan- made, although his introduction to the thoscelides obtectus, although larvae and paper is a discussion of pollen and nectar adults utilize different types of food (that nutrition. Johnson (1977d) listed eight is, seed endosperm vs. pollen grains), species of plants from which adult Acan- carbohydrate compositions of the two food thoscelides pauperculus were collected, sources are remarkably similar. Glyco- but precise observations on actual feed- sidase activity in the guts of larvae and ing were not made. He suggested that adults are likewise similar. feeding on pollen is necessary for mating

6 For an excellent discussion of bruchid 3. Scattered seed guild, in which eggs are physiology and nutrition, see Johnson and laid on mature seeds that have fallen to Kistler (1987). the ground from a fully dehisced pod. (Examples: Stator sordidus, Zabrotes Oviposition spp.) Huignard et al. (1990) observed that most Acanthoscelides obtectus females bite holes Bruchidae are specialists developing on a in green Phaseolus spp. pods through limited number of host species and host which they insert eggs to fall freely within selection is accomplished by females ovi- the pod. In later generations, they insert positing on pods or fruits that are only eggs in old emergence holes. Oviposition available during a short period of the year. for this species begins early in the season Neonate (newly hatched) larvae alone can- on immature pods and may end on mature not search for their food substrate but de- pods that have dropped to the ground or pend on the female to locate suitable host on naked seeds in storage. plants or seeds upon which to oviposit. Bruchid eggs are usually either cemented Physiology to the pod or seed of a suitable larval food No attempt is made here to discuss physi- plant, dropped into a pod after the female ological aspects of bruchids, but the follow- cuts a hole in the pod wall, or tucked into ing references will provide an introduction surface cracks or old emergence holes. to the literature. Discussions of Acanthos- The adhesive used originates either in the celides obtectus, Callosobruchus chinen- epithelium of the egg follicles (Snodgrass sis, C. maculatus, Caryedon serratus, and 1935) or in accessory glands (Wigglesworth Zabrotes subfasciatus include additional 1947) and not only cements the egg to references on the subject. For a discus- the substrate but also covers it. In some sion of nutritional ecology of bruchids, see species, strands of adhesive resembling Johnson and Kistler (1987). guy wires extend from the edge of the egg covering, apparently to aid in holding the Chromosomes.—Takenouchi 1955, 1971a, egg to the pod surface as the pod expands 1971b; Bawa et al. 1974; Smith and Brow- during growth (Forister and Johnson 1970; er 1974; Garaud and Lecher 1982; Garaud Pfaffenberger and Johnson 1976). 1984. Johnson (1981a) classified egg placement Digestion.—Gatehouse et al. 1979; Choud- into three general types, or “guilds”: huri and Paul 1983; Gatehouse and Boul- ter 1983; Gatehouse et al. 1989. 1. Mature pod guild, in which eggs are cemented to the surface of the seed Embryology.—Brauer 1925; Tantawi et al. pod. Neonate larvae in this guild must 1976; Daniel and Smith 1994. bore through the pod wall and into Enzymes.—Sharma and Sharma 1979, the seed. (Examples: Acanthoscelides 1981; Gatehouse and Anstee 1983; Puri chiricahuae, Merobruchus spp., Mi- and Sharma 1984; Sharma and Rai 1984; mosestes spp.) Garg et al. 1990. 2. Mature seed guild, in which eggs Egg production.—Huignard 1970; Sandner are cemented to the surface of seeds and Pankanin 1974; Hinton 1981; Choud- still attached to the inside of a partly huri and Paul 1984; Dick and Credland dehisced pod, so that larvae need 1984; Butare and Biémont 1987. only bore through the seed integument. (Examples: Stator limbatus, S. Genetics.—Breitenbacher 1925; Sano-Fujii pruininus.) 1986; Messina 1987, 1990. Lipids.—Nwanze et al. 1976; Sidhu et al. 1984.

7 Nervous system.—Ogijewicz 1948. incidentally included illustrations of struc- Overwintering.—Brauer 1942; Hodek et al. tures for several genera, including Ameri- 1981. can forms, but the field has been almost totally neglected by American workers. Pheromones.—Hope et al. 1967; Halstead 1973; Rup and Sharma 1978; Tanaka et A moderately large resource of morpho- al. 1982; Qi and Burkholder 1985; Kitch logical papers has recently emerged from and Murdock 1986; Sakai et al. 1986. several Old World laboratories. Singh (1973, 1978a,b, 1981a,c, 1982, 1983), for Temperature effects.—Larson and Simmons instance, published on morphological char- 1924a; Menusan 1934b, 1936; Menusan acteristics of several Old World bruchid and MacLeod 1938; Sano 1967; Kistler species. Many of the papers on Old World 1985. bruchids are cited at the end of this section. Morphology of the Bruchidae Johnson and Kingsolver (1973) introduced the terms lateral, lateroventral, ventral, and Adults dorsomesal to designate metatibial carinae Little attention has been paid to the com- in the genus Sennius. Here, I am modifying parative morphology of New World bruchid lateroventral to ventrolateral to be consis- adults. Descriptions have been written tent with dorsomesal. These metatibial with little thought to consistency of no- terms are valid for the majority of genera menclature of body parts, causing confu- in the Bruchinae; however, the positions sion when papers of various authors are of the tibial carinae are different in the compared. Most references to morphologi- Pachymerinae and in the Megacerini and cal characters have been incidental in ge- will be further elaborated in the descriptive neric or specific descriptions. Such papers sections for those taxa. as Doyen’s “Skeletal anatomy of Tenebrio For the purposes of this handbook, only molitor” (1966), Snodgrass’ 1908 and 1909 external morphology and male genitalia papers on the insect thorax, or Hopkins’ and considered. A thorough morphological venerable but valid “Contributions toward analysis is sorely needed in order to make a monograph of the bark-weevils of the ge- comparisons between various sections of nus Pissodes” (1911) are useful for general the family and to provide sets of characters reference but, of course, do not explain for phylogenetic analyses. Only those body morphological peculiarities of the bruchids. parts that can be seen without dissection Approximately 275 taxonomic papers have or dismemberment of the body (except been written about Western Hemisphere male and female genitalic parts, wings, and bruchids during the past 50 years, but mouthparts) will be discussed and illus- only two compared a structure of a body trated here. One species is illustrated as a part (male genitalia) in several genera of basic reference, and important variations this family (Teran 1967, Kingsolver 1970a). are illustrated. Although nomenclature used in the two Because Bruchidae develop in a restricted papers differed, a basis for subsequent space—in most instances the excavated studies was established. In addition to interior of a single seed—and must emerge offering a nomenclature for various sclero- through a circular hole excavated by the tized parts, Kingsolver included a diagnosis larva, evolution has favored a smoothly of the genitalic musculature. No one has rounded body lacking protuberances or surveyed the various genera for female spines that would hinder escape from the genitalia, wing venation, elytral form, head interior of the seed. Most bruchids are form, legs, or antennae. Luk’yanovich and ovate or subelliptical in outline and gener- Ter-Minassian (1957) and Borowiec (1987) ally circular or oval in cross section. Legs

8 and antennae are capable of being tucked usually carries a transverse row of long close to the body, and the head hangs setae. vertically and usually rests on the proster- The compound eye is surrounded by a num. In many species, the dorsal profile narrow, setiferous channel, the supra- is arched, which leads to a shortening of ocular sulcus (Kingsolver 1980a:230), sternal areas of the thorax and consequent which may correspond to the ocular su- reduction of sclerites in this part of the ture found in more generalized insects body. In addition, the elytra are abbreviat- (Snodgrass 1935:109) or may be second- ed, exposing the heavily sclerotized seventh ary. tergum (pygidium). The antennal socket is situated on the Head.—The bruchid head at rest is either ventral rim of the eye at the base of an vertical (hypognathous) or reflexed so that emargination, the ocular sinus. The sinus it rests on the prosternum (opistho- gna- varies in depth from a slight emargination thus) (figure 17). In a frontal view (figures (as in Amblycerus and Caryobruchus) to 18 and 68), the head is obovate with the an almost complete division of the eye (as subspherical base inserted into the ante- in Meibomeus). The antennal socket car- rior foramen on the front of the prothorax. ries on its ventral rim a small projection, In lateral aspect, the head tapers toward the antennifer, forming an articular point the apices of the mandibles. Compound upon which the bulbous base of the an- eyes are usually protuberant but in some tennal scape pivots. The scape is usually species follow the contour of the lateral longer than the second segment, the pedi- margin of the head. cel (figures 6–13); theflagellum is invari- Most of the sutures delimiting sclerites of ably composed of nine segments and may the head in primitive beetles are lost in be uniformly serrate, gradually clavate, Bruchidae, and only general areas can be strongly eccentric, or even pectinate. defined. Thevertex is the area of the head On the ventral side of the head, the eyes above the upper limits of the eyes, some- are prominent on either side of a flat, times set off by a transverse sulcus (figure central submentum (figure 21). The latter 18). The frons lies between the eyes and area is continuous with the gena in front sometimes carries a vertical, median fron- of and behind the eye and is limited be- tal carina. The ventral margin of the frons hind by a deep, transverse sulcus. At the is the transverse frontoclypeal suture. In posterior end of the head is a large open- some genera (for example, Caryobruchus ing, the occipital foramen, through which and Amblycerus) the frontogenal suture the digestive and nervous systems pass extends from the inner corner of the eye into other parts of the body. On the ventral to the frontoclypeal suture separating the part of the rim of the foramen is a quadrate frons from the cheek portion of the head or trapezoidal sclerite, the gula. Along the (the gena) but in most bruchids this suture lateral margins of the gula are the posterior is lost and the frons and gena are continu- tentorial pits, which are the attachments of ous beneath the eye. The dorsal portion of the internal skeleton of the head. Attached the gena carries the antennal socket. to the anterior margin of the submentum The clypeus is a quadrangular sclerite, is the mentum (figure 21), an emarginate, sometimes elongated, usually rounded transverse sclerite to which the premen- basally, and truncate apically. The fronto- tum is attached. The prementum is the clypeal suture limits it dorsally, the lateral base for the paired, three-segmented labial margins extend to the mandibles, and the palpi and the ligula, which is divided into apical boundary is the clypeolabral suture. paired paraglossae. On either side of the The clypeus partly covers the mandibles mentum and submentum is a deep emar- and maxillae. The labrum is attached gination, the maxillary fossa, into which to the apical margin of the clypeus and the maxilla is inserted. The maxilla (figure 19) is composed of five parts: thecardo ,

9 the stipes, the three-segmented maxillary The prosternum is usually short in front of palpus, the galea, and the lacinia. Both the procoxae because of the ventral com- the galea and the lacinia are provided with pression of the body. In Spermophagus, an long curved setae, which, in some species, Old World genus, it is reduced to a mem- are plumose. These specialized setae are branous bridge. The prosternal process apparently used to rake pollen into the oral may be flat as inCaryobruchus and some cavity. The inner surface of the labium and Amblycerus (figure 41) or narrow as in of the labrum is also provided with rows of other Amblycerus (figure 45), or it may be setae to assist in ingesting pollen. an internal, vertical lamella partly hidden The mandibles project beyond the labrum, by the coxae (Acanthoscelides). but their attachment is on each side of The attachments of the fore and middle the head with the dorsal pivots, or articu- coxae generally take two forms in bru- lations, at each end of the frontoclypeal chids. In the generalized form (as in Ambly- suture and the ventral articulations at the cerus and Caryobruchus), the coxa freely lateral corner of the maxillary fossa. The articulates with the trochantin, which in apex of each mandible is acute, and the turn articulates with the body wall (figure inner margin forms a sharp, entire cutting 23). In the more advanced form (such as edge. In some species, the inner margin Acanthoscelides and Sennius), the trochan- is fringed with long setae. A membranous tin is replaced by a straplike extension of flap, theprostheca , and a flattened plate the sternum forming a socket that allows near the base of the mandible, the mola the coxa to rotate freely through about 90 (figure 20), are characteristic of all Bruchi- degrees of arc but limits lateral movement dae. The face of the mola is covered with of the leg (figure 24). The triangular tro- short spines to form a grinding surface. chanter is rigidly attached to the basal end Prothorax.—The prothorax, the first leg- of the femur. bearing segment, is composed of two Mesothorax and Metathorax.—The two sclerites: the prosternum (figures 22, 23, wing-bearing segments in Bruchidae are and 41) is a Y-shaped sclerite bounded rather rigidly fused together to provide by the front coxal cavities, the pleuroster- strength. Internal sutural ridges (phragma, nal suture, and the anterior foramen, and or ingrowths from the body wall) provide the pronotum is a large, continuous dor- attachment points for the massive muscles sal sclerite making up the remainder of required to operate the mesothoracic ely- the body of the prothorax (figure 41). The tra, the metathoracic flight wings, and the pronotum consists of a dorsal surface, legs. Only the external sclerites, however, the disk, and the hypomeron (the lateral, will be identified here. concave area dorsal to the coxa). The disk Each of the thoracic segments is divided is limited in some species by a distinct into a ventral sternal region, a lateral pleu- lateral carina but more often is continuous ral region, and a dorsal notal region. Except laterally with the hypomeron itself, marked for the apex of the scutellum protruding only by the concave surface. The epimeron, between the bases of the elytra (figures 30 an extension of the hypomeron but not a and 60), the notal sclerites are concealed separate sclerite, projects behind the front beneath the elytra and have not been coxae to meet the apex of the intercoxal included in descriptions of bruchids. The piece. flight wings, concealed beneath the elytra In some species in the Pachymerinae and (figures 14 and 15), are folded longitudi- Amblycerinae, the margin of the pronotal nally and transversely. disk is distinctly sulcate and corners of The mesothorax is the shortest of the three the disk bear one to three setiferous punc- thoracic segments. It consists of three vis- tures. ible plates—the mesosternum, the mesepis-

10 ternum, and mesepimeron (figure 28). The tion of the sternal and pleural sclerites is visible mesosternum is a Y-shaped plate distinctly different from that of the me- lying in front of and between the middle sothorax. The metasternum is rigidly at- coxae and is separated laterally from the tached to the mesosternum and to the ventral end of the mesepisternum by the horizontal metepisternum (figure 28) and sternopleural suture. The intercoxal process carries the median articulation for the rests on the corresponding extension of the metacoxa as well as the attachment of the metasternum. The mesepi abdomen. It also forms the posterior wall of sternum and mesepimeron are adjoining the mesocoxal cavity and the anterior wall vertical plates extending from the elytral of the metacoxal cavity. The pleurosternal margin to the coxal cavity. In more gener- suture separates it from the metepister- alized bruchids (figure 25)(Caryobruchus, num. The postmesocoxal sulcus follows the Amblycerus), these two plates are approxi- posterior margin of the mesocoxal cav- mately the same size, but in more special- ity and in some genera bends sharply to ized forms the mesepimeron is reduced in parallel the pleurosternal suture (parasu- size and shape by the encroachment of the tural sulcus), but in other, more specialized mesepisternum (figures 26 and 27). genera it ends at or near this suture. Along the anterior border of the metacoxal cav- The coxal cavity lies between the metaster- ity extends a narrow, transverse sclerite num, the mesosternum, and the ventral variously termed the antecoxal piece or the end of the two pleural sclerites. The attach- precoxal strap. ment of the leg is essentially the same as described for the prothorax. The metepisternum is an elongated, horizontal sclerite, set between the mesepi- The form of the elytra in beetles is a radical meron and the dorsal one-half of the modification of the metathoracic wings to metacoxal cavity, and carries the dorsal form a protective cover for the notal areas articulation of the metacoxa. In some more of the mesothorax, the metathorax and generalized genera, the face of this sclerite abdomen, and the flight wings. At rest, the bears an angulate sulcus—the metepister- elytra meet on the midline with a tongue- nal sulcus (figure 28)—which extends to- and-groove interlock mechanism. During ward and bends to parallel the pleuroster- flight, the elytra are rotated laterally on nal suture and forms the corresponding the mesopleural wing process to allow the parasutural sulcus to that of the metaster- flight wings to unfold and operate freely. num. This sulcus is not present in more Although the elytra in bruchids exhibit specialized genera. The metapleural suture many modifications of surface sculpture separates the metepisternum from the nar- and configurations of the striae, especially row, horizontal metepimeron that, except in the basal area, 10 striae are always for the posterior end, is hidden beneath the present. margin of the elytra. Venation of the flight wings throughout The external face of the hind coxa is an the family is not detailed here (except in elliptical or reniform plate rotating on a figures 14 and 15). The wings of only a transverse axis between the ventral coxal few species have been illustrated, and a articulation and the dorsal articulation. thorough survey of venation in the family The coxa, however, is roughly triangular has not been made. References to papers in cross section and projects into a cavity that include bruchid wing patterns can be between the metasternum and the abdo- found at the end of this section. men. The coxa can rotate on its axis about The metathorax is considerably larger 30 degrees, and the trochanter can rotate than the mesothorax to accommodate the in its fossa near the ventral end of the hind volume of muscles that operate the flight coxa more than 180 degrees. Extending wings and legs. The size and relative posi- laterad from the trochanteral fossa is a

11 thin suture usually bordered by a polished, Modifications of the visible sterna include impunctate strip. I am here applying the a tuft of setae or a pit on the basal ster- term “fossula” to this suture (figure 28). num of males in some genera, the terminal sternum in many genera being more deeply The trochanter is more or less rigidly at- emarginate in males than in females, or tached to the proximal end of the femur the abdominal segments being telescoped. (figure 29). The latter segment of the hind leg is laterally compressed and often dor- The membranous pleural areas of the soventrally expanded. The ventral margin abdomen that carry the spiracles for each is often furnished with denticles of various segment are hidden beneath the elytra, sizes and numbers (figures 40, 47, 49, 50, except that the spiracle for the 7th tergum 53, and 63–65), a primary source of sub- (the pygidium) is located at the extreme family and generic characters and, in some upper corner of the tergum. The pygidium instances, specific characters. Despite in this family is always exposed beyond the the obvious resemblance to the saltato- apices of the elytra and is always heavily rial, or jumping, hind legs in other fami- sclerotized; however, in Kytorhinus the 5th lies and orders of insects (such as fleas and 6th terga are also sclerotized (figure and fleabeetles), legs of bruchids are not 51). The sculpture, shape, and setation equipped for jumping. The development of of the pygidium is often a good source of the internal musculature is opposite that generic and specific characters. The male for jumpers; that is, the greatest develop- pygidium is often more convex and inflexed ment of muscle bands is in those that flex apically than is that of the female. the metatibia against the ventral margin of Genitalia.—Zacher (1930) was the first the metafemur. The hind leg thus becomes worker to explore the usefulness of male a grasping organ, probably for holding to genitalia in the classification of bruchids, edges of leaves or perhaps used in mating. but his descriptions of the various parts See Furth and Suzuki (1990). were meager and poorly illustrated. Muker- The proximal end of the metatibia hinges ji and Chatterjee (1951) compared genitalia between two terminal plates of the metafe- of species in five genera and identified the mur and has a range of movement of about various parts using the nomenclature of 120 degrees in the same plane as the Snodgrass (1935). Teran (1967) illustrated metafemur. Terminal spurs are found in genitalia of species in nine additional gen- only a few genera, the most common termi- era but used a different set of morphologi- nal armament being a ventral spine (mucro) cal terms. Kingsolver (1970a) made a study and several smaller terminal denticles (co- of male genitalia from an evolutionary rona) (figures 44, 47, 49, 50, 53, and 54). standpoint, including the identification of genitalic muscle strands and a theoretical Abdomen.—The abdomen in Bruchidae explanation of the mechanism for everting consists of five externally visible, heavily the internal sac. His nomenclature, dif- sclerotized sternal segments, but the basal ferent from that of Mukerji and Chatterjee sternum is anatomically the 3rd segment, or that of Teran, followed Sharp and Muir the 1st and 2nd sternites being reduced to (1912) and has generally been employed by lightly sclerotized or membranous relicts recent workers in the family. Most of the forming the vertical anterior wall of the taxonomic papers published since 1960 abdomen behind the metacoxae. For con- use and illustrate male genitalia for species venience, the visible sternal segments are discrimination. Several papers have re- numbered from 1 to 5. The terminal seg- cently been published on male genitalia of ments, including those transformed into Old World species of Bruchidae. These are sex organs, are radically modified and are referenced at the end of this section. withdrawn into the end of the abdomen.

12 Little has been published in the United • The median lobe (aedeagus) (figure 30) States on the female genitalia of Bruchi- is a sclerotized tube with an enlarged, dae. See below for a discussion of these spoon-shaped base (cucullus) modi- structures. fied from ventral struts found in related Male Genitalia.—Male genitalia provide the families. most consistent source of generic and spe- • The eversible internal sac (figures 30 cific identifying characters in this family. and 34) normally lies retracted inside Moreover, the combination of character- the median lobe and is attached to the istics of the genital complex is sufficiently apical orifice of the median lobe. distinctive to warrant separation of the These various parts are highly modified family from other Chrysomeloidea. derivatives of the terminal segments of the The male reproductive organs are of both abdomen and are enclosed in the elongate mesodermal and ectodermal origin. The intersegmental membranes of the 7th, 8th, testes and vasa deferentia originate in the and 9th segments. mesoderm, whereas the ejaculatory duct, The apical orifice (figure 30a), which is the accessory glands, and intromittent organs attachment for the base of the internal sac, are ectodermal. Ectodermal parts contain is located near the apex of the median lobe. chitin and will withstand treatment in The basal orifice (figure 30b) is located at sodium hydroxide to remove muscle fibers, the point at which the basal piece sur- whereas endodermal parts will dissolve in rounds the median lobe. Although the rigid the same solution. apex of the median lobe of the bruchid As is true of most Coleoptera, the median subfamilies Pachymerinae and Rhaebinae lobe, lateral lobes, internal sac, and associ- is similar to that found in the Sagrinae and ated parts are completely invaginated into the Cerambycidae, the apex in the Am- the end of the abdomen and are normally bly-cerinae, Kytorhininae, and Bruchinae not visible without dissection. To examine has developed into a separate sclerite, the details of the various components, caustic ventral valve (figure 30c). Some groups of treatment is usually necessary to remove bruchids also have a sclerotized or largely extraneous body tissues (see Materials and membranous, hoodlike dorsal valve (figure Methods). Significant characters for spe- 30d) above the apical orifice. The configu- cific and sometimes generic identification ration of the ventral valve is often one of are usually found in the parts resistant to the specific diagnostic characters. the caustic solution. The lateral lobes are attached to the dorsal Male genitalia of bruchids are modified rim of the basal piece and lie on the dorsal from the cucujoid type (Crowson 1955) side of the median lobe. They take various with a basic form similar to those found in forms, sometimes fused into an apically the phytophagous families Cerambycidae, bilobed straplike structure, or are vari- Chrysomelidae, and Curculionoidae. The ously divided, sometimes nearly to their nearest approach of bruchids to another attachment with the basal piece. The basal family appears to be that of the Pachym- piece surrounds the middle of the median erine genera Caryoborus and Caryobruchus lobe and is attached to the ventral rim of to the Sagrine genus Carpophagus. the basal orifice by a sclerotized mem- The genital complex comprises three prin- brane. The tegminal strut extends beneath cipal parts (figures 30–34): the cucullus and may be flat or keeled. Between the rim of the cucullus and the • The tegmen (figure 31) comprises the tegminal strut extend thick muscle bands lateral lobes (parameres), the basal (figure 32) bridging the space between the piece (tegmen ring) surrounding the two parts. The membrane between the median lobe, and the ventral strut. basal piece and basal orifice effectively

13 seals the cavity formed by the muscle Withdrawal of the sac from the vagi- bands, and compression of the muscles nal tract begins at its apex. Fine muscle converts the structure into a pump to evert strands attached to saccal sclerites and the internal sac during copulation. inserted on the internal walls of the medi- Associated sclerotized structures are the an lobe contract to effectively peel the sac Y-shaped, ventrally positioned 8th sternite from inside the vagina and restore it to its (spiculum gastrale) (figure 32), to which original position inside the median lobe. muscle bands that move the genital com- Female Genitalia.—Female genitalia gener- plex in and out of the body are attached, ally have not been illustrated nor described and the modified8th tergite, a hood- due to their apparent lack of taxonomic shaped sclerite lying beneath the pygidium significance. This facet of bruchid mor- and dorsad of the lateral and median lobes. phology, however, should be more fully The internal sac is the primary intromit- explored for potential species group or tent organ (figures 30, 33, and 34). It is generic characters. attached at the apical orifice near the apex Kingsolver and Johnson (1978) published of the median lobe and is essentially an illustrations of ovipositors for North Ameri- extension of that lobe. For that reason, can species of Mimosestes, and theirs is the terminology applied to the internal the only American paper illustrating fe- sac seems contradictory but is logical if it male genitalia for all of the species treated. is remembered that it is the median lobe No one has made a comparative study extended. When the sac is invaginated of female genitalia for the Bruchidae in within the median lobe in repose, its base this country. Nilsson and Johnson (1990) is nearest the apex of the median lobe and included drawings for two species of Caryo- its apex is nearer the base of the median bruchus, and Donahaye (1974) sketched lobe (figure 30). Inflation of the sac is -ac genitalic parts for seven species of Bru- complished through pumping action of the chus. Mukerji and Bhuya (1937) detailed basal pump. Various sclerites (spicules, the reproductive system of both Calloso- spines, denticles, serrata) (figure 35) at- bruchus maculatus and C. chinensis. Tan- tached to the sac emerge on the external ner (1927) included Acanthoscelides (as surface of the sac as it is evaginated into Mylabris) obtectus in his survey of female the vaginal tract and undoubtedly act as ovipositors of Coleoptera. Johnson et al. holdfasts to keep the sac in place during (1989) included various female characters copulation. The arrangement of this saccal in New World Stator. Several papers detail- armature is, in many instances, the source ing ovipositors and internal reproductive of characters for specific identification. organs have been published for Old World Because of the difficulty of everting the sac bruchids, including those for Bruchidius of prepared specimens, illustrations of the mackenziei Kingsolver from Australia (fig- male genitalia usually show the sac in the ure 36), and for several species of Bruchid- invaginated position. ius by Iablokoff-Khnzorian and Karapetian The vas deferens (figure 32) extends (1973). References to these papers are through the muscle strands of the pump listed at the end of this section. and is attached to the closure valve, or The female genital complex, as in the male, transfer valve, of the internal sac. During has its origin in both the mesoderm and inflation of the sac, it follows the sac apex the ectoderm. Products of the mesoderm through the lumen of the median lobe and are the ovaries, the lateral oviducts, and finally through the lumen of the sac. perhaps part of the common oviduct. Of Leopold (1941) discussed the role of male ectodermal origin are the vagina, the bursa accessory glands in reproduction. copulatrix, the spermatheca and spermathecal glands, part of the common oviduct,

14 and the ovipositor. These are shown in Surtees 1961; Iablokoff-Khnzorian 1966, figure 36. 1967; Teran 1967; Huignard 1968; King- In certain groups of bruchids (such as Ac- solver 1970a; Arora 1971; Singh 1973; anthoscelides and Callosobruchus), a pair Iablokoff-Khnzorian and Karapetian 1973; of structures first described by Mukerji Spirina 1974; Ahmed et al. 1976; Leopold and Bhuya (1937:200) are probably glan- 1941; Thukral 1976; Pawar and Verma dular in function and are positioned one 1977; Singh 1978a,b; Cassier and Huig- on each side of the vaginal tube at the en- nard 1979; Singh 1979; Ahmad and Murad trance of the bursa copulatrix (figure 36). 1980a,b; Kasap and Crowson 1980; Kumar These vary from being merely convex evagi- and Verma 1980; Monga and Sareen 1980; nations into the lumen of the vagina to Hamon et al. 1982; Singh 1982; Crowson oviform, membranous sacs nearly occlud- 1984; Kasap and Crowson 1985; Kasap ing the lumen. Their attachment is ringlike and Crowson 1988. with a central opening through which pass tracheae into the gland. These are lacking Larval Characters in Bruchus, but a pair of “glands” are posi- The destructive feeding period in the tioned at the anterior end of the bursa. The Bruchidae occurs during the larval stages, function of these structures is not known. yet bruchid larvae are relatively poorly A family-wide survey of their position and known and efforts to improve on their structure may offer phylogenetic clues. identification are in their infancy. Papers on Adult Morphology G.S. Pfaffenberger, who has contributed more to the study of bruchid larvae than Head morphology.—Stickney 1923; Ferris anyone else in the United States, kindly 1942; Tandon 1960; Mathur and Dhadial supplied the following description of larval 1961, 1963; Singh 1981c, 1982, 1983. characteristics. Brain.—Satija et al. 1975; Sandhu and Bruchid larvae have four instars, the first Neena 1982. being significantly different from the oth- Eyes.—Schmitt et al. 1982. ers (figures 37 and 38). Body sizes of first Thorax.—Snodgrass 1909; Hlavac 1972. instars of various species vary significantly but are characterized by the presence of a Metendosternite.—Crowson 1938, 1944. sclerotized X or H-shaped prothoracic plate Wings.—Forbes 1922; Kempers 1923; Mar- (figure 16) and a sclerotized abdominal cu 1939; Chen 1940; Seeliger 1943; Jolivet spine immediately above the spiracle of the 1957; Suzuki 1969; Wallace and Fox 1975, first abdominal segment. These structures 1980; Singh 1981a. are used to gain leverage to penetrate the integument of the seed or pod. Append- Alimentary canal.—Kasap 1978b; Mann ages, when present, are highly variable in and Crowson 1983a. length and form. Ventral nerve cord.—Kasap 1978a; Mann Integument.—Cream colored except pro- and Crowson 1983b. notum yellowish brown. Chaetotaxy of the Sense organs.—Pouzat 1981. first instar may consist of noticeably long Tracheal system.—Pajni 1969. or short sensilla, but they are distributed in recognizable patterns. Reproductive systems.—Kannan 1923; Tanner 1927; Zacher 1932; Zia 1936; Head.—Hypognathous and retractable; yel- Mukerji and Bhuya 1937; Mukerji 1949; lowish brown with mouthparts more darkly Mukerji and Chatterjee 1951; Srivas- pigmented. tava 1953a,b; De Luca 1959; Pajni 1959; Antenna.—With one to three telescopic segments. A single elongate sensillum chaeti-

15 cum and at least one sensillum basiconi- papers on Argentine larvae are also listed cum located distally (Pfaffenberger 1985b). by Pfaffenberger (1985a). Clypeolabrum.—Clypeal portion with single sensillum trichodeum near each lateral Materials and Methods border, subtended medially by sensillum ampullaceum. Labral portion rounded dis- Collecting and Rearing tally, single or double arc of sensilla tricho- The common name “seed beetles” for the dea located distally among mat of variable family Bruchidae is appropriate since length microtrichia; lateral sensillum trich- the larvae destroy the host seeds by their odeum and medial sensillum ampullaceum internal feeding and since the pupal stage located laterally near proximal margin of in most cases is passed within the feeding labrum (Pfaffenberger 1985b: figure 34). cavity. The adult, once it emerges from the Epipharnyx.—Epipharyngeal groove bor- seed, probably has nothing to do with the dered laterally by one to three pairs of seed until time for the female to deposit decurved sensilla trichodea. her eggs. During this free period before mating and oviposition, the adult may feed Mandible.—Monocondylic, with gouge- on nectar and pollen of whatever plant is in shaped mesal face. bloom at the time and may even be found Maxilla.—Cardo, stipes, palpifer, palpus, on flowers of plants that would not support and lacinia present. Membranous stipes development of the larvae. with variable number of sensilla trichodea. By far the most productive collection meth- Lacinia with five contiguous, distomedial, od, both in numbers of specimens and in spatulate spines. Palpus with dorsal, re- associating the species with its true host, cessed sensillum placodeum (Pfaffen-berg- is the collection of fruits containing seeds, er 1985b: figure 36). Distal end of palpus or the seeds themselves, and confining with variable number (usually greater than them until the adults emerge. This method 10) of retractible sensilla basiconica. presupposes some knowledge of potential Labium.—Labial palps absent. Submen- host plants but is the most likely to build tum transversely elongate. Base of men- a useful collection. Larvae and pupae can tum with lateral pair of sensilla trichodea, also be associated with the adult stage mental sclerite with paired islets midway by carefully opening some of the fruits or toward distal end. Islets usually with single seeds and extracting and preserving them sensillum trichodeum. Glossa and para- while awaiting emergence of the adults glossa narrowed distally. from the remaining seeds. Larvae are best Anal cleft.—Transverse in all Bruchinae preserved by dropping them in simmer- examined; Y-shaped in other subfamilies. ing hot water for a few minutes to kill the internal body bacteria, then transferring Pfaffenberger’s papers are listed in Refer- them to 70-percent alcohol for permanent ences. The “Literature Cited” section of one preservation. of his papers (Pfaffenberger 1985a) is an excellent source of pertinent works. Adults may be collected by sweeping flowers, but the usual result is small numbers Other papers on larval morphology include of specimens. Furthermore, sweep records Kannan 1923; Anderson 1943; Srivas- are unreliable as indicators of the true tava 1966; Prevett 1971; Vats 1972, 1973, larval host since adults are ubiquitous in 1974a,b, 1976a,b; Pfaffenberger and Jan- their feeding habits. Some species of bru- zen 1984; Hassan et al. 1987. Teran’s four chids diapause or aestivate in forest duff, birds’ nests, or spanish moss, under bark, or in similar habitats. A few species are attracted to light, but collecting at light is usually unprofitable.

16 Bottimer (1961) advocated the use of pint for treatment in caustic solution. The fol- fruit jars covered with cheesecloth held lowing procedure will produce consistent in place with a screwtop ring for rearing preparations: chambers. Johnson (1970, 1990c) essen- 1. Thoroughly relax the specimen either in tially recommended the same procedure a humidity chamber or in hot water or except that he keeps the jars in a room alcohol. with controlled temperature and humidity. Pyeomitid mites and small ants can invade 2. Gently lift the pygidium with fine forceps containers of seeds and destroy emerging to determine the sex of the specimen. adults. To avoid this, Johnson covers rear- Males have a hoodlike structure, the ing jars with paper towels treated with 1- eighth tergite, lying beneath the pygidi- percent Kelthane in acetone and covers the um; females do not have this structure. shelves on which the jars are placed with 3. Sever the membranes inside the terminal similarly treated paper (1990c:299). sternal segment and on the ventral side of D. Janzen (personal communication, 1972) the pygidium and remove the entire mass places the seeds or seed pods in a small of musculature and fat bodies along with plastic bag from which he exhausts the the genital parts, the eighth tergum, and air, then suspends that bag inside a larger the Y-shaped eighth sternite. inflated bag. Any bruchid that penetrates 4. Occasionally, depending on the preser- the smaller bag has difficulty escaping due vation technique, the fat bodies will be to the slippery inner surfaces of the outer hardened and will need to be further bag. softened with alcohol before removal of Bottimer (1961) recommended mainte- the genital parts. nance of a seed collection and an accom- 5. Immerse the genital complex in a 10- to panying herbarium for reference. Johnson 15-percent solution of potassium or so- follows this practice. Keeping green plant dium hydroxide until the muscle fibers parts in a confined situation invites the de- are softened and dissolved. velopment of mold. Plant specimens should 6. Transfer the genitalia to distilled water be dried immediately to avoid this prob- or 70-percent alcohol and wash thor- lem, but collections of seeds from which oughly. a worker wishes to rear bruchids cannot be subjected to the heat necessary to dry 7. With bent forceps, or bent needles, gently plants. press the liquified muscle tissue so that the genitalia are cleared of all debris. A Preparation of Male Genitalia second wash may be necessary to further neutralize the action of the hydroxide. Species identification in bruchids often depends on comparisons of characters of For temporary observation or for illustra- the male genitalia. Proper technique in tion, the parts may be mounted in glycerin preparing these organs for examination on a microscope slide beneath a cover slip. is essential for consistent results. Those Best results are attained when the median parts of the genitalia used in identification lobe, lateral lobes, eighth sternite, and are ectodermal in origin, and thus chitin- eighth tergite are separated on the slide. ous, and are connected by muscle bands to Definitive characters are most often found each other and to the body wall. To see the in the apex and internal sac of the median internal details clearly, this musculature lobe and in the form of the lateral lobes. must be dissolved in a potassium hydrox- Genital parts can be permanently stored in ide or sodium hydroxide solution. minute plastic vials partly filled with glyc- The genital complex is invaginated within the abdominal apex and must be removed

17 erin and closed with neoprene stoppers. Type Collection Abbreviations Such vials can be impaled on the pin below The following list includes all of the mu- the specimen for convenient association. seums wherein primary type specimens of American bruchid species are deposited. Measurements These abbreviations, with the exception of Certain measurements are consistently USNM, are from A Catalog of the Coleoptera used in species descriptions for compari- of America North of Mexico (U.S. Depart- son: ment of Agriculture, 1978–97). Head: BMNH: British Museum (Natural His- tory), London Ocular index: Greatest width across eyes divided by narrowest width between eyes CASC: California Academy of Sciences, measured on a horizontal plane. San Francisco Ocular sinus (emargination of eye): Mea- CNCI: Canadian National Collections, sured along the axis of the sinus, length Ottawa, ON of eye divided by the length of the sinus. FMNH: Field Museum of Natural History, Antennal length: Measured with head in Chicago normal hypognathous position and an- ICCM: Carnegie Museum, Pittsburgh tenna fully extended along side of body. MCZC: Museum of Comparative Zoology, Body length: Measured from anterior margin Harvard University, Cambridge, of pronotum to apex of abdomen. Sexual MA differences of the abdomen may cause some discrepancies. MNHP: Museum National d’Histoire Natural, Paris Body width: Measured across greatest width of elytra. NHRS: Naturhistoriska Riksmuseet, Stockholm, Sweden Legs: Mucro length measured against length of basitarsus, but measured separately OSUC: Ohio State University Collec- because of angular attitude of basitar- tions, Columbus, OH sus. PURC: Purdue University Collections, Pecten or denticle length: Measured at right West Lafayette, IN angles to metafemur, then compared with UCDC: University of California, Davis, width of tibia at its base. CA USNM: National Museum of Natural His- Resources tory, Washington, DC The Washington, DC, area was the most UZMC: University Zoological Museum, logical place to compile this handbook be- Copenhagen, Denmark cause the most comprehensive bruchid col- ZMUM: Zoological Museum, University of lection in the New World is at the National Moscow, Russia Museum of Natural History (Smith-sonian Institution). The most extensive library resources, holding many thousands of books and periodicals, are also located in the Washington area: National Agricultural Library, Smithsonian Institution Library, Library of Congress, National Library of Medicine, and several local universities.

18 Taxonomy of the Bruchidae of classification of North American bruchids. Principal contributors during this period the United States and Canada have been L.J. Bottimer, J.C. Bridwell, C.D. Johnson, J.M. Kingsolver, G.S. The family Bruchidae in the New World Pfaffenberger, A.L. Teran, and D.R. White- is well-developed in numbers of species, head. The most recent world catalog of spe- and 36 of the 41 genera of Bruchidae are cies was compiled by S. Udayagiri and S.R. presently considered endemic to the hemi- Wadhi (1989). A key to world genera was sphere (Kingsolver 1990a). published by S. Udayagiri and S.R. Wadhi Seven of the 24 genera in the United in 1982. States and Canada include accidently or deliberately introduced species. These are Acanthoscelides, Borowiecius, Bruchidius, History of the Family Name Bruchus, Callosobruchus, Caryedon, and The confusion over the correct name for Lithraeus. Some of the most notorious the family Bruchidae originated in the stored-product pests belong to these gen- application of the type genus and was era. All species now found in Hawaii are partly due to duplication of certain generic introduced (appendix I). names. Recent decisions by the Interna- The early history of North American bru- tional Commission on Zoological Nomen- chidology (late 18th century to early 19th clature, stimulated by applications submit- century) was dominated by European tax- ted by Borowiec, have clarified and stabi- onomists, and the type specimens for most lized nomenclatural situations not only in of the species described during this period the Bruchidae but also in the Meloidae. are in museums in London, Stockholm, A brief history of each of the involved Copenhagen, Moscow, and Paris. names follows: Kingsolver (1990a) outlined the history of Bruchus Linnaeus (1767), with pisorum New World bruchid taxonomy, dividing it Linnaeus (1767) as type species, is a into four broad periods with considerable large valid bruchid genus. At one time, overlap. These periods illustrate the evolu- most of the described species were tion of the current classification from ran- placed in either Bruchus or Spermopha- dom descriptions to a reasonable stability gus, but many more genera have been and from the generalist describing species proposed in the past 50 years, frag- in many families to the specialist concen- menting these older taxa. trating in one family or in one genus. The early taxonomists described species in the Bruchus Geoffroy 1762 is not available genera Bruchus, Caryoborus, and Sper- since it was proposed in a work placed mophagus. on the Official Index of Rejected and In- valid Works in Zoological Nomenclature. Early generalists in the United States Mylabris Geoffroy (1762) is not avail- include Fall, Horn, LeConte, Say, and able for the same reason. Schaeffer. Horn (1873) and Fall (1910) made the first attempts at classification Bruchelae was the first validly proposed by dividing all of the species then known family group name for the Bruchidae into species groups. Bridwell (1929c, 1932, (by Latreille in 1802) based on the 1946) laid the foundation for a world clas- species Bruchus pisorum (Linnaeus). sification of the family, and his work has The form “Bruchidae,” with the “-idae” not basically changed in the intervening ending, was first used by Stephens in years. 1829. Pope (1956) was of the opinion that since this usage was the first us- Taxonomic activities during the past 60 ing this ending, it should be the correct years have established a reasonably stable name for the family.

19 Laria Scopoli 1763 is not available as a Fossil Bruchidae bruchid generic name because Bridwell (1932) fixed the type species asLaria The only fossil Bruchidae known from dulcamarae Scopoli, a species in the North America were found in Florissant Nitidulidae. Bedel (1901) used Lariidae shale and White River fossil beds in the in error for the family name, and sev- western United States and were described eral papers subsequently used Lariidae. between 1876 and 1917 by S.H. Scudder and H.F. Wickham. At that time, nearly all Mylabris Müller (1764) validated Mylabris Bruchidae were placed in either Bruchus or Geoffroy (1762), and Bridwell (1932) Spermophagus; however, these two gen- fixed the type species asBruchus piso- era have subsequently been restricted to rum (Linnaeus), making Mylabris Müller species primarily found in the Old World. synonymous with Bruchus Linnaeus. The only fossil specimens I have examined Several authors followed Heyden et al. so far belong to the type series of six spe- (1883) in using Mylabridae, based on cies described by Wickham. These species Mylabris Müller, for the seed beetles. names were placed in the genus Oligobru- Mylabris Fabricius (1775) is a large valid chus Kingsolver (1965b) with Bruchus flo- genus in the Meloidae. To have two rissantensis Wickham designated the type large genera in different families with species. The remaining species are here left the same name is contrary to the rules, in the original genera until type specimens but most workers in the seed beetles and other material in the Colorado Mu- rejected Mylabridae and accepted Bru- seum and Museum of Comparative Zoology chidae as the correct name, and meloid can be examined. specialists recognized which specific The latest dating techniques place both names belonged to their family. For White River and Florissant formations in many years no one moved to petition the Oligocene near the transition to the the Commission to stabilize the names. Eocene period (about 35 million years ago), Pope (1956) suggested that Mylabris Müller although the Florissant beds were thought be placed on the Official Index of Rejected to be Miocene when Wickham described and Invalid Names but apparently never this species. submitted the application. Oligobruchus belongs to the subfam- Not until 1988 was an official proposal ily Pachymerinae and closely resembles submitted by Borowiec to not only have the Pachymerus, whose geographical range names Mylabris Müller 1764 and Bruchus does not now extend north of Costa Rica. Müller 1764 suppressed for Principle of A checklist of U.S. fossil Bruchidae can be Priority and Principle of Homonymy, but found in appendix II. also to have the names Mylabris Fabricius 1775 and Bruchus Linnaeus 1767 conserved by the International Commission, and to have the family name Bruchidae Latreille 1802 (type genus Bruchus Lin- naeus 1767) and the specific namepisorum Linnaeus 1767 (type species of the genus Bruchus Linnaeus) conserved. This proposal was approved by the International Com- mission in Opinion 1809, June 30, 1995 (International Commission on Zoological Nomenclature 1995).

20 Key to the Bruchidae of the United Pygidium only sclerotized and exposed beyond elytra States and Canada, Including Hawaii (figures 346 and 347); antenna of various shapes; mesepimeron either very narrow between mesepisternum 1. Pronotum flat or slightly convex with at least basal and and metepisternum (figure 26) or reduced to triangular lateral margins delimited by distinct submarginal sulcus sclerite on dorsal margin of mesepisternum (figure 27)...6 (figure 39); large species with metafemur enlarged, metatibia curved to fit closely to ventral margin of femur 6(5) Lateral pronotal margin with single large tooth projecting (figure 40); antenna serrate; eye shallowly emarginate: horizontally (figure 46); metafemur with tooth or angula- Pachymerinae...... 2 tion on ventrolateral margin lying outside metatibia when legs are closed (figure 47)...... Bruchus Pronotum convex, without submarginal sulcus except on anterolateral margin behind eye, other characters variable Lateral pronotal margin without large tooth but may be set ...... 3 with small denticles (figure 48); metafemur with (figure 50) or without external tooth, lobe, angulation, or row of 2(1) Prosternum flat between procoxal apices (figure 41); ely- denticles on ventrolateral margin...... 7 tra uniformly red to black; metatibia lacking lateral carina (figure 40)...... Caryobruchus 7(6) Ventrolateral margin of metafemur with a lobe, an angulation, a tooth, or a row of denticles (figures 50, 53, and 54) Prosternum short, triangular, acute, not separating apices ...... 8 of procoxae; elytra yellowish brown speckled with dark brown; metatibia with lateral carina (figure 42); Hawaii Ventrolateral margin of metafemur without lobe, angula- and Florida...... Caryedon tion, tooth, or row of denticles...... 11 3(1) Metatibia with two apical movable spurs (figures 4, 29, 8(7) Ventrolateral margin of metafemur with row of denticles and 43); metacoxal face broader than metafemur (figure (figure 53), ventromesal margin with several denticles; 4): Amblycerinae...... 4 pronotum gibbous; female pygidium with glabrous median spot...... Metatibia lacking movable apical spurs but may have Gibbobruchus fixed apical spines or denticles (figure 44); metacoxal Ventrolateral margin angulate or strongly lobed (figures face as wide as or narrower than metafemur (figure 44)...5 50 and 53), ventromesal margin with single denticle...... 9 4(3) Eye feebly emarginate at antennal socket (figure 1); 10th 9(8) Lateral pronotal margin with a distinct, arcuate carina...... stria reaching nearly to apex of elytron; prosternum sepa- ...... Stator rating coxae at apex (figure 45); metatibia lacking carinae; Lateral pronotal margin lacking distinct carina...... 10 tibial spurs unequal in length (figure 29) ...... Amblycerus 10(9) Hind legs red, or reddish brown with at most partial darker clouded area, especially along ventral margin of Eye deeply emarginate (figure 17); 10th stria abbreviated hind femur...... Callosobruchus opposite metacoxa; procoxae not separated by prosternum; tibial spurs equal in length (figure 43); metatibia Hind legs entirely black except tarsi yellow...... carinate (figure 43)...... Zabrotes ...... Borowiecius 5(3) Pygidium and preceding two tergal sclerites fully sclerotized and exposed beyond apices of elytra (figure 51); male antenna strongly pectinate (figure 84), female antenna serrate; mesepimeron not strongly narrowed at ventral end, broadly reaching coxal cavity: Kytorhininae...... Kytorhinus

21 Metafemur with multiple denticles on ventromesal margin 11(7) Tenth elytral stria ending opposite metacoxa (figure 55) (figure 64); mucro variable in length...... 21 (except M. impiger); frontal carina sharp; eyes of male 16(15) Metafemoral denticle triangular, serrate on caudal margin enlarged; male antenna pectinate (figure 6), female an- (figure 65); in seeds of malvaceous plants; east of 100th tenna serrate (figure 7); metafemur not enlarged, ventral meridian...... Althaeus face flat between ventral carinae; ventro-mesal carina smooth (figure 56) or serrate (figure 52)...... Megacerus Metafemoral denticle usually small, not serrate, or absent; host plants various...... 17 Tenth stria extending to apical margin; frontal carina various; eyes dimorphic or not; antenna various; metafemur 17(16) Metatibia lacking lateral and ventral carinae (figures 66, usually moderately or strongly enlarged with ventral face 335, and 714); metafemoral denticle absent or extremely taking varied forms...... 12 minute...... 18 12(11) Front of head with V- or Y-shaped, glabrous boss be- Metatibia with at least one carina, usually two to four; tween dorsal margins of eyes (figure 57); male metafemur metafemur with one denticle...... 19 with setose pocket on ventral margin (figure 58); mucro 18(17) Elytral color uniform, without maculae; elytra together as short, usually not as long as lateral denticle wide as long; body length less than 2 mm; South Texas...... Mimosestes In seeds of Abutilon...... Abutiloneus Front of head otherwise; male metafemur lacking pocket; Elytra maculate, longer than wide; body length 2.4 mm; mucro variable...... 13 Hawaii...... Lithraeus 13(12) Fourth stria only abbreviated at base and ending in small 19(17) Pronotum moderately or strongly gibbous (figures 67 and denticle (figure 59); metatibia strongly bent at base; mu- 824); southwestern United States...... Neltumius cro minute; pecten of 5–7 minute denticles (figure 729) Pronotum uniformly convex, not gibbous; widely distrib- ...... Meibomeus uted...... 20 Fourth stria not abbreviated; with more than one stria 20(19) Body and appendages all black except two basal antennal denticulate at base, or stria not denticulate; metatibia usu- segments sometimes reddish brown; metafemoral den- ally slightly arcuate at base, sometimes with entire tibia ticle minute, hidden among marginal setae (figures 288 arcuate; mucro of variable length...... 14 and 294)...... Bruchidius 14(13) Scutellum 1 1/2 to 2 times as long as wide (figure 60); Body and appendages, in part or wholly, red or reddish female pygidium with two deep, polished sulci (figure orange; metafemoral denticle easily visible, sometimes 61); male genitalia with H-shaped sclerite in armature of as long as width of metatibia at its base (figures 864 and internal sac (figure 62)...... Algarobius 877)...... Sennius Scutellum quadrate or transverse; female pygidium 21(15) Head long (figures 68 and 678); width across eyes equal lacking polished sulci; male genitalia lacking H-shaped to distance from top of eyes to end of clypeus; 3rd and sclerite...... 15 4th elytral striae ending in prominent single basal gib- 15(14) Metafemur with single, sometimes minute, denticle on bosity (figure 69); pronotum with prominent median and ventromesal margin (figures 63 and 714), or without lateral gibbosities; Florida...... Caryedes denticle or denticles; mucro usually no longer than lateral Head short (3:1); elytral striae usually ending in basal denticle...... 16 denticles, sometimes on slight swelling; pronotum not gibbous; widely distributed...... 22

22 22(21) Antenna extremely long (figures 13 and 971), that of male extending beyond apices of elytra, that of female extending to 1st or 2nd abdominal segment; mucro slender, curved, one-half as long as basitarsus (figures 70 and 972); 3rd and 4th striae prominently denticulate (figure 968); pecten with three minute, slender denticles (figure 972); eye deeply emarginate (figure 969)... Stylantheus Antenna not exceptionally long, never extending beyond middle of elytra; mucro various; striae denticulate or not; pecten variably formed...... 23 23(22) Elytral striae 3 and 4 denticulate (figure 71 and 72), denticles sometimes on transverse ridge or slight swelling (except M. major with denticles on 3 to 6); anal notch of female 5th sternum deep, usually laterally flanged (figure 73); metafemur strongly swollen; tibia strongly arcuate (figure 64); ventral valve of male genitalia broadly rounded or truncate (figures 74 and 761); armature of internal sac usually includes variant of forked sclerite (figures 74 and 761)...... Merobruchus Denticles of elytral striae various, sometimes absent, never on basal gibbosity; metafemur usually not strongly swollen; tibia various; female 5th sternum not notched or flanged; male genitalia of various forms, seldom with forked sclerite...... Acanthoscelides

Some species of Merobruchus resemble certain species of Acanthoscelides, and the distinction between the two genera remains nebulous at various points. Unequivocal definitions of these two genera are not possible at this time.

23 Subfamily Pachymerinae Pachymerus gonager: Chujo 1937a:82. Bridwell 1929c Caryedon gonagra: Herford 1835:22; Southgate and Pope 1957:669; Dav- Tribe Caryedontini Bridwell 1929c ey 1958:385; Cancela da Fonseca 1964:633; Prevett 1965; Donahaye et Bridwell (1929c) proposed the tribal name al. 1966; and many other references. Caryedini (emended to Caryedontini by De- Pachymerus acaciae: Lepesme 1944:215; celle 1968) to include only Caryedon, but Cancela da Fonseca 1956. Decelle described four additional African Pachymerus longus Pic 1913a:8. genera in this tribe. Bruchus fuscus Bedel 1901:341. Taxonomy.—Nilsson and Johnson 1993a. Pachymerus sibutensis Pic 1924:42. Caryoborus gonagra: Gyllenhal 1833:96. Genus Caryedon Schoenherr Pachymerus gonagra: Pic 1913a:7. Caryedon Schoenherr 1823:1134; See Davey (1958) and Decelle (1966) for Bridwell 1929c:144; Southgate and additional taxonomic combinations. De- Pope 1957; Johnson 1966:162; Bot- celle succinctly summarized the taxonomic timer 1968c:1043; Cancela da Fon- history of Caryedon serratus. seca 1975:71; Johnson and King- solver 1982:409; Singh 1982; Borow- Color.—Integument reddish brown to pice- iec 1987:48; Udayagiri and Wadhi ous; elytra with irregular piceous macula- 1989:233. Type species: Bruchus serrations (figure 75); pygidium mostly dark tus Olivier 1790:199, by monotypy. brown with median line and apex yellow. Pubescence reddish brown to yellow. Medium-sized bruchids (2.2–6.7 mm long). Structure.—Body elongate, strongly convex Frontal carina prominent; eye facets hemi- (figures 1 and 75); vertex finely imbricate; spherical, eye shallowly emarginate (figure frons minutely punctate, sparsely foveo- 1); antenna serrate (figure 1); pronotum late; frontal carina prominent; ocular index flat, lacking lateral carina; prosternum 7.2:1 in males, 6.5:1 in females; ocular short, not separating procoxal apices; sinus shallow, one-eighth length of eye mesepisternum broader than mesepi- (figure 1); facets of eye coarse, hemispheri- meron; metafemur swollen dorsoventrally, cal; antenna elongate (figure 1), coarsely pecten with 10–15 denticles (figures 2 and serrate, male antenna reaching metacoxa, 78); metatibia arcuate, curved to fit ventral female antenna reaching middle of met- margin of tibia (figures 1 and 78). episternum. Pronotum trapezoidal, lateral margins rounded anteriorly, disk slightly Caryedon serratus (Olivier) convex, without asperities or sulci, surface (Figures 1, 38, 42, 75–78) densely microfoveolate, interspaces punctulate; lateral carina blunt, extending only Bruchus serratus Olivier 1790:1134; halfway to apex; cervical sulcus completely Southgate and Pope 1957:669. encircling anterior foramen; prosternum Caryedon serratus: Schoenherr 1823:1134; short, triangular, not separating procoxae Decelle 1966:1; Kingsolver 1967:900, at their apices. Scutellum rectangular, flat 1970c:303; Velez-Angel 1972:71; (figure 75); mesepisternum broader than Kingsolver et al. 1977:114; John- mesepimeron. Elytra 1.5 times as long as son 1986:264; Udayagiri and Wadhi wide, strongly convex; striae subparallel, 1989:233; Kingsolver 1992a. two and three slightly divergent near base, Bruchus gonagra Fabricius 1798:159. shallow, punctures elongate, narrow; in- Caryoborus gonagra (Fabricius): Swezey terstices minutely imbricate; metepimeron 1912:167. with angulate sulcus; metacoxa uniformly

24 punctulate except glabrous near trochan- Natural enemies.—World list: Bracon kirk- teral fossa; hind leg (figure 78) swollen dor- patricki, Bracon sp.; Entedon sp.; Eupelmus soventrally; pecten with 14 small denticles swezeyi; Goniozus emigrata; Heterospilus in arcuate row; metatibia strongly arcuate prosopidis; Oedaule stringifrons; Para- to fit against ventral margin of femur, with sierola emigrata; Sclerodermus immigrans; one lateral, one mesal, and three ventral Stenocorse bruchivora; Urosigalphus bruchi; carinae (figures 42 and 78); mucro blunt, Uscana caryedoni, U. semifumipennis. See slightly concave on dorsal face; tibial apex Gagnepain and Rasplus (1989) for African truncate, lacking denticles. Abdomen not parasitoids. modified except male 5th sternum emar- Immatures.—Prevett 1967a (larva); Arora ginate; pygidium narrow, apically rounded, 1978:45 (larva); Wightman and Southgate disk shallowly imbricate, densely setose. 1982 (egg). Male genitalia.—As in figures 76 and 77; Discussion.—This genus comprises a num- median lobe apically broadened; ven- ber of African and Asian species that at- tral valve subtriangular but basal angles tack seeds of Leguminosae and Combret- rounded, dorsal valve subtriangular, less aceae. Caryedon serratus has an artificial acute than ventral valve; internal sac with tropicopolitan distribution because of its a mass of fine spicules, a pair of stout, association with Tamarindus indica. This hooklike denticles each with an auxiliary bruchid is known from Florida and Hawaii denticle near apical orifice, two pairs of in the United States but is found in Mexi- slender, hooked sclerites with associated co, Colombia, Guyana, and the West Indies fine spicules at apical two-thirds, and a (Johnson 1966, 1986; Kingsolver 1970c; mass of microspicules and a tubelike valve Velez Angel 1972; Kingsolver 1992a). at apex; lateral lobes broad, caudal faces concave, median cleft shallow. Because of its importance as a storage pest of groundnuts in Africa, various biologi- Size.—Body length 3.5–6.8 mm; width cal aspects of this species have received 1.8–3.0 mm. considerable attention by Old World work- Type depository.—MNHP. ers. The following references will serve as a Type locality.—Senegal. guide to the literature: Distribution.—FL, HI; Mexico, Haiti, Ja- Chromosomes.—Yadav 1973; Smith and maica, Virgin Islands, Dominica, Curaçao, Brower 1974; Thiara et al. 1988. Venezuela; Central Africa, Madagascar, Distribution.—Kingsolver 1970c, 1992a. Middle East, Indian Subcontinent, South- Egg morphology.—Wightman and South- east Asia. gate 1982. Host plants.—World list: Acacia farnesiana, Larval digestive system.—Vats 1976b. A. seyal, A. spirocarpa, A. tortilis; Adenan- thera pavonina; Arachis hypogaea; Bauhin- Larval malpighian tubules.—Vats 1976a. ia malabarica, B. monandra, B. racemosa, Larval morphology.—Prevett 1967a. B. reticulata, B. thonningi, B. rufescens, B. tomentosa; Caesalpinia pulcherrima; Caja- Larval tracheal system.—Vats 1972. nus cajan; Cassia afrofistula, C. arereh, C. Life history.—Lepesme 1944; Donahaye et fistula, C. grandis, C. javanica javanica, al. 1966; Cancela da Fonseca 1965; Con- C. javanica indo-chinensis, C. sieberiana; way 1983. Phaseolus vulgaris; Prosopis grandulosa Ovaries.—Monga and Sareen 1980. grandulosa, P. pallida; Senna obtusifolia; Tamarindus indica. Physiology.—Garg et al. 1990. Population dynamics.—Amaro et al. 1958.

25 Reproduction.—Robert 1985; Boucher and concave on dorsal face; apical spurs ab- Huignard 1987; Boucher and Pierre 1988. sent. Reproductive system.—Mukerji and Chat- terjee 1951; Mukerji et al. 1957; Southgate Caryobruchus gleditsiae (Linnaeus) and Pope 1957; Prevett 1967b; Arora 1977; (Figures 2, 12, 14, 16, 18–23, 25, 28, Singh 1982. 39–41, 79–82) Spermatheca.—Surtees 1961. Dermestes gleditsiae Johansson and Lin- Spermathecal gland.—Thukral 1976. naeus 1763:9. Bruchus gleditsiae: Linnaeus 1767:605. Taxonomy.—Southgate and Pope 1957; Davey 1958; Prevett 1965; Decelle 1966; Pachymerus gleditsiae: Pic 1913a:7. Singh and Yadav 1979. Caryobruchus gleditsiae: Bridwell 1929c:155; Lepesme 1947:572; Black- Tribe Pachymerini Bridwell 1929c welder and Blackwelder 1948:44; Craig- head 1950:279; Bottimer 1968c:1038; Genus Caryobruchus Bridwell Woodruff 1968:1; Fonseca 1981:71; Johnson and Kingsolver 1982:409; Caryobruchus Bridwell 1929c:148; Bridwell Borowiec 1987:43; Udayagiri and Wa- 1946:53; Kingsolver 1965b:29; dhi 1989:239; Nilsson and Johnson Prevett 1966a:181, 1966b:81; Bottimer 1990:50; Johnson et al. 1995:31 1968c:1037,1039; Johnson and King- solver 1982:409; Borowiec 1987:43; Bruchus arthriticus Fabricius 1801:398. Udayagiri Caryoborus arthriticus: Schoenherr and Wadhi 1989:238. Type species: 1833:93; Horn 1873:312; Schwarz Dermestes gleditsiae Linnaeus, by origi- 1878:457; Cushman 1911:504–505; nal designation. Forbes 1922 (wing venation). Large bruchids (5.0–25.0 mm). One species Bruchus fuscus Goeze 1777:332. in the United States. Caryobruchus arthriticus: Zacher 1952:468. Head subtriangular, strongly constricted Color.—Integument piceous, sometimes behind eyes, transverse sulcus distinct, reddish in teneral specimens. Vestiture eyes coarsely faceted, shallowly emarginate gray, short, inconspicuous, without at antennal insertion; antenna serrate from pattern except for scattered, white ely- 4th segment, terminal segment elongate- tral spots (figure 79). oval. Pronotum quadrate, anterior angles Structure.—Vertex densely punctulate, rounded (figure 39); disk flat, coarsely frons punctulate with scattered foveae, foveolate, submarginal sulcus bounding frontal carina sometimes present (fig- disk; lateral carina distinct, anterior corner ure 18); clypeus sparsely punctate with with two or three setose punctures; pro- scattered umbilicate foveolae; venter of sternum narrow, flat, completely separat- head as in figure 21. Prothorax as in ing procoxae (figure 41); protrochantins generic description. Scutellum quad- and mesotrochantins distinct; mesepi-ster- rate (figure 79). Elytra longer than wide, num and mesepimeron subequal in width; convex, striae parallel, except deflected metafemur dorsoventrally broad (figure laterally near base, 1st stria curved 40), ventral margin with 10–12 prepecten around scutellum, strial punctures denticles, pecten with one long and 9–11 shallow, elongate; interstices moderate- shorter denticles in arcuate row in apical ly convex, minutely punctate-imbricate; one-half of mesoventral margin, denticles mesotrochantins visible; postmesocoxal lying mesad to tibia in closed position; sulci deep, joined at midline, laterally metatibia strongly curved fitting closely parallel to pleurosternal suture nearly to ventral margin of metafemur, tibia with to margin of antecoxal piece (figures 28 three ventral carinae; mucro short, acute,

26 and 80); metepisternal sulcus deep, an- was being used as a hibernaculum, and gulate, bent mesad and extending par- in wood rat nests. allel to pleurosternal suture; metacoxal Natural enemies.—None recorded. face minutely punctate, fossula deep, Immatures.—Cushman 1911 (egg); Pfaffen- extending to lateral articulation of coxa; berger and Johnson 1976 (egg, 1st 5th abdominal sternum of male broadly larval instar); Pfaffenberger 1991:564 emarginate; pygidium subtriangular, (larva). apex rounded in male, broadly truncate in female; discal surface with irregular, Discussion.—This is the largest species of lunate punctures nearly concealed by Bruchidae in the United States; some vestiture. individuals reach 10 mm in length. Specimens reared from Sabal by Fon- Male genitalia.—As in figures 81 and 82; seca (1981) in Veracruz, Mexico, are median lobe broad (figure 81), apex probably this species. The original host ogival, tip acute, not separated into was mistakenly identified asGleditsia— a movable ventral valve, dorsal valve hence the specific name of the beetle. broadly rounded; middle one-third of internal sac lined with minute den- This species has been reported to be capa- ticles, armature consisting of a pair ble of jumping considerable distance since of falcate sclerites near base, a pair of the hind legs are greatly enlarged, but this elongate, serrate sclerites enclosed in feat does not seem logical since the depres- lateral pockets of sac near apex, a pair sor muscles in the femur are much more of elongate, lightly sclerotized lobes greatly developed than the levator muscle, between serrate sclerites, and a large, just the opposite of that in a saltatorial leg. alate, lightly sclerotized, apical struc- Stimulation of the hind leg of live beetles ture (transfer apparatus?); lateral lobes can bring about a grasping and pinching (figure 82) fused basally into straplike action, possibly a form of defense. structure, apices fused into broad, alate The wing venation (figure 14) was previ- lobe briefly cleft on midline, apical mar- ously illustrated by Forbes (1922). gin setose. Size.—Body length 5.1–10.0 mm; width 3.3–4.8 mm. Body size depends largely on the size of the host seed. Type depository.—BMNH. Type locality.—”America septentrionalis.” Distribution.—AR, FL, LA, NC, SC, TX; West Indies, Mexico. Woodruff (1968) mapped collections in Florida. Host plants.—Sabal etoni, S. mexicana, S. minor, S. palmetto, S. uresana; Serenoa repens; Washingtonia filifera. Woodruff (1968) and Johnson et al. (1995) list palms from which this bruchid has been reared or on whose seeds eggs have been found: Coccothrinax argentata, C. martii; Livingstona chinensis; Phoenix sylvestris; Sabal yapa, S. parviflora, S. minor, S. glaucescens, S. mauritiaeformis, S. palmetto; Serenoa repens. Specimens have been collected from Tillandsia usneoides, which apparently

27 Subfamily Kytorhininae 1968:1269, 1976a:50; Decelle 1971:106; Johnson and Kingsolver Bridwell 1932 1982:411; Borowiec 1987:68; Udayagiri and Wadhi 1989:223. Kytorrhinae Hoffman 1965:63 Kytorrhinus prolixus: Teran 1967:307 (male Kytorrhininae Luk’yanovich and Ter- genitalia). Minassian 1957 Color.—Integument dark brown to black. Genus Kytorhinus Fischer Vestiture of short, white setae, sometimes von Waldheim with scattered golden brown setae; scutellum white (figure 83). Kytorhinus Fischer von Waldheim Structure.—Vertex and frons reticulately 1809:298; Bridwell 1932:101,105, punctate, frontal carina evanescent, fron- 1946:53; Blackwelder and Blackwelder tal lobes carrying antennal fossae alate, 1948:45; Brown 1952:342; Bottimer prominent; eyes more prominent in male 1961:293, 1968c:1039,1068; John- than in female; ocular index 8:1 in males, son 1968:1269; Decelle 1971:106; 3.5:1 in females; ocular sinus four-fifths Johnson and Kingsolver 1982:411; length of eye; male antenna strongly pec- Borowiec 1987:68; Udayagiri and Wa- tinate (figure 84), extending beyond apex dhi 1989:223. Type species: Kytorhi- of elytron, female antenna serrate, extend- nus karasini Fischer von Waldheim ing to 1st abdominal segment. Pronotum 1809:298, by subsequent designation, bell-shaped, lateral margins sinuate (figure Crotch 1870:222. 83); disk densely, finely imbricate, setae Cytorhinus Agassiz 1846:115. Unnecessary short, recumbent, disk shallowly depressed emendation of Kytorhinus. near posterior corners; lateral carina Pygobruchus Sharp 1886b:38. Type spe- evanescent in some specimens, absent in cies: Pygobruchus scutellaris Sharp others; cervical sulcus deep, extending 1886:38 (=Kytorhinus sharpianus nearly to meson; prosternum short, trian- Bridwell 1932:106, not Kytorhinus scu- gular, procoxae contiguous. Protrochantin tellaris (Fabricius) 1792), by monotypy. and mesotrochantins present. Scutellum Kytorrhinus Baudi 1886:10; Luk’yanovich longer than wide, bifid apically; mesepim- and Ter-Minassian œ1957:181; Teran eron narrow but extending to coxal cavity. 1967:307. Elytra 1.3 times as long as wide, evenly Kytorhynus Motschulsky 1874:204. convex, humeri prominent; striae parallel, slightly deflected laterad near base, shal- One species in the United States and lowly impressed, punctures elongate, two Canada. Because this at present is a to six ending basally in transverse carina; monotypic genus, the generic and specific interstices finely imbricate; metacoxal face diagnoses are combined. of females entirely punctate (except fos- Redescription: Johnson 1976a:50. sula), males with mesal one-third glabrous; metafemur slender, unarmed; metatibia Kytorhinus prolixus (Fall) slender, mucro a small tooth. Abdomen not especially modified, sterna of male (Figures 51, 83–86) telescoped; 5th and 6th terga (prepygidial) Mylabris (Bruchus) prolixus Fall 1926:204. sclerotized (figure 51), similar in texture to Kytorhinus prolixus: Bridwell 1932:101, pygidium, discal surface finely imbricate. 1946:53; Blackwelder and Blackwelder Male genitalia.—As in figures 85 and 86; 1948:45; Brown 1952:342; Bottimer median lobe moderately broad; ventral 1961:293, 1968c:1036,1039; Johnson valve triangular, lateral margins incurved, apex acuminate; internal sac armed with

28 complex of brushes, clusters of spines, a pair of elongate, clavate sclerites, a pair of lateral, hooklike spines articulated to median complex of recurved spines and paired setal brushes; lateral lobes fused, apex expanded, apex with pair of digitate processes. Size.—Body length 1.9–2.7 mm; width 1.0–1.4 mm. Type depository.—MCZC, lectotype No. 25059, by Johnson (1976a). Type locality.—Alaska, McKinley Park Sta- tion. Distribution.—AB, AK, BC, ID, MB, NT, SD, SK, YT. Host plants.—Hedysarum alpinum americanum, H. boreale (neither plant confirmed as a larval host record). Natural enemies.—None recorded. Immatures.—Not described. Discussion.—This species is the only North American representative of the genus; all other species are Asian. The combination of characters—two sclerotized, prepygidial terga; strongly pectinate male antenna; prominent eyes in male; uniformly dark brown or black integument; unarmed metafemur; and lack of tibial carinae—easily characterize this species.

29 Subfamily Amblycerinae (calcaria) (figure 87); protibia and mesotibia not calcarate; mesepisternum and Bridwell mesepimeron subequal in width, attaining coxal cavity. This subfamily is worldwide in distribution, and includes the following genera: In addition, the dorsal and ventral valves of Amblycerus Thunberg, Zabrotes Horn, and the male genitalia are separate and mov- Spermophagus Schoenherr. able (figure 30), and the bases of the lateral lobes are fused into a flat, straplike struc- Characteristics of the subfamily (modified ture (figure 31). from Bridwell 1932) are metafemur not strongly swollen, only one-half as broad Both Amblycerus and Zabrotes are restrict- as metacoxa, channeled and longitudi- ed to the Western Hemisphere (including nally and ventrally bicarinate with carinae the Galapagos Islands) except Zabrotes usually unarmed, never with more than a subfasciatus (Boheman), which has be- single blunt denticle; trochantins of fore come a tropicopolitan tramp spread by legs and mid legs visible; metacoxa broad, commercial shipments of Phaseolus bean broader than 1st length of abdominal seeds either for propagation or for food. segment; metatibia straight, not mucro- This species probably originated in Mexico nate but with two movable terminal spurs or Central America.

Key to Genera of North American Amblycerinae

1 Emargination of eye shallow, depth less than Emargination of eye deep, one-half or more one-third diameter of eye (figures 109 and of diameter of eye (figure 17); parasutural 122); metathorax with parasutural sulci on sulci absent; procoxae contiguous or sepa- either side of pleurosternal suture; procoxae rated by a thin vertical lamella; metatibia with well separated by intercoxal process (figure one or more prominent lateral carinae (figure 45); metatibia without prominent lateral cari- 196)...... Zabrotes nae...... Amblycerus

The Old World genus Spermophagus as Tribe Amblycerini Bridwell now defined is similar in form to Zabrotes Amblycerinae Bridwell 1932:103 but can be separated from it by the following: Tenth elytral stria extending nearly to Amblycerini: Borowiec 1987:60 apex of elytron (extending only halfway to Genus Amblycerus Thunberg apex in Zabrotes), eye in repose partly concealed by anterior corner of pronotum (eye Amblycerus Thunberg 1815:121; Hatton free of pronotum), apex of ventral margin 1895:290; Bridwell 1930:29,1946:53; of metafemur with short, vertical sulcus Blackwelder 1946:762; Bradley (ventral margin entire). 1946:97; Blackwelder and Blackwelder 1948:44; Bottimer 1968c:1012; John- Borowiec (1987:60–63) proposed to divide son 1968:1268; Kingsolver 1970a:471, the Amblycerinae into two tribes: Amblyc- 1980a:230; Johnson and Kingsolver erini Bridwell to include only Amblycerus, 1982:410; Borowiec 1987:60; Udaya- and Spermophagini Borowiec to include giri and Wadhi 1989:15; Romero et al. Zabrotes and Spermophagus. 1996. Although Spermophagus has been included Anthotribus Gistel 1856:815. in New World species catalogs, no species Spermophagus, of authors. in that genus is now known to exist in the Western Hemisphere. See Kingsolver and Small to medium size bruchids (U.S. spe- Borowiec (1988) for discussion. cies 2.6–6.0 mm).

30 Body subovate, arched in profile (figure 4). Head subtriangular, strongly constricted Key to U.S. Amblycerus behind eyes; eyes large, convex, facets 1. Pronotum and elytra with scattered, black, setiferous coarse; emargination at antennal fossa shallow; frons finely carinate to smooth; foveae (figure 88); integument mostly reddish brown antenna strongly serrate from 4th segment to black; apex of female pygidium trilobed (figure 89). (figure 4). Pronotum bell-shaped, trapezoi- Widespread in U.S.A...... robiniae (Fabricius) dal, or semicircular, moderately convex, Pronotum and elytra lacking black foveae; integument red without asperities, with traces of submar- or reddish brown; female pygidium evenly rounded...... 2 ginal sulcus on basal margin and anterolateral corners; procoxae distinctly separated 2. Elytral striae moderately deep, coarsely punctate, inter- by intercoxal process of prosternum. Scu- stices uniformly convex (figure 90); pronotum densely tellum distinct, with characteristic shape punctate; metepisternum with elliptical stridulatory file for many species. Elytra with striae not (figure 91); metafemur with blunt tooth on mesoventral distorted, 10th stria reaching elytral apex; margin. Florida...... eustrophoides (Schaeffer) metasternum and metepisternum with Elytral striae with fine elongate punctures, sometimes parasutural sulci parallel to pleurosternal suture; metatibia lacking carinae. Pygidium hidden by vestiture (figure 92); metepisternum lacking oblique to vertical, disk nearly flat. stridulatory file; metafemur lacking ventral tooth...... 3 Armature of internal sac of median lobe 3. Scutellum short, triangular; pygidium with prominent characteristic for each species. median line of setae; pronotum and sometimes elytra with Thunberg included eight species names stripes of yellowish setae...... 4 with his description of Amblycerus—three of Scutellum rectangular or elongate; pronotum and elytra them new. The remaining five names were lacking stripes...... 5 cited without author—nebulosus, robiniae, reticulatus, scabrosus, and varius, but he 4. Male genitalia with paired prominent, thorn-like sclerites did not designate a type species. Crotch near base of internal sac (figure 101). Florida (1870:227) designated nebulosus, attribut- ...... nigromarginatus (Motschulsky) ing the name to “Fabr.” Bridwell (1930:29) Male genitalia lacking prominent, thorn-like sclerites but rejected Crotch’s designation on the may have small denticles near base of internal sac (figure grounds that nebulosus could not be identi- 106). Texas...... obscurus (Sharp) fied with any of the three species available then to Thunberg—Anthribus nebulosus 5. Size 2.6–3.7 mm; body uniformly red except eyes black. Forster (1771), Bruchus nebulosus Olivier Known only from Arizona ...... vitis (Schaeffer) (1795), or Macrocephalus nebulosus Olivier Size 4.5 mm or longer; color various...... 6 (1795). Bridwell then selected Bruchus robiniae Fabricius (1781), to replace nebulosus 6. Scutellum trilobed at apex; elytral vestiture uniformly as the type species. Bradley (1946) argued silvery gray; pygidium with distinct median stripe; body that Crotch’s designation was valid and length 4.5–5.0 mm; male genitalia as in figures 98 and that Bridwell’s action was unjustified since 99. Known only from Texas the name robiniae as Thunberg listed it could have referred to any of three species ...... ireriae Romero, Johnson, and Kingsolver by that name. Bridwell’s concept of Ambly- Scutellum tapered to acute apex; elytral vestiture vaguely cerus was further developed in later papers mottled; pygidium occasionally with diffuse stripe; body (1932,1946) and was adopted by Bottimer length 6–8 mm; male genitalia as in figures 114–117. In (1968c), Kingsolver (1970b and subsequent U.S.A. from Florida Keys papers), and Romero et al. (1996). Other ...... schwarzi Kingsolver authors (see A. robiniae citations) have used the name in Bridwell’s concept.

31 Amblycerus eustrophoides (Schaeffer) bia with apical margin diagonal, finely denticulate, lateral tibial spur two-thirds (Figures 87, 90, 91, 93, 94) length of basitarsus, ventral spur one-half Spermophagus eustrophoides Schaef- as long as lateral spur (figure 87). Abdo- fer 1904:228; Schaeffer 1907:293; Pic men unmodified except 5th segment of 1913a:59. male broadly emarginate; pygidium convex, Amblycerus eustrophoides: Bottimer foveolate, apex of male rounded, of female 1968c:1012; Johnson 1968:1268; King- subtruncate. solver 1970b:474; Johnson and King- Male genitalia.—As in figures 93 and 94; solver 1982:410; Udayagiri and Wadhi median lobe moderately broad (figure 93); 1989:9; Kingsolver 1992b; Kingsolver et ventral valve triangular, apex acute, lateral al. 1993; Romero et al. 1996:48. margins incurved, dorsal valve rounded Color.—Body and appendages uniformly apically; armature of internal sac consist- reddish yellow to dark red with margins ing of paired, acute basal sclerites with of pronotum and scutellum piceous, frons finely serrate basal margins, middle of sac sometimes dark brown. Vestiture of uni- with paired, elongate sclerites attached to formly distributed fine silvery gray hairs, V-shaped sclerite, apex of sac with elon- pygidium with narrow median line of hairs. gate, looped closure valve; lateral lobes Y- Structure.—Body fusiform in outline. Vertex shaped (figure 94), median cleft rounded. and frons evenly convex, densely punctate, Size.—Body length 6.0.–6.25 mm; width frontal carina lacking but some specimens 3.0–3.5 mm. have median impunctate line; ocular index Type depository.—USNM, lectotype No. 3:1, ocular sinus about one-fifth length of 42284 (by Johnson). eye, ocular facets large, spherical; postocular lobe narrow, sparsely fringed; an- Type locality.—Florida, Lake Worth. tenna serrate from 4th segment, reaching Distribution.—FL; Cuba, Mexico. metacoxa. Pronotum trapezoidal, lateral Host plants.—Drypetes lateriflora; “huilo- margins nearly straight, apex truncate, tillo” (Mexico). disk evenly convex, densely but irregularly microfoveolate, intervals micropunctate; Natural enemies.—None recorded. lateral carina extending to anterior corner, Immatures.—Not described. submarginal sulcus extending across midline; prosternum narrow, flat. Scutellum Discussion.—This species is easily distin- longer than wide, tridentate, slightly con- guished from the other U.S. species by the stricted on lateral margins. Elytra together combination of uniform coloration, distri- 1.3 times as long as wide (figure 90), evenly butional pattern, size, and the presence convex except slightly depressed around of the fusiform stridulatory node on the scutellum, striae parallel, evenly spaced, metepisternum. The scraper is a cluster deeply impressed, strial punctures deep, of crescentic, ridgelike tubercles, probably evenly spaced, interstices convex, finely modified setal sockets, on the mesal face punctate; metepisternal sulcus sharply of the metafemur. Rotation of the femur curvate, extending posteriad and bordering positions the femoral tubercles to scrape a transversely striate, fusiform node (fig- across the metasternal node. ure 91), the file of an apparent stridulatory Amblycerus eustrophoides belongs to a organ; metacoxa sparsely microfoveolate group of Neotropical species, and is the in lateral two-thirds, cluster of punctures only representative of the group in the near trochanteral fossa; metafemur with United States. angulate tooth on ventral margin (figure Drypetes lateriflora is a West Indian eu- 87) (scraper for stridulatory organ), metati- phorb shrub found in southern Florida.

32 Amblycerus ireriae Romero, Johnson, spur nine-sixteenths as long as lateral and Kingsolver spur. Abdomen unmodified except posterior margin of 5th ventral segment of male (Figures 95–100) broadly emarginate; pygidium gently con- Amblycerus ireriae Romero, Johnson, and vex, slightly more so in male, apical margin Kingsolver 1996:58. truncate, that of male with a deep submar- Color.—Body and appendages uniformly ginal puncture. dark red, eyes usually black, sometimes Male genitalia.—As in figures 98 and 99. silvery in older specimens. Vestiture of Median lobe 3.5 times as long as wide, recumbent silvery gray setae intermixed broadest at base of ventral valve; base of with slender golden setae that on elytra are ventral valve broad tapering to acute apex twice as long as gray setae, elytral vestiture with lateral margins sharply incurved; base arranged in distinct stripes; pronotum and of dorsal valve about one-half as wide as pygidium each with indistinct gray median ventral valve, apex broadly rounded; arma- stripe. ture of internal sac consisting of two small Structure.—Body subfusiform in dorsal sclerites of varying size near base of dorsal aspect, strongly arched in lateral aspect valve, a median flask-shaped sclerite with (figure 96). Vertex and frons evenly convex, basal end acute and bent laterad, apical densely, finely punctate, lacking median end blunt, paired elongate sclerites bear- carina; ocular index 3:1, ocular sinus ing dentate ridges (figure 98), sac between about one-fifth length of eye, ocular fac- elongate sclerites with several small thorn- ets coarse, spherical, postocular lobe very like denticles; apical end of internal sac narrow, sparsely fringed; antenna (figure with U-shaped transfer valve with long 96) reaching metacoxa with segments 4–10 filamentous processes, membrane sur- moderately serrate, eleventh elliptical. Pro- rounding transfer valve with many min- notum trapezoidal (figure 95), sides nearly ute spicules; lateral lobes (figure 99) with straight in basal two-thirds then strongly broad emargination between strongly arcuate to anterior angle, apical margin setose lobes. broadly emarginate; disk densely foveo- Size.—Body length 4.3–5.3 mm; width late, intervals micropunctate, lateral carina 2.8–3.3 mm. extending from posterior corner to cervical Type depository.—USNM. sulcus, cervical sulcus short; prosternum narrow, constricted between procoxae. Type locality.—Texas, Pecos Co. Scutellum (figure 100) 1.25 times as long Distribution.—TX. as wide, apex evenly rounded. Elytra together slightly longer than wide, broadest Host plants.—Unknown. behind middle, strongly, evenly convex; Natural enemies.—None recorded. striae shallowly impressed, evenly spaced; Immatures.—Not described. interstices flat, densely pubescent concealing dense punctation; metepisternal sul- Discussion.—The arched body form and cus obtusely curvate, extending posteriad details of the male genitalia ally this spe- two-thirds length of pleurosternal suture; cies with A. vitis (compare figures 123– metacoxal face densely pubescent, moder- 124). Only eight specimens of this species ately densely punctate in lateral four-fifths, have been seen. Nothing is known of its glabrous near trochanteral insertion, with biology or food plant associations. dense cluster of punctures; hind leg as in figure 96, lateral spur four-fifths as long as basitarsus (figures 96 and 97), ventral

33 Amblycerus nigromarginatus (Motschulsky) Color.—Body and appendages usually dark red, occasionally piceous; eyes black; py- (Figures 92, 101–103) gidium with large, median, piceous macu- Spermophagus nigromarginatus Motschul- lation divided by stripe of pale setae. Vesti- sky 1874:249; Pic 1913a:60; ture (figure 104) of yellowish-brown, acicu- Bondar 1936:31. late setae evenly distributed over body Amblycerus nigromarginatus: Bridwell except three indistinct stripes on pronotal 1944:135; Blackwelder 1946:763; disk; elytra faintly striped in some speci- Udayagiri and Wadhi 1989:12. mens and with three or four evanescent, sinuate brown bands, or rows of brown In color and size, this species closely re- maculae in apical one-half; pygidium with sembles A. obscurus (Sharp) but basal narrow line of yellowish setae. sclerites in the male genitalia are thornlike and are much larger than those of A. Structure.—Body subelliptical, widest at obscurus (compare figures 101 and 106). middle of elytra, dorsal outline arcuate The only U.S. records of A. nigromarginatus in lateral aspect. Vertex and frons con- are from Florida, but the type specimen vex, mesal margins strongly convergent, is from Surinam. Records are also known sparsely punctulate; frontal carina lacking; from Brazil. See discussion of Amblycerus eyes laterally convex, facets coarse, indi- obscurus. vidually hemispherical; ocular index 4.3:1; ocular sinus one-fifth length of eye; postoc- Type depository.—ZMUM. ular lobe lacking; antenna with 4th to 10th Type locality.—Surinam. segments slightly eccentric, apex reaching Distribution.—FL; Surinam, Brazil. middle of metepisternum. Pronotum semicircular, middle one-third strongly convex, Host plants.—Senna corymbosa (Lamk.) Ir- lateral areas flat; disk densely punctulate win and Barn. (Florida). Other extralimital on median convexity, lateral punctures in- hosts listed in Romero et al. (1996:68) are termixed with microfoveolae; lateral carina Senna alata, S. bicapsularis bicap-sularis, evident in posterior one-half of margin, S. occidentalis, S. splendida, S. uniflora; evanescent anteriorly; cervical sulcus fine, Caesalpinia spp.; Cassia spp. nearly hidden by vestiture; hypomeron Natural enemies.—None recorded. deeply concave; prosternum Y-shaped, narrowed to a vertical lamella between Immatures.—Not described. procoxae; mesosternum tongue-shaped. Discussion.—The genitalia of males from Scutellum (figure 105) slightly longer than the Florida collections match precisely wide, moderately attenuated, truncate and those of the type specimen of Amblycerus channeled apically. Elytra together 1.4 nigromarginatus. Its presence in Florida is times as long as wide, moderately convex, likely an introduction and establishment suture elevated, 3rd interstice slightly el- since several collections have been made in evated, striae parallel, deep, nearly hidden the State. by vestiture; interstices convex, densely setose, minutely imbricate-punctate; me- Amblycerus obscurus (Sharp) tepisternal sulcus right-angled; postme- (Figures 45, 104–108) socoxal sulci continuous across midline, extending along pleurosternal suture. Spermophagus obscurus Sharp 1885:495; Metacoxal face with lateral three-fifths and Pic 1913a:60. posterior margin densely setose, fossula Amblycerus obscurus: Blackwelder with thin line of setae, mesal area around 1946:763; Johnson and Kingsolv- trochanteral fossa densely punctate; lateral er 1982:410; Udayagiri and Wadhi tibial spur six-tenths as long as basitarsus, 1989:12; Romero et al. 1996:69. ventral spur one-half length of lateral spur.

34 Fifth sternum of male slightly emarginate, der denticles, sometimes only on one side. apex of 8th tergite visible in emargination; The middle sclerites vary as illustrated in pygidium finely punctate, punctures con- shape and sinuation. cealed by vestiture. This species is closely related to A. nigro- Male genitalia.—As in figures 106–108; marginatus (Motschulsky) described from median lobe moderately broad (figure 106); Surinam, recorded from Brazil, and ap- base of ventral valve broad, lateral mar- parently introduced into Florida. Further gins sinuate to long, acute apex; armature studies with adequate material from Mex- of internal sac consisting of two small, ico, Central America, and northern South thornlike, basal spines, two spiny pads, a America may confirm suspicions that the cluster of fine spines in middle, two flat, names A. obscurus and A. nigromarginatus sinuate blades, and small, slightly hooked apply to one variable species. sclerite between their bases; closure valve These two species as a group are related U-shaped with long posterior arms; lateral to another group of two South American lobes short (figures 107 and 108), broad, species—A. hoffmanseggi (Gyllenhal) and A. shallowly cleft. The two small basal spines submaculatus (Pic) (Ribeiro 1989). of the sac may be minute and difficult to see. Amblycerus robiniae (Fabricius) Size.—Body length 4.7–6.0 mm; width (Figures 3, 4, 29–34, 88, 89, 109–112) 2.7–3.5 mm. Bruchus robiniae Fabricius 1781:75; Fa- Type depository.—BMNH. bricius 1787:41, 1792: 70; Linnaeus Type locality.—Mexico, Belize, Guatemala, 1788:1734; Gmelin 1790:1781; Olivier Panama (lectotype not selected). 1790:198, 1795:7; Fabricius 1801:397; Distribution.—TX (new U.S. record); Mexico Dejean 1821:78. south to Costa Rica and Panama. Spermophagus robiniae: Gyllenhal 1833:104; Dejean 1833:233; Schoen- Host plants.—No host records are known herr 1833:104; Melsheimer 1853:99; from the United States. Other records, Gemminger and Harold 1873:3218; mostly from Mexico: Senna alata, S. bicap- Horn 1873:312; Riley and Howard sularis bicapsularis, S. hirsuta leptocarpa, 1892c:166; Wickham 1895b; Britton S. obtusifolia, S. pendula advena, S. pendu- 1897; Blatchley 1910:1235; Cush- la ovalifolia, S. uniflora. These names may man 1911:504; Pierce 1912:74; Pic be found in the older literature under the 1913a:61; Leng 1920:306; Zacher generic name Cassia. 1952:468,472. Natural enemies.—None recorded. Amblycerus robiniae: Thunberg Immatures.—Not described. 1815:106,109,121; Bridwell 1930:29, 1932:106; Bissell 1938:536; Bradley Discussion.—The geographic range of Am- 1946:97; Bridwell 1946:56; Craighead blycerus obscurus extends from Texas to 1950:279; Dillon and Dillon 1961:734; Costa Rica and probably to Panama if the Bottimer 1968c:1012; Kirk 1969:99; record from the original description proves Kingsolver 1970a:371,377, 1970b:476; to be A. obscurus. Mathwig 1971:1, 1972:200; Baker Sclerites of the internal sac exhibit some 1972:137; Gibson 1972:99; Kingsolver variation in size and form but external 1975b:35; De Luca 1977:8; Kingsolver characters appear to be uniform. Some 1979a:341; Pfaffenberger 1979:231; specimens have two basal, thornlike scler- Johnson and Kingsolver 1982:410; ites, whereas these may be entirely lacking White 1983:306; Pfaffenberger 1985a:2; in others or may be present as small slen- Borowiec 1987:60; Udayagiri and Wadhi 1989:14; Romero et al. 1996:81.

35 Spermophagus hoffmannseggi [sic]: Boeving together one and one-third times as long 1927:139; Brimley 1938; Kirk 1969:99; as wide, gently convex except slightly de- Wallace and Fox 1980 (wing). pressed in scutellar area; striae composed Chrysomela gleditsiae Castiglioni of lines of shallow punctures in deep sulci; 1790:253; Kingsolver 1979a:341. interstices slightly convex, sutural interstice slightly elevated apically, each black Spermophagus hoffmanseggi or hoff- macula with transverse setal base; wing mannseggi, of authors venation as in figure 110; metepi-sternum Color.—Body and appendages dark red; with angulate sulcus extending for short pronotum, elytra, pygidium, metasternum, distance along sternopleural suture; post- and abdomen with scattered, setiferous mesocoxal sulcus not contiguous across black spots (figure 109); metasternum midline, extending along sternopleural with some diffusion of black; eyes black. suture to margin of metacoxa; metacoxa Vestiture of golden aciculate hairs and sparsely foveolate, punctulate, and setose white slender hairs; pronotum with pre- in distal four-fifths; fossula and proximal dominantly white hairs, some with golden one-fifth glabrous, cluster of punctures sheen, pair of white hair patches one-third near trochanteral insertion; hind leg as the distance from apex; elytra with fine, in figure 29, femur carinate on mesal and golden hairs in setal punctures, rows of lateral ventral margins, channeled between white hairs in striae, interstices mostly carinae; metatibia slightly dilated, lack- golden but with scattered clusters of white; ing carinae; lateral spur seven-tenths as pygidium clothed as on pronotum; ventral long as basitarsus, ventral spur one-half areas of body evenly white except abdomen as long as lateral spur. Abdominal sterna tessellated. not modified except posterior margin of Structure.—Body elliptical, moderately 5th slightly emarginate; pygidium slightly arcuate in lateral aspect (figure 4). Vertex convex in both sexes, posterior margin of and frons finely, densely, evenly punctate pygidium evenly rounded in male, trilobed with a few slightly larger punctures, dense- in female (figure 89). ly setose; frontal carina prominent in some Male genitalia.—As in figures 3, 30–34, specimens, in others evident only as an 111, and 112; median lobe moderately impunctate line; eye facets coarse; ocular broad, constricted in middle; base of ven- index 4:1; ocular sinus one-fifth as long as tral valve broad with sides rounded and eye (figure 109); postocular lobe crescentic, margins sinuate to acute apex; dorsal valve densely setose; antenna (figure 4) elongate, short, broad, arcuate; armature of internal serrate, apex reaching anterior margin of sac consisting of a U-shaped sclerite near metacoxa. Pronotum nearly semicircular base, a pair of triangular plates in middle, (figure 109), basal lobe emarginate, disk and a pair of small, spatulate blades at moderately convex, slightly sulcate in basal apex, closure valve complex; lateral lobes one-half, surface densely foveolate, each with apices truncate, concave on mesal foveola with single seta; interspaces finely faces, median cleft broadly V-shaped. imbricate; lateral carina prominent extending from posterior corner to near ante- Structure.—Body length 3.8–7.3 mm; width rior margin then bending sharply dorsad; 2.2–4.1 mm. cervical sulcus nearly hidden by vestiture; Type depository.—UZMC. prosternum Y-shaped, narrowly constrict- Type locality.—”Americae seminibus.” ed between coxae then expanded slightly behind coxae; mesosternum tonguelike, Distribution.—AB, AL, AR, CA, DC, FL, GA, slightly channeled, adjoining bulbous IA, IL, IN, KS, KY, LA, MA, MD, MI, MO, anterior lobe of metasternum; scutellum MS, NJ, NC, ND, NH, NM, NY, OH, OK, OR, elongate-triangular (figure 109), slightly PA, SC, SD, TN, TX, VA, WI, WV. carinate, and cuspidate apically. Elytra

36 Host plants.—Gleditsia triacanthos (honey lacking median carina, eyes subglobular, locust), G. aquatica (water locust); Robinia coarsely facetted; ocular index 8:1; ocular pseudoacacia (probably misidentification of sinus about one-fifth length of eye, post- host). Acacia farnesiana is a questionable ocular lobe absent; antenna (figure 120) record. Tillandsia usneoides is a hibernac- subserrate in segments 4–10, 11th elon- ulum record. gate-subelliptical, antennal apex reaching Natural enemies.—Bracon sp.; Heterospilus metacoxa. Pronotum quadrilateral (figure bruchi, H. prosopidis, H. spermophagi; Uro- 113), margins nearly straight in basal sigalphus bruchi, U. neobruchi; Eurytoma three-fourths, sharply arcuate at anterior sp., E. tylodermatis; Eupelmus cyaniceps; corner, basal margin bisinuate; disk evenly Horismenus missouriensis. convex, densely foveolate, lateral carina arcuate in lateral view, ending dorsad of Immatures.—Patton 1895 (larva); Wickham anterior coxa, not joined with submarginal 1895a (larva); Boeving 1927:143 (mouth- sulcus, the latter extending dorsad nearly parts); Pfaffenberger 1979:231 (first and to midline; prosternum narrow, apex final larval instars). slightly expanded. Scutellum (figure 118) Discussion.—This species is one of the lingulate, slightly constricted medially, most common bruchids in the eastern half tapered to blunt apex. Elytra together 1.5 of the United States, generally following times as long as wide, broadest one-third the distribution of Gleditsia triacanthos, from base, transversely convex, slightly its principal host plant. Records from depressed around scutellum, striae paral- New Mexico, California, and the Pacific lel, lightly impressed, strial punctures dis- Northwest are undoubtedly introductions crete, interstices finely punctate or imbri- through infested seed stocks. Mathwig cate; metepisternal sulcus obtusely angu- (1971, 1972) investigated the biology of A. late, extending one-half distance to meta- robiniae in Kansas. coxa; metacoxal face moderately densely pubescent and sparsely punctate in lateral The black spots on the pronotum, elytra, four-fifths, glabrous medially with dense and other parts of the body are character- cluster of punctures; lateral tibial spur istic of a group of three species, the other one-half length of basitarsus (figure 119), two being found in Mexico and Cuba. ventral spur one-half as long as lateral spur. Abdomen unmodified, sexes similar Amblycerus schwarzi Kingsolver in outline of terminal margin of pygidium (Figures 113–120) and 5th sternum. Amblycerus schwarzi Kingsolver Male genitalia.—As in figures 114-117. Me- 1970b:477; Udayagiri and Wa- dian lobe slender; ventral valve elongate- dhi 1989:14; Kingsolver 1992b:78, acute; dorsal valve rounded apically; arma- 1996:24; Romero et al. 1996:86; Al- ture of internal sac consisting of two burr- varez Marin and Kingsolver 1997:216; like sclerites, a large, elongate, comma- Genaro and Kingsolver, 1997:229. shaped median sclerite split basally, two Color.—Body and appendages uniformly rows of small denticles, two small hooks, dark red; eyes black, sometimes silvery; and a pair of fragile, arcuate clusters of vestiture of fine yellowish gray setae evenly minute spicules; lateral lobes (figures 116 distributed over body except slightly mot- and 117) fused along midline, apices mas- tled on elytra, sometimes with rounded sive, hollow-faced. brown subapical elytral spots, abdominal Size.—Body length 4.7–5.7 mm; width pleura with similar brown spots; pygidium 2.7–3.5 mm. with slightly denser median line of setae. Type depository.—USNM Type #69943. Structure.—Body subfusiform in outline. Vertex and frons convex, frons triangular,

37 Type Locality.—Cuba, Las Villas Prov., na (figure 122) moderately serrate, apex Cuyamas. reaching middle of metacoxa. Pronotum Distribution.—FL; Bahamas, Cuba, Domini- trapezoidal (figure 121), lateral margins can Republic, Grand Cayman, Curaçao, arcuate anteriorly, disk convex, densely, Jamaica, Puerto Rico, Virgin Is. uniformly microfoveolate; lateral carina arcuate, extending from posterior corner to Host plants.—USA: Hippomane mancinella; near anterior border then bending sharply West Indies: Guettarda sp.; Hippomane dorsad delimited by cervical sulcus and mancinella; Tectona grandis; Terminalia extending nearly to dorsal midline; pro- catappa; Ricinis communis. sternum Y-shaped, constricted between Natural enemies.—None recorded. procoxae, slightly expanded behind coxae. Metasternum narrowly rounded. Postme- Immatures.—Not described. socoxal sulci extending across midline, Discussion.—Amblycerus schwarzi is laterally paralleling sternopleural suture widely distributed in the Caribbean area nearly to metacoxal margin; metepisternal and affects a variety of host plant families. sulcus obtusely angulate, extending half- Hippomane and Ricinis are in the Euphor- way along pleurosternal suture. Scutellum biaceae, Guettarda is in the Rubiaceae, tridentate. Elytra together one and one- Tectona is in the Verbeneaceae, and Termi- fourth times as long as wide, evenly con- nalia is in the Combretaceae. Another spe- vex; striae parallel, evenly spaced, deep, cies of Amblycerus in Mexico and Central narrow; interstices flat, finely imbricate. America, A. spondiae Kingsolver, is also as- Metacoxal face sparsely foveolate, inter- sociated with hosts in four different plant spaces minutely imbricate, dense patch families (Kingsolver 1980a:241). of fine punctures near trochanteral fossa; Amblycerus schwarzi is related to A. dispar hind leg as in figure 122, femur channeled (Sharp) whose distribution extends from ventrally between carinae; metatibial apex Mexico to Chile. diagonal, finely denticulate, lateral spur three-eighths as long as basitarsus (figure Amblycerus vitis (Schaeffer) 122), ventral spur two-thirds as long as lateral spur. Abdomen normal, male 5th (Figures 121–124) sternum broadly emarginate, female 5th Spermophagus vitis Schaeffer 1907:293; sternum evenly rounded; pygidium in both Cushman 1911:505; Pic 1913a:63; sexes finely punctate, sculpture concealed Leng 1920:306; Essig 1958:487. by vestiture. Amblycerus vitis: Bottimer 1968c:1012, Male genitalia.—As in figures 123 and 124; 1012,1038; Johnson 1968:1268; John- median lobe constricted in middle; base of son and Kingsolver 1975:321; Cen- ventral valve broad, lateral margins sinu- ter and Johnson 1976:201; Johnson ate to elongated, acute apex; dorsal valve and Kingsolver 1982:411; Udayagiri subelliptical; armature of internal sac and Wadhi 1989:17; Romero et al. consisting of large, thornlike median scler- 1996:102. ite set in transverse band of fine spicules, Color.—Body and appendages uniformly and on each side a curved, denticulate reddish brown except eyes black. Vestiture sclerite with concave face; apex of sac with yellow, uniformly distributed over body. U-shaped closure valve with long slender Structure.—Subovate in outline (figure posterior arms; lateral lobes (figure 124) 121). Vertex and frons densely punctulate broadly Y-shaped, densely setose. with scattered larger punctures; frontal Size.—Body length 2.6–3.7 mm: width carina usually lacking, sometimes with 1.6–2.5 mm. short, impunctate ridge; eyes coarsely Type depository.—USNM, lectotype No. faceted; ocular index 2.5:1; ocular sinus 42337 (by Johnson). shallow; postocular lobe lacking; anten-

38 Type locality.—Arizona, Huachuca Moun- spurs dark red. Vestitural pattern relative- tains. ly constant for each species, occasionally Distribution.—AZ. dimorphic. Host plants.—Vitis arizonica Englem. Vertex and frons usually finely punctate, sometimes with mixed larger punctures, Natural enemies.—Urosigalphus sp. frontal carina usually prominent; ocular Immatures.—Not described. index 3:1 to 5:1; ocular sinus three-fourths to seven-eighths length of eye (figures 17 Discussion.—This species is known only and 127); postocular lobe narrow, sepa- from Arizona and Texas and is the smallest rated from gena by short sulcus; antenna species of Amblycerus in the United States. elongated, narrow, laterally compressed, It lacks the piceous elytral spots of A. rob- segments 4 to 10 subtriangular, terminal iniae, the metepisternal stridulatory node segment elongate-parallel (figures 125– of A. eustrophoides, and differs by its tri- 126). Anterior margin of pronotum semi- denticulate scutellum and uniform pygidial circular, apex sometimes truncate, base vestiture from the triangular scutellum broadly angulate, basal margin gently ar- and faintly striped pronotum and elytra of cuate from midline to posterior angles, disk A. obscurus and A. nigro-marginatus. It is evenly convex except basal lobe sometimes probably most closely related to A. ireriae marked by short, diagonal sulci delimit- as evidenced by details of the male genita- ing slight median dome; lateral surfaces of lia and by body form. disk sparsely foveolate (except Z. amplis- Johnson and Kingsolver (1975) detailed the simus), intervals between foveolae finely biology of this species. punctate, median one-third usually finely Essig (1929a) gave the common name “wild punctate; lateral carina sharp, anterior grape seed weevil” to A. vitis. end hooked toward foramen; hypo-meron deeply concave to receive fore legs in repose; prosternum short, triangular, but Tribe Spermophagini Borowiec seldom visible because of reflexed position of head; mesosternum rounded or cari- Spermophagini Borowiec 1987:62 nate, sometimes protruding above level of Genus Zabrotes Horn metasternum; mesepisternum and mesepimeron subequal in width; post-mesocox- Zabrotes Horn 1885b:156; Leng 1920:306; al sulcus extending across metasternum. Bridwell 1932:106, 1946:53,56; Scutellum narrowly triangular, densely Blackwelder 1946:763; Bottimer pubescent. Elytra together slightly wider 1968c:1021; Kingsolver 1970b:469; than long, nearly flat in middle two-thirds, Johnson and Kingsolver 1982:411; convex laterally; striae regular in course, Borowiec 1987:62; Udayagiri and Wa- usually narrow; strial punctation varying dhi 1989:31. Type species: Zabrotes from broad and shallow to deep and nar- cruciger Horn 1885b, by subsequent row, each puncture or foveola with seta designation, Zacher 1930. emerging from beneath its anterior rim; in- Most species are small—1.0–2.5 mm—but terstices sparsely punctate (except densely one species reaches 3.7 mm in length. Spe- foveolate in Z. amplissimus). Metacoxal face cies of Zabrotes vary little in many of their reniform, broader than metafemur, sparse- morphological structures. The following ly foveolate and densely pubescent in later- characteristics appear to be fairly constant al one-half (except Z. amplissimus), median in the species found in the United States. one-half to one-third either with cluster of about 20 fine punctures (figure 129), Integument of body and appendages black or polished and impunctate with deep pit except basal one or two segments of an- adjacent to insertion of trochanter (figure tenna usually red or reddish brown; tibial 130); hind leg as in figure 128; metafemur

39 with ventrolateral carina gently sinuate to Abdomen with sterna telescoped, 5th ster- subacute apex, ventromesal carina more num of male deeply emarginate for apex of strongly sinuate to rounded apex; metati- pygidium; pygidium broader than long in bia straight, ventrolateral, ventral, and both sexes, convex with apex more strongly ventromesal carina each with row of short, reflexed in male, disk microfoveolate. stiff setae, dorsolateral face with row of Lateral lobes of male genitalia fused basal- setiferous punctures; lateral spur shorter ly, straplike, expanded apically. than mesal spur; terminal margin of tibia oblique, finely denticulate; basitarsus Characters that constitute the basis for with two ventral, parallel, setose carinae. specific differentiation are found in the color pattern of the pronotum, elytra,

Key to United States species of Body length 2.1–2.9 mm; vestiture dark brown with Zabrotes strongly contrasting white pattern (figure 151); male genitalia as in figures 153 and 154...... cruciger Horn 1 Medial one-half of metacoxal face with cluster of coarse punctures near trochanteral insertion (figure 129). Sec- 6(5) Male genitalia as in figures 157 and 158; lateral margin of tion I: Punctate metacoxae...... 2 pronotum with diffuse white patch . cynthiae Kingsolver Medial one-half of metacoxal face smooth and polished; Male genitalia as in figures 184 and 185; lateral margins some species with deep pit surrounding trochanteral of pronotum with sharply defined white patch...... condyle (figure 130). Section II: Smooth metacoxae. ....10 spectabilis Horn 2(1) Dorsal color nearly uniformly gray except apex of elytra 7(4) Elytra mostly yellowish orange with white maculae; male brown...... 3 pronotum white with orange stripes, elytra with white sutural stripe, pygidium white; female pronotum orange Dorsum of different color combination ...... 4 with lateral and basal spots white, pygidium orange with 3(2) 3rd and 5th elytral interstices striped in basal one-fourth white median stripe (figure 144). ...chandleri Kingsolver (figure 186); male genitalia as in figures 188 and 189 Coloration otherwise...... 8 ...... stephani Kingsolver 8(7) Pronotum and elytra reddish brown or mixed gray and Interstices not striped, uniformly gray (figure 177); male brown; lateral surfaces of abdominal sternites brown genitalia as in figures 179 and 180 Horn planifrons mottled; pygidium with indistinct median and lateral gray 4(2) Pronotum, elytra and pygidium with strongly contrasting stripes sometimes connected by transverse band (figure white maculae on dark brown or golden brown back- 148) ...... chavesi Kingsolver ground; males with sutural stripe on pronotum and elytra Pronotum and elytra brown or gray; abdominal sternites (figures 151 and 181)...... 5 uniformly gray; pygidium with or without median stripe . 9 Coloration otherwise...... 7 9(8) Pygidium usually with gray median stripe (figure 209); 5(4) Body length 1.3 to 1.6 mm; golden brown grading to dark male genitalia as in figures 210 and 211; Texas and brown vestiture with white pattern; male genitalia as in Mexico figures 153 and 184...... 6 ...... victoriensis Kingsolver

40 pygidium, and ventral areas of the meta- This genus is found naturally only in the thorax and abdomen; punctation of the Western Hemisphere from the United pronotum, elytral striae and interstices, States to Brazil. Host plant associations metacoxal face, and pygidium; shape of the are known for only a few species but those antenna and mesosternal apex; and form known are with leguminous seeds. of median and lateral lobes of male genita- Revision: Kingsolver 1990b (U.S. species). lia.

Pygidium usually unicolorous (figure 134), occasionally curved elytral maculae (figure 191), pygidial pattern vari- with narrow evanescent median line; male genitalia as in able (figures 193 and 194)...... 14 figures 135 and 136; eastern United States. ...arenarius Metasternum convex or flat, with only a median suture. 15 (Wolcott) 14(13) Median lobe of male genitalia with armature of internal 10(1) Vestiture of pronotum and elytra either uniformly gray or sac as in figure 198; internal armature of female bursa white sometimes intermixed with reddish brown or with (arrow) as in figure 207...... subfasciatus (Boheman) vaguely defined median band extending from border to border on elytra ...... 11 Median lobe of male genitalia as in figure 204; internal armature of female bursa (arrow) as in figure 206 Elytra with distinct maculae of different color from back- ...... sylvestris Romero and Johnson ground (figure 191) or with mottled pattern (figure 190) ...... 13 15(13) Male genitalia with flask-shaped median sclerite and small “stopper” sclerite (figures 161 and 174)...... 15 11(10) Body length 3.6 mm; pronotum, elytra, and pygidium finely, densely foveolate (figures 131 and 132); pygidium Genitalic armature differently shaped (figures 165 and with small, white, subbasal spot either side of midline 170)...... 16 ...... amplissimus Kingsolver 16(15) Body length 2.0–2.5 mm; male genitalia as in figures 174 Body length less than 3 mm; if foveolae present, not and 175; female 8th sternite as in figure 176; Texas and evenly distributed; pygidium lacking white spots...... 12 Arizona...... obliteratus Horn 12(11) Elytra with vaguely defined, gray median band, remainder Body length 1.6 mm; male genitalia as in figures 161 and of elytra uniformly gray or gray intermixed with reddish 162; southern California...... densus Horn brown setae (figure 137); Texas 17(15) Body length 1.7 mm; male genitalia as in figures 170 and ...... bexarensis Kingsolver 171; northern California Elytra lacking band, vestiture uniform (figure 200); east- ...... humboldtae Kingsolver ern United States...... subnitens Horn Body length 2.2 mm; male genitalia as in figures 165 and 13(10) Metasternum with elongate, pubescent sulcus on midline 166; female 8th sternite as in figure 167; Arizona (figure 195); male with mottled elytral pattern (figure ...... eldenensis Kingsolver 190), pygidium uniformly clothed, or with indistinct median stripe (figure 192); female with transverse, white,

41 Zabrotes amplissimus Kingsolver genitalia must be delayed until additional collections that include males have been (Figures 131, 132) made. The genitalia of the undes-cribed Zabrotes amplissimus Kingsolver Mexican sister species places it near Z. 1990b:152. subnitens. Color.—Vestiture white, uniformly distributed over entire body except pair of con- Zabrotes arenarius (Wolcott) densed spots near base of pygidium. (Figures 133–136) Structure.—Punctation of vertex and frons Bruchus arenarius Wolcott 1912:162; Pic nearly concealed by vestiture, frontal ca- 1913a:162. rina fine but distinct; transverse sulcus at Mylabris arenarius: Leng 1920:305. upper level of eyes; ocular index 3:1, eyes angular on lateral margin, ocular sinus Megacerus arenarius: Bradley 1947:37. five-sixths length of eye; antenna slender Zabrotes arenarius: Bottimer 1956:67, (female), reaching middle of metacoxa. 1968c:1013,1038; Johnson 1968:1271; Pronotum (figure 131) nearly semicircu- Johnson and Kingsolver 1982:411; lar but lateral margins subparallel near Kingsolver 1990b:141. basal angles; disk convex, slightly domed Color.—Vestiture evenly distributed of uni- in middle, surface evenly microfoveolate, formly gray to mixed gray and dark brown, foveolae mostly discrete; lateral carina occasionally with white patch on basal lobe arcuate. Mesosternum flat. Elytra together of pronotum, and a marginal maculation slightly wider than long; arcuately convex; on each elytron (figure 133). densely microfoveolate as on pronotum, striae traceable only in center of disk, eva- Structure.—Antenna not dimorphic, reach- nescent on lateral areas. Pygidium micro- ing middle of metacoxa; head densely foveolate with slightly larger foveolae than punctulate, frontal carina prominent. on elytra. Metacoxa with lateral one-half Pronotum subtrapezoidal, apex truncate, densely foveolate, medial one-half smooth lateral margins arcuate, base broadly an- and polished. Metepisternum and lateral gulate; disk sparsely microfoveolate, foveo- areas of abdominal sterna microfoveolate lae shallow, inconspicuous; lateral carina (figure 132). prominent. Scutellum narrowly triangular. Elytra together as long as wide, slightly Male genitalia. —Not described. (Only one depressed medially in basal one-half, striae female specimen is available.) shallow, inconspicuous, strial punctures Size.—Body length 3.7 mm; width 2.7 mm. approximate; interstices flat, finely, densely punctate-imbricate. Metacoxal face sparse- Type depository.—USNM. ly, shallowly foveolate in lateral one-half, Type locality.—New Mexico, near Grants. medial one-half with cluster of punctures. Distribution.—NM. Abdomen not modified (figure 134) except the usual reflexed male pygidium. Host plants.—Phaseolus metcalfi (now P. ritensis). Male genitalia.—As in figures 135 and 136; median lobe slightly constricted in middle; Natural enemies.—None recorded. ventral valve subtriangular; dorsal valve Immatures.—Not described. with slender, ligulate process and strongly recurved apicolateral margins; internal sac Discussion.—This species and a yet unde- with pair of ovate sclerites at apical orifice, scribed species from Mexico are the largest folds of internal sac densely setose; middle known species of Zabrotes. The distinc- of sac with pair of reniform sclerites; apico- tive foveolate elytra and pygidium and lateral sac lobes spinose; lateral lobes as in large size easily separate them from any figure 136. other described species. Description of the

42 Size.—Body length 1.5–2.1 mm; width segment; 1st and sometimes 2nd segment 1.2–1.4 mm. dark red; pronotum truncate apically, Type depository.—FMNH, holotype No. 234. disk minutely punctate with scattered micro-foveolae; elytra with striae narrow, Type locality.—Illinois, Havana, Devil’s strial punctures deep, elongate, interstices Hole. minutely punctate; apex of mesosternum Distribution.—AL, DC, FL, GA, IL, IN, MD, flat, truncate; metacoxal face with 15–20 NJ, NC, OK, SC, TX, VA. shallow foveae in lateral one-half, medial one-half impunctate with deep pit near in- Host plants.—Swept from Cassia fasci-cula- sertion of trochanter; metasternum slightly ta Michx. flowers at Havana, but the larval depressed in posterior one-half, vestiture host is not known. Bridwell collected it on extending obliquely from median sulcus, strawberry flowers in Maryland. pygidium minutely punctate and foveate, Natural enemies.—None recorded. foveae slightly larger than on pronotum. Immatures.—Not described. Male genitalia.—As in figures 140 and 141; Discussion.—The male genitalia are nearly apical one-third of median lobe slightly identical to those of Z. spectabilis; how- expanded; ventral valve evenly arcuate; ever, the color pattern is uniformly gray or dorsal valve ogival with apex produced; brown with or without faint elytral maculae internal sac with pair of flat, lightly sclero- but lacking the bold pattern of Z. spec- tized plates at apical orifice; middle of sac tabilis. Specimens from Havana are uni- with about 25 spicules and an ovate struc- formly brown or gray, whereas the elytra ture enclosing pair of sinuate spines; apex from other localities are usually maculate. of sac abbreviated; lateral lobes strongly Zabrotes stephani is also gray but with expanded, apical margin with row of flat- some faint striping on the elytra and with a tened setae (figure 141). distinctively different male genitalia (fig- Size.—Body length 1.6–1.9; width 1.1–1.5 ures 188 and 189). mm. Type depository.—CNCI. Zabrotes bexarensis Kingsolver Type locality.—Texas, Bexar Co. (Figures 137–141) Host plants.—Vicia sp. and Vicia leaven- Zabrotes sp. 1, Johnson 1979a:124. worthii Torr. and Gray. Zabrotes bexarensis Kingsolver 1990b:153. Natural enemies.—None recorded. Color.—Vestiture of head white, sparsely distributed; pronotum uniformly white to Immatures.—Not described. white intermixed with brown, sometimes Discussion.—Although this species has patchy; elytra (figure 137) ranges from the smooth metacoxal face, it seems to be uniformly white to white intermixed with isolated from the others in this group. The brown to brown with white maculation, broad, ogival ventral valve, the cluster of maculation usually vaguely defined; pygid- slender spines and paired aciculate spines ium (figure 138) uniformly white or white in the middle of the sac, and the alate with evanescent median stripe or gray with lateral lobes with flattened setae are unlike brown lateral patches; ventral surfaces any other Zabrotes species. uniformly white. The evanescent elytral pattern, long male Structure.—Frontal carina ridgelike; ocu- antennae, and the uniformly clothed pygid- lar index 6:1; ocular sinus three-fourths ium with a faint median line are additional length of eye; antennae dimorphic (figure salient characters for this species. 139), that of male reaching beyond apices of elytra, that of female to 1st abdominal

43 Zabrotes chandleri Kingsolver lobes lined with fine spicules; lateral lobes slightly expanded, apices rounded (figure (Figures 142–146) 146). Zabrotes chandleri Kingsolver 1990b:142. Size.—Body length 2.4–2.5 mm; width Color.—Basal antennal segment red or red- 1.7–1.8 mm. dish brown. Vestiture of orange and white setae in dimorphic pattern; males with Type depository.—UCDC. head, pronotum, pygidium and ventral ar- Type locality.—California, 3 mi north of Big eas white, elytra orange with median trans- Pine. verse bars and sutural stripe white (figure Distribution.—CA. 142), pronotum sometimes with vague orange spots on middle of disk; females with Host plants.—Not known. head white, pronotum orange with white Natural enemies.—None recorded. spot on basal lobe and lateral margins, Immatures.—Not described. elytra orange, each elytron with elliptical, transverse spot (figure 143), pygidium Discussion.—The orange and white vesti- orange with white median stripe expanded tural pattern is immediately recognizable. at apex, venter white except coxa, a lateral This pattern and the form of the male spot on 1st abdominal sternum and at genitalia place this species near Zabrotes apex of 5th segment orange. cruciger and Z. spectabilis. The species is not known outside California. Structure.—Vertex and frons minutely punctate, clypeus more coarsely punctate, Zabrotes chavesi Kingsolver frontal carina prominent; ocular index 5:1; ocular sinus four-fifths length of eye; (Figures 147–150) antenna slightly dimorphic, that of male Zabrotes chavesi Kingsolver 1980a:229; reaching 1st abdominal segment, that Janzen 1980:949; Johnson and King- of female reaching metacoxa. Pronotum solver 1982:411; Romero and Johnson semicircular, disk convex, slightly domed, 1997:74. minutely punctate with larger punctures in lateral areas and along base; lateral ca- Zabrotes vandykei Kingsolver 1990b:150; rina gently arcuate. Scutellum triangular, Romero and Johnson 1997:74. densely pubescent; mesosternum project- Color.—Basal antennal segment and metat- ing, carinate. Elytra slightly wider than ibial calcaria dark red. Vestiture of brown, long, disk nearly flat; striae nearly con- gray, and reddish-brown slender setae (fig- cealed by vestiture; interstices flat, puncture 147); head uniformly gray; pronotum ulate. Pygidium 1.4 times as wide as long with mixed brown and gray, lateral areas (figure 144), convex, apex in male strongly paler; elytra with mixed brown and gray reflexed into 5th sternum. Metacoxal face setae, usually with pale transverse band, with lateral four-fifths densely punctu- darker apically; pygidium (figure 148) with late and 30–35 shallow foveolae, median reddish-brown basal band, remainder one-fifth smooth, polished, with cluster of mixed gray and brown with reddish-brown coarse punctures set in slight depression. patches; ventral side of body mostly gray with reddish-brown patches along dorsal Male genitalia.—As in figures 145 and 146; margin of abdominal segments, coxa red- median lobe broad, apex broadly ovate; dish brown. ventral valve arcuate; dorsal valve ogival, apex moderately produced; internal sac Structure.—Vertex and frons punctulate, with prominent folds; middle of sac with frontal carina evanescent; ocular index pair of large, L-shaped spines attached to 4.5:1; ocular sinus seven-eighths length semicircular structure; apicolateral sac of eye; antenna not dimorphic, reaching posterior margin of metacoxa. Pronotum

44 semicircular, moderately convex, slightly Romero and Johnson (1997) recognized domed in middle one-third; disk punctu- that the description of Z. chavesi from late on dome, more coarsely punctate on Costa Rica was identical to the description lateral surface of disk; lateral carina not of Z. vandykei from Arizona, and compari- arched. Mesosternum flat. Elytra together son of the type specimens confirmed the slightly wider than long; striae deep, evi- synonymy. Locality records from Arizona, dent in basal one-half, nearly hidden by Mexico, Honduras, Nicaragua, El Salvador, vestiture apically, strial punctures con- Costa Rica, and Venezuela indicate an ex- tained within striae; interstices densely tensive and probably continuous distrbu- punctulate. Metacoxa shallowly foveolate tion. in lateral one-half, lateral margin densely setose, medial one-half with cluster of Zabrotes cruciger Horn about 30 punctures set in depression near (Figures 151–154) trochanteral insertion. Pygidium 1.3 times as wide as long; disk shallowly foveolate, Zabrotes cruciger Horn 1885b:157; Leng interspaces about the diameter of one fo- 1920:306; Bottimer 1968c:1013,1038; veola. Johnson 1968:1271; Johnson and Kingsolver 1982:411; Kingsolver Male genitalia.—As in figures 149 and 150; 1990b:143. median lobe suddenly expanded in apical one-fifth; ventral valve triangular; dorsal Spermophagus cruciger: Pic 1913a:59. valve with elongate beaklike process, api- Zabrotes cruciatus: Kingsolver 1990b:143 colateral margins evenly arcuate; internal (misspelling). sac with pair of small sclerites near apical Color.—Basal segment of antenna and orifice, pair of triangular sclerites in mid- tibial spurs dark brown. Vestitural pattern dle; apex of sac lined with fine denticles; dimorphic: In male, pronotum dark brown lateral lobes as in figure 150. dorsally with lateral patches and median Size.—Body length 1.6–2.2 mm; width stripe white, elytra with sutural stripe and 1.3–1.6 mm. lateral, oblique maculae white; in female median and sutural stripes lacking, oth- Type depository.—USNM (chavesi); CASC erwise as in male (figure 151); vestiture (vandykei). white ventrally except coxal face in both Type locality.—Costa Rica, Guanacaste sexes and lateral patch on 1st abdominal Prov., Santa Rosa National Park (chavesi); segment brown; pygidium white with large, U.S.A., Arizona, 6 mi north of Chiricahua dark brown patches either side of midline National Monument (vandykei). (figure 152). Distribution.—AZ; Mexico to Venezuela Structure.—Frontal carina prominent; (Romero and Johnson 1997). ocular index 5:1; ocular sinus four-fifths length of eye; antenna reaching posterior Host plants.—Senna hirsuta leptocarpa margin of metacoxa. Pronotum semicir- (Benth.) Irwin and Barn. in AZ to Costa cular apically, broadly angulate basally, Rica; Senna spectabilis DC. in Venezuela. evenly convex, basal lobe marked by short, Natural enemies.—None recorded. lateral sulci; surface sparsely foveolate, Immatures.—None described. foveolae elliptical, each with seta emerging from its anterior rim; intervals finely punc- Discussion.—This species closely resembles tate; lateral carina sharp; mesosternum Zabrotes stephani and Z. victoriensis, but carinate, protruding above level of meta- the lateral abdominal areas are mostly sternum. Scutellum triangular, densely brown in Z. chavesi, contrasted with mostly pubescent. Elytra together as long as wide, gray in the other two species. nearly flat in middle two-thirds, convex laterally; striae regular in course, consisting

45 of large, elliptical foveolae, each with seta mens nearly uniformly white with scat- emerging from beneath its anterior rim, tered golden setae, female usually darker interstices sparsely punctate. Metacoxal brown than male; venter of body mostly face sparsely foveolate in lateral two-thirds, white with golden brown on metacoxae and intervals finely punctate, median one-third brown patch on lateral areas of abdomen, with cluster of deep punctures near inser- especially on 5th sternum; legs white. First tion of trochanter. antennal segment red, remaining segments Male genitalia.—As in figures 153 and 154; black. median lobe strongly expanded apically; Structure.—Frontal carina prominent; ventral valve subtriangular; dorsal valve ocular index 4:1; ocular sinus three- with short process; internal sac with small, fourths length of eye; male antenna slightly paired sclerites leading to thickened me- broader and longer than that of female, dian folds; apex of sac with pair of tapered, the male’s reaching pygidial margin, sinuate sclerites in oval, saclike structure; the female’s reaching metacoxa. Prono- apicolateral sac lobes each with patch of tum punctulate with scattered foveolae; minute denticles; lateral lobes as in figure pronotal disk semicircular with apex 154. subtruncate; lateral carina normal for Size.—Body length 2.1–2.9 mm; width genus. Scutellum minute, broadly trian- 1.5–2.1 mm. gular. Elytra together slightly wider than long; striae shallowly foveolate, interstices Type depository.—MCZC, holotype No. densely punctulate. Lateral two-thirds of 3899. metacoxal face extremely finely punctu- Type locality.—”Colorado.” late and sparsely pubescent, median one- third smooth except for depressed cluster Distribution.—AR, CO, NM, OK, TX. of about 30 punctures near trochanteral Host plants.—Not known. insertion. Natural enemies.—None recorded. Male genitalia.—As in figures 157 and 158; Immatures.—Not described. median lobe broadly expanded at apex; ventral valve broadly triangular, lateral Discussion.—This large, handsome species margins slightly sinuate; dorsal valve with could hardly be confused with any other ligulate median process, apicolateral mar- Zabrotes except Z. spectabilis. The bold, gins evenly arcuate similar to those of Z. sexually dimorphic color pattern of the spectabilis except shoulders of dorsal valve dorsal surface is similar to that of Z. spect- rounded, not recurved (compare figures abilis and Z. chandleri, wherein the sutural 157 and 184); internal sac with prominent and pronotal stripes distinguish the male, folds near apical orifice; middle of sac with and their absence, the female. pair of small, triangular sclerites; apicolateral sac lobes each with patch of fine Zabrotes cynthiae Kingsolver denticles; lateral lobes as in figure 158. (Figures 155–158) Size.—Body length 1.7–1.9 mm; width Zabrotes cynthiae Kingsolver 1990b:144. 1.1–1.3 mm. Color.—Vestiture mostly golden brown dor- Type depository.—USNM. sally, head sparsely white, pronotum with Type locality.—California, San Bernadino white median stripe from base to, or nearly Co., Providence Mountains. to, apex, lateral margins sometimes with diffuse white patches; elytra with white Distribution.—CA. sutural stripe, each elytron with oblique, Host plants.—Swept from Asclepias sp., elongate white bar (figure 155); pygidium Cupressus macnabiana, Eremocarpus sp., variable, usually brown with white median stripe (figure 156) but some speci-

46 Lotus sp., and Quercus sp., but with no placed foveolae; lateral carina arcuate. rearing records. Scutellum small, equilaterally triangular. Natural enemies.—None recorded. Elytra together slightly wider than long; striae not impressed, strial punctures Immatures.—Not described. elongate, interstices punctulate. Pygidium Discussion.—Four species found in Cali- 1.25 times as wide as long, convex, densely fornia—Zabrotes cynthiae, Z. densus, Z. but uniformly foveolate. Metacoxal face humboldtae, and Z. spectabilis—are similar sparsely setose, sparsely foveolate in later- in size and color pattern and can easily be al one-half, smooth, impunctate in median confused with one another. Two of the spe- one-half. cies—Z. densus and Z. humboldtae—are in Male genitalia.—As in figures 161 and 162; the group with smooth metacoxae, whereas median lobe moderately slender, margins the remaining two are in the punctate coxa subparallel; ventral valve ovate; dorsal group. Differentiation of Z. cynthiae and Z. valve nearly evenly rounded; armature of spectabilis can be made by examination of internal sac consisting of flask-shaped male genitalia (figures 157 and 184); the sclerite with small, ovoid sclerite (stop- apical margin of the dorsal valve is evenly per) at small end, base of sclerite enclosing arcuate in Z. cynthiae but recurved in Z. cluster of fine spicules; lateral lobes as in spectabilis. The pronotal lateral margin of figure 162. Z. spectabilis carries a lunate white mark (figure 181), but the margin ofZ. cynthiae Size.—Body length 1.1–1.7 mm; width is without markings (figure 155). 0.9–1.3 mm. Type depository.— MCZC, holotype No. Zabrotes densus Horn 3003 (Figures 159–162) Type locality.—”California.” Zabrotes densus Horn 1885b:158; Leng Distribution.—AZ, CA, OR. 1920:306; Bottimer 1968c:1013,1038; Host plants.—None recorded. Johnson 1968:1271; Johnson and Kingsolver 1982:411; Kingsolver Natural enemies.—None recorded. 1990b:154. Immatures.—Not described. Spermophagus densus: Pic 1913a:59. Discussion.—This species closely resembles Color.—Vestiture dark brown usually with Z. cynthiae Kingsolver. The smooth coxal faint grayish or light brown striping on face and male genitalia (figure 161), how- elytra (figure 159), some specimens with a ever, ally it with Z. obliteratus although gray bar on each elytron; head brown; pro- the elytral and pygidial patterns are not notum gray on basal lobe, sometimes with as contrasting as in that species. Zabrotes faint mottling on disk; pygidium (figure humboldtae is similar in appearance to Z. 160) mostly brown, basal band and me- densus and Z. cynthiae, but the male geni- dian stripe gray; venter of body gray except talia are of a different type nearly identical coxal face and sometimes lateral abdomi- to those of Z. spectabilis. The three spe- nal spot brown. cies—Z. cynthiae, Z. densus, and Z. hum- Structure.—Vertex and frons punctulate, boldtae—are found almost entirely within frontal carina prominent; ocular index 3:1; California. ocular sinus three-fourths length of eye; antenna reaching 1st abdominal segment Zabrotes eldenensis Kingsolver in both sexes. Pronotum semicircular but (Figures 163–167) with apex truncated; disk convex, slightly Zabrotes eldenae: Kingsolver 1990b:139 domed medially, punctulate with sparsely (misspelling). Zabrotes eldenensis Kingsolver 1990b:155.

47 Color.—Vestitural color of both sexes simi- without resorting to examination of the lar to that of female Z. subfasciatus; males male genitalia and the 8th sternite of the with broad white spot on basal lobe of pro- female genitalia. Sufficient variation ex- notum (figure 163), females with median ists in associated females that—unless white stripe; elytral maculation a solid bar, the elytra and pygidium are contrastingly not composed of short stripes, the remain- black with white markings with no brown der of elytra uniformly dark brown except streaks to differentiate Z. eldenensis—iden- some males have short, basal stripe on 5th tification of the female’s species is prob- interstice, females occasionally have nar- lematical without examining the genitalia. row, white median stripe; pygidium (figure In males, the genitalia are clearly diag- 164) of each sex dark brown with white nostic (compare figures 165 and 174). The basal band and narrow median stripe; only broad ventral valve and arched sac sclerite 1st antennal segment dark red. of Z. eldenensis contrasts with the narrow ventral valve and lageniform sclerite of Z. Structure.—Metasternum flat or slightly obliteratus. depressed, not sulcate. Female 8th sternite with broad basal process (figure 167). Male genitalia.—As in figures 165 and 166; Zabrotes humboldtae Kingsolver ventral valve of median lobe broad, lateral (Figures 168–171) margins subparallel, spicules around central sclerite smaller than in Z. subfasciatus; Zabrotes humboldtae Kingsolver dorsal valve with broad, rounded process; 1990b:156. apical margin of each lateral lobe slightly Color.—Dorsal vestiture predominantly emarginate (figure 166). dark brown except median stripe of pronotum and of pygidium white (figure 168), Size.—Body length 2.3–2.5 mm; width elytra with white lateral maculae, pygidium 1.5–1.9 mm. (figure 169) with median stripe and basal Type depository.—USNM. band white; venter predominantly white Type locality.—Arizona, Coconino Co., but with dark brown patches on metacoxal Flagstaff. face and lateral angle of 1st abdominal sternum; head and legs white. Distribution.—AZ. Structure.—Vertex micropunctate, punc- Host plants.—None recorded. tures more elongate on frons, frontal ca- Natural enemies.—None recorded. rina prominent; antenna not dimorphic, reaching middle of metacoxa. Pronotum Immatures.—Not described. semicircular but with apex truncate; disk Discussion.—Females of this species can micropunctate with coarser punctures easily be mistaken for Zabrotes subfas-cia- toward lateral margins, disk evenly convex. tus unless the metasternum is examined, Scutellum triangular. Elytral striae narrow, but males are easily differentiated by the distinct in basal two-thirds changing to color pattern of the pygidium. In both faint lines of punctures apically; interstices sexes of Z. subfasciatus, the metasternum sparsely punctate. Metacoxal face sparsely, is longitudinally sulcate and clothed with shallowly foveolate in lateral two-thirds, conspicuous yellowish-gray setae, whereas impunctate medially; metatibia lacking the metasternum of Z. eldenensis is con- dorsolateral row of fine punctures. 5th ab- vex or slightly depressed and is uniformly dominal sternum nearly divided by pygidial clothed; 1st and 2nd antennal segments notch; pygidium sparsely microfoveolate. are dark red in Z. subfasciatus, but only Male genitalia.—As in figures 170 and the 1st segment is red in Z. eldenensis. 171; median lobe only slightly expanded Separation of this species from the partly in apical one-fourth; ventral valve subtri- sympatric Z. obliteratus is more difficult

48 angular with apex narrowly rounded; pygidium (figure 173) with basal band of dorsal valve with median, ligulate process, white or white mixed with light brown con- apicolateral margins strongly recurved; in- nected to narrow median white stripe, re- ternal sac with pair of small sclerites near mainder of pygidium uniformly light brown apical orifice leading to thickened folds in in males but dark brown to black in fe- basal one-half of sac; middle of sac with males; head and ventral areas gray except pair of small, sickle-shaped sclerites; api- metacoxal face and row of vague, brown colateral sac lobes densely, finely denticu- patches along lateral margins of abdomen. late; lateral lobes apically expanded (figure Structure.—Frontal carina sharp but hid- 171). The genitalia are nearly identical to den by vestiture. Antenna reaching 1st ab- Zabrotes spectabilis. those of dominal segment. Pronotum nearly semi- Size.—Body length 1.3 mm; width 1.0 mm. circular, not truncated apically, disk evenly All specimens seen are of uniform size. convex, basal lobe slightly depressed, Type depository.—USNM. surface of disk sparsely micro-foveolate, interspaces finely punctate; mesosternal Type locality.—California, Humboldt Co., apex broad, truncate in both sexes. Elytral Bair’s Ranch, Redwood Creek. striae narrow, nearly hidden by vestiture, Distribution.—CA. set with recumbent, curved setae; 1st stria deflected laterad near scutellum; interstic- Host plants.—None recorded. es flat, punctate; metacoxal face with shal- Natural enemies.—None recorded. low, lunate foveolae in lateral two-thirds, Immatures.—Not described. medial one-third nearly impunctate but with deep pit adjacent to trochanteral in- Discussion.—The combination of the sertion; pygidia of both sexes convex, male smooth metacoxal face and Zabrotes spec- more strongly reflexed apically. tabilis-type male genitalia is puzzling since other species with this configuration of Male genitalia.—As in figures 174 and 175; male genitalia have a puncture cluster on median lobe with lateral margins subpar- the metacoxa. Likewise, those species with allel; ventral valve broadly triangular; dor- smooth metacoxa have the genital form of sal valve rounded, lacking ligulate process; Z. obliteratus (figure 175),Z. subfasciatus internal sac with pair of thickened strands (figure 198), orZ. bexarensis (figure 140). extending from apical orifice; middle of sac with flask-shaped sclerite and “stopper” Zabrotes obliteratus Horn sclerite similar to that of Z. densus (figure 161); apex of sac abbreviated; lateral lobes (Figures 172–176) strongly expanded as in figure 175. Zabrotes obliteratus Horn 1885a:158; Leng Female 8th sternite as in figure 176. 1920:306; Bottimer 1968c:1013; John- son 1968:1271; Johnson and Kingsolv- Size.—Body length 1.9–2.5 mm; width er 1982:411; Kingsolver 1990b:157. 1.5–1.7 mm. Spermophagus obliteratus: Pic 913a:60. Type depository.—MCZC, holotype No. Color.—Basal two segments of antenna red- 3901. dish brown. Vestiture of pronotum usually Type locality.—“Arizona.” dark brown with light brown patches and Distribution.—AZ, TX; Mexico. streaks, median stripe and posterior corners white (figure 172); elytra dark brown Host plants.—Rhynchosia sp. to black with indistinct light brown stripes, Natural enemies.—None recorded. each elytron with a lateral, white, sinuate band fading to light brown marginal patch;

49 Immatures.—Not described. cal one-half of elytra confusedly punctate. Discussion.—See also the discussion of Metacoxa with scattered, shallow foveo- Zabrotes eldenensis. Z. obliteratus could lae in lateral one-half, cluster of 30–40 easily be confused with Z. subfasciatus but fine punctures near trochantin. Abdomen for the distinctive deep metasternal sulcus with male 5th sternum deeply emarginate; of the latter species and the male genita- pygidium micropunctate, sparsely microfo- lia. The female 8th sternite is slender and veolate. slightly expanded at the tip of the strut Male genitalia.—As in figures 179 and 180; (figure 176). The dimorphic pygidium in median lobe relatively short, broad; ven- Z. obliteratus serves to separate the sexes, tral valve rounded apically, lateral margins but the degree of dimorphism is not as slightly sinuate; dorsal valve triangular; striking as in Z. subfasciatus. The disjunct internal sac abbreviated, sparsely lined distribution (Kerrville, TX, and southern with fine spicules and two pairs of small, Arizona) probably is an artifact of random median sclerites; lateral lobes fused into collecting. ligulate median process with lateral, alate processes (figure 180). Zabrotes planifrons Horn Size.—Body length 1.3–1.5 mm; width (Figures 177–180) 0.8–1.1 mm. Zabrotes planifrons Horn 1885b:158; Leng Type depository.—MCZC, lectotype No. 1920:306; Bottimer 1968c:1013,1038; 3904 (by Johnson 1968). Johnson 1968:1271; Johnson and Type locality.—Arizona. Kingsolver 1982:411; Kingsolver 1990b:146. Distribution.—AZ, NM; Mexico. Spermophagus planifrons: Pic 1913a:61. Host plants.—Cassia leptadenia Greenm. Color.—Vestiture mostly gray but with in- (Johnson 1979a) (now Chamaecrista nicti- termixed reddish-brown setae, especially tans mensalis). along lateral margins of elytra, occasionally Natural enemies.—None recorded. on pronotum; apex of each elytron with Immatures.—Not described. large patch of reddish-brown setae (figure 177); pygidium with basal band of inter- Discussion.—This species is usually easily mixed gray and reddish brown (figure 178); separated by its small size, uniformly gray scutellum white. Antennal segments one appearance except for a large brown spot and two dark red. Vestiture on pronotal at the apex of each elytron, the cluster of disk directed toward midline. punctures on the metacoxal face, and the distinctive male genitalia (figure 179) with Structure.—Vertex densely micropunctate, a broad, rounded ventral valve, massive punctures becoming slightly coarser on apical frame around the base of the valve, frons; frontal carina feeble or evanescent; and fused lateral lobes with alate lateral ocular index 4.5:1; ocular sinus seven- appendages (figure 180). It is generally eighths length of eye; antenna not sexu- similar to Zabrotes stephani, n.sp., in ex- ally dimorphic, reaching posterior margin ternal appearance. These two species form of metacoxa. Pronotum semicircular, not a subgroup in which the metasternum is truncate apically, disk uniformly micro- flat, the metacoxa is punctate, and the punctate, lateral areas microfoveolate, bas- male genitalia are reasonably similar. al lobe slightly swollen. Scutellum triangular. Elytra wider than long, lateral margins Zabrotes spectabilis Horn arcuate; striae shallow, well-marked basally, evanescent in apical one-half, strial (Figures 129, 181–185) punctures elongate; interstices with single Zabrotes spectabilis Horn 1885b:157; Pic or double row of shallow punctures; api- 1913a:62; Leng 1920:306; Bottimer

50 1968c:1013,1038; Johnson 1967:1271, Size.—Body length 0.7–1.6 mm; width 1979a:124; Johnson and Kingsolver 0.5–0.9 mm. 1982:411; Kingsolver 1990b:147. Type depository.—MCZC, lectotype No. Spermophagus spectabilis: Pic 1913a:62. 3000 (by Johnson 1968). Color.—Basal antennal segment red; basic Type locality.—Nevada (lectotype). vestitural color varying from gray to golden brown to dark brown; head gray in both Distribution.—AZ, CA, NM, NV, OR, UT, TX. sexes; white maculations as follows: male Host plants.—Five species have been cited pronotum with median white stripe (figure for this bruchid: Senna roemeriana; S. du- 181), lateral margin with white spot; each rangensis durangensis; S. lindheim-eriana elytron with lateral white maculation and from Big Bend National Park, TX; Senna white sutural stripe, male pygidium (fig- bauhinioides; and Senna covesii from Ari- ure 183) with white basal band and me- zona. Plant names have been updated fol- dian stripe; female pronotum with basal lowing Irwin and Barneby (1982). Records and marginal spots white, lacking sutural from Astragalus, Acacia, Condalia, and stripe (figure 182); elytra without sutural Stanleya cited in Kingsolver (1990b) are stripe; female lacking basal band but medi- probably incidental and will need confir- an stripe narrower than for male; venter of mation by rearing procedures. male gray except brown on metacoxal face, Natural enemies.—None recorded. female with brown coxal spot and variable patch along lateral margin of abdomen. Immatures.—None described. Structure.—Frontal carina fine, short; ocu- Discussion.—The pattern of the dark brown lar index 5:1; ocular sinus seven-eighths form is similar to that of Zabrotes cruciger, length of eye; antenna not dimorphic, but the size and different armature in the reaching posterior margin of metacoxa. lobes of the internal sac (compare figures Pronotal margins semicircular but trun- 153 and 184) clearly separate them. In cated apically, punctation concealed by California, the golden brown forms could vestiture; lateral carina slightly arcuate. be confused with more distinctly marked Mesosternal apex carinate. Elytra slightly forms of Z. cynthiae, but in the latter wider than long; striae nearly concealed by species, the sexes are not dimorphically vestiture, strial punctures coarse, shallow, marked, the pronotum lacks the marginal striae discrete basally but becoming con- white patches, and the shoulders of the fused apically, interstices finely punctate; dorsal valve are not recurved. Two similar metacoxal face finely punctate (figure 129) California species, Z. densus and Z. hum- except area around medial cluster of punc- boldtae, can easily be separated from Z. tures, lateral one-third of face shallowly spectabilis by their smooth metacoxae. foveolate; pygidium foveolate, sculpture nearly hidden by vestiture. Zabrotes stephani Kingsolver Male genitalia.—As in figures 184 and 185; (Figures 186–189) apical one-fourth of median lobe expanded; Zabrotes stephani Kingsolver 1990b:149. ventral valve broadly triangular, apex narrowly rounded, lateral margins sinuate; Color.—Vestiture on head sparse, very fine; dorsal valve with ligulate median process, pronotum and elytra with mixed brown apicolateral margins strongly recurved; in- and gray fine setae, elytron often with faint ternal sac with prominent folds; middle of brown macula behind humerus and at sac with pair of small, lunate sclerites at- middle of lateral margin (figure 186); in- tached to ovate structure; apicolateral sac terstices, especially the 5th, faintly striped lobes lined with fine spicules; lateral lobes with gray in basal one-half, pygidium gray as in figure 185.

51 with denser median line to mixed brown Zabrotes subfasciatus (Boheman) and gray with gray median line (figure (Figures 17, 43, 125–128, 130, 190–199, 187). Venter of body gray. 207) Structure.—As usual for genus; ocular in- Spermophagus subfasciatus Boheman dex 6:1; ocular sinus three-fourths length 1833:111; Pic 1913a:62. of eye; antenna not dimorphic, reaching posterior margin of metacoxa; frontal Zabrotes subfasciatus: Zacher 1930:236; carina prominent; apex of pronotum not Harper 1966:93; Bottimer 1968c:1013; truncated. Elytral striae lightly impressed, Kingsolver 1970b:487; Howe 1973:572; strial punctures elongate. Metacoxa with Pfaffenberger and Johnson 1976:42; cluster of about 30 punctures set in Kingsolver 1990b:158. smooth medial one-half of metacoxal face, Spermophagus musculus Boheman lateral one-half punctulate and sparsely 1833:112. foveolate. Spermophagus semifasciatus Boheman Male genitalia.—As in figures 188 and 1839:137; Lucas 1858:28; Bottimer 189; median lobe narrowed then suddenly 1968c:1015. expanded at apex; ventral valve shield- Bruchus nesapius Fahraeus 1839:108. shaped; dorsal valve ligulate with lateral Amblycerus semifasciatus: Blackwelder margins incurved; internal sac abbrevi- 1946:763. ated, prominently folded near apical orifice; Zabrotes semifasciatus: Bottimer middle of sac with slender sclerite in oval 1968c:1015; Kingsolver 1970b:487. membranous structure; apicolateral sac lobes each with small cluster of spicules; Bruchus cingulatus Suffrian 1870:169; lateral lobes broadly expanded, apical mar- Sharp 1885:493. gin emarginate (figure 189). Bruchus leucogaster Sharp 1885:493. Size.—Body length 1.4–1.7 mm; width Spermophagus (Zabrotes) pectoralis Sharp 0.9–1.3 mm. 1885:492; Chittenden 1902:103; Pic 1913a:60. Type depository.—USNM. Spermophagus pectoralis: Back and Duck- Type locality.—Oklahoma, Latimer Co. ett 1918:16; Wolcott 1936:286; Back Distribution.—OK. 1940:7. Host plants.—Unknown. Zabrotes pectoralis: Blackwelder 1946:763 (as synonym of Z. subfasciatus). Natural enemies.—None recorded. Spermophagus semicinctus Horn 1894:411. Immatures.—None described. Zabrotes semicinctus: Bottimer 1968c:1013 Discussion.—This species most closely re- (in synonymy with subfasciatus); John- sembles Zabrotes chavesi and Z. planifrons son 1968:1271. in the form of the male genitalia, flat meso- Spermophagus dorsopictus Lepesme, in sternum, and punctate metacoxa. All three Vayssière and Lepesme 1941:201. species have a fairly uniform vestiture with Spermophagus minusculus Pic 1943:148 muted pattern—vestiture is gray in Z. pl- (misspelling of musculus). anifrons but with the elytral apices brown, the elytra of Z. stephani are usually gray Zabrotes obtectus: Kingsolver with some faint basal stripes, and those of 1990b:169,171. Z. chavesi are brown with yellowish-gray Spermophagus basicornis Pic 1928:298. lateral maculae. The male genitalia, al- Color is sexually dimorphic: though similarly constructed for the three species, are distinctive. Males.—Vestiture of head brown, pronotum and elytra with indistinct pattern of yellowish-brown and dark brown setae

52 (figure 190), pygidium uniformly yellow- rounded by clusters of short, stout spines; ish brown with indistinct, narrow median lateral lobes as in figure 199. stripe, sometimes mottled (figure 192); Size.—Body length: males 2.0–2.3 mm; metacoxa and lateral areas of 1st abdomi- females 2.2–2.8. Body width: males 1.4–1.5 nal segment dark brown, otherwise yel- mm; females 1.3–1.4 mm. lowish brown; integument of 1st and 2nd segments of antenna and of mouthparts Type depositories.—NHRS (subfasciatus, reddish brown, fore legs suffused with red- semifasciatus, musculus); Havana, Cuba dish brown. (cingulatus); BMNH (leucogaster, pectoralis); MCZC (semicinctus); unknown (dorsopic- Females.—Vestiture of head gray, prono- tus). tum with variable pattern of gray and dark brown, usually with median white stripe; Type localities.—subfasciatus, “Brasilia, elytra (figure 191) with curved, median, Insula Saint-Barthelemi”; musculus, Brazil; white to gray maculation composed of semifasciatus, St. Barthelemy; cingulatus, short stripes, remainder of elytra light Cuba; leucogaster, Central America; semi- to dark brown in variable pattern, usu- cinctus, Baja California; dorsopictus, Mada- ally with vague stripes, usually with short gascar. stripe on second interval near scutellum; Distribution.—Tropicopolitan. Bottimer pygidium varying from yellowish brown (1968c) reported it as being established with dark spots (figure 194) to dark brown in Texas. Blanc (1965) reported finding a with basal band yellowish brown and me- single adult on cotton at Blythe, CA, but dian line white (figure 193); ventral areas, its establishment in that State is doubtful. antenna, mouthparts, and legs as in male. Harper (1966) also reported it from Califor- Structure.—Antenna sexually dimorphic nia. In recent years, the species has be- (figure 197), that of male reaching 1st come widespread in South America, Africa, abdominal segment, that of female reach- and India (Southgate 1964). ing metacoxa. Apex of pronotum truncate; Host plants.—Cajanus cajan; Cicer ari- disk moderately convex, basal lobe slightly etinum; Dipogon lignosus; Glycine max; prominent; pronotum extremely densely Lablab purpureus; Phaseolus acutifolius, P. punctulate, lateral areas of pronotal disk angularis angularis, P. coccineus, P. luna- with sparse microfoveolae; strial punctures tus, P. vulgaris; Pisum sativum arvense; of elytra elongate, deep, interstices dense- Vicia faba; Vigna angularis, V. mungo, V. ly punctulate, each with a single row of subterranea, V. unguiculata sesquipedalis, slightly coarser punctures; metasternum of V. unguiculata unguiculatus. A record for both sexes with a median, depressed sul- Sebastiania palmeri is questionable since cus, the sulcus densely setose (figure 195); that species belongs in the Euphorbiaceae. lateral one-half of metacoxal face with 40 Natural enemies.—I have been unable to to 50 lightly impressed, crescentic foveolae, find reports of any parasites reared from medial one-half impunctate except for deep this bruchid in the United States, but at pit near trochanter (figure 130); pygidium least 13 species are reported from other densely punctulate with scattered, ellipti- parts of the world: Anisopteromalus ca- cal foveolae. landrae; Bruchocida orientalis; Charito- Male genitalia.—As in figures 198 and 199; podinus swezeyi, C. terryi; Choetospila median lobe moderately broad, margins elegans; Dinarmus laticeps, D. vagabun- subparallel; ventral valve subconical with dus; Entedon sp.; Heterospilus prosopidis; margins slightly curvate, dorsal valve with Pseudocatolaccus bruchocida; Tetrastichus short, ligulate process; internal sac with sp.; Triaspis thoracica; Uscana mukerjii, U. hourglass-shaped median sclerite sur- semifumipennis; Pyemotes ventricosus.

53 Immatures.—Kannan 1923 (egg); Pfaffen- Wittich (1940) reported that in Minnesota berger and Johnson 1976; Steffan 1945 Z. subfasciatus caused allergic reactions (first larval instar); Arora 1978, Zacher in humans as a result of handling infested 1930, Peterson 1951 (final instar); Pfaffen- beans. berger 1991:568 (larva). Recent studies by Osborn et al. (1988) Discussion.—Zabrotes subfasciatus is usu- have isolated a seed protein in wild strains ally easily distinguished from other spe- of Phaseolus vulgaris, arcelin, that is toxic cies of Zabrotes by the sulcate, tomentose to Z. subfasciatus larvae. Experimental metasternum and by the sexual dimor- transfer of this trait to lines of cultivated phism in antennae, color pattern, and form beans has resulted in a high degree of re- of pygidium (but see Z. sylvestris). Zabrotes sistance to the bruchid. eldenensis closely resembles Z. subfas- Common names: Mexican bean bruchid, la ciatus but lacks the metasternal sulcus bruche brasilienne, der Brasilenbohnen- (although in some specimens the metaster- käfer. num is slightly depressed) and the sexes are similar, especially in the color pattern Because of its importance as a stored le- of the pygidium. The male genitalia are gume pest, references to studies on various also diagnostic to separate these two spe- aspects of the morphology and life history cies. of Zabrotes subfasciatus are listed. Full citations will be found in “References.” This species undoubtedly originated in Central America where it has been found Adult illustration.—Bondar 1936; Kingsolv- breeding in “wild” Phaseolus vulgaris and er 1990b. P. lunatus. It was described from Brazil in Adult morphology.—Zacher 1930; Seeliger the early nineteenth century but through 1943. commerce, mainly in shipments of dried Chromosomes.—Smith and Brower beans, it has become tropicopolitan in 1974:322. distribution. Various species and varieties of Phaseolus seem to be preferred hosts, Eclosion.—McFarlane and Wearing 1967 but in the past 20 years, it has become (pupal window). a storage pest of cowpeas in Africa (Meik Egg morphology.—Bondar 1936; Wightman and Dobie 1986). Some of the hosts listed and Southgate 1982. above are probably experimental or adven- titious. Female genitalia.—Singh 1973; Monga and Sareen 1980 (ovarioles); Romero and John- Zabrotes subfasciatus is a continuous son 1999. breeder in stored seeds, a logical result of its trait of ovipositing only on seeds that Host plant change.—Pimbert 1985b; Meik have fallen from a pod or that are still at- and Dobie 1986. tached inside a partly dehisced pod. The Larval digestive system.—Vats 1976b. bruchid apparently does not oviposit on Larval malpighian tubules.—Vats 1976a. the seed pod itself. Detection of infested seeds is difficult unless the prepupal stage Larval parasites.—Kistler 1985. has been reached, in which case the larva Life histories.—Zacher 1930; Lepesme has excavated to the inner surface of the 1944; Steffan 1945; Howe and Currie integument and has partly cut a “window” 1964; Carvalho and Rosetto 1968; Davies to facilitate emergence of the adult after 1972; Singh et al. 1979; Jarry and Bonet eclosion. Pierre and Pimbert (1981) report- 1981; Pimbert 1985c; Pajni and Jabbal ed on reproductive activity of this species. 1986; Pfaffenberger 1990. Male accessory glands.—Singh 1978b.

54 Male genitalia.—Kingsolver 1970b (figures shaped sclerite, apex of sac abbreviated; 48 and 49), 1990b:174 (figures 74 and 75). lateral lobes as in figure 203. Mating and reproduction.—Pierre 1980; Size.—Body length 2.1–2.3 mm; width Pierre and Pimbert 1981; Pimbert and 1.5–1.8 mm. Pierre 1983; Pimbert 1985b. Type depository.—MCZC, lectotype No. Nervous system.—Kasap 1978a; Sandhu 3902 (by Johnson 1968). and Neena 1982 (brain). Type locality.—”Georgia.” Oviposition.—Howe and Currie 1964; Utida Distribution.—AZ, AL, FL, IL, KY, MS, NJ, 1967; Pimbert 1985a. OK, SC, TX, VA. Physiology.—Sharma and Sharma 1979 Host plants.—Unknown. (proteins); Minney et al. 1990. Natural enemies.—None recorded. Pheromones.—Pouzat et al. 1989. Immatures.—Not described. Spermatheca.—Surtees 1961. Discussion.—Salient characters for this Wing venation.—Singh 1981a. species include the smooth medial one- half of the metacoxa, evenly distributed Zabrotes subnitens Horn white body setae, dimorphic antennae, and (Figures 200–203) male genitalia with a narrow ventral valve and lageniform median sclerite, a pair of Zabrotes subnitens Horn 1885b:158; Bot- spiny, bladelike sclerites, and a pair of timer 1968c:1015; Johnson 1968:1271; small, curved, spiny sclerites. This species Johnson and Kingsolver 1982:411; is most similar to Z. bexarensis, but the Kingsolver 1990b:160. elytra of the latter species is faintly macu- Spermophagus subnitens: Pic 1913a:62. late, and the male genitalia are distinctive Color.—First antennal segment sometimes (compare figures 140 and 202). dark red beneath, metatibial spurs dark red. Vestiture white, uniformly distributed Zabrotes sylvestris Romero and Johnson over body (figure 200). (Figures 204–206) Structure.—Antenna dimorphic, that of Zabrotes sylvestris Romero and Johnson male reaching margin of pygidium, that of 1999:87–98 female reaching 1st abdominal segment; Color.—Body black, head with basal an- pronotal disk sparsely microfoveolate, each tennal segments yellowish. Vestiture of foveola elliptical, interspaces densely punc- intermixed yellow and white pubescence, tate; elytral striae deep and narrow, strial yellowish brown spot of setae on vertex, punctures elongate, interstices minutely pronotum with scattered minute brown punctate; pygidium (figure 201) foveolate and yellowish spots; scutellum white; ely- as on pronotum; apex of meso-sternum tron with variable small brown and yellow truncate; lateral one-half of metacoxal face spots; pygidium yellowish with median with about 25 shallow, crescentic foveae, stripe; abdomen white except for yellowish medial one-half polished, impunctate ex- spot either side of first sternum; metacoxa cept for trochanteral pit. brown; metepisternum with small yellow- Male genitalia.—Of the Z. obliteratus type ish spot; abdomen with mixed yellow and (figures 202 and 203); lateral margins of white pubescence. median lobe subparallel; ventral valve Female color.—Similar to male except has subtriangular with apex narrowly rounded; contrasting white band across middle of dorsal valve likewise rounded; internal sac elytra. with two elongate, spinous structures, two small spinous sclerites, and median, flask- Structure.—Antenna seven-eights length of body; ocular sinus two-thirds length

55 of eye; head with evanescent median ca- the entire disk and the metatibia has only rina. Pronotum semicircular; disk densely a ventral carina, whereas in Z. subfasciatus punctulate and microfoveolate. Scutellum the pronotum is foveolate only on lateral small, triangular. Elytral disk micropunc- areas of disk and the metatibia has a faint tulate, striae deep, basal punctures deep; lateral carina in addition to the ventral metasternum with mesal fossa; metacoxal carina. face foveolate in lateral one-half, medial Differences in the male and female genita- one-half smooth and polished, finely stri- lia are illustrated (figures 204 and 198 for ate, cavernous at trochanteral insertion; male; figures 206 and 207 for female). metatibia with only one ventral carina. Female structure.—Ocular sinus two-thirds Zabrotes victoriensis Kingsolver to three-quarters length of eye; antenna (Figures 208–211) shorter than male, three-fifths to three- fourths length of body. Otherwise as male. Zabrotes victoriensis Kingsolver 1990b:151. Male genitalia.—As in figures 204 and 205; Color.—Vertex with extremely fine, white median lobe moderately broad, margins setae, clypeus densely clothed with white; subparallel; ventral valve (figure 204) with dorsal vestiture mostly reddish brown but base broad, apex broadly rounded; dorsal intermixed with white; white patches on valve ogival; internal sac with long bifid basal lobe, lateral margins of pronotum, process near base; median sclerite broadly scutellum, and in middle of 6th and 7th circular with two horn-like processes, sur- interstices of elytra (figure 208); pygidium rounded with fine spicules, a larger apical predominantly white with some intermixed cluster of fine spicules; lateral lobes as in reddish-brown setae either side of narrow figure 205. median white stripe (figure 209); venter of body densely clothed with white setae ex- Female genitalia.—Genitalia with large, cept metacoxal face reddish brown. spiny sclerite in bursa copulatrix. Structure.—Vertex finely, shallowly micro- Size.—Body length of males 1.5–2.0 mm; of punctate, punctures slightly deeper on females 2.1–2.5 mm. frons; frontal carina prominent, minutely Type depository.—Collección Entomológica granulose; ocular index 5:1; ocular sinus del Instituto de Fitosanidad, Colegio de three-fourths length of eye; antenna not Postgraduados, Montecillo, Mexico. sexually dimorphic, reaching middle of Type locality.—Mexico, Tehuacan, Puebla. metacoxal face. Pronotum semicircular, slightly swollen medially, minutely punc- Distribution.—CA; Mexico. tate on swelling, lateral areas microfove- Host plant.—Phaseolus vulgaris. olate. Scutellum minute, triangular; meso- Natural enemies.—Not known. sternum acute and carinate. Elytra together about as long as wide; striae shallow, Immatures.—Not known. distinct in basal four-fifths but becoming Discussion.—Zabrotes sylvestris and Z. confused near apex, strial punctures elon- subfasciatus are similar in size, shape, and gated; interstices flat, each with a single color, so positive identification should ulti- row of punctures. Metacoxal face micro- mately be made by comparing male genita- foveolate in lateral one-half, medial one- lia of the two species (figure 204,sylvestris, half with cluster of 20–25 punctures near and figure 198,subfasciatus ). Zabrotes trochanter. Male 5th abdominal sternum sylvestris is described as lacking the spot deeply emarginate; pygidium punctulate of yellowish brown setae found on the ver- with two sizes of punctures, male pygidium tex of Z. subfasciatus, but this should be strongly reflexed. relied on with caution because of abrasion. Male genitalia.—As in figures 210 and 211; In addition, the pronotum is foveolate over median lobe with lateral margins sub-par-

56 allel; ventral valve triangular with margins incurved; dorsal valve arcuate, lacking median process; internal sac with flask- shaped sclerite with “stopper” sclerite at narrow end; sac abbreviated; lateral lobes as in figure 211. Size.—Body length 1.5–1.7 mm; width 1.2–1.4 mm Type depository.—USNM. Type locality.—Texas, Victoria. Distribution.—TX. Host plants.—The host cited on the holotype label, Xanthoxylum clava-herculis, is doubtful since other species of Zabrotes for which the host is recorded prefer legumes whereas “Xanthoxylum,” (now Zanthoxy- lum) is in the Rutaceae. Natural enemies.—None recorded. Immatures.—None described. Discussion.—This species most closely resembles Zabrotes chavesi Kingsolver, but that species is mostly gray with some reddish mottling on the dorsal surfaces and the pygidium, the mesosternum is flat, and the male genitalia is of a different type (compare figures 149 and 210).

57 Subfamily Bruchinae Pic 1913a to form a hoodlike structure covering most of the median lobe with the apical portion of the fused lobes demarcated by a trans- Tribe Megacerini Bridwell verse ridge bearing setae or pores or small Megacerini Bridwell 1946:53; Bottimer sensory structures (figure 217). Lateral 1968c:1033,1039,1042; Teran and margins of the lobes extend ventrad partly Kingsolver 1977:27. Type genus: enveloping the median lobe. Megacerus Fahraeus 1839. Since Megacerus is the only included ge- Elytra with 10th stria not extending past nus, the above description applies to it as epipleural lobe (except M. impiger); front well. carinate; antenna serrate in females (figures 7 and 231), strongly pectinate in Genus Megacerus Fahraeus males (figures 6 and 230); pronotum Megacerus Fahraeus 1839:34; Bridwell conical, lateral margin straight or slightly 1929b:112, 1932:105, 1946:54; Bottim- convex, lateral carina sharp, sinuate; er 1968c:1033; Teran and Kingsolver elytra depressed along suture, interstices 1977:27. Type species Bruchus pescap- variable in width, 4th and 5th striae ab- rae Fahraeus 1839, monotypic. breviated apically; metafemur (figure 215) moderately thickened, ventral face flat or Pachybruchus Pic 1912:92; described as slightly convex, both margins carinate, a subgenus of Bruchus for “coryphae mesoventral carina smooth or dentate or Ol., pescaprae et voisins,” Bridwell with subapical angular tooth; metatibia 1929a:113; Bottimer 1968c:1034; with four longitudinal carinae—dorsolat- Teran and Kingsolver 1977:140. Type eral and ventrolateral on lateral face (figure species Bruchus coryphae Olivier, 215), dorsomesal and ventromesal on me- by subsequent designation, Bridwell sal face1; pygidium flat or with slight relief 1929b:113. or vertical or slightly oblique. Serratibruchus Teran and Kingsolver 1977:174 (as subgenus of Megacerus). The structure of the male genitalia in the Type species: Bruchus maculiventris Megacerini is unlike that of any other Fahraeus 1839, by original designation. bruchid tribe. The median lobe consists of an apical ring to which is attached the Nine species of Megacerus are known from ventral valve (figure 216). The ventral part the United States. of the ring is narrowed to a long, bifurcate Using the manuscript name Megacerus ventral rod expanded at the anterior end to pescaprae, Faldermann sent a bruchid form the cucullus. In most other bruchids, to Schoenherr for naming. Fahraeus the body of the median lobe is sclerotized described it as Bruchus pescaprae cit- to some degree, whereas in Megacerus the ing Megacerus in synonymy. Bridwell middle of the median lobe is a membra- (1929b:112) regarded Megacerus as a valid nous tube attached to the apical ring and generic name based on “Opinion No. 4” of to the cucullus with the rod providing ri- the International Code. Under Article 11e gidity. Pressure to evert the internal sac is of the third edition of the Code, the name accomplished by the usual muscle masses is valid since it was used by Bridwell prior connecting the rim of the cucullus and to 1961. The name Pachybruchus was the ventral strut to form a bulblike pump. taken by Pic from a Jekel manuscript. Lateral lobes are fused along the midline Morphology of this genus is described under tribe Megacerini above. ______Revision: Teran and Kingsolver 1977. 1Nomenclature herein proposed for the metatibial carinae is different from that proposed by Johnson and Kingsolver (1973) for Sennius, which is applicable to most of the Acanthoscelidini.

58 Key to U.S. Megacerus [The following key includes subgeneric as well as specific distinctions.]

1 Metafemur serrate on mesoventral margin (figure 52); Elytra with complete or interrupted stripes of white metacoxal face foveolate or punctate and setose in lateral setae especially on 3rd, 5th, and 7th interstices (figure one-half, polished on mesal one-half; mucro of metatibia 243)...... leucospilus (Sharp) shorter than width of tibial apex (figure 249) (subgenus 5(3) Pygidium white with horizontal black line, sometimes Serratibruchus)...... 2 broken into two segments (figure 225); elytra red with Metafemur not serrate on mesoventral margin (figure black sutural stripe (figure 224).. cubicus (Motschulsky) 215); metacoxal face foveolate or punctate over entire Pygidium and elytra otherwise...... 6 surface (except fossula), setose or not (figure 237); mucro short or long...... 3 6(5) Elytra black with large red maculation in basal one-half (figure 228)...... discoidus (Say) 2(1) Vestiture uniformly white on body (figure 252) except pygidium with four round, black spots (figure 253); 5th Elytra reddish brown with black lateral maculations sternum unicolorous...... schaefferianus Bridwell (figure 235)...... 7 Vestiture mostly black on elytra with tufts of white setae, 7(6) Anterior margin of male metacoxa with broad, polished especially on 3rd interstice (figure 247); pygidium with area, extending posteriorly onto coxal face (figure 237); two or four spots (figure 248); 5th sternum with one or apex of elytra extending past base of pygidium (figure two spots on each side of pygidium maculiventris 237); epipleural lobe slightly sinuate (figure 237), 10th (Fahraeus) stria extending to 1st abdominal segment; elytra with faint diagonal ridge extending from prominent humeral 3(1) Strial punctures large, deep (figure 212); interstices gibbosity...... impiger (Horn) convex or ridgelike; metacoxa with large but shallow punctulate foveae (figure 215); pygidium white with black Anterior margin of male metacoxa narrow (figure 220); or dark red spots; male with hooklike mucro on protibia apex of elytra not reaching past base of pygidium (figure (subgenus Pachybruchus)...... 4 220); epipleural lobe strongly sinuate (figure 220), 10th stria ending abruptly opposite metacoxal angle; Strial punctures fine (figure 218); interstices flat; elytra without diagonal ridge extending posteriorly from metacoxa finely punctate; pygidium various (subgenus humeral gibbosity...... 8 Megacerus)...... 5 8(7) Size larger than 2.3 mm long X 1.4 mm wide 4(3) Elytra red, usually with median sutural patch of white ...... cubiculus (Casey) hairs, remainder of elytra with short, scattered, inconspicuous hairs (figure 212)...... coryphae (Olivier) Size smaller than 1.7 mm long X 1.2 mm wide; Florida ...... ripiphorus (Fahraeus)

Megacerus (Pachybruchus) coryphae Megacerus coryphae: Bridwell 1929b:112; (Olivier) Blackwelder and Blackwelder 1948:44; Bottimer 1968c:1034,1039,1042; John- (Figures 6, 7, 212–217) son and Kingsolver 1982:412; Borowiec Bruchus coryphae Olivier 1795:16, no. 79; 1987:72; Udayagiri and Wadhi 1989. Fabricius 1801:402; Gyllenhal 1833:42; Megacerus (Pachybruchus) coryphae: Teran Fahraeus 1839:39; Jekel 1855:1; Horn and Kingsolver 1977:153. 1873:321; Blatchley 1919:66; Bondar Color.—Head and antenna black, thoracic 1936:43; Johnson 1969c:56. sterna and legs dark red to black, pro-no- Bruchus (Pachybruchus) coryphae: Pic tum and elytra dark red (figure 212), apex 1913a:23; Teran 1967:312. of pronotum and elytral suture usually Mylabris coryphae: Leng 1920:305.

59 piceous; pygidium dark red but thickly Size.—Body length 3–3.5 mm; width 2–2.5 covered with white, tomentose pubescence mm. except pair of black marginal subapi- Type depository.—Type apparently lost. cal spots and small, black spiracles near dorsal angles (figure 214). Vestiture white, Type locality.—”Amerique septentrionale.” short and sparse on head, pronotum, and Distribution.—AL, FL, LA, MS, SC, TX. elytra except for common, tomentose patch Host plants.—Ipomoea pes-caprae, I. sagit- at middle of suture; ventrally tomentose tata. except metacoxal face bare. Natural enemies.—None recorded. Structure.—Head finely, densely punctate; ocular index 20:1 in males, 10:1 Immatures.—Not described. in females; male antenna as in figures 6 Discussion.—The dark red, nearly bare pro- and 213, female as in figure 7; pronotum notum and elytra except for white median subpentagonal, lateral margins sinuate, sutural patch, densely white pygidium with disk strongly convex, densely but dis- two subapical black spots, deep, rounded cretely foveate, sulcate along basal margin strial punctures, smooth ventral carinae of either side of scutellum, basal lobe shal- metafemur, foveolate metacoxa, and long lowly sulcate, lateral carina short, serrate. mucro should be sufficient to distinguish Scutellum prominent, densely pubescent. this species from others in this handbook. Elytra together as long as wide; sulcate along suture; strial punctures deep, round, The other U.S. species in this subgenus, encroaching on sinuate interstices, crest of M. cubicus, superficially resemblesM. each interstice with small, shallow foveo- coryphae but differs in that in addition to lae. Metacoxal face finely punctate between the median sutural patch of vestiture, the large, flat-bottomed but umbilicate foveae simply punctate interstices are alternately (figure 215); hind leg as in figure 215, ven- striped with white hairs and the apices of tral carinae entire; metatibia with dorso- the 2nd and 3rd interstices are pubescent lateral and ventrolateral carinae divergent (figure 224). These two species are distin- toward apex, ventromesal and dorsomesal guished from others in the genus by char- carinae complete; mucro three-tenths as acters in the key separating the subgenus; long as basitarsus, apical margin crenu- however, M. cubicus with similar coloration late. Pygidium nearly flat in both sexes; can be separated by the presence of a hori- male 5th sternum emarginate, 8th tergum zontal black pygidial line (figure 225) in protruding between emargination and py- addition to the subapical elytral spots. gidium. Megacerus (Megacerus) cubiculus (Casey) Male genitalia.—As in figures 216 and 217; (Figures 218–223) median lobe slender, rodlike, expanded into ring apically; ventral valve ogival with Bruchus cubiculus Casey 1884:183; Horn apex acute; armature of internal sac con- 1885a:110; Pic 1913a:23; Schaeffer sisting of cluster of fine spicules at apical 1907:295. orifice, a pair of amorphous structures Mylabris impiger var. cubiculus: Leng apicad of cluster; another smaller cluster 1920:305. in middle of sac at one end of double row Megacerus cubiculus: Bottimer of 20–22 elongate denticles, sac sparsely 1968c:1034,1039; Johnson 1969e:237; lined with minute denticles; lateral lobes Johnson and Kingsolver 1982:411; (figure 217) fused into broad, troughlike Udayagiri and Wadhi 1989:209. structure, apex evenly rounded, setose, transverse row of minute setae near apex, Megacerus (Megacerus) cubiculus: Teran interior surface with many circular pores. and Kingsolver 1977:50.

60 Color.—Body mostly reddish brown with Male genitalia. —As in figures 221 and vertex and sternal regions black, metacoxal 222; median lobe rodlike, Y-shaped, with face and portions of the ventral margin of broad, terminal ring; ventral valve nar- the metafemur black, fore legs and mid rowly triangular, apex acute; armature of legs reddish brown; middle of pronotum internal sac consisting of cluster of fine usually with two quadrate black patches spicules at apical orifice, an amorphous, (figure 218), elytra usually with elongate saclike structure; apex of sac with 20–22 black spot on humerus, at middle of 7th slender denticles and many fine, acute interstice, and rounded subapical spot denticles, a pair of denticles near apex; (figure 218); variably maculate with black; lateral lobes fused, hoodlike, apex bluntly pygidium usually with two subapical triangular, porous, delimited by transverse black spots (figure 219). Antenna usually porous, setose ridge (figure 222). with apical four or five segments dusky or Size.—Body length 2.5 mm; width 1.7 mm. black. Vestiture white or yellowish white; head mostly yellow with white patch above Type depository.—USNM, holotype No. eye and adjoining frontal carina; pronotum 49229. yellow with white curved stripes; scutel- Type locality.—”Arizona.” lum white; elytra with stripes of alternating Distribution.—AZ, FL, GA, LA, MD, OK, SC, white and yellow; pygidium densely pubes- TX, VA; Mexico. cent on lateral borders and median stripe but more sparse between; ventral areas of Host plants.—Ipomoea batatas, I. lacunosa, body white. I. purpurea, I. trifida; Merremia quinquefolia. Records in the literature for Senecio Structure.—Vertex punctulate, male eyes salignus (floral record),Baccharis sarathroi- narrowly separated, ocular index 24:1 in des (floral record), andConvolvulus sp. are male, 7:1 in female; male antenna as in probably misidentifications or accidental figure 223; pronotum bell-shaped, convex, floral visits. with shallow sulcus either side near apex and on basal margin opposite 3rd and 4th Natural enemies.—None recorded. elytral striae (figure 218); disk finely -re Immatures.—Not described. ticulate-punctate, lateral margins slightly convex with short segment of carina oppo- Discussion.—This species most nearly site mesepisternum. Scutellum quadrate, resembles M. impiger. See the discussion bifid apically. Elytra together slightly lon- of that species for comparison. Geographi- ger than wide (figure 218), convex, except cal ranges of the two species overlap in the slightly depressed medially, apices not ex- Southwest. tending beyond basal margin of pygidium (figure 220), strial punctures fine, elongate, Megacerus (Megacerus) cubicus not encroaching on margins of interstices; (Motschulsky) interstices flat, finely imbricate; metacoxal (Figures 224–227) face densely microfoveolate (figure 220), foveolae mostly discrete, becoming more Kytorhinus cubicus Motschulsky 1874:206. crowded medially; meta-femur as in figure Bruchus cubicus: Sharp 1885:487. 220; ventral margin of metafemur flat be- Bruchus (Pachybruchus) cubicus: Pic tween two entire ventral carinae; metatibia 1913a:23. with dorsolateral carina absent, ventrolat- Megacerus cubicus: Blackwelder 1946:762; eral, dorsomesal, and ventromesal carina Janzen 1980:948; Pfaffenberger complete; mucro one-seventh length of ba- 1980:350; Johnson and Kingsolver sitarsus. Fifth abdominal sternum broadly 1982:411; Pfaffenberger 1984:24. emarginate; pygidium in both sexes verti- Megacerus (Megacerus) cubicus: Teran and cal. Kingsolver 1977:57.

61 Bruchus leucosomus Sharp 1885:488; Horn two irregular rows of elongate denticles in 1894:345; Fall 1910:164. middle, apex with four elongate denticles, Bruchus (Pachybruchus) leucosomus: Pic closure valve C-shaped; apex of fused 1913a:31. lateral lobes evenly rounded (figure 227), setose, with a pair of membranous alae Mylabris leucosomus: Leng 1920:305. behind apical plate, inner surface of lobes Megacerus leucosomus: Blackweld- porous. er 1946:762; Bottimer 1961:296, 1968c:1035; Udayagiri and Wadhi Size.—Body length 1.7–2.5 mm; width 1989:210. 1.1–1.5 mm. Color.—Integument black except legs red, Type depository.—ZMUM (cubicus); BMNH disk of elytra red with black border. Vesti- (leucosomus). ture white, sometimes shading to yellow on Type locality.—Mexico (cubicus, leucoso- pronotal disk, pronotum with broad, nearly mus). bare median stripe (figure 224) but pubescent on lateral margins, sparse to dense on Distribution.—TX; Mexico to Peru. elytra, tomentose on pygidium and ven- Host plants.—Ipomoea sp. Extralimital tral areas; pygidium with horizontal black hosts: Argyreia nervosa; Calystegia sepium; mark in middle of disk and pair of subapi- Ipomoea arborescens, I. carnea fistulosa, I. cal, marginal black maculae (figure 225). crassicaulis, I. hederifolia, I. leptophylla, I. Structure.—Vertex punctulate, frons with meyeri, I. nil (Janzen 1980:948). No speci- triangular bare area at dorsal end of fron- mens of this bruchid have been reared tal carina; ocular index 18:1 in male, 10:1 from seeds grown in the United States in female; ocular sinus one-half length although I. crassicaulis, I. hederifolia, and of eye in male, four-fifths length of eye I. nil occur here. in female; antenna of each sex as for ge- Natural enemies.—None recorded. nus. Pronotum subpentagonal (figure Immatures.—Pfaffenberger (1980) described 224), lateral margins straight, convergent, the egg and first larval instar and later in- disk convex, medially sulcate, basal mar- cluded the species in a key (1984:24). gin with curved sulcus each side of basal lobe, middle of disk densely, irregularly Discussion.—Characters to distinguish this microfoveolate, each foveola with fine seta species are the black pronotum, black-bor- beneath its anterior rim; lateral carina dered red elytra, and horizontal black line curved, concealed by vestiture. Scutellum on the pygidium (sometimes broken into ovate, convex, apically emarginate. Elytra two short lines). It resembles M. discoidus, together as long as wide; striae impressed, but the red elytral patch extends to, or punctures small, deep; interstices convex nearly to, the apex in M. cubicus, whereas to flat, densely imbricate. Metacoxal face in M. discoidus it extends only two-thirds densely punctulate, lateral two-thirds of to the apex. Pygidial markings of M. discoi- face concealed by dense vestiture; metatib- dus consist of four black dots, whereas in ia lacking dorsolateral carina; mucro short, M. cubicus the mark is a horizontal black curved, one-sixth as long as basitarsus, line (compare figures 225 and 229). lateral and coronal denticles small, incon- Pfaffenberger (1980:354) described oviposi- spicuous. tion for this species. Male genitalia.—As in figures 226 and 227; median lobe slender, apical ring broad, Megacerus (Megacerus) discoidus (Say) ventral valve subelliptical with apex bent (Figures 56, 228–234) ventrad; armature of internal sac consisting of two rows of minute spicules at apical Bruchus discoidus Say 1824:307; Schoen- orifice, a pair of flat plates in middle, and herr 1833:102, 1839:131.

62 Bruchus discoideus: Melsheimer 1853:99, either side of basal lobe (figure 228); disk and authors; LeConte 1859:171; densely foveolate, each foveola with seta Horn 1873:321; Blatchley 1910:1235; under anterior rim; lateral carina short, Cushman 1911:494; Weiss and Dick- curved. Scutellum cuneate, emarginate erson 1919:9; Carr 1920:7; Brimley apically. Elytra as long as wide (figure 1938:232; Johnson 1969c:56. 228); striae impressed, strial punctures Bruchus discoidens: Dury 1879:11. moderately deep; interstices convex in middle of elytra, imbricate; metacoxal face Kytorhinus discoidii: Motschulsky irregularly foveolate in lateral two-thirds 1874:205. with interspersed punctures, medial one- Bruchus (Pachybruchus) discoideus: Pic third punctate, fossula narrow; metafemur 1913a:24. as in figure 232, metatibia with dorsolat- Mylabris discoideus: Leng 1920:305. eral carina intermittent or absent, remain- Megacerus discoideus: Bridwell 1929b:112; ing carinae sharp, mucro one-sixth length Blackwelder and Blackwelder 1948:44; of basitarsus, lateral denticle distinct but Teran 1967:312; Parella and Kok short. Male with apex of 8th tergite visible 1975:44. between apex of pygidium and emargination of terminal abdominal sternum. Megacerus discoidus: Bottimer 1968c:1034; Mohyuddin 1969:143; Male genitalia.—As in figures 233 and 234; Teran and Kingsolver 1977:67; John- median lobe with apical ring broad, angu- son and Kingsolver 1982:411; Wang lar; ventral valve narrowly ogival with sinu- and Kok 1983:109; Pfaffenberger et al. ate side margins, apex acute; armature 1984:2; Pfaffenberger 1985a:4; Wang of internal sac consisting of two elongate, and Kok 1986a:359, 1986b:834; Uday- spiculate processes attached at their bases agiri and Wadhi 1989:210. to two flat, membranous lobes; middle of Color.—Integument black, occasionally with sac with two irregular rows of elongate some reddish suffusion on hind legs, each denticles separated by gap from four simi- elytron with large red spot extending from lar denticles near apex of sac; closure valve lateral margin to 2nd interstice and from C-shaped, entire sac lined with minute posterior margin of humerus two-thirds to- denticles; fused lateral lobes rounded or ward apex leaving narrow black basal and slightly emarginate apically (figure 234), sutural borders, apical one-third of elytra lateral alae angulate, apex of lobe porous, black (figure 228). Vestiture white; head, inner surface with grid of fine, transverse middle of pronotum, and elytra sparsely ridges. pubescent, pronotum usually with nar- Size.—Body length 2.6–3.6 mm; width row median line and flanks more densely 1.8–2.4 mm. pubescent; pygidium and ventral areas Type depository.—Type destroyed. densely pubescent except metacoxal face sparse; 5th abdominal segment with black Type locality.—”Arkansa.” spot at lateral margin; pygidium with four Distribution.—AB, AR, BC, CO, DC, FL, GA, black spots (figure 229). IN, KS, MD, MI, MN, MO, MS, MT, NB, NC, Structure.—Vertex finely, densely punctate; ND, NE, NH, NJ, NM, NS, NY, OH, OK, ON, frontal carina nearly hidden by trans- PA, PQ, SC, SD, SK, TX, VA; Mexico. versely directed setae; ocular index 16:1 in Host plants.—Calystegia macounii, C. males, 8:1 in females; ocular sinus three- sepium, C. sepium repens; Convolvulus fifths length of eye in both sexes; antenna arvensis; Ipomoea leptophylla, I. tiliacea, I. (figures 230 and 231) as for genus. Prono- pandurata. Flowers of Daucus carota and tum subpentagonal, basal margin arcuate, Hibiscus sp. lateral margins slightly sinuate, disk convex, faintly sulcate, basal margin sulcate

63 Natural enemies.—Uscana semifumipennis; Color.—Integument reddish brown with Dinarmus sp. variable darker shading on ventral areas of Immatures.—Pfaffenberger 1984 (egg, first body, on elytra, and on pronotum; hu- and final larval instars). meri and elytral apices piceous, pygidium usually with four irregular, dark blotches Discussion.—This handsome species with sometimes blended into two lateral macu- its quadrate, red elytral maculae can lae or into one large, central, piceous spot hardly be mistaken for any other eastern (figures 239 and 240). Vestiture white, ar- American bruchid. ranged in variable linear striping on elytra, The original spelling of the specific name uniformly distributed on pronotum, pygidi- by Say (1824) is “discoidus,” but Mels- um, and ventral areas. heimer (1853) inexplicably changed it to Structure.—Vertex finely punctate, frons “discoideus,” the form used by LeConte carinate in both sexes, ocular index 25:1 (1859). This spelling was used by sub- in male, 5:1 in female; ocular sinus 0.4 as sequent authors until Bottimer (1968c) long as eye in male, 0.6 as long as eye in reverted to the original. female; male antenna as in figure 238, 3rd Mohyuddin (1969), Wang and Kok (1983, and 4th segments of male antenna un- 1986a,b), and Keeler (1980) have detailed usually elongated. Pronotum bell-shaped, the life history of this species. evenly convex, lateral margins slightly arcuate, disk microfoveolate (figure 235), Adults are commonly found on flowers of lateral carina brief, located in middle of many different plant species. lateral margin. Scutellum broader at apex than at base, apex emarginate. Elytra 1.2 Megacerus (Megacerus) impiger (Horn) times as long as wide (figure 235), male (Figures 235–242) with prominent humeri extended pos- Bruchus ramicornis Boheman 1859:112 teriorly as a low, curved ridge to middle (not Erichson 1848). of elytron, striae subparallel except that two approaches three at base; epipleural Megacerus ramicornis: Blackwelder lobe not prominent, 9th stria reaching 1946:762. posterior margin of metacoxa, interstices Bruchus impiger Horn 1873:323, 1875:151; minutely imbricate, apices of elytra usu- Sharp 1885:486; Schaeffer 1907:295. ally extending beyond base of pygidium Bruchus (Pachybruchus) impiger: Pic in male (figure 237); female evenly convex 1913a:28. except humeri slightly prominent, striae Mylabris impiger: Leng 1920:305. and interstices as in male; metacoxal face of male with large polished extension on Megacerus impiger: Bridwell 1929b:112; anterior margin (figure 237), otherwise Blackwelder 1946:762; Blackwelder finely punctate; female with small fossular and Blackwelder 1948:44; Bottimer spot; metafemur slender; metatibia with 1968c:1035,1039; Johnson 1968:1269, evanescent dorsolateral carina, ventrolat- 1969c:55; Teran and Kingsolver eral, ventromesal, and dorsomesal carinae 1977:86; Schlising 1980:1; Johnson absent; mucro short, bent at angle to ven- and Kingsolver 1982:411; Udayagiri tral margin of tibia, lateral denticle nearly and Wadhi 1989:212. as long as mucro. Male abdomen slightly Megacerus eugenie Bottimer 1968c:1034 telescoped, 5th sternum broadly emargin- (replacement name for ramicornis Bo- ate; male and female pygidia vertical, disk heman 1859); Teran and Kingsolver finely, shallowly punctate. 1977:86. Male genitalia.—As in figures 241 and 242; Megacerus (Megacerus) impiger: Teran and median lobe with apical ring broad; ven- Kingsolver 1977:86. tral valve narrowly triangular; armature

64 of internal sac consisting of small cluster pare figure 218 with figure 235). The dor- of spicules at apical orifice, two ovate, flat somesal carina of the metatibia is absent structures, and 28–30 elongate denticles in in M. impiger but is strong in M. cubiculus. two irregular rows; closure valve C-shaped; These two species belong to a closely re- apex of fused lateral lobes bluntly trian- lated group of reddish-brown species that gular (figure 242), margins setose, inner are variable in color pattern and difficult to surface densely setose; transverse, arcuate separate. band densely setose, middle of lobes porous. Schlising (1980) conducted an intensive study of the life history of M. impiger in Size.—Body length 2.0–3.1 mm; width California. 1.3–1.8 mm. Type depository.—MCZC (impiger), lecto- Megacerus (Pachybruchus) leucospilus type not selected (Johnson 1968); NHRS (Sharp) (ramicornis). (Figures 243–246) Type locality.—California (impiger, ramicor- Bruchus leucospilus Sharp 1885:489. nis). Bruchus (Pachybruchus) leucospilus: Pic Distribution.—AZ, AR, CA, FL, GA, MS, SC, 1913a:31. TX; Mexico. Megacerus leucospilus: Bridwell Host plants.—Calystegia atriplicifolia, C. 1929b:113; Blackwelder 1946:762; occidentalis fulcrata, C. longipes, C. macro- Bottimer 1961:297, 1968c:1035; Teran stegia arida, C. macrostegia cyclostegia, C. and Kingsolver 1977:160; Johnson and macrostegia macrostegia, C. malacophylla Kingsolver 1982:412; Udayagiri and deltoidea, C. malacophylla malacophylla, Wadhi 1989:218. C. malacophylla pedicellata, C. occidenta- Megacerus leucorpilus: Bridwell 1929b:113 lis occidentalis, C. piersonii, C. purpurata, (misspelling). C. soldanella, C. stebbinsii, C. subacaulis; Convolvulus arvensis; Ipomoea sp. Megacerus alternatus Bridwell 1929b:113; Hinckley 1960:260; Bottimer Natural enemies.—None recorded. 1968c:1034; Johnson 1968:1269; King- Immatures.—Not described. solver 1970a:383. Discussion.—This species resembles M. Bruchus (Pachybruchus) coryphae var. cubiculus, but its range is generally west- lineatipennis Pic 1938:23; Hinckley ern with only scattered records east of 1960:260. Texas. Males are easily distinguished from Color.—Head and antennae, occasionally males of M. cubiculus by the shape of the anterior borders of elytra, and metati- antenna, the prominent humeri, the prono- bia black, remainder of body dark red, tal integumental pattern, the larger fos- legs sometimes piceous. Vestiture white sular spot on the metacoxa, and the less to yellow on head, pronotum, and elytra, prominent epipleuron. Elytral apices of M. otherwise white. Pronotal disk with alate impiger often extend beyond the vertical glabrous pattern, often divided by median plane of the pygidium, whereas those of M. line of setae (figure 243); vestiture of elytra cubiculus appear to end at the dorsal mar- striped on alternate interstices, interven- gin, but this character is variable. Females ing interstices glabrous, stripes sometimes of M. impiger are more difficult to separate fractured or lacking. Pygidium uniformly than males from those of M. cubiculus, setose except for two subapical, marginal but the pronotal pattern of M. impiger is spots (figure 244). usually a broad median stripe with lateral Structure.—Vertex finely punctate, frontal “wings,” whereas that of M. cubiculus is carina prominent, transverse sulcus sepa- composed of a broken median stripe (com- rating vertex from frons; eyes dimorphic,

65 male eyes enlarged, ocular index 15:1 in Natural enemies.—None recorded. male, 6:1 in female; antennae as for genus. Immatures.—None described. Pronotum bell-shaped (figure 243), strongly convex, lateral margins slightly convex; Discussion.—This species was evidently disk moderately convex, densely microfo- introduced into Hawaii sometime before veolate; lateral carina indistinct; cervical 1923, when O.H. Swezey first collected it. sulcus lacking. Scutellum elongate, dense- Bridwell compared his Megacerus alter-na- ly pubescent. Elytra striae sinuate, punc- tus to M. leucospilus but considered them tures deep, rounded (figurse 243); inter- to be distinct species. The alternate strip- stices convex, not imbricate; sutural stria ing of pubescence tends to be complete in depressed. Metacoxal face shallowly fove- the Hawaiian specimens but may be frag- ate; metafemoral margins nearly parallel; mentary in Mexican and Central American metatibial carinae distinct and complete, specimens. ventrolateral carina contiguous with dorsal margin of mucro; mucro curved, slender, Megacerus (Serratibruchus) maculiventris one-third as long as basitarsus, similar to (Fahraeus) that of M. coryphae (figure 215). Apex of (Figures 52, 55, 247–251) 8th tergite visible at apex of pygidium. Bruchus maculiventris Fahraeus 1839:38. Male genitalia.—As in figures 245 and 246; median lobe moderately broad at apex; Bruchus (Pachybruchus) maculiventris: Pic ventral valve ogival, apex acute; armature 1913a:33. of internal sac consisting of broad, dense Megacerus maculiventris: Blackwelder cluster of fine spicules attached to ovate 1946:762; Teran and Kingsolver amorphous structures, middle of sac with 1977:196; Janzen 1980:948; Johnson 30–32 curved, elongate denticles, closure and Kingsolver 1982:412; Pfaffen- valve C-shaped; fused lateral lobes with berger 1984:21; Udayagiri and Wadhi apical plate evenly rounded (figure 246), 1989:221. marginal setae stout, inner surface with Kytorhinus quadratus Motschulsky small clusters of fine setae, transverse 1874:205, synonymized by Teran and ridge setose, inner surface anteriad of ridge Kingsolver 1977. finely setose in fanlike pattern and with Bruchus impiger var.: Sharp 1885:487; two clusters of pores. [Schaeffer (1907:295) stated that his Size.—Body length 3.4–4.2 mm; width new species, serratifemur, is the species 2.2–2.6 mm. Sharp illustrated as a variety of impi- Type depository.—BMNH (leucospilus); ger]. USNM holotype No. 41901 (alternatus); Bruchus serratifemur Schaeffer 1907:294; MNHP (lineatipennis). Pic 1913a:49; Johnson 1968:1269, synonymized by Teran and Kingsolver Type locality.—Mexico, Veracruz (leuco-spi- 1977. lus lectotype); Hawaii, Oahu, Barber’s Point (alternatus); Hawaii (lineatipennis). Bruchus (Pachybruchus) kytorrhinensis: Pic (1913a:29) proposed this new name for Distribution.—LA, TX, HI; Mexico. quadratus Motschulsky (not Suffrian Host plants.—Ipomoea cairica, I. carnea 1870) fistulosa, I. crassicaulis, I. leptophylla, I. Mylabris serratifemur: Leng 1920:305. pes-caprae; Merremia aegyptia (Hawaii). Megacerus kytorrhinensis: Blackwelder Janzen (1980) records it from Costa Rica 1946:762. in Ipomoea carnea fistulosa and I. pes-caprae. De Luca (1967a) lists I. biloba without Bruchus mactatus Pic 1932:36, synony- locality or source. mized by Teran and Kingsolver 1977.

66 Acanthoscelides mactatus: Blackwelder tra 1.1 times as long as wide (figure 247), 1946:760. convex but sulcate in basal one-half along Bruchus (Pachybruchus) triangularifer Pic suture; striae narrow, deeply impressed, 1942:10, synonymized by Teran and punctures elongate; interstices finely, Kingsolver 1977. densely imbricate. Metacoxal face densely Bruchus (Pachybruchus) triangularifer var. punctate in lateral two-thirds, smooth and multisparsus Pic 1942:10, synonymized impunctate medially; metafemur serrate on by Teran and Kingsolver 1977. ventromesal margin (figure 52); metatibia with ventrolateral, dorsomesal, and ven- Pachybruchus triangularifer: Pic 1954:184. tromesal carinae complete, dorsolateral Pachybruchus triangularifer var. multispar- carina intermittent or evanescent, ventral sus: Pic 1954:184. face of tibia with partial carina ending in Bruchus kytorrhinensis Pic 1913b:110. mucro, mucro short, scarcely longer than Megacerus (Serratibruchus) maculiventris: lateral denticle, coronal denticles minute Teran and Kingsolver 1977:196. (figure 249). Pygidial punctation concealed by vestiture. Color.—Integument varying from red to black; head always black, antenna varying Male genitalia.—As in figures 250 and 251; from basal seven and terminal segments median lobe slender, apical ring broad; red to black with some reddish suffu- ventral valve narrowly triangular, apex pro- sion; pronotum all black to black median duced, margins slightly sinuate; armature shield with red flanks (figure 247); elytra of internal sac consisting of small cluster varying from all black to red with black of fine spicules at apical orifice attached basal patch; venter of body variable with to bilobed, amorphous structure, middle red and black patches; legs red to black. of sac with dense, V-shaped cluster of fine Vestiture of head mostly yellow with white spicules, apex of sac with eight curved, patch behind eye; pronotum mixed white elongate denticles in two irregular rows; and yellow in pattern shown in figure 247; closure valve angulate; lateral lobes with elytral vestiture varying from black with apical plate rounded but slightly emargin- scattered white spots in those specimens ate (figure 251), densely setose on margin, with uniformly black elytra to black with transverse row of pores caudad of plate, white patches on red integument; pygidium ridge extended laterally into small, mem- uniformly yellow, uniformly white, or yel- branous alae; main body of lobes porous low with white patches, always with paired and marked with diagonal lines. subapical black spots, sometimes with Size.—Body length 2.0–2.5 mm; width another pair at one-third from base (figure 1.1–1.6 mm. 248); venter of body with variable-sized patches of red, white, yellow, and red vesti- Type depository.—NHRS (maculiventris); ture in no consistent pattern. ZMUM (quadratus); MNHP (mactatus, triangularifer, multisparsus); USNM, holotype Structure.—Vertex finely, densely punctate, No. 42338 (serratifemur). frontal carina prominent; ocular index 9:1 in males, 5:1 in females; ocular sinus Type locality.—Venezuela, Caracas (macu- three-fifths as long as eye in males, one- liventris); Brazil (quadratus, multisparsus); half as long as eye in females; antennae as Arizona (serratifemur); Colombia (macta- for genus. Pronotum subpentagonal (figure tus); Peru (triangularifer). 247), lateral margins nearly straight, disk Distribution.—AZ, TX; Mexico, Central convex, basal lobe slightly elevated and America, northern South America. sulcate, surface microfoveolate on basal Host plants.—No hosts are known for lobe, interspaces punctate, lateral areas the United States records. Extralimital more finely punctate; lateral carina curved. hosts are Ipomoea pes-caprae, I. purpurea, Scutellum rectangular, pubescent. Ely-

67 I. marginisepala, I. aristolochiaefolia, I. ru- Discussion.—The two specimens Blatchley briflora, I. nil. described as Megacerus wheelocki appear Natural enemies.—None recorded. to be identical to the type specimen of M. ripiphorus, but the presence of this species Immatures.—Pfaffenberger 1980 (Egg, first in Florida is puzzling since the other three and final larval instars). specimens examined are from Venezuela Discussion.—The serrate ventromesal and Colombia. No additional records are margin of the metafemur will immediately known from the United States. Mislabel- separate this species from all other spe- ing of the specimens seems unlikely since cies in this report except M. schaefferi- Blatchley collected them. Since the range anus, from which M. maculiventris may be of M. cubiculus extends from eastern Unit- distinguished by its mottled integumental ed States to Panama, it and M. ripiphorus and vestitural pattern (no pattern in M. may eventually prove to be one species. schaefferianus), by the two or four small, Horn (1885a) synonymized cubiculus with black pygidial spots (four large spots in M. impiger, but the problem of the correct M. schaefferianus), and by the fine setae application of the three names needs fur- on the metacoxal face (tuft of coarse white ther study. setae in M. schaefferianus). Megacerus The small size of this species will for the maculiventris is widespread in the Western present separate it from M. cubiculus and Hemisphere. M. impiger, the other two reddish-brown species in the United States. Megacerus (Megacerus) ripiphorus (Fahraeus) Megacerus (Serratibruchus) schaefferianus Bruchus ripiphorus Fahraeus 1839:35; Pic (Bridwell) 1913a:44. (Figures 252–255) Acanthoscelides ripiphorus: Blackwelder Bruchus crenatus Schaeffer 1909:385; Pic 1946:761. 1913a:23; Johnson 1968:1269 (lecto- Megacerus ripiphorus: Teran and King- type designation). solver 1977:133; Johnson and King- Megacerus schaefferianus Bridwell solver 1982:412; Udayagiri and Wadhi 1929b:112 (as new name for B. cre- 1989:215. natus Schaeffer, not Thunberg 1791); Mylabris wheelocki Blatchley 1930a:35, Blackwelder and Blackwelder 1948:44; 1930b:59. Bottimer 1968c:1035; Teran and King- Megacerus wheelocki: Bottimer solver 1977:206; Johnson and King- 1968c:1035; Johnson 1969e:237. solver 1982:412; Udayagiri and Wadhi Except for its smaller dimensions, this spe- 1989:221. cies is indistinguishable from M. cubiculus Megacerus crenatus: Johnson 1968:1269. (Casey). Megacerus (Serratibruchus) schaefferianus: Size.—Body length 1.7 mm; width 1.2 mm. Teran and Kingsolver 1977:206. Type depository.—NHRS (ripiphorus); PURC Color.—Integument entirely black. Vestiture (wheelocki). white, uniformly distributed (figure 252), denser on ventral parts of body. Pygidium Type locality.—Colombia (ripiphorus); with four black spots (figure 253), 5th Florida, Royal Palm Park (wheelocki). ventral abdominal segment with black spot Distribution.—FL; Colombia, Venezuela. either side of emargination. Host plants.—Ipomoea alba (cited by Structure.—Vertex finely, densely punctate; Blatchley as I. bona-nox). frontal carina prominent; ocular index 3.25:1; ocular sinus one-half length of Natural enemies.—None recorded. eye. Pronotum (figure 252) subpentago- Immatures.—None described. 68 nal, apex rounded, posterior angles acute; Tribe Bruchini Bridwell basal lobe broadly sulcate between gibbosities; disk strongly convex in lateral aspect; Genus Bruchus Linnaeus lateral carina present in anterior one-half Bruchus Linnaeus 1767:604 (not Bruchus of margin; cervical sulcus brief. Scutellum Geoffroy 1762). Type species: Der- quadrate, densely setose. Elytra 1.5 times mestes pisorum Linnaeus 1758:356, by as long as wide (figure 252); striae regular, subsequent designation, Latreille 1810. punctures deep, scarcely encroaching on Laria Scopoli 1763:21, of authors. interstices, 1st and 2nd deflected laterally at base; interstices finely imbricate. Meta- Mylabris Müller 1764:14. Type species: coxal face irregularly foveolate in lateral Dermestes pisorum Linnaeus 1758, by one-half, polished and impunctate in me- subsequent designation, Bridwell 1932 dial one-half. Metafemur serrate on ventro- (not Mylabris Fabricius 1775). lateral margin. Metatibia with dorsolateral, Bruchoides Ramos 1978:317. Type spe- ventrolateral, ventromesal, and dorsome- cies: Bruchus tesselatus Mulsant and sal carinae present; mucro short, not as Rey 1858:38, by original designation (as long as lateral denticle. Fifth abdominal subgenus, not North American). segment of both sexes deeply emarginate; Acanthobruchus Ramos 1978:317. Type pygidium convex. species: Bruchus venustus Fahraeus Male genitalia.—As in figures 254 and 255; 1839:75, by original designation (as median lobe slender, apical ring broad; subgenus, not North American). ventral valve triangular with margins For a full explanation of the involved ap- straight; armature of internal sac con- plications of the names Bruchus and Myla- sisting of small cluster of fine spicules at bris, see Pope (1956) and Borowiec (1988). apical orifice connected to bilobed struc- Donahaye (1974) illustrated male and ture; middle of sac with short, V-shaped female genitalia for seven of the most com- cluster of sharp spines, apex with another mon stored-product Bruchus species. V-shaped cluster of eight spines, and a Although Laria was substituted for Bru- median rod, closure valve C-shaped; lat- chus by numerous authors, it cannot eral lobes with fused apices rounded (fig- be used in the Bruchidae since Bridwell ure 255), densely setose, transverse ridge (1932) designated Laria dulcamarae Sco- membranous with rows of sensory pores, poli, a nitidulid, as the type species. middle of lobes porous. Three species have become established in Size.—Body length 3.9 mm; width 2.2 mm. continental North America, two of these Type depository.—USNM, lectotype by also in Hawaii. Johnson 1968 (as Bruchus crenatus). Integument black with legs sometimes red Type locality.—Arizona, Nogales. or reddish orange. Vestiture of white or yellowish setae in relatively consistent dorsal Distribution.—AZ; Mexico. and pygidial patterns. Host plants.—Ipomoea longifolia (flowers) Head lacking frontal carina, antenna sub- Natural enemies.—None recorded. serrate; pronotum transverse, apex trun- Immatures.—Not described. cate, lateral margin with median tooth. Elytral striae straight, lacking basal denti- Discussion.—The large size, uniformly gray cles; interstices subequal in width; middle vestiture, and four pygidial spots should tibia of male with terminal armature char- distinguish this species easily from others acteristic of species; ventrolateral margin in this handbook. Available specimens are of metafemur with distinct subapical angle, remarkably uniform in size.

69 or tooth, ventromesal margin with or with- Bruchidae described in both the Old World out subapical denticle. and the New World. Schilsky (1905) described Acanthoscelides and Bruchidius Male genitalia elongate, slender. and transferred several species names to Most species of Bruchus—including all Bruchidius. Subsequent authors trans- of the species described herein—are uni- ferred the New World “Bruchus” species voltine (producing one generation a year) to Acanthoscelides. The three New World because eggs are laid only on green pods of species belonging to the “true” Bruchus— the host legume (Southgate 1981). These pisorum (Linnaeus), rufimanus Boh., and species do not breed in storage; however, brachialis Fahraeus—were introduced from they may emerge under storage conditions. the Old World. Bruchus lentis Froelich has This Old World genus is diverse as it now been reported from the Pacific Northwest stands and needs a thorough review. Char- as an escape from imported seeds but ap- acters used to distinguish Bruchus are the parently has not become established in the lateral pronotal margins each with a single United States or Canada. tooth (figure 46), or denticle; ventrolateral References to Bruchus are extensive in margin of the metafemur with a denticle Old World literature but are limited in the or angular tooth (figure 47); ventromesal New World. See Luk’yanovich and Ter- margin of metafemur with or without a Minassian (1957:70) for a summary of Old denticle; and bases of elytral striae lacking World references to that date. Aitken (1975) denticles. lists most of the commercially important The taxon Bruchus was used for many species of Bruchus and their host plants years as the repository for new species of and distribution.

Key to U.S. Bruchus 1 Hind femur with long external tooth near apex; mucro of remainder black; mesofemur of male inflated (figure hind tibia shorter than lateral denticle (figure 47); protibia 276); mesotibia of male sinuate with sharp terminal hook; and apical one-half of mesotibia and tarsus red 3.6–3.8 mm long...... rufimanus Boheman ...... pisorum (Linnaeus) Mucro of metatibia straight, short, no longer than lateral Hind femur with only an angulation in ventral border of denticle (figure 257); basal five and apical antennal femur (figure 257); mucro short or long; at least part of segments red in female, entirely yellow in male; protibia fore legs red, mid legs black...... 2 and mesotibia curved in male; mesotibia with flat, bifid, toothlike terminal process in male 2(1) Mucro of metatibia set at angle, twice as long as lateral ...... brachialis Fahraeus denticle (figure 274); basal four antennal segments red,

Bruchus brachialis (Fahraeus) and Gillespie 1958:401; Steinhauer (Figures 256–263) 1959:955; Bottimer 1968c:1033,1039; U.S. Department of Agriculture Bruchus brachialis Fahraeus 1839:79; 1971:736; Johnson and Kingsolv- (American references only follow) Bot- er 1982:416; Udayagiri and Wadhi timer 1931:347; Bridwell and Bot- 1989:183. timer 1933:739; Annand and Pinckney Bruchus pallidicornis Mulsant and Rey 1936:1; Bottimer 1936:807, 1937:379; 1858:33. Pinckney 1937:621; Vayssière and Lepesme 1941; Osborn 1949:159; Laria brachialis: Bedel 1901:357. Neilson and Handford 1954; Randolph Mylabris brachialis: Baudi 1886a:13.

70 Color.—Body and appendages black ex- spine bifid (figure 258). First abdominal cept four basal antennal segments reddish segment of male concave with setae ar- orange (female) or entire antenna reddish ranged in circular pattern; pygidium (figure orange (males), no more than apical one- 259) slightly convex, surface foveolate. half of profemur and all of protibia reddish Male genitalia.—As in figures 260 and 261; orange (females) or fore leg entirely reddish median lobe slender (figure 260), slightly orange (males). Dorsal vestiture intermixed expanded toward apex; ventral valve ovate, black and brown with irregular rows of apex broadly rounded, dorsal valve round- white spots elytra, pronotum mostly brown ed; armature of internal sac consisting with white on posterior corners, usually of a pair of small setal patches at apical along lateral borders and on basal lobe; orifice, sac lined with fine spicules for -en scutellum white; pygidium mostly white tire length, closure valve undifferentiated; with scattered brown setae and two vari- lateral lobes elongate (figure 261), slender, able black subapical spots; venter of body cleft one-half their length, apices strongly white with some brown intermixed with curved, each with setose, alate process; white on metepisternum, row of condensed 8th sternite bilobed, apices conjoined; male white spots near lateral border of abdo- spiculum gastrale Y-shaped. Eighth tergite men. of male hoodlike, apical margin thickened, Structure.—Vertex and frons densely punct- setose (figure 262); female 8th tergite hood- ulate tending in places to be imbricate, like, apical margin bilobed, serrate and frontal carina absent, dorsal margin of setose; 8th sternite (figure 263) elongate, frons with small, glabrous boss; ocular in- T-shaped. dex 3.2:1; ocular sinus three-fifths length Size.—Body length 2.5–2.9 mm; width of eye; antenna with club segments broad, 1.7–1.7 mm. terminal segment ovate, extending to humerus. Pronotum trapezoidal (figure 256), Type depository.—NHRS. lateral margins strongly sinuate, marginal Type locality.—Germany. denticle prominent, disk convex, basal Distribution.—AL, AR, BC, CA, CO, CT, DC, lobe sulcate, margin depressed either side DE, FL, GA, ID, IA, IN, KS, LA, MA, MD, of basal lobe; surface densely foveolate, MI, MO, MS, NC, NE, NJ, NY, OH, OK, asperate, interspaces imbricate; lateral ca- ON, OR, PA, SC, TN, TX, VA, WA, WY; Old rina brief at lateral denticle; cervical sulcus World. hidden in vestiture. Scutellum quadrate, deeply emarginate. Elytra together slightly Host plants.—Lathyrus sativus; Vicia ben- longer than wide; convex, depressed near ghalensis, V. caroliniana, V. cracca, V. gran- scutellum; striae narrow, deep, parallel; diflora, V. pannonica, V. sativa nigra, V. sa- interstices densely punctate-imbricate in tiva sativa, V. villosa dasycarpa, V. sepium, basal one-half, more sparsely punctate to- V. villosa villosa. Waterworth (1986) listed ward apex; metacoxal face densely punctu- several other host records of this species late to imbricate, fossula glabrous; hind leg found in seeds imported from Europe. as in figure 257; metafemur with ventrolat- Adults found on Daucus carota flowers. eral carina angulately toothed near apex, Natural enemies (Old World and New ventromesal margin neither carinate nor World).—Anisopteromalus calandrae; Di- denticulate, mesal face minutely tubercu- brachys cavus; Dinarmus acutus; Eupelmus late; mesotibia (figure 257) slightly arcuate amicus, E. cyaniceps; Eurytoma obtusa, and dilated; lateral, ventral, and dorsome- E. tylodermatis; Habrolepoidea tarsalis; sal carinae present, ventrolateral carina Laelius utilis; Lariophagus distinguendus; absent, mucro short, subequal to lateral Macroneura vesicularis; Microdontomerus denticle, three coronal denticles small; anthonomi; Pteromalus sp.; Tetrastichus male mesofemur slightly dilated medially; male mesotibia slightly arcuate, apical

71 bruchivorus, T. bruchophagi; Triaspis tho- 1956:45; Kingsolver 1964b:3; Bot- racica; Zatropis incertus. timer 1968c:1033,1039,1042; De- Immatures.—Pfaffenberger 1977 (final lar- celle 1975:115; Johnson and King- val instar), 1991:564 (larva); see below for solver 1982:416; Borowiec 1987:70, description of the egg. 1988:194; Udayagiri and Wadhi 1989:197; Decelle and Lodos 1989:172. Discussion.—The “vetch bruchid” was first Mylabris pisorum: Baudi 1886a:16; Leng detected in 1931 in pods of Vicia villosa in 1920:305; Kannan 1923:27; Essig New Jersey, Delaware, and Maryland and 1929a:484. reported by Bottimer (1931). A distribution map was published by USDA (1971), Mylabris cruciger Geoffroy 1785:112. and the species has now spread to 33 of Laria salicis Scopoli 1763:22. the 48 contiguous states and British Co- Bruchus sparsus Fabricius 1801:398. lumbia and Ontario in Canada. Downie Bruchus intermedius Motschulsky 1854:10. (1950) found specimens of B. brachialis from Washington State dated 1931, the Bruchus lunarius Rey 1893:3. same year as the first detection on the East Bruchus pisorum var. unifasciatus Rey Coast. 1893:3. This species is smaller than either Bruchus Bruchus fabae Brulle 1873:3228. pisorum or B. rufimanus. Male antennae Mylabris pisorum: Baudi 1886a:16. and fore legs are entirely yellow. The male Color.—Integument black with four basal mesotibial spine is bifid contrasted to the antennal segments, protibia and tarsi, single hook of the other two species. Use- part or all of mesotibia, and tarsus reddish ful characters are found in female as well orange. Vestiture white, yellowish brown, as male genitalia for this species. and dark brown or black in pattern of Eggs of Bruchus brachialis are 0.5–0.6 mm figure 264 on pronotum and elytra; head long, white, cylindrical, slightly tapered white or white mixed with yellowish brown; with ends rounded, and are elevated above pronotum mostly yellowish brown with in- the pod surface on an elongated basal termixed white, scattered white patches on “foot” cemented to the pod. disk, basal lobe with intense white patch; scutellum white; elytra mostly yellowish Bruchus pisorum (Linnaeus) brown with dark brown lateral maculae separated from dark brown apical patches (Figures 46, 47, 264–271) by diagonal row of white patches, small Dermestes pisorum Linnaeus 1758:356. white patches on 3rd interstice, on anterior Bruchus pisi: Linnaeus 1767:604; Gmelin margin of maculae, and near apices; pygid- 1790:1767; Jacquet 1888. ium mostly white, apex yellowish brown, basal and subapical patches black; venter Bruchus pisorum: Linnaeus 1758:34; mostly white but intermixed with yellow- Latreille 1810:192; Horn 1873:315; ish brown on metepisternum and sides of Hubbard and Schwarz 1878:660; abdomen. Abeille de Perrin 1888; Riley 1892:185; Blatchley 1910:1235; Fall 1910:161; Structure.—Vertex, frons, and clypeus Cushman 1911:507; Pic 1913a:41; densely foveolate; frontal carina absent; Bridwell 1932:104; Bissell 1938:536; ocular index 3.6:1; ocular sinus three- Larson et al. 1938; Metcalf and fifths as long as eye; postocular lobe Flint 1939:808; Back 1940:7; Vays- wedge-shaped, densely setose; antenna sière and Lepesme 1941:198; Hoff- moderately eccentric, club segments ovate man 1945:46; Blackwelder 1946:758; in cross section, extending to humerus. Bridwell 1946:52,54; Brindley et al. Pronotum transverse (figure 265), broadly 1946, 1952; Zacher 1952:461; Pope semicircular; each lateral margin with

72 small tooth at midpoint; disk densely Distribution.—Cosmopolitan. AB, AL, BC, foveolate, interspaces punctulate; lateral CA, CO, CT, DC, FL, GA, ID, KS, KY, MA, margin beneath with fine, polished sulcus MB, MI, MN, MO, MS, MT, NB, NC, NF, from posterior corner to tooth; cervical sul- NH, NJ, NM, NS, NY, ON, OR, PA, PE, PQ, cus fine, indistinct. Scutellum transverse SC, SD, SK, TX, UT, WA. (figure 264), densely setose; mesepi-meron Host plants.—Lathyrus sativus; Pisum narrowly reaching mesocoxa. Elytra to- sativum arvense, P. elatius; Vicia sp., Vicia gether 1.1 times as long as wide, depressed faba; Vigna radiata var. radiata. The Vicia or slightly sulcate along suture, laterally and Vigna records may be the result of convex; striae deep, narrow, subparallel, misidentifications. lacking basal denticles; interstices coarsely but sparsely foveolate and imbricate; hu- Natural enemies (Old World and New meri coarsely imbricate; apex of mesotibia World).—Anisopteromalus calandrae; Apro- acuminate (figure 266); metacoxae densely, stocetus aethiops, A. claviger; Bruchobius finely punctate; hind leg as in figure 47; laticeps; Bruchocida orientalis; Dinarmus ventrolateral margin with long, tapered laticeps, D. magnus; Eupelmus amicus; denticle; ventromesal margin with small, Eurytoma sp.; Gastrancistrus undulatus; blunt denticle; metatibia slender, slightly Microdontomerus anthonomi; Pteromalus dilated toward apex, lateral, ventral, and leguminis, P. micans, P. sequester; Py- dorsomesal carinae distinct; mucro short, emotes ventricosus; Staphylinus nigrellus; equal to lateral denticle, three minute coro- Systasis encyrtoides; Tetrastichus claviger, nal denticles. Abdomen not modified, male T. nerio; Triaspis gibberosus, T. thoracica; 5th ventral slightly emarginate; pygidial Trichomalopsis leguminis; Uscana semifu- disk (figure 267) densely foveolate, on male mipennis, U. senex. apex truncate exposing apex of 8th tergum, Immatures.—Riley 1892 (1st larval instar); on female apex evenly rounded. Middle Pfaffenberger 1977 (final larval instar), tibia of male with short, curved hook. 1991:564 (larva). Male genitalia.—As in figures 268 and 269; Discussion.—Bruchus pisorum is one of median lobe slender, cucullus scarcely the most widely distributed species in the widened; ventral valve subtriangular, world. Old World literature is voluminous strongly curved; dorsal valve evenly round- on its control (see Luk’yanovich and Ter- ed; armature of internal sac consisting of Minassian 1957). The species is most de- double row of fine spicules at apical orifice, structive to the common varieties of Pisum an elongate mass of fine, acute spicules in sativum and usually emerges in storage. middle, a pair of curvate masses of spic- It does not, however, reinfest stored pea ules at apex enclosing a large patch of seeds as does Acanthoscelides obtectus and imbricate, scalelike setae; apical closure some other species. Because it must ovi- valve membranous; lateral lobes elongate, posit on green pods in the field, it has only slender, cleft nearly to base, apices strong- one generation each year. ly bowed, finely setose; male 8th sternite Larson et al. (1938) and Pajni (1981) de- reduced to paired lobes, apex of 8th tergite tailed the life history of this species. Smith bluntly rounded, spiculum gastrale (fig- and Brower (1974:322) recorded the karyo- ure 270) T-shaped; female 8th tergite (t8) type. Back (1940:9) and Brindley et al. sinuately lobed; 8th sternite (s8) as shown (1952) discussed field infestations. in figure 271. The common name for this bruchid is “pea Size.—Body length 3.9–4.9 mm; width weevil.” 2.3–2.8 mm. Type depository.—Uncertain. Possibly destroyed.

73 Bruchus rufimanus (Boheman) glabrous boss, each foveola round and setate; frontal carina absent; ocular index (Figures 272–280) 3:1; ocular sinus four-fifths as long as eye; Bruchus rufimanus Boheman 1833:58; antenna slightly eccentric from 5th seg- Horn 1873:315; Hamilton 1892b:162; ment, terminal segment ovate, reaching Pic 1913a:45; Campbell 1920:1; Kan- to metacoxal border. Pronotum trapezoi- nan 1923:28; Essig 1929a:485; Back dal (figure 273), lateral margins sinuate, 1940:7; Vayssière and Lepesme 1941; with denticle at midpoint of margin, apex Decelle 1975:115; Pfaffenberger truncate; disk convex, slightly depressed 1977:133. Many other references can near posterior corners, basal lobe with be found in Luk’yanovich and Ter- triangular sulcus; surface densely, ir- Minassian 1957:100 and Udayagiri and regularly punctate; lateral carina absent; Wadhi 1989:200. cervical sulcus hidden in vestiture. Scutel- Laria rufimana: Chittenden 1912a:59. lum quadrate, emarginate. Elytra slightly Mylabris rufimana: Baudi 1886a:14. longer than wide, moderately convex; striae deep, narrow, subparallel, slightly sinuate Mylabris rufimana var. velutina: Baudi toward base; interstices flat, punctate-im- 1886a:31. bricate in basal one-third, densely punctu- Mylabris rufimanus: Leng 1920:305; Essig late in apical two-thirds, humeri coarsely 1929a:485. imbricate; metacoxal face densely, finely Bruchus granarius, of authors (not Lin- punctate, fossula glabrous; hind leg as in naeus 1767). figure 274; ventrolateral margin with angu- Bruchus fabae Motschulsky 1854:6. late subapical denticle, ventromesal margin with minute subapical denticle; meso- Bruchus rufimanus var. velutinus Mulsant tibia straight, slightly dilated, ventral and and Rey 1858:27. dorsomesal carinae distinct, lateral carina Bruchus rufimanus var. diversipubens Pic intermittent, ventrolateral carina absent; 1930b:14. mucro one-fifth as long as basitarsus; male Bruchus rufimanus var. rufimanus mesofemur strongly dilated and sulcate Luk’yanovich and Ter-Minassian (figure 276), ventral margin of mesotibia 1957:100. sinuate, apex with small, angulate hook. Color.—Body and appendages black except Fifth abdominal sternum shallowly emar- four basal antennal segments and fore ginate in male, straight in female; pygidi- legs reddish orange. Vestiture variable; um of both sexes convex (figure 275), apex head and pronotum usually with yellow- of male truncate with apex of 8th tergite ish-brown vestiture, lateral pronotal mar- visible, apex of female evenly arcuate. gin and discal spots white, basal lobe with Male genitalia.—As in figures 277 and 278; white, triangular patch; scutellum white; median lobe elongate, apical one-eighth elytra varying from pattern of white spots slightly expanded, cucullus scarcely ex- on black background with short, yellowish- panded; ventral valve broadly ogival, apex brown sutural stripe (figure 272) to inter- narrowly acute; dorsal valve not differenti- mixed yellowish brown, black, and white, ated; armature of internal sac consisting with variably distributed white spots, an of convergent pair of fine spicules at apical occasional specimen with elytra entirely orifice, a mass of fine, acute spicules in white; pygidium varying from white to basal one-half, imbricate scales and scale- brownish yellow, two subapical black spots like setae in apical one-half, a curved, den- bordered with brown; venter of body white, tate sclerite near apex; closure valve undif- usually with some yellowish brown on met- ferentiated; lateral lobes short, cleft one- episternum. half their length, apices strongly incurved; Structure.—Vertex and frons densely fo- male 8th sternite bilobed with apex narrow veolate in circular pattern around median,

74 and truncate (figure 279), spiculum gas- Tribe Bruchidiini Bridwell trale Y-shaped; female 8th tergite narrowly thickened (figure 280) on apical margin; Genus Borowiecius Anton 8th sternite as shown in figure 280 (S8). Borowiecius Anton 1994:108. Type spe- Size.—Body length 3.1–4.4 mm; width cies: Bruchus ademptus Sharp 1886, by 2.2–2.6 mm. original designation. Type depository.—NHRS. Small beetles, 1.1–3.0 mm long. One spe- Type locality.— Mediterranean. cies in southeastern United States; origin eastern Asia. Distribution.—CA, HI, LA, MB, NH, ON, PQ, SK. Head with prominent frontal carina; eyes and antennae not sexually dimorphic; an- Host plants.—Vicia faba and V. sativa tenna subserrate; pronotum bell-shaped, sativa in the United States. Twenty other disk evenly convex except shallowly sul- legumes have been reported as hosts in the cate on basal lobe and slightly depressed Old World, but many of these are question- along basal margin; lateral carina absent; able. prosternum short, triangular, not dividing Natural enemies (Old World and New coxae; scutellum short, quadrate; elytra World).—Bruchocida orientalis; Charito- ovate, striae sharply defined, 2nd and 3rd podinus swezeyi, C. terryi; Choetospila ending basally in dentate gibbosity; ventral elegans; Chremylus elaphus; Dinarmus margin of metafemur sulcate with single acutus, D. laticeps, D. magnus; Eurytoma acute ventromesal denticle, ventrolateral watchli; Lariophagus distinguendus; Py- carina with shallow sinus; male pygidium emotes tritici, P. ventricosus; Triaspis for- reflexed at apex; female pygidium vertical. besii, T. gibberosus, T. luteipes, T. pallipes, Male genitalia broad; internal sac with T. similis, T. stictostiba, T. thoracica; Zelus apparent hinge sclerites on either side of renardii. apical orifice; basal strut of tegmen ring Immatures.—Chittenden 1912a (egg, 1st lacking ventral keel; lateral lobes slender, larval instar); Pfaffenberger 1977 (final lar- deeply cleft. val instar), 1991:564 (larva). This genus closely resembles Calloso-bru- Discussion.—This species is similar in size chus, especially in the configuration of and general appearance to Bruchus piso- the hind leg and the basal elytral gibbos- rum, but the lateral denticle of the meta- ity found in some Callosobruchus, but the femur is much smaller than in B. pisorum internal sac of species of Callosobruchus is (compare figures 47 and 274) and the armed with burr-like sclerites, the number black subapical spots on the pygidium are and shape being characteristic of the spe- smaller and less pronounced than in B. pi- cies, whereas the internal sac in species of sorum. Both species are much larger than Borowiecius is devoid of such sclerites. In B. brachialis. contrast to the species of Callosobruchus found in the United States and Canada, The swollen mesofemur and sinuate meso- which have reddish brown or reddish yel- tibia of the male is a good key character. low hind legs at most clouded with darker Campbell (1920) investigated the biology areas, the of B. rufimanus, and Smith and Brower hind legs of B. ademptus are entirely black, (1974:322) recorded the karyotype profile. except that the tarsi are yellow. Back (1940:11) discussed field infestations. The common name for this bruchid is Borowiecius ademptus (Sharp) “broadbean weevil. (Figures 281–286) Bruchus ademptus Sharp 1886:36; Pic 1913a:13.

75 Callosobruchus ademptus: Chujo 1937a:35; Male genitalia.—As in figures 285 and 286; Bridwell 1938a:75; Bottimer 1961:295; median lobe 5 times as long as wide; ven- Kingsolver 1969a:7; Johnson and King- tral valve broad, semicircular, apex slightly solver 1982:412; Udayagiri and Wadhi produced, middle of internal sac with coni- 1989:186; Morimoto 1990:132. cal mass of elongate spicules, apical one- Borowiecius ademptus: Anton 1994:108. half of sac lined with fine, elongate spic- Color.—Body black except fore legs and mid ules, apex with ring. Lateral lobes simple, legs, antenna and apical three or four ab- elongate, expanded apically. dominal segments yellow, hind legs black Size—Body length 2.5–2.9 mm; width except tarsi yellow. Vestiture of black, 1.6–1.9 mm. white, and yellow hairs in pattern shown Type depository.—BMNH. (figure 281). Head sparsely yellow; pronotum yellow on flanks and basal lobe, other- Type locality.—Japan, Yuyama and Naga- wise black; scutellum white; elytra with su- saki. tural interstices yellow, 3rd interstice with Distribution.—AL, DC, MD, NC; Japan, Ko- elongate, white basal, median, and apical rea, China. spots, lateral interstices with alternating Host plants.—Pueraria montana lobata white and yellow fragmentary stripes and (kudzu). spots; pygidium uniformly yellow except for basal triangular patch and subapical Natural enemies.—None recorded. paired bare spots. Immatures.—Not described. Structure.—Vertex, frons, and clypeus Discussion.—This species was formerly coarsely, densely punctate; frontal carina assigned to Callosobruchus because of the prominent; ocular index 2.5:1; ocular si- structure of the hind femur, which bears a nus one-half length of eye; antenna (figure denticle on the ventromesal margin as well 282) not dimorphic, reaching middle of me- as a blunt angulation on the lateroventral tepisternum. Pronotum subconical (figure margin. The male genitalia, however, are 281), convex, densely foveolate; basal lobe broad, not long and slender as in Callo- slightly domed; lateral carina obscured by sobruchus, and the internal sac lacks the vestiture. Scutellum appearing bilobed. burrlike sclerites characteristic of Callo- Elytra about as long as wide, evenly con- sobruchus. The dorsal pattern resembles vex; striae subparallel, slightly sinuate, 3rd that of many Bruchidius, and the late B.J. and 4th deflected laterad basally but not Southgate (personal communication, 1978) reaching basal margin; interstices densely expressed the opinion that ademptus imbricate; metacoxal face densely fossulate would be better placed in that genus rather except fossula smooth; metafemur with than in Callobruchus. ventrolateral margin carinate, with subapical angulation (figure 283), ventromesal Genus Bruchidius Schilsky margin carinate, with single sharp denticle as long as the ventrolateral angulation; Bruchidius Schilsky 1905:B; American metatibia with lateral carina prominent references: Bridwell 1932:104; Hoff- ending in sharp denticle, ventrolateral man 1945:61; Bottimer 1968a:139, carina sometimes reaching base of mucro, 1968c:1028; Udayagiri and Wadhi ventral and dorsomesal carinae complete; 1989:114. Many additional references mucro short, less than one-sixth length of to the genus can be found in the Eu- basitarsus. Pygidium not modified, apex of ropean literature: Hoffman 1945; male reflexed into notch in 5th abdominal, Luk’yanovich and Ter-Minassian 1957; disk sculpture hidden by vestiture (figure Borowiec 1987, 1988; Udayagiri and 284). Wadhi 1989. Type species: Bruchus

76 quinqueguttatus Olivier 1795, by subse- Bruchidius is comparable to the New World quent designation Bridwell 1932. Acanthoscelides in numbers of described Sparteus Bridwell 1946:55. Type species: species and complexity and needs a thor- Bruchus villosus Fabricius, by original ough monographic treatment. designation. Bruchidius villosus could be difficult to Spermophagus: Arnett 1962:957 separate from melanistic forms of Sennius, (error). especially Sennius cruentatus, in the east- Small bruchids (1.4–2.8 mm). Two species ern United States unless male genitalia are have been introduced into the North Amer- dissected. An important character marking ican fauna. Sennius is the presence of “hinge sclerites” at the apical orifice of the male genitalia. Body and appendages entirely black or piceous, basal segments of antenna some- Bruchidius cisti (Fabricius) times reddish yellow. Vestiture white, sparse. (Figures 287–291) Frontal carina lacking or briefly indicated; Bruchus cisti Fabricius 1775:65. eyes deeply emarginate; antenna only Bruchidius cisti: Southgate 1963:796; slightly eccentric. Pronotum convex, punc- Decelle & Lodos 1989:182. tate, lacking lateral carina. Elytra with stri- Bruchus unicolor Olivier 1795:7. ae parallel, with or without basal denticles. Bruchidius unicolor var. debilis: Schilsky Metafemur with single, minute subapical 1905:41. denticle; mucro short or lacking. Bruchidius unicolor: Bottimer 1968a:143; Median lobe of male genitalia lacking hinge Hewitt and Burleson 1976. sclerites. Bruchus canus Germar 1824:183. This genus consists of about 300 described Bruchus debilis Gyllenhal 1833:53. species and is widespread in the Old World. It is characterized principally by the Bruchidius villosus var. pubescens: minute single denticle on the ventromesal Luk’yanovich and Ter-Minassian margin, absence of denticles on the ventro- 1957:143. lateral margin of the metafemur, and lack Bruchus ater Curtis 1839: vol. 16, p. 745. of lateral marginal denticles on the pro- Bruchus ater var. pubescens: Jacquet notum. These characters are also descrip- 1888:19. tive of several other genera (for instance, Bruchus autumnalis Motschulsky Sennius). 1874:226. Bruchus nugax Motschulsky 1874:226. Bruchus pulverulentus Motschulsky Key to North American Bruchidius 1874:218. Bruchus rostratus Motschulsky 1874:232. [Adapted from Bottimer 1968a] Bruchus tibiellus Stephens 1829:266. 1 Body and appendages entirely black; 4th stria with Bruchus villosus var. pubescens: Pic prominent, subbasal denticle (figure 287); mucro absent; 1913a:56. western United States and Canada...... cisti (Fabricius) Mylabris villosa var. pubescens: Baudi Body black with basal four antennal segments reddish 1886a:100. yellow, legs sometimes piceous; striae lacking denticles Spermophagus cisti: Schilsky 1905:3. (figure 292); mucro one-eighth as long as basitarsus; See Aldridge and Pope (1986:182) for eastern United States and Canada.....villosus (Fabricius) additional discussions of the taxonomic history of this species.

77 Color.—Body and appendages entirely Size.—Body length 2.3–3.4 mm; width black (figure 287). Vestiture white with 1.4–1.9 mm. golden sheen on dorsal surfaces, white on Type depository.—BMNH (cisti, lectotype ventral areas, pygidium with evanescent by Aldridge and Pope 1986:183); Halle white basal triangle. (canus, lectotype by Zampetti 1981:404); Structure.—Vertex and frons densely unknown (unicolor and debilis). punctulate, punctures crowded, coales- Type locality.—Great Britain. cent, frontal carina evanescent or absent; ocular index 7:2; ocular sinus seven- Distribution.—BC, MT; Europe, Asia. eighths length of eye; antenna extending to Host plants.—Cynoglossum sp.; Onobry- humerus, segments only slightly eccentric. chis viciifolia; Spartium junceum. Pronotum subconical (figure 287), apex Natural enemies.—Dynarmus acutus; rounded, basal margin sinuate, lateral Pteromalus sequester; Pyemotes tritici; Us- margins straight; disk evenly convex, cana semifumipennis, U. senex. faintly sulcate on basal lobe, basal margin impressed either side of basal lobe; surface Immatures.—Not described. densely, irregularly foveolate-punctate, Discussion.—Because of its totally black foveolae crowded, coalescent; lateral ca- integument and restricted range in the rina absent; cervical sulcus distinct, deep. United States, this species should pres- Scutellum quadrate, emarginate. Elytra 1.2 ent no problem in identification. The lack times as long as wide, evenly convex ex- of a substantial mucro and the prominent cept depressed in middle of basal margin; denticle at the base of the 4th stria will striae deep, narrow, slightly sinuate; 4th separate it from Bruchidius villosus. stria with prominent basal denticle; interstices flat, minutely imbricate; metacoxal Bruchidius cisti was first reported in the face densely punctate except fossula gla- United States (Montana) in 1976 (Hewitt brous; metafemur with minute denticle on and Burleson, as Bruchidius unicolor). ventromesal margin (figure 288); metatibia nearly straight, slightly dilated toward Bruchidius villosus (Fabricius) apex; lateral carina usually present, some- (Figures 292–296) times intermittent; dorsomesal carina fine; Bruchus villosus Fabricius 1792:373. ventrolateral and ventral carinae absent; mucro minute, lateral denticle and coronal Sparteus villosus: Bridwell 1946:55. denticles minute, subequal in size. First Bruchidius villosus: Zacher 1952:464. abdominal segment of male flattened, bear- Frick 1962; Southgate 1963:795. ing long, silky setae; 5th sternum deeply Bruchus cisti sensu Paykull 1800 (not Fa- emarginate; pygidium of male strongly re- bricius 1775). flexed, that of female oblique; disk densely Bruchus ater Marsham 1802:236. punctulate and setose (figure 289). Bruchidius ater: Southgate 1963:795; Par- Male genitalia.—As in figures 290 and 291; nell 1964:73; De Luca 1967b; Bottimer median lobe slightly expanded apically 1968a:142; Frankenhuyzen and Per- (figure 290), apex of dorsal valve rounded, quin 1971, 1972. fringed with fine setae; ventral valve trian- Bruchus fasciatus: Steffan 1946 and oth- gular, strongly arched in lateral aspect; ar- ers (not Olivier 1795). mature of internal sac consisting of paired, spiny, ovoid sclerites near base of sac with Spermophagus fasciatus (Olivier) of Arnett remainder of sac elongate, densely lined 1962:957. with small, thornlike denticles; lateral Mylabris villosus: Baudi 1886a:44. lobes as in figure 291, deeply cleft. Color.—Body and appendages black except antennal segments 1–4 reddish brown.

78 Vestiture uniformly white with golden Type locality.—Germany (villosus, lecto- sheen on dorsal surfaces, white on ventral type); Great Britain (ater, lectotype). surfaces. Distribution.—DE, MA, MD, NJ, NY, ON, Structure.—Vertex and frons densely punct- VA; Europe. ulate; frontal carina short, not prominent; Host plants.—Cytisus scoparius; Laburnum ocular index 2.5:1; ocular sinus four-fifths alpinum, L. anagyroides; Petteria ramen-ta- as long as eye; antennal segments slightly cea; Spartium junceum. Several other host eccentric from 5th segment, extending to plants are recorded from Europe (Frick humerus. Pronotum bell-shaped (figure 1962). Adults feed in flowers ofCytisus sco- 292), lateral margins straight except con- parius. stricted at apex by cervical sulcus; disk convex, basal lobe shallowly sulcate, basal Natural enemies.—Dinarmus acutus; margin impressed either side of basal lobe; Pteromalus sequester; Torymus persicariae; surface irregularly punctulate-foveolate; Triaspis sp.; Trichogramma sp. lateral carina absent; cervical sulcus dis- Immatures.—Frick 1962 (oviposition, eggs); tinct, extending across prosternum. Scu- Parnell 1964 (all instars—larvae, prepupa, tellum subquadrate, emarginate. Elytra pupa; illustrated). together slightly longer than wide (figure Discussion.—Bruchidius villosus has ap- 292), convex, lightly impressed around peared in literature under several names, scutellum; striae narrow, deep, subparal- and Aldridge and Pope (1986) have sum- lel, 3rd, 4th, and 5th each ending in deep marized the complexities. Southgate (1963) basal pit, the rim of which is sometimes concluded that Bruchidius ater was the extruded into blunt denticle; interstices proper name, and Bottimer (1968a) fol- densely imbricate; metacoxal face densely lowed his interpretation. punctulate, fossula glabrous; hind leg as in figure 294; ventro-mesal margin with one This Old World species was first reported minute, subapical denticle nearly hidden in the United States by Olsen (1918), who in pubescence; metatibia slightly dilated found it in Massachusetts. It was subse- toward apex, lateral and ventral carinae quently found in Virginia and several other sometimes evanescent; mucro short, acute, eastern states and provinces. See Bottimer one-tenth as long as basitarsus, lateral (1968a) for a discussion of early history of and coronal denticles subequal. Fifth its invasion. abdominal sternum broadly emarginate; Characters in the key to genera and key pygidium (figure 293) densely punctate and to Bruchidius species will separate it from pubescent. other species. Male genitalia.—As in figures 295 and 296; median lobe slender; dorsal valve membra- Genus Callosobruchus Pic nous; ventral valve broad at base, strongly Bruchus (Callosobruchus) Pic 1902:6. Type narrowed, apical portion subparallel; species: Curculio chinensis Linnaeus internal sac armed with round or truncate 1758, by subsequent designation, denticles in basal one-third, two parallel Bridwell 1929a. patches of elongate spicules at midpoint, and a large, single patch of spicules near Callosobruchus Pic 1912:92; Bridwell apex; lateral lobes slender, deeply cleft (fig- 1929a:40; Bottimer 1968c:1029,1042 ure 296). (history of usage of the name Calloso- bruchus). Size.—Body length 2.4–3.2 mm; width Small beetles, 1.1–3.0 mm long. Four spe- 1.5–2.2 mm. cies are found in the United States and Type depository.—UZMC (villosus); BMNH Canada. (ater). Integument black, red, or yellow.

79 Head with prominent frontal carina; eyes 310); metatibia with four complete carinae. and antenna sometimes dimorphic; anten- Male genitalia usually long, slender, lateral na subserrate (figure 300) to strongly pecti- lobes slender, deeply cleft. nate (figure 299); pronotum bell-shaped, disk usually medially sulcate, sometimes Callosobruchus differs from Bruchus prin- moderately gibbous; elytra with or without cipally by lack of a tooth on each lateral dentate basal gibbosity; striae distinct, margin of the pronotum (compare figures generally parallel; ventral margin of meta- 265 and 297) and lack of a hooklike ap- femur sulcate, ventrolateral and ventrome- pendage on the middle tibia of males. sal carinae dentate or angulate (figure

Key to U.S. Callosobruchus and at apex (figure 301), female with more intense patches sometimes connected to form paired crescents [Also see Haines (1989) for a key to Callosobruchus species] (figure 302); antennae as in figures 299 and 300...... 1 Elytral striae three and four each with prominent subbasal chinensis (Linnaeus) denticles on slight gibbosity (figures 281 and 297)...... 2 3(1) Pronotum yellowish with brown or black longitudinal Elytral striae extending to basal margin, lacking prominent lines (figure 321); elytral pattern as in figure 321; eye with denticles or gibbosity (figure 305)...... 3 narrow posterior fringe...... phaseoli (Gyllenhal) 2(1) Pattern of pronotum and elytra as in figure 327; pygidium Pronotum dark red to black, lacking longitudinal lines in both sexes only slightly convex, nearly immaculate (figures 305 and 306); elytral pattern variable (figures except for subapical, paired spots (figure 331); antennae 305–307); eye with posterior margin lobed . maculatus as in figures 328 and 329...... pulcher Pic (Fabricius) Pattern of pronotum and elytra as in figure 297; pygidium in both sexes convex, male with vague patches at middle

Callosobruchus chinensis (Linnaeus) Bruchus chinensis: Cushman 1911:505; Pic 1913a:20. (Figures 8, 9, 297–304) Bruchus pectinicornis Linnaeus 1767:605. Curculio chinensis Linnaeus 1758:386. Bruchus rufus De Geer 1775:281. Pachymerus chinensis: Schilsky 1905, Bruchus scutellaris Fabricius 1792:372. no. 99; Chittenden 1912b:1; Reitter 1912:226. Bruchus bistriatus Fabricius 1801:402. Mylabris chinensis: Harold 1878:86; Baudi Bruchus biguttatus Fabricius 1801:402. 1886b:45; Leng 1920:305. Bruchus biguttellus Schoenherr 1833:42. Callosobruchus chinensis: Bridwell Bruchus bistriotus: Bridwell 1929a:41 (er- 1918:467; Herford 1935:5; Chujo ror). 1937a:38; Back 1940:7; Vayssière Bruchus (Callosobruchus) chinensis: Pic and Lepesme 1941; Bottimer 1944:23; 1912:92. Lepesme 1944:209; Hoffman 1945:87; Bruchus elegans: Sturm et al. 1843:3222. Southgate 1958, 1964; Bottimer 1968c:1029; Howe 1973:572; Borowiec Bruchus rufobrunneus: Wollaston 1870:25. 1987:132; Furusawa 1987:388; Udaya- References are numerous for this species giri and Wadhi 1989:167. because of its wide distribution and im- Bruchus (Callosobruchus) chinensis: Zacher portance as a pest of leguminous grains. 1930:282. Selected references are listed under Dis-

80 cussion. Consult Luk’yanovich and Ter- except fossula polished. Male pygidium Minassian (1957:67) and Pic (1913a:20) for strongly reflexed nearly to apical margin of older literature pertaining to Calloso-bru- basisternum; hind leg as shown in figure chus chinensis. Synonyms listed above are 298; mucro one-fourth as long as basitar- extracted from Southgate (1958). sus. Color.—Integumental color red to black; Male genitalia.—As in figures 303 and 304; vertex red with broad median stripe black, median lobe elongate (figure 303), ventral frons black; antenna of male mostly black valve elongate, apically triangular, internal with two basal segments and dorsal face sac with two burrlike sclerites near apex; of one or more additional segments yel- lateral lobes elongate (figure 304), divided low, antenna of female dark red with distal nearly to base. segments gradually darkening. Pronotum Size.—Body length 2.7–3.0 mm; width black to red; if black, with red median line. 1.8–2.3 mm. Elytra red to black but usually with diagonal yellow band in basal one-half, only Type depository.—Not known, possibly faintly visible on red specimens. Venter of BMNH. body usually black, sometimes partly red. Type locality.—”China.” Male pygidium black; female usually yel- Distribution.—Tropicopolitan. HI, AL, CA, low laterally, red median stripe with four DC, FL, GA, HI, LA, MD, MS, OH, PA, SC, black or red spots sometimes connected VA. in curvate maculae (figure 302); fore legs and mid legs yellow, hind legs black with Host plants (Old World and New World).— red to yellow dorsal margin. Vestiture of Cajanus cajan; Cicer arietinum; Cyamopsis head yellow; of pronotum mostly yellow tetragonoloba; Glycine max; Lablab pur- but with small, white spot either side of pureus; Lathyrus sativus; Lens culinaris; midline, basal lobe with elongate bilobed Nelumbo nucifera; Phaseolus lunatus, P. spot of waxy, matted white and golden vulgaris; Pisum sativum; Psophocarpus setae (figure 297). Scutellum white. Elytra tetragonolobus; Vicia faba; Vigna aconiti- golden in basal one-half (figure 297), 3rd folia, V. angularis, V. mungo, V. radiata interstice with elongate, white patch and radiata, V. unguiculata unguiculata. All irregular, diagonal line of white patches at host plants are in the Leguminosae except apical one-third; male pygidium white with Nelumbo, which is in the Nelumbonaceae vague to strong dark apical and median (Kingsolver 1979b; Furusawa 1987). spots (figure 301). Venter of body white, Natural enemies (Old and New World).— condensed into spots on metepisternum Anisopteromalus calandrae; Bruchocida and lateral margins of abdominal sterna orientalis; Choetospila elegans; Dinarmus (figure 298). colemani, D. laticeps, D. vagabundus; Structure.—Frontal carina prominent; Heterospilus prosopidis; Lariophagus dis- ocular index 4:1; ocular sinus one-half tinguendus; Pteromalus sp., P. schwenkei; length of eye; male antenna (figure 299) Pyemotes tritici, P. ventricosus; Stenocorse pectinate, female antenna serrate (figure bruchivora; Urosigalphus bruchi; Uscana 300). Pronotum convex, basal lobe promi- mukerjii, U. semifumipennis. nent, medially sulcate, laterally bounded Immatures.—Wightman and Southgate by sulci; disk minutely punctate-reticulate; 1982 (egg); Pfaffenberger 1991:564 (larva). lateral carina lacking. Scutellum minute, appearing bilobate. Elytra together slightly Discussion.—Although this species is vari- longer than wide; striae deep, strial punc- able in the intensity of its color pattern, tures elongate, 3rd and 4th ending in basal the characters given in the key and de- gibbosity; interstices slightly convex, imbri- scription should separate it easily. Salient cate; metacoxa evenly punctate (figure 298) characters are the waxy white pad of setae on the pronotal basal lobe, the white patch

81 on the lateral margin of the abdomen, the Pawar and Verma 1977 (musculature); Ah- 3rd and 4th elytral striae each ending in mad and Murad 1980a. a basal gibbosity, and the pectinate male Malpighian tubules.—Rahman and Ameen antenna. 1990. Common names for this species are cow- Oviposition.—Srivistava and Bhatia 1959; pea weevil, gorgojo del caupi, Kundekäfer, Avidov et al. 1965; Nakamura 1968; Os- mung bean bruchid, tonchio chinese, hima et al. 1973; Bhattacharya et al. 1977; chinesischer Bohnenkäfer, and la bruche Tikku et al. 1978; Gokhale et al. 1990; chinoise. Yamamoto 1990. Callosobruchus chinensis, like its congener Parasites.—See “Natural Enemies” above; C. maculatus, has been the laboratory ani- Utida 1961; Roomi et al. 1973; Fujii and mal for a multitude of research projects. Khin Mar Wai 1990. Selected references pertaining to various Pheromones.—Tanaka et al. 1981, 1982; Qi aspects of biology and morphology are and Burkholder 1985. listed below: Sexual dimorphism.—Shukla and Pandey Adult feeding.—Shinoda and Yoshida 1984, 1977. 1987a. Spermatheca.—Surtees 1961. Adult illustration.—Hoffman 1945:87; Temperature effect.—Choudhuri and Paul Luk’yanovich and Ter-Minassian 1957; 1984. Kingsolver 1969a; Arora 1977. Head morphology.—Singh 1981c. Callosobruchus maculatus (Fabricius) Chromosomes.—Smith and Brower 1974. (Figures 48, 50, 305–320) Digestion.—Choudhuri and Paul 1983. Bruchus maculatus Fabricius 1775:65 Digestive tract.—Mansour 1934. Callosobruchus maculatus: Bridwell Dimorphism.—George and Verma 1997. 1929a:40; Herford 1935:5; Larson Eclosion.—Southgate 1984. and Fisher 1938:5; Back 1940:7; Bis- sell 1940:846; Vayssière and Lepesme Egg morphology.—Howe and Currie 1964; 1941; Hoffman 1945:89; Blackwelder Wightman and Southgate 1982. 1946:761; Utida 1954:161; Southgate Host resistance.—Fernandez and Talekar et al. 1957:79; Southgate 1958:591, 1990; Kitamura et al. 1990. 1964:277; Howe 1973:572; Borow- Identification.—Vats 1974a,b. iec 1988:175; Udayagiri and Wadhi Larval development.—Podoler and Apple- 1989:169. baum 1971; Rahman and Ameen 1986. Bruchus quadrimaculatus Fabricius Larval illustration.—Arora 1978:4; Begum 1792:371; Olivier 1795:19; Horn et al. 1982. 1873:317; Sharp 1885:478; Blatchley 1910:1237; Cushman 1911:493; Pic Life history.—Back 1940:7; Lepesme 1944; 1913a:44; Larson and Fisher 1924:297; Sawaf 1956; Howe and Currie 1964; Fu- Brauer 1942; Lepesme 1944:212. jii 1965; Applebaum et al. 1968; Raina 1970; Arora and Singh 1971; Okamoto Mylabris quadrimaculatus: Baudi 1887:36; 1971; Bato and Sanchez 1972; Begum et Leng 1920:305. al. 1978; Singh et al. 1980; Singh 1981b; Pachymerus quadrimaculatus: Schilsky Giga and Smith 1983; Bellows and Hassell 1905:MM1. 1984; Shinoda and Yoshida 1987b; Yoshi- Bruchus ornatus Boheman 1829:103; da 1989. Baudi 1886b:47. Male genitalia.—Mukerji and Chatterjee Bruchus vicinus Gyllenhal 1833:36. 1951; Kingsolver 1969a:13; Arora 1977:28; Bruchus litteratus Schoenherr 1833:102.

82 Bruchus ambiguus Gyllenhal 1839:11; Fall bursae couplatrices (figures 319 and 320) 1910:163. are consistently diagnostic (Spirina 1974). Bruchus sinuatus Fahraeus 1839:8. The flightless form, having normal wings, Bruchus arachidis Fahraeus 1839:10. is capable of flight but prefers to remain in Laria quadrimaculata: Bedel 1901:349 on the substrate of seeds in relative dark- ness. A population buildup with the ensu- Pachymerus ornatus: Schilsky 1905:MM. ing crowded conditions in a granary situa- Bruchus (Acanthoscelides) trabuti Caillol tion can, due to metabolic overheating by 1919:236. larval activity, trigger the development of Callosobruchus ornatus: Hoffman 1945:90; the flight form that readily flies and seeks Zacher 1952:466. light. Females of the flight form are lighter Callosobruchus sinuatus: Zacher 1952:466. in weight, their ovaries are poorly developed, and their abdomens are filled with Callosobruchus quadrimaculatus: Shomar fat bodies; also, they develop few fertile 1963:182. eggs. Testes of the flight males are imma- Acanthoscelides sinuatus: Blackwelder ture at emergence but gradually develop as 1946:761. the individual ages (Utida 1981). For fur- Southgate et al. (1957) presented a thor- ther information, see Utida (1981) and his ough discussion of the synonymy of Callo- included bibliography. sobruchus maculatus, and there is no need Illustrating the many variants in color to repeat it here. pattern would require more space than is Color and Structure here justified. The following descriptions Callosobruchus maculatus is variable in and associated illustrations can give only a coloration, leading to confusion with other hint of the variety. Other references to this species of Callosobruchus. Part of the phenomenon are listed in the discussion variation is due to the presence of “two below. distinct adult forms, which differ not only in their morphology, but in physiology, Flightless or normal form: behaviour, etc.” (Utida 1981). Utida (1954) Male (figures 305 and 306).—Integument named these “flightless form” and “flight mostly black with red areas on abdomen, form,” which Caswell (1960) later termed legs, and elytra; head black, red spot be- “normal” and “active,” respectively. Utida hind eye; antenna mostly dark red (fig- (1981) discusses at length the basis of the ure 309); pronotum piceous with reddish two forms. Striking differences in the male margins; elytra red with small maculae on genitalia of the two forms were illustrated humerus and lateral margin; hind leg as by Spirina (1974). Other references to this in figure 310; pygidium black with reddish phenomenon are listed in the discussion suffusion (figure 311); legs reddish yel- below. low. Vestiture yellowish on dorsal surfaces, Utida (1972) illustrated male and female venter white; densely white patch on basal patterns he claimed would distinguish lobe of pronotum, faint median line on the two forms. In my own examinations pygidium. of several long series of both the flight- Male genitalia as in figures 315 and 316; less and flight forms, however, color pat- median lobe parallel-sided; ventral valve terns were so variable as to be unreliable triangular, apex rounded, bent ventrad; for distinguishing the forms, although the internal sac armature consisting of a patterns were moderately consistent within small mass of spicules at apical orifice, each series. Characteristics of both male two lunate masses of scalelike, flat spines genitalia (figures 315 and 317) and female continuous with two dense masses of large spicules; apical closure valve not well

83 defined; lateral lobes slender, parallel-sid- portions of elytra, and pygidium yellow, ed, cleft to base, apices each with membra- darker portions of elytra black, border of nous lobe and mesally directed setose pad. lateral maculae not prominent; 3rd inter- Female.—Head red to black, red spot be- stice with elongate white stripe,; venter hind eye, antenna red to piceous (figure white. 309); pronotum as in male except with Bursa copulatrix as in figure 320. median line; elytra red with basal border, Size.—Body length: males 2.7–3.1 mm; large lateral and apical maculae piceous or females 3.3–3.8 mm. Width: males 1.5–1.9 black; pygidium black; venter red to black; mm; females 1.5–1.9 mm. legs mostly red with some darker suffusion. Vestiture yellow on head, pronotum Type depository.—UZMC (maculatus, black with yellow to white spot on basal quadrimaculatus); NHRS (ornatus, littera- lobe and on flanks; elytra black on darker tus, vicinus, ambiguus, sinuatus); MNHP maculae, white bordering maculae, brown (tabuti); NHRS (arachidis). along suture; pygidium black with white Type locality.—”America” (ambiguus, macu- median stripe, or yellow with apical spots. latus, sinuatus); Senegal (vicinus); Sierra Venter and legs white. Leone (ornatus); Timbuctu (trabuti); Brazil Bursa copulatrix as in figure 319. and Greece (arachidis); “North, Central, and South America” (quadrimaculatus). Flight or abnormal form: Distribution.—Cosmopolitan. U.S. and Ca- Male (figure 307).—Similar to flightless nadian records: AK, AR, AZ, CA, DC, FL, form except basic integumental color of GA, HI, IA, ID, IN, KS, LA, MD, MO, MS, venter darker, distal seven or eight anten- NC, NM, NY, OH, OK, ON, PA, SC, SD, SK, nal segments piceous; hind legs piceous TN, TX, VA. Southgate (1964) reasoned with dorsal margin brownish yellow, fore that the most likely region of origin was on legs and mid legs yellow; elytral maculae the African continent. larger than in flightless form; pygidium as in figure 312. Host plants.—Cajanus cajan; Cicer arietinum; Glycine max; Lablab purpureus; Male genitalia as in figures 317 and 318; Lathyrus aphaca, L. clymenum, L. sativus; median lobe slender, parallel-sided; ventral Lens culinaris; Phaseolus acutifolius, P. valve triangular, apex slightly produced; lunatus, P. vulgaris; Pisum sativum arvense; armature of internal sac consisting of di- Vicia faba, V. lutea, V. sativa sativa; Vigna vided mass of spicules connected by trans- mungo, V. radiata radiata, V. unguiculata verse sclerite at apical orifice, two slen- unguiculata. der masses of mixed, minute scales and spines, two elongate masses of slender, Natural enemies.—Anisopteromalus ca- thornlike spines, two semicircular thorny landrae, A. pratti; Bruchobius laticeps; sclerites, and two clusters of spines near Bruchocida vuilleti (Terrasse and Rojas- apex; closure valve circular; lateral lobes Rousse 1986, bionomics); Cephanolo- as for normal form. mia gallicola; Chaetostricha mukerjii; Choeto-spila elegans; Dinarmus laticeps, Female.—Head black, antennal color as in D. vagabundus; Heterospilus prosopidis; male, pronotum black; elytra dark red ex- Lariophagus distinguendus, L. texanus; Oe- cept basal marginal spots, lateral and api- daule magnus; Pyemotes ventricosus; Sen- cal maculae black; pygidium dark red with egalella acythopoensis; Uscana lariophaga, lateral maculae black; venter black except U. marilandica, U. mukerjii, U. semifumipen- abdominal sutures red; fore legs and mid nis, U. senex. legs dark red, hind legs usually piceous, sometimes with reddish suffusion. Vesti- Immatures.—Larson and Fisher 1938:23– ture of head, pronotum, dark red 27 (larva and description of egg); Mukerji 1938 (1st larval instar); Lepesme 1944

84 (larva); Prevett 1971 (larva); Vats 1974a,b Egg morphology.—Wightman and South- (larva); Arora 1978 (larva); Wightman gate 1982. and Southgate 1982 (egg); Pfaffenberger Egg development.—Kannan 1923; Brauer 1991:564 (larva). 1925, 1942, 1946; Tantawi et al. 1976; Discussion.—This cosmopolitan species is Miyamoto and Van Der Meer 1982. one of the best known and most univer- Enzymes.—Gatehouse and Anstee 1983; sally destructive of the Bruchidae. Like its Gatehouse and Boulter 1983; Gatehouse et sister species, Callosobruchus chinensis, it al. 1985; Kitch and Murdock 1986. has the ability to infest and reinfest many Eye.—Schmitt et al. 1982. species of stored legumes, resulting in Female genitalia.—Mukerji and Bhuya enormous post-harvest losses. In recent 1937; Pajni 1968b; Monga 1972; Spirina years, much research has focused on the 1974; Monga and Sareen 1980 (ovaries). morphological and biological aspects of C. maculatus, especially in a laboratory at Genetics.—Breitenbacher 1925; Wasser- Tours, France. The species has proven to man and Futuyma 1981; Messina 1990; be an important laboratory animal in ge- Taper 1990. netic studies. Hormones.—Tantawi et al. 1976. The common name for this bruchid is Host plants.—Kingsolver 1979b; Staneva “cowpea weevil.” 1982. Identifying characters are found in the key. Host resistance.—Janzen et al. 1976; Gate- house et al. 1979; Dobie 1981; Gatehouse Selected references pertaining to various and Boulter 1983; Brewer and Horber aspects of biology and morphology are 1984; Birch et al. 1985; Kitamura et al. listed below: 1990. Adult emergence.—Yadav 1977; Highland Larval development.—Arora and Pajni 1986 (package penetration). 1959; Fujii 1965; Mookherjee and Chawla Adult illustration.—Larson and Fisher 1965; Satija and Kaur 1967a,b; Pajni 1938; Johnson 1983a. 1968b,c; Satija and Kaur 1968; Yadav and Larval alimentary canal.—Vats 1976b. Pant 1975; Janzen 1977b; Osuji 1980; Aldana Alfonso 1983; Sano-Fujii 1984; Antenna.—Rup 1988. Boughdad et al. 1987; Credland and Dick Body chemistry.—Nwanze et al. 1976; 1987. Sharma et al. 1983; Sidhu et al. 1984; Larval illustration.—Larson and Fisher Wasserman 1987; Gatehouse et al. 1990. 1938; Vats 1974b; Arora 1978. Chromosomes.—Smith and Brower 1974. Larval malpighian tubules.—Vats 1976a. Digestive tract.—Neelgund and Kumari Larval penetration of seeds.—Nwanze and 1983; Vats 1976b. Horber 1976; Janzen 1977b. Dimorphism.—Utida 1954; Southgate et Larval salivary glands.—Pajni 1965. al. 1957; Arora and Pajni 1959; Caswell 1960; Arora et al. 1967; Sano 1967; Utida Life history.—Chittenden 1912b; Paddock 1969; Gill et al. 1971; Bawa et al. 1972; and Reinhard 1919; Wade 1919; Larson Utida 1972; Bawa et al. 1974; Taylor 1974; and Simmons 1923, 1924a,b; Larson and Taylor and Agbaje 1974; Taylor and Aludo Fisher 1924; Back 1940:7; Schoof 1941; 1974; Utida 1974; Staneva 1980; Utida Lepesme 1944; Sawaf 1956; Mookherjee 1981; Nwanze and Horber 1975; Staneva and Chawla 1965; Raina 1970; Chok- 1982; Sano-Fujii 1984, 1986; Messina ouhian 1973; Ahmed et al. 1978; Saplina 1987. 1980; Singh et al. 1980; Giga and Smith 1983; Staneva 1983; Roche et al. 1985; Eclosion.—Mukerji 1938. Credland et al. 1986; Germain et al. 1987;

85 Messina 1990; Mitchell 1990; Moller et al. Blackwelder 1946:761; Luk’yanovich 1990. and Ter-Minassian 1957:67; Bot- Male genitalia.—Mukerji and Bhuya 1937; timer 1961:296, 1968c:1032; Decelle Pajni 1968a; Arora 1971; Gill et al. 1971 1981:195; Wightman and Southgate (testes); Spirina 1974; Taylor and Agbaje 1982:95; Borowiec 1988:178; Udayagiri 1974; Taylor and Aludo 1974; Ahmed et al. and Wadhi 1989:171. 1976; Pawar and Verma 1977 (muscula- Bruchus figuratus Gyllenhal 1839:12, Ne w ture); Kasap and Crowson 1979; Messina Sy n o n y m y . and Renwick 1985. Bruchus conicicollis Fairmaire 1898:247. Mating and reproduction.—Credland 1986; Color.—Integument reddish brown except Rup 1986. eyes black, variable black patches on elytra Oviposition.—Brauer 1944; Nakamura (figure 321), vertex sometimes black; legs 1968; Gokhale and Srivastava 1975; reddish brown; distal antennal segments Mitchell 1975; Monga and Sareen 1980; brown to piceous except apical segment Wasserman 1981; Dick and Credland red. Vestiture of yellow, medium brown, 1984; Gupta and Bhaduri 1984; Messina dark brown, and white hairs; head yellow, 1984; Singh et al. 1984; Giga and Smith pronotum mostly yellow with dark brown 1985; Messina 1985; Wasserman 1985; paired stripes and white basal lobe; elytra Messina et al. 1987a,b; Ofuya 1987; Wil- mostly yellow with elongate white patches son 1988; Ofuya 1989; Ofuya and Agele at base and at middle of 3rd interstice; lat- 1989; Credland and Wright 1990; Gokhale eral and apical parts of disk with curvate et al. 1990; Yamamoto 1990; Fox and Ta- dark brown and black maculation. Pygid- tar 1994. ium (figure 324) uniformly yellow, or with Parasitoids.—See “Natural Enemies,” dusky sublateral spots. above; Utida 1961; Taylor and Aludo 1974; Structure.—Vertex and frons finely punc- Mizell and Nebeker 1982. tate; frontal carina fine but evident; ocular Pheromones.—Sakai et al. 1986; Yamamoto index 6:1 in male, 4:1 in female; ocular 1986; Pouzat et al. 1989. sinus about one-half length of eye; anten- Population studies.—Bellows 1982. na dimorphic (figure 322). Pronotum subconical, lateral margins slightly sinuate; Respiration.—Seurat 1900; Yadav and disk unevenly convex; basal lobe domed, Singh 1977. vestiture not matted; lateral carina absent. Spermatheca.—Surtees 1961. Scutellum narrow, parallel. Elytra together Spermiogenesis.—Brauer 1928; Mickey one-fifth longer than wide, subdepressed 1935; Bawa and Kanwar 1975; Bawa et al. along suture, convex laterally; striae sub- 1971, 1975; Eady 1994. parallel, slightly sinuate, 2nd to 5th reach- Tracheal system.—Pajni 1969; Vats 1972. ing basal margin, 2nd and 3rd nearly conjoined and with minute basal denticles; Wing venation.—Suzuki 1969. interstices flat, finely imbricate-punctate. Callosobruchus phaseoli (Gyllenhal) Metacoxal face finely, evenly foveolate except fossula smooth; metafemur as in (Figures 321–326) figure 323, ventrolateral carina of metafe- Bruchus phaseoli Gyllenhal 1833:37; Pic mur with distinct toothlike angulation, 1913a:40; Swezey 1936:201; Mukerji ventromesal carina with short, acute den- and Chatterjee 1951:8 (male genitalia). ticle about as long as lateral angulation; metatibia (figure 323) dilated toward apex, Mylabris phaseoli: Baudi 1886b:46. with lateral carina sinuate apically, ventral Pachymerus phaseoli: Bridwell 1919:17. and dorsomesal carinae complete, ventro- Callosobruchus phaseoli: Herford 1935:6; lateral evanescent apically; mucro short, Bridwell 1938b:75; Hoffman 1945:88; only slightly longer

86 than lateral denticle; pygidium (figure 324) nae reddish yellow; pronotum all black or slightly tumescent at apex in both sexes. with reddish apical margin; elytra dark red Male genitalia.—As in figures 325 and 326; to black; pygidium and most of abdomen median lobe elongate, about 8 times as red; fore legs and mid legs red, hind legs long as wide; ventral valve hastate, pro- black with dorsal one-half red. Vesti-ture duced into slender apex, base constricted; of black, white, and yellow hairs in variable internal sac with small bundle of spic- dorsal pattern (figure 327); head yellow. ules at apical orifice, two pairs of burrlike Pronotum yellow with black apical and sclerites in middle, and pair of spine clus- lateral diagonal streaks (figure 327), basal ters near apex; most of sac lined with fine lobe with waxy secretion on matted hairs. spicules; lateral lobes elongate (figure 326), Scutellum white. Elytra with basal yel- cleft to base, and curved inward at tips. low band leading medially to yellow patch surrounding scutellum, 3rd interstice with Size.—Body length 2.3–2.9 mm; width elongate yellow patch, sutural interstice 1.4–1.6 mm. with narrow black streak reaching nearly Type depository.—NHRS. to scutellum, lateral one-half of each elytron with narrow lines of white hairs. Py- Type locality.—”Brasilia.” gidium in both sexes uniformly white with Distribution.—Tropicopolitan. FL, HI. one pair of subapical spots (figure 331). Host plants.—Cajanus cajan; Cicer arieti- Venter of body mostly white, lateral mar- num; Crotalaria spectabilis; Lablab purpu- gin of abdomen, anterior spot on metepi- reus; Phaseolus lunatus, P. vulgaris; Pisum sternum, and mesepimeron golden yellow, sativum arvense; Sesbania sp.; Vicia faba; white patches on lateral margin of 3rd, Vigna angularis, V. radiata radiata. 4th, and 5th abdominal segments; small, white tuft on apical margin of 5th sternum. Natural enemies.—Anisopteromalus calandrae; Heterospilus prosopidis; Uscana Structure.—Head with prominent frontal semifumipennis. carina; eyes dimorphic, ocular index 9:1 in male, 6:1 in female; antennae dimorphic Immatures.—Wightman and Southgate (figures 328 and 329). Pronotum narrowly 1982 (egg). trapezoidal, lateral margins incurved, disk Discussion.—This species is easily sepa- gibbous, medially sulcate. Scutellum trian- rated from other Callosobruchus by the gular, bifid apically. Elytra subde-pressed; yellowish appearance with dark brown to striae slightly impressed, sinuate, punc- black, lunate elytral maculae, red terminal tures discrete, 3rd and 4th striae ending antennal segment, enlarged eyes in male, basally in subbasal gibbosity; interstices 2nd and 3rd striae nearly conjoined, and imbricate. Pygidium (figure 331) with slight slightly tumescent pygidium. Male genitalia tumescence between subapical spots. are distinctive. Metacoxa densely punctate; metafemur (figure 330) sulcate ventrally, ventrolat- Callosobruchus pulcher Pic eral carina straight with subapical notch, ventromesal carina with small, acute den- (Figures 327–333) ticle; metatibia with lateral, ventrolateral, Callosobruchus pulcher Pic 1922:15; U.S. ventral, and dorsomesal carinae complete; Department of Agriculture 1973:149 [as mucro one-fourth as long as basitarsus. C. albocallosus (Pic)], 1973:171 (cor- Male genitalia.—As in figures 332 and 333; rected to pulcher); Udayagiri and Wadhi median lobe elongated, ventral valve tri- 1989:171. angular, middle of internal sac with two Color.—Integument dark red to black, oc- large, curvate and dentate sclerites, apical casionally purple. Head black, labrum red, vertex with red spot behind eye; anten-

87 one-half of sac lined with minute, aciculate Tribe Acanthoscelidini Bridwell, 1946 spicules, apical orifice with conspicuous bundle of spicules, apical one-third of sac Genus Abutiloneus Bridwell pleated, lined with minute, overlapping, semicircular scales; lateral lobes slender, Abutiloneus Bridwell 1946:55; Blackwelder expanded suddenly at apex, divided to at- and Blackwelder 1948:45; Kingsolver tachment with tegmen (figure 333). 1965a:125; Bottimer 1968c:1016,1038; Johnson 1968:1267; Johnson and Size.—Body length 2.3–3.1 mm; width Kingsolver 1982:416; Johnson 1.4–1.9 mm. 1983d:378; Udayagiri and Wadhi Type depository.—MNHP. 1989:34. Type species: Abutiloneus idoneus Bridwell (=Bruchus flavicornis Type locality.—Philippine Is., Luzon. Sharp 1885, not Fabricius 1792), by Distribution.—Hawaii (introduced); Philip- original designation. pine Islands. Borowiec (1987) synonymized this genus Host plants.—Cajanus cajan; Cicer ari-eti- with Acanthoscelides, but Johnson and num; Phaseolus vulgaris, Vigna angularis; I agree that this action was premature. flowers ofCajanus cajan. Johnson (1990c:306) resurrected Abutil- oneus to generic status. Natural enemies.—None recorded. Because this at present is a monotypic ge- Immatures.—Not described. nus, the generic and specific diagnoses are Discussion.—According to specimen data, combined. this handsome Philippine species was Taxonomy was summarized by Kingsolver first collected in Hawaii in 1965 by N.L.H. (1965a). Krauss on Maui. It was subsequently collected at light at Honolulu, Manoa, and Abutiloneus idoneus Bridwell Ewa (all in Oahu) and at Haiku, Maui. It was reared from Cajanus cajan at Ewa in (Figures 66, 334–339) 1965. Bruchus flavicornis Sharp 1885:480; This species would most likely be con- Townsend 1903:88; Schaeffer fused with Callosobruchus chinensis, but 1907:296; Zacher 1952:462. the lateral extensions of each of the male Bruchus (Abutiloneus) idoneus: Zacher antennal segments are broader in C. pul- 1952:462. cher (figure 328), the ventral vestiture is Acanthoscelides flavicornis: Blackwelder all white, and the lateroventral tooth of the 1946:759. metafemur is more blunt (compare figures 298 and 330). The key characters will suf- Abutiloneus flavicornis: Kingsolver fice for differentiation from other species of 1965a:125 (misidentification). Callosobruchus. Abutiloneus idoneus Bridwell 1946:55 (new name for flavicornis Sharp); Black-welder and Blackwelder 1948:759; Bottimer 1968c:1016,1038; Johnson 1968:1267; Johnson and Kingsolver 1982:416; Johnson 1983d:378; Borowiec 1987:88; Udayagiri and Wadhi 1989:34. Althaeus idoneus: De Luca 1967a:16. Color.—Integument black except antenna, labrum, and labial and maxillary palpi bright yellow. Pubescence on pronotum

88 and elytra evenly yellowish gray; that on Type locality.—Texas, Brownsville (idon- pygidium and ventral areas of body ashy eus); Mexico, Guanajuato (flavicornis). gray. Distribution.—TX; Mexico. Structure.—Body short and broad (figure Host plants.—Abutilon berlandieri (=califor- 334). Vertex and frons minutely punctate; nicum), A. abutiloides (=lignosum); Allowis- frontal carina lacking but frons marked sadula holosericea. Johnson (1983d) sum- by impunctate median line; ocular in- marized the host plant data. dex 3:1; ocular sinus two-thirds length of eye; antenna (figure 336) not dimorphic. Natural enemies.—Zatropis incertus (Ash- Pronotum nearly semicircular in outline, mead). evenly convex, disk sparsely microfoveo- Immatures.—Not described. late, surface densely pubescent, minutely Discussion.—The small size, bright yellow imbricate; lateral carina ridgelike, arched; appendages, subapical femoral denticle cervical sulcus short, deep; fore coxae minute or lacking, and distinctive male contiguous at apices. Scutellum slightly genitalia will distinguish this species. Of longer than wide, bidentate apically. Elytra similar size and shape and also breeding in as long as wide, striae parallel, 2nd and Abutilon spp. are Acanthoscelides aequalis 3rd slightly bent laterad near base, 2nd to and A. subaequalis, but in both the legs 5th minutely denticulate on basal margin; are dark brown, at least in part, and male interstices minutely imbricate; metacoxal genitalia are distinctly different. face densely imbricate-punctate, densely setose in lateral one-half; hind leg (figure Kingsolver (1965a:125) erroneously listed 335a) usually without subapical denticles, Abutiloneus idoneus as a synonym of Bru- sometimes with one minute, black denticle chus flavicornis Sharp. (figure 335b); metatibia slightly dilated toward apex, ventrolateral carina absent, lat- Genus Acanthoscelides Schilsky eral carina evanescent (figure 335a); mucro short, scarcely longer than lateral and the Acanthocelides (sic) Schilsky 1905: four coronal denticles. Abdomen not modi- IV,41F,41C,41L. fied, male 5th sternum broadly emargin- Acanthoscelides Schilsky 1905:95–95a; ate; pygidium of both sexes convex, male’s Pic 1913a:13 (as subgenus of Bru- more strongly reflexed than female’s. chus); Leng 1920:304; Bridwell Male genitalia.—As in figures 337–339; 1929a:39, 1932:104, 1946:52; Black- median lobe slender; ventral valve ogival, welder 1946:758; Luk’yanovich and acute, strongly curvate; internal sac armed Ter-Minassian 1957:170; Bottimer with paired, elongate clusters of slender 1968c:1016,1038,1040; Johnson spicules in basal one-half of sac, pair of 1968:1267, 1970:1–116, 1981b:73; small, thornlike spines in middle, and pair Johnson and Kingsolver 1982:409–422; of larger, thornlike spines near apex; later- Johnson 1983a:1–370, 1990c:297–618. al lobes slender, slightly expanded apically, Acanthoscelides was described in 1905 by extreme apex acute and recurved, cleft Schilsky for Bruchus irresectus Fahraeus, between lobes nearly to base. now a synonym of Acanthoscelides obtectus (Say). Schilsky included species under two Size.—Body length 1.5–1.8 mm; width different spellings—Acanthoscelides and 1.0–1.2 mm. Acanthocelides; however, Bottimer (1968c), Type depository.—USNM (idoneus, holotype followed by Johnson (1970), adopted Acan- No. 5766); BMNH (flavicornis). thoscelides as the spelling Schilsky probably intended, with Acanthocelides regarded as a lapsus.

89 Pic, in his world catalogue (1913a:13), list- times alternately variable in width, surface ed Acanthoscelides as a subgenus, but the usually minutely imbricate. Metasternum name did not come into general use until convex, postmesocoxal sulci prominent, Bridwell’s papers in the 1920–1930 era in median sulcus extending half-way to ante- which he transferred some of the U.S. spe- rior border; metacoxal face usually punc- cies from Bruchus into Acantho-scelides. tate and at least partly setose, fossula al- Blackwelder, in his Neotropical checklist ways glabrous and impunctate; metafemur (1946), transferred 322 species names, expanded dorsoventrally, ventral margin mostly from Bruchus, into Acanthoscelides. often shallowly sulcate; subapical ventral Some of these names have subsequently margin usually with cluster of one to four been transferred to other genera. acute denticles (pecten); metatibia straight The following description is modified from to moderately arcuate, dilated toward apex, Johnson (1983a) to apply to North Ameri- ventral and dorsomesal carinae always can species of Acanthoscelides. present, lateral and ventrolateral carinae varying in extent; apex with acute spine Small bruchids (1.1–3.5 mm). (mucro) on ventral margin extending one- Vertex and frons usually punctate, inter- tenth to nearly same length of basitarsus, spaces impunctate, punctate, or granu- a lateral denticle at end of lateral carina, lose, frons sometimes with median vertical and two to four coronal denticles. First carina or impunctate line; eyes protruding abdominal sternum often modified in male from lateral margins of head, often sexually with median setose pit, sometimes flat- dimorphic with eyes larger in male than tened or concave, or with posterior marin female, posterior border of eye usually gin lobed and fringed with long setae, 5th separated from side of head by postocular sternum moderately to deeply emarginate sulcus, eye partly divided by setose ocular in male for reception of pygidial apex; 1st sinus; antennal insertion at anterior mar- sternum of female not modified, 5th usu- gin of eye; antenna subserrate from 4th or ally slightly emarginate; pygidium broadly 5th segment except 11th segment elliptical, obovate, arcuate in lateral aspect, male often sexually dimorphic with male an- pygidium usually strongly inflexed at apex, tenna longer or broader. Pronotum bell- female pygidium usually vertical at apex; shaped or subconical, disk convex without disk microfoveolate or punctulate. elevations, usually with slight depressions Male genitalia usually elongate, median either side of basal lobe and near poste- lobe lacking distinct dorsal valve, ventral rior corners, basal lobe usually with brief valve with base usually broad, apex of sulcus; lateral carina absent or represent- various shapes, usually diagnostic; base ed by blunt ridge; cervical sulcus usually of median lobe spoon-shaped; internal conspicuous; prosternum /T/- or /Y/- eversible sac armed with variously shaped shaped, triangular between procoxae but spines, spicules, and denticles (figure 35) not separating their apices; protrochan- whose arrangement is usually species-spe- tin absent, replaced by trochantinal flap cific; lateral lobes variously shaped, sepa- (figure 24); mesepimeron reduced to small rated by cleft of various lengths, apices triangular sclerite by encroachment of with sensory setae. mesepisternum (as in figure 27). Scutellum quadrate or slightly elongated, apical mar- The genus Acanthoscelides is a large and gin bidentate with angulate emargination diverse, poorly defined aggregate of mostly between points. Elytra together as long or small species of Bruchidae. C.D. Johnson longer than broad, striae varying in width is the leading authority on this genus with and depth, often with basal or subbasal his many papers published on taxonomy denticles; interstices usually flat, some- and host associations (1968, 1969a,b,

90 1970, 1974, 1977a,b,c,d, 1979a,b, 1981b,e, 1983a, 1988b, 1990a,c,d). His studies on western United States, Mexico- Central American, and northern South American species of Acanthoscelides have failed to provide a wholly satisfactory definition of the genus but have resulted mostly in defining species groups within the genus. None of these species groups appear to be sufficiently distinct to warrant separation as a new genus.

91 Key to Species of Acanthoscelides This key is based on a combination of integumental and vestitular color and morphological features. In using color, especially in beetles, one must be cautious because of variation in intensity of melanism. Beetles that have recently emerged are often red or brown but darken after a few days.

1 Integument of body and appendages entirely black except 7. Body length 0.9–1.2 mm; ventral valve broadly triangular basal two or three antennal segments red or reddish (figure 355) yellow, some species with reddish suffusion on legs in ...... atomus (Fall) teneral specimens, or with red postocular patch...... 2 Body length 1.4–1.7 mm; ventral valve narrowly triangu- Body black or red or mottled; appendages entirely or in lar (figure 446) part red or reddish orange...... 21 ...... helianthemum Bottimer 2(1) Metatibia lacking lateral carina (figures 539 and 444) or 8(2) Elytral vestiture ocellate, each rounded, glabrous spot carina evanescent (figure 378); ventrolateral carina absent centered on a single, curved seta (figure 345) ...... 3 ...... alboscutellatus (Horn) Metatibia with distinct lateral carina (figure 348)...... 8 Elytral vestiture not ocellate...... 9 3(2) Elytral vestiture uniform or maculate (figures 377 and 9(8) Elytral vestiture maculate (figure 517)...... 10 482)...... 4 Elytral vestiture uniformly distributed (figure 555)...... 12 Elytral vestiture ocellate, small glabrous spots each 10(9) Bases of 3rd and 4th striae each with prominent denticle; centered on fine, curved seta (figure 443)...... 6 vestiture of dark brown, yellowish-brown, and white setae 4(3) Dorsal vestiture uniform with no pattern (figure 377); in mottled pattern (figure 517); great denticle of metafe- scutellum contrasting white; body length 3.0–3.2 mm; mur longer than tibial width at base (figure 518); body male genitalia as in figures 380 and 381; east of 100th length 2.1–2.9 mm meridian...... calvus (Horn) (part) ...... obsoletus (Say) Dorsal vestiture with faint to distinct pattern (figure 538); Strial bases lacking denticles, or denticles minute, nearly scutellum not strongly contrasted; body length 2.0 mm or hidden in vestiture; great denticle of pecten small, shorter less; western United States...... 5 that width of tibial base (figures 551 and 570)...... 11 5(4) Metatibial mucro slender, curved; about one-third length 11(10) Mucro of metatibia long, slender, curved, slightly longer of basitarsus (figure 539); male genitalia as in figures 541 than apical width of tibia (figure 551)...... and 542; 1st abdominal sternite of male with small pit ...... pedicularius (Sharp) ...... pauperculus (LeConte) Mucro of metatibia short, not as long as tibial width Mucro less than one-fourth length of basi-tarsus (figure (figure 570)...... pullus (Fall) 483); male genitalia as in figures 485 and 486; 1st ab- 12(9) Strial punctures rounded or ovate, encroaching on dominal sternite of male with prominent lobe on posterior adjacent interstices (figure 555); pronotum subconical, margin...... margaretae Johnson (part) densely, coarsely, punctate; pecten of metatibia with two 6(3) Body length 2 times greatest width across elytra; mucro minute denticles (figure 557); 1st abdominal sternum of slightly longer than lateral denticle (figure 633); pecten male concave with median pit and long fringe of golden with one large and two minute denticles; median lobe setae...... perforatus (Horn) (figure 635) without large spines Strial punctures elongate; other characters varied...... 13 ...... tenuis Bottimer 13(12) Lateral tibial carina evanescent, sometimes with portions Body length 1.6 to 1.8 times width; mucro long, slender visible (figure 378); pronotum subconical (figure 377); (figures 353 and 444); pecten with two or three minute 2nd stria curved laterad to meet 3rd at basal margin; denticles; male genitalic armature includes two large, thornlike spines...... 7

92 1st denticle of pecten 2–3 times as long as 2nd and 3rd 19(18) Eye in lateral aspect protruding below plane of gula; (figure 378); 1st abdominal sternum of male not modified lateral margins of ventral valve straight (figure 404); body ...... calvus (Horn) (part) covered with dense, white setae giving pruinose appear- Lateral carina visible for entire length; other characters ance...... daleae Johnson varied...... 14 Eye in lateral aspect not protruding below plane of gula; 14(13) Elytra with striae shallow, especially in lateral one-half, lateral margins of ventral valve usually incurved (figure sometimes with only elongate punctures visible; 1st 430); vestiture of body white to yellow...... 20 denticle of pecten two times as long as 2nd or 3rd (figure 20(19) Vestiture of moderately dense to extremely dense white 433); 1st abdominal sternum of male not modified setae; body and appendages usually all black except ...... fumatus (Schaeffer) basal four antennal segments reddish orange, sometimes Elytra with striae narrow, deep, distinct; femoral denticles with elytral apices and apical portions of legs reddish subequal in length; 1st sternum of male modified by a orange; male genitalia as in figure 430...... lobe or pit (except compressicornis)...... 15 ...... fraterculus (Horn) 15(14) Metafemur with three or four denticles on ventral margin If integument of head, body, and legs mostly black, then between trochanter and pecten (figure 390); pronotum vestitural color usually yellow; if integumental color paler, with tuft of setae opposite base of 5th stria; 1st abdominal then vestiture white, yellow, or intermixed white and sternum of male not modified; male antenna broad and golden; male genitalia as in figure 360 elongated (figure 391) ...... aureolus (Horn) complex...... compressicornis (Schaeffer) 21. Metatibia lacking lateral carina, or carina evanescent Ventral margin of metafemur with denticles present only (figure 528)...... 22 in pecten (figure 465); pronotum lacking setal tuft; 1st ab- Metatibial carina present and distinct dominal segment modified; male antenna not especially (figure 477)...... 24 broad 22(21) Metafemur lacking ventromesal denticles (pecten) (figure ...... 16 528)...... 23 16(15) Male with setose lobe and rounded depression on 1st Metafemur with one or more ventromesal denticles (figure abdominal segment; pygidium with lateral margins nearly 454)...... inquisitus (Fall) straight (figure 464)...... lobatus (Fall) 23(22) Pronotal lateral margins sinuate (figure 527); metatibia Male lacking lobe although posterior margin of 1st seg- yellow; apical one-third to one-half of each elytron yel- ment sinuate and with large rounded depression; female low; 1st abdominal sternite of male with small pit enclos- pygidium with lateral margins more rounded (figure 427) ing setal tuft; male genitalia as in figure 530 ...... 17 ...... oregonensis Johnson 17(16) Apex of metatibia with deep clefts between coronal Pronotum nearly semicircular (figure 504); legs dark denticles (figure 606); male genitalia with acute spine in red to black; elytra black with faint golden maculae; 1st internal sac (figure 609) abdominal sternite of male convex, without pit; male ...... seminulum (Horn) genitalia as in figure 507...... napensis Johnson Apex of metatibia with shallow intervals between denticles 24(21) Elytral integument basically red, or reddish brown, some- (figure 575); internal sac differently armed...... 18 times with darker maculae, or darker marginal or sutural 18(17) Male genitalia with small, U-shaped sclerite in internal stripe...... 25 sac (figure 577); elytra maculate (figure 574)...... Elytral integument basically black, sometimes with red- ...... pusillimus (Sharp) dish stripe, or with apices reddish yellow...... 41 Internal sac differently armed; elytral vestiture uniform...... 19

93 25(24) Third and 4th elytral striae strongly bent laterad at base Head red with at most small, black postocular spots; male (figures 474 and 493), nearer to each other than to adja- genitalia as in figures 614 and 615...... cent striae, sometimes nearly merging, each stria ending ...... speciosus (Schaeffer) in prominent subbasal denticle, sometimes with denticles 32(25) Pronotal integument uniformly black in contrast to red merging into bidentate ridge...... 26 elytra; pronotum sometimes with pale, thin, median line Third and 4th striae parallel, equidistant from each other of setae (see also A. kingsolveri)...... 33 and from adjacent striae, with or without basal denticles... Pronotal and elytral integument similarly colored, some- ...... 32 times with darker brown, broad, median pronotal stripe.... 26(25) Pronotum lacking distinct pattern...... 27 ...... 35 Pronotum with distinct pattern...... 28 33(32) Internal sac of male genitalia with one thornlike and one 27(26) Mucro longer than lateral denticle of meta-tibia (figure serrate sclerite (figure 399); lateral lobes cleft nearly one- 477); eyes prominent, ocular index of males 14:1 (figure half their length (figure 400); Texas...... 475), of females 6.25:1 (figure 476); frontal carina promi- ...... comstocki Johnson nent...... Internal sac with one large tapered sclerite (figure 536); ...... macrophthalmus (Schaeffer) lateral lobes cleft about one-fourth their length (figures Mucro equal to or shorter than lateral denticle (figure 537 and 425)...... 34 494); ocular index 6:1 or less; frontal carina evanescent... 34(33) Lateral margins of ventral valve of male genitalia incurved ...... modestus (Sharp) (figure 536); mostly west of 100th meridian 28(26) Head red with frons black, black area extending dorsad ...... pallidipennis (Motschulsky) (part) from median spot on vertex; thoracic sterna, humeri, ely- Lateral margins of ventral valve straight (figure 424); east tral margins and apex, and leg bases piceous, abdomen of 100th meridian...... floridae (Horn) red; elytral denticles nearly confluent on common gibbos- 35(32) Vestiture of pronotum, elytra, and pygidium bright golden ity (figure 543)...... pectoralis (Horn) yellow, uniformly dense; venter of body densely white; 1st Head red or black; thoracic sterna usually black, some- sternite of male with pit; male genitalia as in figure 536.... times red or piceous; elytral pattern of dark maculae ...... pallidipennis (part) extending toward suture from middle of lateral margins; Dorsal vestiture otherwise; ventral vestiture various .....36 elytral denticles basal or subbasal (figures 561 and 367)...... 29 36(35) Pronotum and sometimes pygidium with broad, median stripe usually divided by narrow stripe or line of pale 29(28) Scutellum elongate, nearly 2 times as long as wide (figure setae ...... 37 561); frontal carina absent; elytral denticles subbasal; Dorsal vestiture otherwise...... 44 2nd and 3rd striae divergent laterad at base (figure 561); pronotum without pattern 37(36) Body slender, ratio of length to width 2:1 (figure 592); ...... prosopoides (Schaeffer) pronotum usually with broad, brown stripe divided by Scutellum quadrate or transverse (figure 367); frontal narrow stripe of pale setae; ocular sinus nearly dividing carina present; elytral denticles on basal margin; 2nd and eye; male antenna extending to 1st abdominal segment; 3rd striae subpar-allel (figure 367)...... 30 male genitalia as in figure 596...... schaefferi (Pic) Body broader, ratio 1.67–1.78:1; pronotal and dorsal 30(29) Head red, or red with median, black maculation...... 31 vestiture various; ocular sinus no deeper than two-thirds Head black; male genitalia as in figures 370 and 371...... length of eye; male antenna various...... 38 ...... bisignatus (Horn) 38(37) Metatibia usually lacking lateral carina, or carina 31(30) Head red with median black maculation; pygidium with evanescent (figure 628); male genitalia as in figure 630 three basal patches of white setae; male genitalia as in (compare A. floridae, figure 424)...... figures 409 and 410...... Johnson desmanthi ...... submuticus (Sharp) (part)

94 Metatibial lateral carina distinct; male genitalia otherwise. Pygidial integument and abdominal segments black, legs ...... 39 variously colored...... 47 39(38) Vestiture of elytra and pygidium dense, pale yellow, uni- 46(45) Antenna unicolorous from 5th segment to apex; mucro formly distributed, pygidium with at most a pale, median slender, base with adjacent deep sinus (figure 588); male line of setae; frontal carina distinct...... 40 with rounded pit on 1st abdominal sternite; male genitalia Vestiture of elytra striped or maculate; pygi-dium faintly as in figure 590...... rufovittatus (Schaeffer) or strongly striped; frons with at most a bare, impunctate Antenna with apical segment red; mucro short, triangular, median line...... 44 scarcely longer than lateral denticle, lacking deep sinus (figure 523); 1st sternite without pit; male genitalia as in 40(39) Dorsal coronal denticle of metatibia prominent, separated figure 525...... obtectus (Say) from adjoining denticle by deep cleft (figure 417); 3rd and 4th striae lacking basal denticles; 1st abdominal sternite 47(45) Hind legs entirely black, fore legs and mid legs reddish of male without pit; male genitalia as in figure 419...... yellow; metacoxa with impunc-tate median area (figure ...... flavescens (Fahraeus) 384); 3rd and 4th striae ending in common, or approxi- Dorsal coronal denticle not prominent, similar to adjacent mate, basal denticles (figure 382)...... 48 denticles (figure 439); male 1st abdominal sternite with Legs variously colored; metacoxa uniformly punctate; prominent pit...... 41 strial denticles in various conformations...... 49 41(24,40) Vertex with black integumental spot; male internal sac 48(47) Impunctate area of metacoxa finely striate (figure 384); with forked sclerite (figure 441) apical sclerites of internal sac with five to eight serrations ...... griseolus (Fall) (figure 387)...... chiricahuae (Fall) Vertex lacking black spot; internal sac densely lined with Impunctate area not striate (figure 600); apical sclerites needle-like spicules (figure 515)...... with 9–13 serrations (figure 602). ...schrankiae (Horn) ...... male obrienorum Johnson 49(47) Mucro at least one-half length of basitarsus (figure 470); 42(36) Pronotum and elytra with prominent lateral maculae; elytra maculate (figures 469 and 616); 1st abdominal pygidium as in figure 512...... sternum of male with setose pit; 4th stria and sometimes ...... female obrienorum Johnson 3rd with basal tubercle...... 50 Pronotum, elytra, and female pygidium striped, or with Mucro no more than one-third length of basitarsus; 1st small, inconspicuous lateral spots...... 43 sternum pitted or not; 4th stria tuberculate or not ...... 52 43(42) Vestiture of 3rd elytral interstice uniform throughout its 50(49) Male genitalia with many small spines lining internal sac length (figure 579); male genitalia as in figures 584 and (figure 472); base of mucro without deep sinus (figure 585...... quadridentatus (Schaeffer) 470); distribution east of 100th meridian...... Vestiture of 3rd elytral interstice interrupted by two brown ...... longistilus (Horn) spots (figure 637); male genitalia as in figures 641 and Male genitalia with either forked sclerite and cluster of 642...... tridenticulatus Bottimer small spines (figure 620) or with large, acuminate sclerite and row of thornlike spines (figure 375); base of mucro 44(39) Basal lobe of pronotum and base of pygidium each with with deep sinus (figure 373); distribution Arizona, New intensely white setal patch (figure 498); 2nd, 3rd, and Mexico, Texas, Mexico...... 51 4th striae deflected laterad at base, 4th with subbasal denticle; male genitalia as in figure 502...... 51(50) Male genitalia with forked sclerite and cluster of small ...... mundulus (Sharp) spines (figure 620)...... stylifer (Sharp) Pronotum and pygidium not so marked; striae not deflect- Male genitalia with large, acuminate spine and row of ed, or 3rd and 4th slightly deflected and denticulate.....45 thornlike spines (figure 375)...... biustulus (Fall) 45(44) Pygidial integument red, abdominal segments and legs usually partly red...... 46

95 52(49) Pronotal and elytral vestiture with brown and white pat- genitalia as in figure 461.... kingsolveri Johnson (part) tern (figure 411); male genitalia as in figure 414; male 56(53) Body elongate ovate (figure 340), length 1.7–2.2 mm; antenna strongly serrate (figure 413), extending to 1st mucro aciculate, about one-fourth as long as basitarsus abdominal sternite...... distinguendus (Horn) (figure 341)...... 57 Vestiture not with distinct pattern; male antenna not Body elliptical (figure 362), length 1.3–1.7 mm; mucro strongly serrate, not extending beyond metacoxa...... 53 short, slightly longer than lateral denticle (figure 363). 59 53(52) Elytra with red integumental stripe of variable width, 57(56) Hind legs mostly reddish yellow with only basal one- sometimes extending nearly to lateral margin, or apical fourth black...... 58 one-third or more of elytra red, or reddish yellow...... 54 Hind legs black or with apex of metafemur and tibia dark Elytral integument entirely black...... 56 red; male genitalia as in figure 451...... herissantitus 54(53) Apical one-third or more of elytral integument red or red- Johnson dish yellow, coloration occasionally extending narrowly 58(57) Male genitalia as in figure 343...... aequalis (Sharp) to elytral base; male genitalia as in figure 491...... Male genitalia as in figure 625...subaequalis Johnson ...... mixtus (Horn) 59(56) Lateral margins of pronotum nearly straight but con- Each elytron with narrow to broad, median integumental vergent toward apex (figure 362); pecten of metafemur stripe...... 55 composed of two or three minute, subequal denticles 55(54) Lateral margins of pronotum nearly straight in basal two- (figure 363); red stripe faint in some specimens; male thirds but convergent toward apex (figure 362); pecten of genitalia as in figure 365...... metafemur composed of two or three minute, subequal ...... baboquivari Johnson (part) denticles (figure 363); red stripe faint in some specimens; Lateral margins of pronotum moderately arcuate (figure male genitalia as in figure 365...... 458); pecten with basal denticle two times as long as ...... baboquivari Johnson (part) distal denticles (figure 459); red stripe narrow to broad; Lateral margins of pronotum moderately arcuate (figure male genitalia as in figure 461...... 458); pecten with basal denticle 2 times as long as distal ...... kingsolveri Johnson (part) denticles (figure 459); red stripe narrow to broad; male

Acanthoscelides aequalis (Sharp) to piceous, fore and middle legs reddish orange with basal one-fourth to one-third (Figures 340–344) and trochanter reddish brown, hind leg Bruchus aequalis Sharp 1885:481; Schaef- with at least trochanter and basal one- fer 1907:304; Fall 1910:173; Pic third of femur reddish brown, remainder 1913b:13. of hind leg reddish orange except tarsi Mylabris aequalis: Leng 1920:305. sometimes brown. Vestiture white, evenly distributed. Acanthoscelides aequalis: Moreno and Bibby 1943:23; Blackwelder Structure.—Vertex and frons micropunctate 1946:758; Bridwell 1952:50; King- except glabrous median frontal line; ocular solver 1965a:128; Bottimer 1968c:336; index 2.5:1; ocular sinus one-half length Johnson 1970:15,16,18,87; John- of eye; antenna as in figure 342, reaching son and Kingsolver 1982:414; John- middle of metepisternum. Pronotum bell- son 1983a:24; Udayagiri and Wadhi shaped (figure 340), disk microfoveolate; 1989:35. lateral carina obsolete; cervical sulcus Color.—Body black; 1st to 4th antennal well-defined. Scutellum rectangular. Elytra segments red, 5th to 11th dark brown convex, lateral margins moderately arcu-

96 ate (figure 340), striae distinct, parallel and with three denticles will separate it except 2nd slightly divergent at base, 3rd, from A. idoneus with golden vestiture, all 4th, and 5th striae each with minute basal legs red or yellow, and metafemur lacking denticle, interstices flat, microimbricate. denticles or at most with one small den- Metacoxal face densely micropunctate, ticle. with distal pubescent patch; hind leg as in figure 341; pecten with one large and Acanthoscelides alboscutellatus (Horn) two minute denticles; metatibia gradually (Figures 345–351) dilated toward apex, lateral, dorsomesal, and ventral carinae complete, ventrolateral Bruchus alboscutellatus Horn 1873:334; carina absent; mucro short, acute (figure Hubbard and Schwarz 1878:660; Casey 341), lateral denticle and coronal denticles 1884:184; Hamilton 1892b:253; Ri- minute, acute. Fifth abdominal sternum of ley and Howard 1892c:165; Townsend male deeply emarginate to receive apex of 1895:277; Fall and Cockerell 1907:201; pygidium, of female nearly straight; pygi- Schaeffer 1907:303; Blatchley dium more strongly convex in male. 1910:1239; Fall 1910:168; Cush- man 1911:505; Pic 1913a:14; Bridwell Male genitalia.—As in figures 343 and 344; 1918:494. median lobe with ventral valve subelliptical (figure 343); internal sac armed with many Bruchus albiscutellaris: Ashmead 1894:328 fine denticles near base, then in order (misspelling) toward apex two small burrs, two clublike, Bruchus alboscutellaris: Zacher 1952:461 spiny sclerites, 10–11 burrs, two slen- (misspelling). der, thornlike spines, a lightly sclerotized, Mylabris alboscutellatus: Leng 1920:305; spinose structure; apex finely denticulate, Essig 1926:485, 1958:486. closure valve C-shaped; lateral lobes flat Acanthoscelides alboscutellatus: Bottimer (figure 344), spatulate, setose, cleft about 1935:128; Bridwell 1935:186; Bissell one-half their length. 1940:846; Bradley 1947:40; Black- Size.—Body length 1.2–2.0 mm; width welder and Blackwelder 1948:44; Peck 0.9–1.3 mm. 1963:956; De Luca 1965:1017,1038; Johnson 1969c:55, 1970:15, 1974:272; Type depository.—BMNH. Center and Johnson 1974:1097; John- Type locality.—Mexico, Guanajuato, Tox- son 1976b:260; Kingsolver 1979a:341; pam. Johnson 1981b:79; Johnson and King- Distribution.—AZ, TX; Mexico to Guate- solver 1982:414; Johnson 1983a:29; mala. Udayagiri and Wadhi 1989:36; Ott 1991:641; Ott and Lampo 1991. Host plants.—Abutilon berlandieri; Allowissadula holosericea, A. lozani. Bruchus conspersus Motschulsky Johnson (1983a:25) lists four additional 1874:224; Bottimer 1968c:1040. host plants in Mexico. Acanthoscelides conspersus: Kingsolver 1979a:341. Natural enemies.—None recorded. Bruchus abutilonis Motschulsky 1874:232; Immatures.—Not described. Zacher 1952:461,468. Discussion.—This is one of the small- Acanthoscelides abutilonis: Kingsolver est bruchids in the United States, rivaled 1979a:341. only by Abutiloneus idoneus, with which it Color.—Body usually entirely black ex- is sometimes associated, and A. atomus, cept basal four antennal segments red- which is found on the east coast. The dish brown on ventral face, some teneral evenly distributed white vestiture and the specimens piceous with legs and antenna metafemur with black base and red apex

97 dark red. Vestiture white, more densely set Type depository.—MCZC (alboscutellatus, on lateral parts of pronotal disk than in lectotype No. 8202, by Johnson 1968), middle (figure 345), dense on mesepimeron ZMUM (abutilonis, conspersus). and caudal margin of metepisternum (fig- Type locality.—"Georgia, Louisiana, and ure 347), conspicuous on scutellum; elytra Missour" (alboscutellatus); America Borea- with regular rows of white patches of setae lis (abutilonis, conspersus). (figure 345); pygidium slightly mottled. Distribution.—AL, AR, CO, CT, DC, DE, FL, Structure.—Vertex and frons convex, lack- GA, IL, IN, KS, LA, MD, MI, MO, MS, NC, ing frontal carina, micropunctate, setae of NJ, NY, OH, OK, PA, SC, TN, TX, VA, WV; punctures directed toward center of frons; Mexico. ocular index 3:1; ocular sinus one-half length of eye; antenna as in figure 349, Host plants.—Daucus carota (floral record); extending to anterior margin of metacoxa. Glycyrrhiza lepidota; Ludwigia alternifolia, Pronotum strongly convex, bell-shaped L. palustris. (figure 345), disk microfoveolate, foveolae Natural enemies.—Dinarmus acutus; Eupel- circular, usually discrete but sometimes mus amicus; Uscana semifumipennis. coalescent. Scutellum cordate, densely Immatures.—Not described. pubescent. Elytral striae deep, narrow, set with short, curved setae, without basal Discussion.—Salient characters for this denticles, arising basally from simple species are the mottled elytra, promi- punctures; interstices flat, of uniform nent white scutellum, convex pronotum, width, microimbricate; metacoxa uniformly bulbous male pygidium, and host plant punctate; hind leg as in figure 348, pecten specificity. This species might be confused with one curved denticle followed by one with A. helianthemum, but that species or two minute denticles; metatibia gradu- averages smaller (1.4–1.7 mm), the scutel- ally dilated, lateral, ventral, and dorsome- lum, although white, is not prominent, the sal carinae complete to apex, ventrolateral male pygidium is not bulging, and the male carina evanescent in apical one-half; mu- genitalia are distinctive (compare figure cro short, acute, about one-seventh length 446). The pygidium of A. tenuis is slightly of basitarsus; lateral denticle and three convex, and the elytra are mottled as in coronal denticles acute. Pygidium of male A. alboscutellatus, but the average size is bulbous (figure 346), that of female moder- smaller and the male genitalia are distinc- ately convex (figure 347). tive (compare figures 350 and 635). These three species breed in seeds of three differ- Male genitalia.—As in figures 350 and 351; ent plant families, none of which is legumi- median lobe medially constricted; ventral nous. valve triangular with lateral extensions at base; dorsal valve membranous; armature Acanthoscelides atomus (Fall) of internal sac consisting of, in order, a section of fine denticles, a section of broad, (Figures 352–356) rounded, serrate denticles (see inset, figure Bruchus atomus Fall 1910:188; Pic 350), another section of fine denticles, two 1913a:17. small thornlike denticles; apex finely spiculate; closure valve C-shaped; lateral lobes Mylabris atomus: Leng 1920:306; Frost slender, slightly spatulate toward apex, 1931:3. mesal faces concave, bordered with fine Abutiloneus atomus: Bradley 1947:41; setae, deeply cleft. Blackwelder and Blackwelder 1948:45. Size.—Body length 1.1–2.3 mm; width Acanthoscelides atomus: Bottimer 0.8–1.4 mm. 1935:128, 1968c:1017; Johnson 1968:1267; Bottimer 1969a:981;

98 Johnson and Kingsolver 1982:414; Type depository.—MCZC, holotype No. Udayagiri and Wadhi 1989:37. 25045. Color.—Body black with reddish highlights Type locality.—Massachusetts, Hyannis. on antenna and legs. Vestiture of slender, Distribution.—FL, IA, MA, MD, ME, NH, NJ. white, scalelike hairs uniformly distributed over body except elytral interstices with Host plants.—Lechea racemulosa, L. tenui- elongate white patches separated by setif- folia (Bottimer 1969a). Swept from Hudso- erous black spots forming ocellate pattern nia ericoides (Frost 1931). (figure 352). Natural enemies.—None recorded. Structure.—Vertex and frons micropunc- Immatures.—Not described. tate, sparsely setose; frons with glabrous median line; ocular index 2.3:1; ocular Discussion.—This species can hardly be sinus three-fifths length of eye; antenna mistaken for any other eastern species (ex- as in figure 354, reaching middle of met- cept A. helianthemum) because of its min- episternum. Pronotum bell-shaped (figure ute size, ocellate elytral vestiture, and lack 352), evenly convex, disk micro-punctate, of lateral and ventrolateral tibial carinae. sparsely setose; lateral carina absent; cer- The male genitalia are distinctive. In A. vical sulcus well defined. Scutellum quad- atomus, the ventral valve is broadly tri- rate, pubescence not contrasting. Elytral angular, whereas in A. helianthemum this margins arcuate (figure 352); dorsal sur- sclerite is narrowly triangular. Body length face strongly convex; striae parallel, nar- for A. atomus ranges from 0.9–1.2 mm; for row, 2nd through 5th arising from minute A. helianthemum, the range is 1.4–1.7 mm. basal denticles; interstices flat, minutely, Host plants for both A. atomus and A. heli- sparsely punctulate. Meta-coxa shallowly anthemum are in the family Cistaceae. micropunctate, sparsely pubescent; metafemur (figure 353) moderately swollen, Acanthoscelides aureolus (Horn) complex pecten with two minute, separate denticles; (Figures 357–361) metatibia gradually dilated toward apex, mucro slender, one-third length of basitar- Bruchus aureolus Horn 1873:328; Ri- sus with deep sinus at base; ventral and ley and Howard 1892c:166; Horn dorsomesal carinae present, lateral and 1894:345; Fall 1901:29; Fall and Cock- ventrolateral carinae absent. First sternum erell 1907:201; Schaeffer 1907:299; unmodified; 5th sternum slightly emargin- Fall 1910:173,184; Cushman ate in male, margin straight in female; py- 1911:505; Pic 1913a:17. gidium convex in both sexes, apex of male Mylabris aureolus: Leng 1920:305. more deeply reflexed than female. Acanthoscelides aureolus: Blackwelder Male genitalia.—As in figures 355 and 356; 1946:758; Bottimer 1968c:1017,1038; median lobe not constricted in middle; ven- Johnson tral valve subtriangular, apex with small 1968:1267, 1969c:55, 1970:33; John- nipple; dorsal valve evenly rounded; arma- son and Kingsolver 1982:414; Johnson ture of internal sac consisting of cluster of 1983a:38; Boe spicules at apical orifice, a dense cluster et al. 1988; Udayagiri and Wadhi of semicircular, serrate sclerites, a pair of 1989:38. slender spines, and a spinose, bifurcate Bruchus fraterculus: Fall 1910:184 (not structure; closure valve V-shaped; lateral Horn 1873). lobes slender (figure 356), flat, slightly Bruchus pauperculus: Schaeffer 1907:299 bowed, deeply cleft. (not LeConte 1857). Size.—Body length 0.95–1.2 mm; width Mylabris pauperculus: Leng 1920:305 (not 0.6–0.8 mm. LeConte 1857).

99 Acanthoscelides pauperculus: Bottimer or concave, with median pubescent pit and 1968c:1020. several large punctures, posterior mar- Bruchus pulicarius: Motschulsky 1874:233. gin of sternum sinuate with fringe of long Bruchus rufus: Motschulsky 1874:238. hairs; 5th male sternum deeply emarginate; 1st sternum of female slightly curvate, Color.—Body reddish orange to black; head 5th only slightly emarginate; pygidium of black, sometimes with dark red spot be- male strongly reflexed into sternal emar- hind eye, antenna usually black with basal gination, that of female less strongly con- four or five segments red, rarely all red; vex, disk in both sexes finely, densely pronotum varying from all red to all black; punctate. elytra varying from all red to all black, often with base black and apex red; sternal Male genitalia.—As in figures 360 and 361; areas variable; legs red to black, some- median lobe slightly constricted at middle; times bicolored. Vestiture white or grayish ventral valve triangular; internal sac with yellow if integumental color of head, body, large, tapered sclerite; apical one-half with and legs mostly black; vestiture usually paired, setose, slender structures; apex white but may appear yellow or intermixed with circular closure valve; lateral lobes white and golden on yellow or red integu- fused at base for two-thirds their length, mental background. apices spatulate, mesal faces setose. Structure.—Head finely punctate, or, in Size.—Body length 1.1–2.7 mm; width some forms, minutely granulose; frons 0.9–1.5 mm. evenly punctate or with median impunc- Type depository.—MCZC, lectotype No. tate line; ocular index 2:1 to 2:7; ocular 8199, by Johnson 1968. sinus one-half to three-fifths length of Type locality.—California. eye; antenna as in figure 359, extending to middle of metepisternum. Pronotum Distribution.—AB, AZ, BC, CA, CO, ID, MB, bell-shaped (figure 357), disk convex, MN, MT, ND, NE, NM, OR, SD, SK, TX, UT, slightly depressed either side of basal lobe WY; Mexico. and near posterior corners, lateral mar- Host plants.—Astragalus allochrous, A. gins gently sinuate, slightly constricted by amphioxys, A. asymmetricus, A. bisulcatus, cervical sulci near apex, surface densely A. calycosus, A. crotalariae, A. douglasii, A. microfoveolate, interspaces minutely im- drummondii, A. fucatus, A. grayi, A. humi- bricate, lateral ridge extending half-way to stratus, A. lancearius, A. lentiginosus, A. procoxal cavity. Scutellum slightly longer lonchocarpus, A. mollissimus mollissimus, than wide, bidentate apically. Elytra to- A. oxyphysus, A. pattersonii, A. praelongus, gether slightly longer than wide (figure A. pterocarpus, A. racemosus, A. thurberi, 357), lateral margins convex, disk slightly A. trichopodus antisellii, A. trichopodus depressed; striae narrow, indistinct be- phoxus, A. trichopodus trichopodus, A. cause of dense vestiture, 3rd and 4th wootoni; Glycyrrhiza lepidota; Lotus pur- with minute basal denticle; interstices shianus, L. scoparius; Oxytropis lambertii, flat, densely setose, minutely imbricate. O. sericea. Meta-coxal face minutely, densely punctate, fossula glabrous; hind leg as in figure Natural enemies.—Eupelmus sp.; Hori- 358, pecten with three minute denticles of smenus missouriensis; Microdontomerus nearly equal size; metatibia bent only at anthonomi. base (figure 358), slightly dilated toward Immatures.—Pfaffenberger and Johnson apex, lateral, ventral, and dorsomesal 1976:21 (first larval instar). carinae distinct, ventrolateral carina lack- Discussion.—Johnson (1970) treated this ing; mucro one-fourth length of basitarsus, as a complex of forms, perhaps cryptic lateral and coronal denticles of equal size. species, varying in size, color, and vesti- First abdominal sternum of male flattened ture somewhat related to host plant and

100 geographical distribution. To further com- bell-shaped (figure 362), lateral margins plicate the picture, the A. aureolus complex gently arcuate, disk strongly convex, sur- is part of a closely related species group face closely set with setiferous microfo- that includes Acanthoscelides pallidipen- veolae; lateral carina ridgelike; cervical nis (Motschulsky), A. fraterculus (Horn), sulcus well-defined. Scutellum quadrate, A. seminulum (Horn), A. perforatus (Horn), bifid apically. Elytra widest behind middle A. lobatus (Fall), A. pullus (Fall), A. mixtus (figure 362); striae parallel, 3rd, 4th, and (Horn), A. daleae Johnson, A. oregonen- 5th with minute basal denticles; interstices sis Johnson, A. margaretae Johnson, and flat. Metacoxal face densely, finely punc- A. floridae (Horn). The male genitalia are tate except fossula glabrous; hind leg as in similar in form in species included in this figure 363; pecten with one small denticle group, and some species are difficult to and one or two minute denticles; metatibia distinguish. (figure 363) dilated toward apex, lateral, ventral, and dorsomesal carinae complete, Nelson and Johnson (1983a,b) described ventrolateral carina evanescent apically; the effect of selenium on this bruchid feed- mucro acute, slightly curved, with deep ing in three species of Astragalus. sinus at base, one-third length of basitarsus, lateral denticles and coronal denticles Acanthoscelides baboquivari Johnson subequal in size. Male with small tuft of setae in middle of 1st abdominal sternum; (Figures 362–366) pygidium strongly convex in male, slightly Acanthoscelides baboquivari John- convex in female. son 1974:268; Center and Johnson Male genitalia.—As in figures 365 and 366; 1976:196; Johnson 1976b:260; John- median lobe not constricted at middle; son and Kingsolver 1982:414; John- ventral valve transverse with apical nipple; son 1983a:40; Udayagiri and Wadhi armature of internal sac consisting of clus- 1989:38. ter of spicules at apical orifice, basal one- Color.—Body black except reddish yel- third of sac lined with fine denticles and low or piceous as follows: ventral sides of a slender, serrate sclerite, middle of sac 1st to 4th antennal segments, postocular armed with two pairs of thornlike spines, spot on head, occasionally on each lateral of which one pair is setose at basal end, one-third of pronotal disk, often with broad a curvate, spinose median sclerite, and a median stripe on each elytron, fore legs, pair of lightly sclerotized structures; apex distal one-half of mesofemur, mesotibia, with elongate, bifid closure valve; lateral and metatarsus, often on middle of py- lobes flat (figure 366), cleft about one-half gidial disk. Vestiture of slender, white setae their length, setose on mesal faces and on evenly distributed over body except often apices. condensed into faint median stripe on pronotal disk; elytral interstices regularly Size.—Body length 1.7–2.2 mm; width interrupted by small, rounded, setiferous 0.9–1.4 mm. spots; pygidium with faint to obvious basal Type depository.—USNM, holotype No. triangle and median stripe; mesepimeron 71401. and metepisternum usually prominently Type locality.—Arizona, Pima Co., Kitt setose. Peak, 5,500 ft. Structure.—Vertex finely, densely punc- Distribution.—AZ; Mexico. tate, intervals minutely granulose, frons often with median impunctate line, setae in Host plants.—Indigofera platycarpa, I. circular pattern; ocular index 2.5:1; ocu- sphaerocarpa. lar sinus about two-thirds length of eye; Natural enemies.—None recorded. antenna subserrate (figure 364), reaching middle of metepisternum. Pronotum

101 Immatures.—Pfaffenberger and Johnson those of female; antenna (figure 369) with 1976 (first larval instar). 5th to 10th segments eccentric, 11th el- Discussion.—Johnson (1983a) tentatively liptical, those of male reaching metacoxa, placed this species in his flavescens group. those of female reaching middle of met- Salient characters are the black body episternum; Pronotum bell-shaped (figure with red elytral stripe, ocellate interstices; 367), lateral margins strongly sinuate, curved, slender mucro with a deep sinus at disk evenly convex, without depressions or its base; dark red elytral stripes; and red protuberances except shallow sulcus on legs. It resembles Stylantheus macrocerus basal lobe; disk densely, evenly microfove- but the black body and appendages of that olate. Scutellum small, quadrate, densely species, its elongated antennal segments, pubescent. Elytra evenly convex, widest and the male genitalia easily distinguish at middle (figure 367); striae sinuate in the species. middle of elytra, 2nd and 3rd bent laterad near base, 3rd and 4th approximate at Acanthoscelides bisignatus (Horn) base, each with prominent black denticle; interstices flat, imbricate. Metacoxa evenly (Figures 367–371) punctate except fossula glabrous; hind leg Bruchus bisignatus Horn 1873:334; Fall as in figure 368; pecten with three slender and Cockerell 1907:201; Schaeffer denticles; metatibia arcuate basally, with 1907:302; Fall 1910:170,174; Cush- lateral, ventral, and dorsomesal carinae man 1911:505; Pic 1913a:19; Bridwell complete, ventrolateral carina evanescent 1918:479; Zacher 1952:461. near apex of tibia; mucro short, slender, one-tenth length of basitarsus, lateral Mylabris bisignatus: Leng 1920:305. denticle and three coronal denticles min- Acanthoscelides bisignatus: Glick 1939:36; ute or absent. Fifth abdominal sternum of Peck 1963:956; De Luca 1965:53; male broadly emarginate to receive apex of Bottimer 1968c:1017,1038; Johnson pygidium, that of female evenly rounded; 1968:1267, 1969c:55, 1970:37; John- pygidium of both sexes convex, apex more son and Kingsolver 1982:414; Udayagiri reflexed in male. and Wadhi 1989:40. Male genitalia.—As in figures 370 and 371; Color.—Body mostly black except greater median lobe slender, not constricted at part of elytra, 1st to 5th and 11th seg- middle; ventral valve ogival with apex at- ments of antenna, fore and middle legs, tenuate; internal sac armature consisting and apical one-half of metafemur and all of fine, acute denticles and truncated den- of tibia and tarsus red; mouth parts dark ticles in basal one-half, middle of sac with brown. Vestiture of white, dark brown, small, thornlike denticles of several sizes, and brassy setae; head black with reddish apex of sac with cluster of fine spicules; postocular spot; pronotum with intermixed closure valve ringlike; lateral lobes slender white and brassy setae and paired dark (figure 371), apically spatulate, cleft about brown spots; elytra with sutural interstice one-half their length. brassy, remainder with intermixed white and brassy, 3rd interstice and apex with Size.—Body length 1.75–2.00 mm; width prominent white spots, lateral marginal 1.1–1.3 mm. dark brown spots prominent; pygidium Type depository.—MCZC, lectotype No. with basal triangle and lateral patches 8203, by Johnson 1968. white, remainder brassy. Type locality.—Kansas. Structure.—Vertex and frons densely punctate, frons with prominent protuberance Distribution.—KS, LA, NM, OK, SD, TX; and median carina; ocular index 5:1 in Mexico. male, 4:1 in female; ocular sinus about one-half length of eye, male’s eyes larger

102 Host plants.—Baptisia sp.; Desmanthus sp. patch; pronotum white with paired, medi- D. illinoensis, D. leptolobus, D. velutinus, D. an, dark brown stripes separated by nar- virgatus virgatus. row white stripe; elytra intermixed white Natural enemies.—Catolaccus hunteri; Eu- and brassy except brown spots as follows: pelmus cyaniceps; Heterospilus proso-pidis; paired spots near scutellum, paired lat- Pteromalus piercei. eral marginal spots at middle of margin, small paired spots on 3rd interstices, api- Immatures.—Not described. cal patches of variable size; pygidium with Discussion.—This species is nearly in- narrow, median, white stripe; ventral areas separable from Acanthoscelides desmanthi of body evenly white except metepisternum Johnson except by comparing the male and lateral margin of abdomen with white genitalia. The head of A. bisignatus is patches. mostly black whereas that of A. desman- Structure.—Vertex densely punctulate; thi is reddish orange with a broad, black frons with median impunctate line; ocular stripe on the vertex and frons. The ves- index 2.5:1; ocular sinus one-half length titure of A. desmanthi is brighter when of eye; antenna as in figure 374, reaching examples of the two species are compared. middle of metepisternum. Pronotum bell- Each species has sexually dimorphic eyes shaped (figure 372), lateral margins nearly and antennae, red terminal antennal straight, apex semicircular; disk evenly segments, and a prominent frontal carina. convex, densely, uniformly microfoveolate. Both species belong to the mexicanus spe- Scutellum quadrate, densely pubescent. cies group according to Johnson (1983a). Elytra convex, widest at apical one-third The geographical range of A. desmanthi (figure 372); striae parallel, narrow, base of (south Texas to Chiapas, Mexico) is con- 4th with small denticle; interstices flat, mi- current with that of Desmanthus virgatus, nutely imbricate. Metacoxa densely, evenly whereas that of A. bisignatus coincides punctate except fossula glabrous; hind closely that of Desmanthus illinoensis. leg as in figure 373, pecten with one long and two shorter denticles; metatibia (figure Acanthoscelides biustulus (Fall) 373) slightly arcuate in basal one-fourth, all carinae complete, mucro curved, about (Figures 372–376) four-tenths as long as basitarsus, sinus Bruchus biustulus Fall 1910:178; Pic at base of mucro deep, lateral denticle and 1913a:19. coronal denticles of equal size. First ab- Mylabris biustulus: Leng 1920:305. dominal sternum of male with small white tuft of setae; pygidium of both sexes con- Acanthoscelides biustulus: Johnson vex, male with apex reflexed. 1968:1267; Bottimer 1968c:1017,1038; Johnson 1970:39; Slobodchikoff and Male genitalia.—As in figures 375 and 376; Johnson 1973:282; Center and John- median lobe broad; ventral valve ogival, son 1976:196; Johnson 1976b:260, acute; armature of internal sac consist- 1977a:64, 1979b:62, 1983a:46; John- ing of angulate cluster of fine spicules son and Kingsolver 1982:414; Udayagiri at apical orifice, a large, pointed sclerite, and Wadhi 1989:40. a median row of thornlike sclerites, two curved, acute sclerites, and apex lined with Color.—Body and appendages black except fine spicules; closure valve circular; lateral apex of fore and middle femur, all of tibia lobes broad (figure 376), truncate, setose and tarsus and basal four segments of an- on mesal faces and on apical borders, cleft tenna red. Vestiture of white, dark brown, about two-thirds of their length. and brassy setae; head sparsely set with white but with intensely white postocular Size.—Body length 1.2–2.3 mm; width 1.0–1.5 mm.

103 Type depository.—MCZC, holotype No. postocular fringe prominent; antenna as 25047. in figure 379; extending to middle of met- Type locality.—Arizona, Santa Rita Moun- epi-sternum. Pronotum subconical, lateral tains. margins straight or slightly incurved (figure 377), disk evenly convex with slight swell- Distribution.—AZ, NM, TX; Mexico. ing anteriad to basal lobe; surface finely, Host plants.—Desmodium sp. D. bato-cau- densely punctulate, each puncture setifer- lon, D. cinerascens, D. grahamii, D. neomex- ous; cervical sulcus prominent; lateral ca- icanus. Johnson (1983a) lists other species rina evanescent. Scutellum subtriangular, of Desmodium in Mexico for this bruchid. slightly wider than long, densely pubescent, prominent. Elytra (figure 377) evenly Natural enemies.—None recorded. convex, humeri prominent; striae parallel Immatures.—Not described. except 2nd bent at base to approach 3rd; Discussion.—Johnson (1983a) places 2nd to 6th striae each with minute basal Acanthoscelides biustulus in the pertinax denticle; interstices flat, minutely imbri- species group. Salient characters are the cate. Metacoxa densely punctulate; hind elongated, curved mucro, red apices of fe- leg as in figure 378; metatibia with lateral, mur and tibia, maculate pattern on elytra, ventral, and dorsomesal carinae complete, fourth stria with prominent basal denticle, lateral and ventrolateral carinae evanes- and male with small, white tuft on 1st cent, mucro slender, curved, one-third as abdominal sternum and tibial carinae all long as basitarsus, lateral denticle slender, present. coronal denticles two or three. First abdominal sternum convex in male but not Acanthoscelides calvus (Horn) modified, 5th male sternum broadly emarginate. (Figures 377–381) Male genitalia.—As in figures 380 and 381; Bruchus calvus Horn 1873:336; Hub- median lobe broad; ventral valve sharply bard and Schwarz 1878:660; Blatch- triangular; dorsal valve evenly rounded ley 1910:1239; Fall 1910:173; Pic with cluster of spicules; armature of inter- 1913a:20; Olsen 1918 (misidentification nal sac consisting of clusters of acute den- of Bruchidius villosus). ticles in basal one-half, two pairs of large, Mylabris calvus: Leng 1920:305. thornlike sclerites, and a spiny structure Acanthoscelides calvus: Bridwell 1925:80; near apex; closure valve circular; lateral Bradley 1947:40; Blackwelder and lobes slender (figure 381), moderately Blackwelder 1948:45; Bottimer spatulate, cleft nearly to base. 1968c:1018; Johnson 1968:1267, Size.—Body length 2.25–3.25 mm; width 1969c:55; Johnson and Kingsolv- 1.5–2.0 mm. er 1982:414; Udayagiri and Wadhi Type depository.—MCZC, lectotype No. 1989:40. 3891, by Johnson 1968. Color.—Body and appendages uniformly black, fore and middle legs occasionally Type locality.—Massachusetts (lectotype). dark brown. Vestiture white, uniformly Distribution.—CT, IL, IN, MA, MD, MI, NC, distributed over body, except scutellum NH, NJ, NY, ON, PA, SC, TN, TX, VT, WI. prominently white. Host plants.—Helianthemum canadense; on Structure.—Head elongate; vertex and frons Aronia flowers. sparsely, evenly punctulate, interspaces Natural enemies.—None recorded. minutely granulate, frons with median carina barely indicated; ocular index 3:1; Immatures.—Not described. ocular sinus three-fourths length of eye, Discussion.—This species will most likely be confused with Acanthoscelides

104 fraterculus with its comparable size, black prominent raised denticle; interstices flat, body and legs usually black, but the prom- minutely imbricate. Metacoxa densely inent white scutellum of A. calvus should punctate and pubescent in lateral one-half, immediately separate them. The 1st ster- median one-half with fine, radiating strigae num of male A. calvus is convex whereas bisected by arcuate ridge (figure 384); hind that of A. fraterculus is concave with a leg as in figure 383; mesoventral face of fringe of long hair along the posterior bor- femur with fine carina (figure 386), possible der of the segment. The male genitalia of scraper over metacoxal ridges; metatibia A. calvus are distinctive and are unlike any (figure 383) arcuate in basal one-third, other eastern species. straight apically; lateral, ventrolateral, ventral, and dorsomesal carinae complete; Acanthoscelides chiricahuae (Fall) mucro short, scarcely longer than lateral denticle, two or three coronal denticles. (Figures 382–388) Male abdomen with sterna telescoped, 5th Bruchus chiricahuae Fall 1910:181. sternum nearly divided by emargination; Bruchus chiracahuae: Pic 1913a:21 (mis- pygidium of both sexes finely punctate, spelling). apex of male strongly reflexed, female py- Mylabris chiricahuae: Leng 1920:305. gidium subvertical. Acanthoscelides chiricahuae: Johnson Male genitalia.—As in figures 387 and 388; 1968:1267; Bottimer 1968c:1018,1038, median lobe slender, slightly constricted 1969b:1187; Johnson 1970:41; Slo- near apex; ventral valve ogival, apex slight- bodchikoff and Johnson 1973:282; ly produced; internal sac densely lined with Center and Johnson 1976:196; Pfaffen- small acute and truncate denticles; two berger and Johnson 1976:24; Ward arcuate, apical sclerites, each with five to et al. 1977:4; Johnson 1979b:62, eight serrations; lateral lobes bowed (fig- 1981b:76; Johnson and Kingsolver ure 388), deeply cleft, arms narrow, apices 1982:414; Johnson 1983a:57; Pfaffen- spatulate. berger 1985b:256; Udayagiri and Wadhi Size.—Body length 1.7–2.4 mm; width 1989:42. 1.1–2.4 mm. Color.—Body black except fore legs, apical Type depository.—MCZC, holotype No. one-half of middle femur, and all of middle 25049. tibia; antenna reddish orange to piceous with apical segment reddish brown. Vesti- Type locality.—Arizona, Chiricahua Moun- ture of fine white setae densely, uniformly tains. distributed on body, pygidium sometimes Distribution.—AZ, NM, TX; Mexico. with median condensed line. Host plants.—Mimosa biuncifera, M. borea- Structure.—Vertex and frons micropunc- lis, M. dysocarpa, M. grahamii, M. laxiflora, tate, densely pubescent; frontal carina M. wherryana. Johnson (1983a) lists five evident, transverse sulcus shallow; ocular additional species from Mexico. Pfaffen- index 3.5:1 in male, 3:1 in female; ocular berger (1985c) lists Acacia greggi. sinus three-fifths length of eye; postocu- Natural enemies.—Eupelmus sp.; Horis- lar fringe narrow but prominent; antenna menus missouriensis; Urosigalphus bruchi- as in figure 385, reaching middle of me- vorus; Zatropis incertus. tepisternum. Pronotum bell-shaped (figure 382), disk strongly convex, lateral margins Immatures.—Pfaffenberger and Johnson arcuate, lateral carina evanescent, cervi- 1976:24 (first larval instar). cal sulcus prominent. Elytra convex with Discussion.—Distinguishing characters lateral margins arcuate (figure 382); striae for this species are the evenly distributed parallel except 2nd and 3rd deflected lat- body setae, the prominent raised denticles erad at base, 3rd and 4th each with at the bases of striae three and four, the

105 telescoped abdominal sterna, and the some specimens; ocular index 3:1; ocular radiating lines on the metacoxal face. This sinus about three-fourths length of eye; species most closely resembles Acantho- postocular lobe with narrow fringe; an- scelides schrankiae, a species found from tenna sexually dimorphic as in figures 391 eastern United States to Venezuela (John- and 392, in the male reaching 1st abdomi- son 1983a). The metacoxa of A. chiricahuae nal sternum, in female middle of metepi- is finely striate in a radiating pattern with sternum. Pronotum trapezoidal (figure a low ridge bisecting the radiating lines, 389), disk strongly convex, microfoveolate, whereas in A. schrankiae the metacoxa is lateral margins straight, lateral carina simply punctate. Each of the serrate api- absent; cervical sulcus deep. Scutellum cal sclerites in the male internal sac (figure elongate, small, densely pubescent. Lat- 387) has five to eight teeth inA. chirica- eral margins of elytra arcuate (figure 389); huae but nine to thirteen in A. schrankiae striae parallel, 2nd to 5th striae minutely (figure 602). denticulate basally; interstices flat, punctation concealed by pubescence. Metacoxal Acanthoscelides compressicornis face evenly punctate except for narrow fos- (Schaeffer) sula; hind leg as in figure 390; metatibia nearly straight with lateral, ventrolateral, (Figures 389–394) ventral, and dorsomesal carinae complete, Bruchus compressicornis Schaeffer mucro slender, curved, about two-fifths 1907:305; Fall 1910:188; Cush- length of basitarsus. Male abdomen with man 1911:506; Pic 1913b:23; Zacher basal sternum unmodified, 5th sternum 1952:461,471. shallowly emarginate; female 5th sternum Mylabris compressicornis: Leng 1920:306. not modified; pygidium convex in both Bruchus subserripes Fall 1910:183; Pic sexes, disk with punctures concealed by 1913a:51. vestiture, apex in male reflexed apically. Acanthoscelides compressicornis: Male genitalia.—As in figures 393 and Muesebeck et al. 1951:509; Peck 394; median lobe slender, elongate; ventral 1963:965; De Luca 1965:54; Bottimer valve semicircular, apex subtruncate; basal 1968c:1018,1038; Johnson 1968:1267; two-thirds of internal sac lined with trun- Bottimer 1969a:977; Johnson 1970:46; cate denticles, middle of sac with clusters Slobodchikoff and Johnson 1973:282; of acute denticles, apex with three pairs of Center and Johnson 1974:1097, thornlike spines; lateral lobes (figure 394) 1976:260; Johnson 1977a:70, fused from base for most of their length, 1981d:74; Johnson and Kingsolver apices divergent, setose. 1982:414; Johnson 1983a:63; Luck- Size.—Body length 1.3–2.3 mm; width ow and Johnson 1987:49; Johnson 0.9–1.4 mm. 1990e:324. Type depository.—USNM (compressicornis, Acanthoscelides subserripes: Bottimer lectotype No. 42348, by Johnson 1968); 1968c:1018,1038; Johnson 1968:1268, MCZC (subserripes, holotype No. 25063). 1970:46; Udayagiri and Wadhi 1989:43. Type locality.—Texas, Brownsville (com- Color.—Body and appendages black except pressicornis); Texas, Ysleta (subserripes). fore and middle legs occasionally reddish. Vestiture of fine white setae uniformly dis- Distribution.—AR, AZ, FL, IA, KS, LA, MO, tributed over body except basal margin of NE, NM, OH, OK, SD, TX; Mexico, Guate- pronotum with small patch opposite base mala. of 5th interstice and faint median line of Host plants.—Desmanthus sp., D. cooleyi, setae on pygidium. D. covillei, D. illinoensis, D. leptolobus, D. Structure.—Vertex and frons punctulate, leptophyllus, D. obtusus, D. subutatus, D. setae directed toward center of vertex; frontal carina usually evident but faint in 106 velutinus, D. virgatus acuminatus, D. virga- Structure.—Vertex and frons punctate- tus depressus, D. virgatus glandu-losus, D. granulose, frontal carina absent; ocular virgatus virgatus; Hoffmanseggia densiflo- index 5:2; ocular sinus one-half length ra, H. drepanocarpa, H. glauca, H. tenella; of eye; antenna (figure 398) extending Mimosa dysocarpa, M. grahamii. John- to middle of metepisternum. Pronotum son (1977a) lists Desmanthus covillei and bell-shaped (figure 395), lateral margins (1983a) D. subulatus from Mexico. slightly sinuate, apex round; disk strongly convex, posterior angles and middle of Natural enemies.—Catolaccus hunteri; Eu- basal lobe depressed, surface densely fo- pelmus cyaniceps; Heterospilus prosopidis; veolate, covered by vestiture; lateral carina Pteromalus piercei. ridgelike, abbreviated; cervical sulcus well Immatures.—Not described. defined. Scutellum one and one-half times Discussion.—Salient characters of this as long as wide, densely setose. Elytra 1.1 species are the black coloration (except times as long as wide (figure 395), evenly basal antennal segments), strial denticles convex; striae narrow, subparallel, hid- at extreme base, slender, curved mucro, den in vestiture, lacking basal denticles; the greatly enlarged, triangular antennal interstices minutely imbricate. Metacoxal segments of the male, and the distinctive face uniformly punctulate except fossula male genitalia. From other all-black spe- glabrous; hind leg as in figure 397; pecten cies in its range, it may be distinguished with three small denticles; metatibia with as follows: A. perforatus by coarse strial lateral, ventral, and dorsomesal carinae punctures and lobate 5th male sternum; A. complete, ventrolateral abbreviated; mucro calvus by the elongate head; A. fraterculus acute, three-tenths as long as basitarsus; by the more elongate body and lobate male lateral denticle and one coronal denticle 5th sternum; A. fumatus by the broad body subequal, dorsal coronal denticle more and distinctive male genitalia. slender, acute. Male basal abdominal segment with small pit, setae arranged Johnson tentatively places this species in in circular pattern, 5th segment broadly his flavescens group. Blatchley (1919:66) emarginate in male, truncate in female; misidentified Florida specimens ofA. heli- male pygidium evenly convex (figure 396), anthemum Bottimer as A. compres-sicornis, female pygidium vertical at apex. but in A. helianthemum the elytral vestiture is mottled similarly to A. alboscutellatus Male genitalia.—As in figures 399 and 400; (Horn). median lobe moderately broad, expanded at apex; ventral valve broadly triangular, Acanthoscelides comstocki Johnson basal angles rounded; armature of internal sac consisting of thornlike spine near base, (Figures 395–400) elongate, serrate sclerite in middle, and Acanthoscelides comstocki Johnson symmetrical pair of elongate, setose scler- 1990a:8. ites near apex; closure valve C-shaped; Color.—Integument black except most of lateral lobes cleft one-third their length elytra, fore and middle legs, apical three- (figure 400), apical lobes obliquely trun- fourths of metafemur, and all of metatibia cate, setose. red. Vestiture mostly white except vari- Size.—Body length 1.3–2.0 mm; width able pronotal stripe golden, elytral margins 0.8–1.2 mm. white, middle and apical one-half yellow Type depository.—CNCI. with reddish-brown lateral maculae, 3rd interstice with anterior and posterior red- Type locality.—Texas, Comstock. dish-brown spots delimiting short, white Distribution.—TX. elongated spot; head, venter of body and pygidium uniformly white. Host plants.—Eysenhardtia texana.

107 Natural enemies.—None recorded. nae complete, ventrolateral carina present Immatures.—Not described. in basal one-half, mucro about one-sixth as long as basitarsus, lateral denticle and Discussion.—Johnson placed this species two coronal denticles slender, acute. First in the puellus group. It most closely re- abdominal sternum of male with median sembles Acanthoscelides pallidipennis in basal pit set in posterior margin of 1st the armature of the internal sac, the pit sternum lobed with fringe of long hairs; on the 1st abdominal segment, and dorsal 5th sternum broadly emarginate; female coloration. lacking modifications of basal sternum, apical margin evenly rounded; male pygidium strongly convex, reflexed apically, Acanthoscelides daleae Johnson that of female moderately convex; discal (Figures 401–405) surface in both sexes densely punctulate, concealed by pubescence. Acanthoscelides daleae Johnson 1970:48; Center and Johnson 1976:196; John- Male genitalia.—As in figures 404 and 405; son and Kingsolver 1982:414; Udayagiri median lobe slender, slightly constricted and Wadhi 1989:44. near apex; ventral valve broadly triangu- Color.—Body and appendages reddish lar; internal sac lined with fine, transverse orange to black; in black specimens, basal clusters of minute setae (see inset, figure two segments of antenna, basal one-half 404), middle of sac with large irregular of fore femur and mid femur, and fore tibia sclerite, apex with paired, slender, spinose and mid tibia reddish; in more reddish structures; closure valve circular; lateral specimens, all appendages reddish orange. lobes fused for nearly their entire length Vestiture of fine, white setae densely but (figure 405), cleft only at apices, apices and uniformly distributed over body giving a mesal faces of lobes setose. pruinose sheen to body. Size.—Body length 1.5–1.75 mm; width Structure.—Vertex and frons densely 1.0–2.0 mm. punctulate, pubescence sparse, frons with Type depository.—USNM, holotype No. impunctate median line; ocular index 3:1; 69686. ocular sinus one-half length of eye, postoc- Type locality.—California, San Bernadino ular lobe narrow; antenna as in figure 403, Co., 8 mi west of Needles. extending to middle of metepister-num. Pronotum bell-shaped (figure 401), lateral Distribution.—AZ, CA, NV, UT; Mexico (Baja margins slightly arcuate; disk evenly con- California). vex except slightly depressed near poste- Host plants.—Dalea sp.; Psorothamnus fre- rior corners, finely sulcate on basal lobe; monti, P. schotti, P. spinosa. surface microfoveolate, foveolae nearly hid- Natural enemies.—Heterospilus prosopidis. den by pubescence; lateral carina marked by slight basal ridge; cervical sulcus well- Immatures.— Not described. defined, deep. Scutellum 2 times as long Discussion.—The extensive range of size as wide, bidentate apically. Elytra moder- and integumental coloration and its re- ately convex (figure 401), lateral margins semblance to Acanthoscelides fraterculus arcuate; striae parallel except 2nd slightly makes this species difficult to character- divergent at base approaching 3rd, 2nd to ize. Johnson (1970) described it as a sepa- 6th striae each with minute basal denticle; rate species entity based principally on interstices flat, minutely imbricate. Meta- its restricted geographical range and host coxa densely, minutely punctate except for plants. The male genitalia are identical to smooth fossula; hind leg as in figure 402; those of A. fraterculus. The elongate scutel- metatibia straight, slightly dilated toward lum, the uniformly gray, pruinose appear- apex, lateral, ventral, and dorsomesal cari- ance, the basal abdominal pit of the male,

108 and the host association are key charac- sternum of male not modified, 5th ster- ters. num slightly emarginate in male, arcuate in female; male pygidium slightly reflexed Acanthoscelides desmanthi Johnson apically. (Figures 406–410) Male genitalia.—As in figures 409 and 410; median lobe slender, elongate; ventral Acanthoscelides desmanthi Johnson valve concave (figure 409a), apex attenu- 1977a:64, 1983a:71; Johnson and ate; armature of internal sac consisting of Kingsolver 1982:414; Luckow and an elongate cluster of denticles near base, Johnson 1987:49; Udayagiri and Wadhi a cluster of elongate rodlike sclerites in 1989:44; Johnson 1990e:324. middle, a cluster of fine, acute denticles, Color.—Body dark red to orange; head dark and seven thornlike spines; apex finely red with vertex and frons black; pronotum spiculate, closure valve circular; lateral mostly red with darker maculations; elytra lobes (figure 410) extremely long, slender, reddish brown with darker red spots and cleft full length, apices scarcely expanded. maculations; pygidium dark red, venter of body black; antenna with basal four and Size.—Body length 1.5–2.1 mm; width terminal segments black; legs red except 0.9–1.2 mm. proximal one-half of metafemur black. Type depository.—USNM, holotype No. Vestiture of white, yellow, dark brown, and 72782. black hairs in dorsal pattern (figure 406), Type locality.—Mexico, Chiapas, 3.5 mi legs and venter of body uniformly white; west of Ocozocoautla. male pygidium uniformly yellow, female pygidium with white basal and lateral Distribution.—TX; Mexico. patches. Host plants.—Desmanthus sp., D. bicor- Structure.—Eye and antenna dimorphic; nutus, D. covillei, D. leptophyllus. Johnson ocular index 6:1 in male, 4:1 in female; (1983a) also lists Desmanthus subulatus, frontal boss prominent, frontal carina hid- D. virgatus virgatus, and D. virgatus de- den by vestiture; ocular sinus two-thirds pressus from Mexico. as long as eye; male lacking postocular Natural enemies.—None recorded. lobe, female with lobe narrow; antenna (figure 408) serrate, reaching 1st abdominal Immatures.—Not described. segment in male, reaching metepisternum Discussion.—This species is closely related in female. Pronotum bell-shaped (figure to Acanthoscelides bisignatus, and the two 406), unevenly convex, lateral margins can only be consistently separated by com- sinuate, apex slightly constricted; disk parison of the male genitalia (figures 370 microfoveolate, lateral carina evanescent. and 410). The antennae and eyes of both Scutellum small, quadrate, bidentate. species are dimorphic and the color pat- Elytra convex, lateral margins subparal- terns are similar. Acanthoscelides specio- lel (figure 406); striae parallel, sinuate in sus resembles A. desmanthi and A. bisigna- middle, 2nd, 3rd and 4th bent laterad at tus but its eyes are not dimorphic and its base, 3rd and 4th with basal denticles. antennae are consistently and entirely red. Metacoxa finely, evenly punctulate except Acanthoscelides speciosus breeds only in fossula glabrous; hind leg as in figure 407; Mimosa species. metatibia slightly arcuate, dilated toward apex; lateral, ventrolateral, ventral, and Acanthoscelides distinguendus (Horn) dorsomesal carinae complete; mucro short, (Figures 411–415) one-third as long as basitarsus, lateral denticle nearly as long as mucro; two Bruchus distinguendus Horn 1873:336; coronal denticles present. First abdominal Hubbard and Schwarz 1878:660;

109 Schwarz 1878:457; Blatchley acute, about one-fourth as long as basitar- 1910:1238; Fall 1910:171; Cush- sus, lateral denticle separated from mucro man 1911:499; Pic 1913a:24; Zacher by deep sinus, lateral and coronal denticles 1952:462. subequal. Fifth abdominal segment deeply Mylabris distinguendus: Leng 1920:305. emarginate; pygidium in both sexes convex, reflexed apically; disk densely microfo- Acanthoscelides distinguendus: Bissell veolate. 1940:846; Bottimer 1968c:1018,1038; Johnson 1968:1267; Bottimer Male genitalia.—As in figures 414 and 415; 1969b:1187; Johnson 1969a:279, median lobe slender, slightly expanded 1969c:55; Kingsolver 1969b:51,54; at apex; ventral valve transverse, arcuate Johnson 1970:18, 1979b:63, 1981a:79; either side of small, median nipple; arma- Johnson and Kingsolver 1982:414; ture of internal sac consisting of fine, acute Johnson 1983a:80; Udayagiri and Wa- denticles in basal one-half, a small cluster dhi 1989:45. of larger denticles at middle, two thornlike Color.—Body mostly dark brown, legs red- sclerites, and subapical, elongate cluster of dish brown, antenna with basal four seg- fine spicules, closure valve circular; lateral ments reddish yellow, remaining segments lobes (figure 415) narrow, elongate, cleft black. Vestiture of white and brown slender nearly to base, apices spatulate, setose. hairs uniformly distributed on body except Size.—Body length 1.9–2.6 mm; width in pattern of alternating brown and white 1.3–1.8 mm. elongate patches on elytral striae, prono- Type depository.—MCZC, lectotype No. tum with broad, dark brown median stripe 8206, by Bottimer 1968b. and lateral white spots (figure 411); pygidium with evanescent median line of hairs. Type locality.—Georgia. Structure.—Vertex and frons densely Distribution.—AL, FL, GA, IN, LA, MI, MS, punctulate, setae directed toward pol- NC, OK, SC, VA, TX; Mexico. ished median boss, frontal carina promi- Host plants.—Mimosa strigillosa; Rhyn-cho- nent; ocular index 3:1; ocular sinus about sia americana, R. difformis, R. erecta, R. three-fifths length of eye; antenna sexually galactoides, R. intermedia, R. latifolia, R. to- dimorphic (figure 413), reaching 1st ab- mentosa. Blatchley (1910) recorded “Crac- dominal sternum in male, reaching middle ca” virginiana (now placed in Tephrosia), of metepisternum in female. Pronotum but this is undoubtedly a misidentification bell-shaped (figure 411), lateral margins of the bruchid, probably of Acanthoscelides straight to slightly arcuate, disk evenly obsoletus (Say). convex, densely microfoveolate, foveolae nearly hidden by vestiture; lateral carina Natural enemies.—None recorded. present only in basal one-third; cervi- Immatures.—Not described. cal sulcus well-marked, deep. Scutellum Discussion.—Salient characters for this one-third longer than wide, bidentate api- species are the brown and white mottled cally. Elytra evenly convex, lateral margins pattern of the pronotum and elytra, the arcuate (figure 411); striae slightly sinu- strongly serrate antenna of the male, the ate in middle, 3rd and 4th striae basally sharp frontal carina, and the distinctive denticulate; interstices flat, finely imbri- male genitalia. This species most closely cate, densely pubescent. Metacoxa densely resembles Acanthoscelides obsoletus but punctulate except fossula polished; hind the following differences apply (A. obsoletus leg as in figure 412; metatibia slightly bent characters in parentheses): Striae 3 and 4 basally, straight and slightly dilated api- lacking distinct strial denticles (prominent cally, with lateral, ventrolateral, ventral, basal denticles); head unicolorous behind and dorsomesal carinae complete; mucro eye (red spot); mucro one-fourth length of basitarsus (one-third length); 1st abdomi-

110 nal sternum of male unmodified (median white spot on 3rd interstice delimited by sulcus and disarrangement of setae). There brown spots; pygidium with faint white are also differences in the male genitalia median stripe and lateral median brown (compare figures 414 and 520).Acan - spots. tho-scelides obtectus has a similar dorsal Structure.—Vertex densely punctulate, pattern to those of A. distinguendus and frons sparsely punctulate, intervals granu- A. obsoletus, but its abdomen is mostly or lose, frontal carina prominent; ocular index entirely red and the apical antennal seg- 3:1; ocular sinus about one-half length of ment is also red. The male genitalia of A. eye; antenna (figure 418) reaching middle obtectus is distinct from either A. obsoletus of metepisternum. Pronotum bell-shaped or A. distinguendus, but the general form (figure 416), lateral margins slightly arcu- of A. distinguendus relates A. distinguen- ate, disk evenly convex with slight depres- dus to A. flavescens. sion each side of basal lobe; lateral carina evanescent; cervical sulcus distinct, deep. Acanthoscelides flavescens (Fahraeus) Scutellum quadrate, emarginate apically, (Figures 416–420) densely pubescent. Elytra together as long as wide (figure 416), lateral margins arcu- Bruchus flavescens Fahraeus 1839:32; Pic ate, disk evenly convex; striae parallel, 1913a:25. indistinct, not deflected at base, 3rd, 4th, Acanthoscelides flavescens: Blackwelder and 5th each with minute basal denticle; 1946:759; Kingsolver 1975a:60; Jan- interstices flat, densely pubescent, minute- zen 1980:941,946; Johnson 1981b:79; ly imbricate. Metacoxa densely punctulate Johnson and Kingsolver 1982:414; except fossula smooth; hind leg as in figure Johnson 1983a:85; Udayagiri and Wa- 417, metatibia with lateral, ventral, and dhi 1989:46. dorsomesal carinae complete, ventrolateral Bruchus ochraceus Schaeffer 1907:303; carina abbreviated apically; mucro acute, Fall 1910:170; Cushman 1911:501,507. slender, one-third as long as basitarsus, Bruchus ochraceicolor Pic 1913b:110 (new separated from acute lateral denticle by name for ochraceus Schaeffer 1907 not deep sinus, dorsal coronal denticles sepa- Baudi 1886b); Pic 1913a:38; Zacher rated by deep sinuses. First abdominal 1952:462. sternum of male smooth, 5th deeply emarginate; 5th of female shallowly emarginate; Mylabris ochraceicolor: Leng 1920:305. pygidium of both sexes strongly convex, Acanthoscelides ochraceicolor: Wolcott punctation nearly hidden by pubescence. 1936:286; Moreno and Bibby 1943:23; Blackwelder 1946:760; Zacher Male genitalia.—As in figures 419 and 1952:462,475; Peck 1963:956; De Luca 420; median lobe moderately broad; ven- 1965:55; Bottimer 1968c:1019,1039; tral valve transverse, broadly arcuate with Kingsolver 1969b:51; Johnson 1970:19; median nipple; armature of internal sac Forister and Johnson 1971:232; De consisting of an elongate cluster of min- Luca 1972:106; Kingsolver 1975a:60. ute denticles in basal one-third, a pair of small, curvate sclerites in middle, a spines- Color.—Body and appendages reddish cent structure and C-shaped closure valve brown except thoracic sterna sometimes in apical one-third; lateral lobes slightly piceous, apical margin of each antennal bowed (figure 420), cleft three-fourths their club segment dusky. Pubescence of white, length. golden, and brown hairs in pattern shown in figure 416, the white and golden hairs Size.—Body length 1.5–2.5 mm; width intermixed over most of body with patches 0.9–1.5 mm. of white on lateral portions of pronotum and stripes and spots on elytra, elongate

111 Type depository.—NHRS (flavescens); sometimes extending as stripe along elytral USNM (ochraceus, lectotype No. 42345, by suture, 1st and sometimes 2nd abdominal Johnson). sterna, base of metafemur; antenna vary- Type locality.—St. Vincent Is. (flavescens); ing from red to black. Vestiture of slender, Texas, Brownsville (ochraceus). white hairs in pattern on elytra and pygidium, intermixed with dark brown and yel- Distribution.—FL, LA, TX; Mexico to low hairs on pronotum and with indistinct Panama, West Indies, South America. white spots either side of midline, often Host plants.—Abutilon hypoleucum; Ga- with faint median white stripe (figure 421); lactia striata; Rhynchosia minima; Vicia pygidium with basal triangle and median sp. Eriosema violaceum and Rhynchosia line white. longeracemosa are extralimital. Structure.—Vertex and frons densely punct- Natural enemies.—Bracon sp.; Eupelmus ulate, setae directed toward impunctate cushmani, E. cyaniceps; Eurytoma sp.; median knob; frontal carina faintly indicat- Heterospilus prosopidis. ed in some specimens; ocular index 2.5:1; ocular sinus about two-thirds length of Immatures.—Not described. eye; antenna (figure 423) reaching middle Discussion.—The reddish-brown integu- of metepisternum. Pronotum bell-shaped ment, yellowish vestiture, prominent fron- (figure 421), evenly convex, lateral margins tal carina, and male genitalia are charac- gently arcuate, disk densely microfoveo- ters to separate this species. Although A. late; lateral carina obsolete; cervical sulcus distinguendus has a more distinct dorsal well-marked. Scutellum quadrate, densely pattern on a piceous integument, the two pubescent. Elytra together slightly longer species are closely related by host prefer- than wide (figure 421), marginal profile ences and form of the male genitalia (com- continuous with pronotal margin; striae pare figures 414 and 419).Acanthoscelides shallow but well-marked, subparallel, 2nd griseolus is similar in external appearance, to 6th minutely denticulate at base, 3rd, but the principal sclerite of the internal sac 5th, and 7th interstices slightly wider than in that species is forked, the geographic alternates, flat, minutely imbricate. Meta- range is more western, and its host plants coxa densely punctulate except fossula are in the genus Sesbania. glabrous; hind leg as in figure 422; metatibia nearly straight, slightly dilated toward Acanthoscelides floridae(Horn) apex, with lateral, ventral, and dorsome- (Figures 421–425) sal carinae complete, ventrolateral carina basal abbreviated apically; mucro short, Bruchus floridae Horn 1873:332; Schwarz acute, one-fourth as long as basitarsus, 1878:457; Fall 1910:170; Bradley lateral denticle short, as long as 1st coro- 1947:40. nal denticle, dorsal denticle one-half as Mylabris floridae: Leng 1920:305. long as mucro. Male with rounded, setifer- Acanthoscelides floridae: Brett 1946:84; ous pit on 1st abdominal sternum; pygidi- Blackwelder and Blackwelder 1948:44; um strongly convex and apically reflexed in Bottimer 1968c:1018; Johnson male, moderately convex with apex vertical 1968:1267, 1970:14,18; Johnson and in female, surface of pygidial disk microfo- Kingsolver 1982:414; Udayagiri and veolate, densely pubescent. Wadhi 1989:46. Male genitalia.—As in figures 424 and 425; Color.—Body and appendages mostly red median lobe moderately broad, slightly ex- with following areas black: Headexcept panded at apex; ventral valve equila-terally red patch above each eye, pronotum in triangular, tip acute; armature of most specimens sometimes with lateral areas of disk red, ventral sclerites of thorax, semicircular patch around scutellum

112 internal sac consisting of transverse rows Acanthoscelides fraterculus: Trelease of minute spicules in basal one-half of sac, and Trelease 1937a:590, 1937b:448; and a large, blunt sclerite about one-third Bridwell 1938a:76; Bradley 1947:40; length of sac, apical one-half with sym- Blackwelder and Blackwelder 1948:45; metrical, setose structure; closure valve Zacher 1952:465; De Luca 1965:54; C-shaped; lateral lobes fused most of their Bottimer 1968c:1018,1038; Johnson length (figure 425), apical lobes setose. 1968:1267, 1969c:55, 1970:50; Green Size.—Body length 2.0–2.5 mm; width and Palmbald 1975; Center and John- 1.25–1.50 mm. son 1976:196; Johnson and Kingsolver 1982:414; Clement and Miller 1982; Type depository.—MCZC, lectotype No. Udayagiri and Wadhi 1989:47. 8201, by Johnson 1968. Bruchus aureolus: Fall 1910:184 (misiden- Type locality.—Florida (lectotype). tification). Distribution.—DC, IA, IL, IN, FL, KS, MI, Color.—Integument black except occa- MO, MS, NC, ND, NE, NJ, NM, OK, SC, SD, sionally with reddish spots on elytra; legs TX, VA, WY; Mexico, West Indies. black, partly red, or entirely red; basal three antennal segments red beneath. Ves- Host plants.—Amorpha fruticosa fruticosa. titure of fine, slender, gray hairs densely, Natural enemies.—Catolaccus sp.; Eupel- uniformly distributed over body. mus amicus, E. inyonesis; Horismenus mis- Structure.—Vertex and frons densely souriensis. punctulate, interspaces minutely granu- Immatures.—Not described. late, midline of frons with impunctate line; Discussion.—This species closely resembles ocular index 3:1; ocular sinus about two- Acanthoscelides submuticus. The only cer- thirds length of eye; postocular lobe nar- tain method of separating the two species row, fringed with white hairs; antenna as is by examining the male genitalia (com- in figure 429, reaching posterior margin pare figures 424 and 630). The presence or of metepisternum. Pronotum bell-shaped, absence of the metatibial lateral carina is strongly convex (figure 426), lateral mar- variable in A. submuticus, but it is always gins nearly straight, discal surface densely present in A. floridae. The ratio of length to microfoveolate, sculpture nearly hidden by width of the female pygidium is 10:9 in A. vestiture; lateral carina ridgelike, reaching floridae and 1:1 in A. submuticus. nearly to procoxal cavity. Scutellum nearly 2 times as long as wide, bidentate api- Johnson (1970) did not include A. floridae cally. Elytra together slightly longer than in his monograph of the western species of wide (figure 426), lateral margins gently Acanthoscelides but he compared it with A. rounded; striae parallel, conspicuous, submuticus and A. collusus [now A. pallidi- slightly sinuate, 2nd stria approaching pennis (Motschulsky)]. 3rd at base, 2nd to 6th striae denticulate on basal margin; interstices flat, minutely Acanthoscelides fraterculus (Horn) imbricate. Metacoxa densely punctulate (Figures 426–431) except fossula glabrous; hind leg (figure 428) nearly straight, slightly dilated toward Bruchus fraterculus Horn 1873:336; Ri- apex; lateral, ventral, and dorsomesal cari- ley and Howard 1892c:165; Baker nae complete, ventrolateral carina evanes- 1895:29; Fall and Cockerell 1907:201; cent in apical one-half, mucro about one- Schaeffer 1907:301; Fall 1910:184; fourth length of basitarsus, lateral denticle Cushman 1911:506; Pic 1913b:26; separated from mucro by deep sinus, two Zacher 1952:462. coronal denticles each one-half as long as Mylabris fraterculus: Leng 1920:184; Essig lateral denticle. Posterior margin of 1st 1929a:486. abdominal sternum slightly lobed

113 and with deep basal pit in male, 5th ster- makes color ambiguous, but differences in num deeply emarginate for apex of py- male genitalia are reasonably consistent. gidium; 1st and 5th sterna of female not The male genitalia of A. daleae are almost modified; pygidium convex in both sexes, identical to those of A. fraterculus, and more strongly reflexed in male, disk dense- coloration of the integument and vestiture ly foveolate (figure 427). are indistinguishable from that species Male genitalia.—As in figures 430 and 431. (except that A. daleae is usually pruinose); Nearly identical to those of A. aureolus however, A. daleae is restricted to the host except cleft between lateral lobes deeper in genus Dalea in the desert areas of Arizona, A. fraterculus. California, Nevada, Utah, and northern Mexico (Johnson 1970). Acanthoscelides Size.—Body length 1.5–3.5 mm; width lobatus is similar to A. fraterculus, but the 1.0–2.0 mm. 1st abdominal segment of the male car- Type depository.—MCZC, lectotype No. ries a rounded, setose lobe that extends 8207, by Johnson 1968. beyond the margin of the segment and the brush-like structure in the male genita- Type locality.—”Plains of Kansas, lia is smaller as is the buttonlike median Nebraske, and Colorado.” sclerite. Acanthoscelides perforatus is also Distribution.—AB, AR, AZ, BC, CA, CO, IA, uniformly black with white vestiture and ID, IN, KS, MB, MO, MT, ND, NE, NJ, NM, the 1st abdominal segment is similar to NT, NV, OK, ON, OR, SD, SK, TX, UT, WA, that of A. lobatus, but the elytral striae are WY. deep and perforate. Host plants.—Astragalus bisulcatus, A. cibarius, A. crassicarpus crassicarpus, A. Acanthoscelides fumatus (Schaeffer) crassicarpus paysonii, A. drummondii, A. (Figures 432–436) falcatus, A. hyalinus, A. missouriensis, A. Bruchus fumatus Schaeffer 1907:302; Fall mollissimus bigelovii, A. mollissimus moll- 1910:183; Pic 1913a:26. issimus, A. pattersonii, A. pectinatus, A. pterocarpus, A. racemosus, A. utahensis; Mylabris fumatus: Leng 1920:305. Glycyrrhiza lepidota; Lotus crassifolius, L. Acanthoscelides fumatus: Johnson mearnsii, L. rigidus, L. scoparius. Hedy-sa- 1968:1267; Bottimer 1968c:1019; rum boreale is reported in the literature but Johnson 1969a:284; Center and John- no authentic rearings are known; it may be son 1976:196; Johnson and King- a flower-visiting record. Zacher (1952:462) solver 1982:414; Udayagiri and Wadhi listed Abrus precatorius, but the highly 1989:47. toxic seeds of this plant support only spe- Color.—Body and appendages entirely cies of Caryopemon in the Old World. black except basal three antennal seg- Natural enemies.—Bracon mellitor; Eury- ments dark red beneath, some specimens toma sp.; Pteromalus sp. with red postocular spot. Vestiture of slender white setae densely, uniformly distrib- Immatures.—Not described. uted over body. Discussion.—This species is difficult to Structure.—Body ovate, lateral margins of separate from A. daleae and A. aureolus. pronotum continuous with elytral margins From A. aureolus, A. fraterculus can usu- (figure 432). Vertex densely punctulate ally be distinguished by its black body medially, reticulate laterally, frons densely and appendages covered by white setae, punctulate, punctures crowded, frontal whereas in A. aureolus, the appendages carina distinct; ocular index 3.5:1; ocular usually have some reddish coloration and sinus two-thirds as long as eye, postocu- the vestiture is usually partly yellowish. lar lobe distinct, fringed with white se- The variation in body color of A. fraterculus tae; antenna (figure 434) reaching elytral

114 humerus. Pronotum bell-shaped (figure femoral pecten elongated, lateroventral 432), lateral margins nearly straight; disk carina of the metatibia complete to apex, strongly convex, lacking depressions, sur- male 1st abdominal sternum not lobed. face densely foveolate; lateral carina ridge- It closely resembles A. fraterculus and A. like. Scutellum quadrate, densely pubes- daleae in general facies, but the frontal cent, bidentate. Elytra moderately convex carina of those species is evanescent, den- (figure 432), lateral margins arcuate; striae ticles of the pecten are small and subequal, shallow, inconspicuous, nearly hidden by the lateroventral carina extends only three- vestiture, parallel, lacking basal denticles. fourths the length of the metatibia, and the Metacoxa entirely punctulate; hind leg male 1st abdominal segment is pitted and as in figure 433; metatibia strongly bent lobed. Acantho-scelides lobatus, another basally, gently curved and slightly dilated evenly gray species, is distinguished from toward apex, lateral, ventrolateral, ventral, A. fumatus by a more slender body, minute and dorsomesal carinae complete; mucro denticles in the femoral pecten, and the slender, acute, curved, one-fifth as long male 1st sternum with a strongly devel- as basitarsus. First abdominal segment of oped lobe. Male genitalia are distinctive for male sometimes with median tuft of setae, all four species. 5th sternum deeply emarginate; pygidium of both sexes convex, that of male strongly Acanthoscelides griseolus (Fall) reflexed apically, disk microfoveolate, (Figures 437–442) nearly concealed by vestiture. Bruchus griseolus Fall 1910:175; Pic Male genitalia.—As in figures 435 and 436; 1913a:27. median lobe moderately broad, slightly expanded toward apex; ventral valve tri- Mylabris griseolus: Leng 1920:305. angular, apex not produced; armature of Acanthoscelides griseolus: Bridwell internal sac consisting of a cluster of fine 1923a:79; Zacher 1952:465; Bibby denticles at apical orifice, a large button- 1961:324; Kingsolver 1965a:128; like sclerite in basal one-fourth; middle of Johnson 1968:1267; Bottimer sac with scattered, fine denticles, apical 1968c:1019,1038; Johnson 1969a:284, one-third with paired, spinose structures 1970:53; Slobodchikoff and John- and fine spicules near C-shaped closure son 1973:282; Center and John- valve; lateral lobes deeply cleft (figure 436), son 1976:196; Janzen 1977a:419, narrow at base, expanded apically, apices 1978:185, 1980:946; Johnson and setose. Kingsolver 1982:414; Johnson 1983a:90; Morimoto 1990:133. Size.—Body length 2.3–3.1 mm; width 1.5–2.0 mm. Bruchus kiotoensis Pic 1913c:14. Type depository.—USNM, lectotype No. Bruchidius kiotensis: Udayagiri and Wadhi 42344, by Johnson 1968. 1989:138. Bruchidius kiotoensis: Chujo 1937b:194. Type locality.—Arizona, Huachuca Moun- tains (lectotype). Acanthoscelides kiotoensis: Morimoto 1990:133. Distribution.—AZ, NM, TX; Mexico. Color.—Integument mostly red with black Host plants.—Rhynchosia texana. areas as follows: spot on vertex, scutel- Natural enemies.—None recorded. lar area sometimes extended into sutural stripe, thoracic sterna, abdominal sterna Immatures.—None described. partly or entirely black, usually with only Discussion.—Characters to separate this 1st and 2nd sterna black. Vestiture yel- species are the distinct frontal carina, lowish white, evenly distributed on body shallow elytral striae, 1st denticle of the except sparse on median pronotal stripe,

115 elytra sometimes with ephemeral pattern of Type locality.—California, near Yuma. spots. Distribution.—AZ, CA, TX; Mexico to Costa Structure.—Vertex densely punctulate and Rica; Japan. setiferous, frons punctulate changing to Host plants.—Sesbania emerus, S. exaltata. imbricate either side of prominent frontal carina; ocular index 3:1, ocular sinus Natural enemies.—None recorded. two-thirds length of eye; antenna extend- Immatures.—None described. ing nearly to metacoxa, serrate from fourth Discussion.—Distinguishing characters for segment (figure 440). Pronotum bell- this species are the dorsal integument red shaped (figure 437), evenly convex, surface with a uniformly yellow vestiture, integu- densely microfoveolate, intervals imbricate; ment often darker on ventral areas, a black lateral carina obsolete; cervical sulcus nar- spot on the vertex, all legs red, and the row but distinct. Scutellum slightly longer male genitalia with a wishbone-shaped than wide, emarginate and biden-tate api- median sclerite. See discussion of A. flave- cally, densely pubescent. Elytra flattened scens. medially, lateral margins arcuate (figure 437); striae parallel, shallow, 3rd and 4th Acanthoscelides helianthemum Bottimer denticulate at base, 2nd with transverse ridge at base. Metacoxa densely punctulate (Figures 443–447) except fossula smooth, polished; hind leg Acanthoscelides helianthemum Bottimer as in figure 439; metatibia sharply bent at 1969a:977; Johnson and Kingsolv- base, straight in apical two-thirds, lateral, er 1982:414; Udayagiri and Wadhi ventral and dorsomesal carinae complete 1989:49. to apex; mucro slender, slightly curvate, Bruchus compressicornis: Blatchley one-half as long as basitarsus, lateral den- 1919:66 (misidentification). ticles separated from mucro by deep sinus. First abdominal sternum of male slightly Color.—Integument mostly black with legs depressed medially with setiferous medial and antenna dark brown. Vestiture of fine, pit, 5th sternum nearly divided by apical white setae in ocellate pattern on elytra, emargination; female abdomen normal; otherwise evenly distributed. male pygidium strongly convex, reflexed Structure.—Vertex and frons minutely apically (figure 438); female with apex ver- imbricate, without frontal carina; ocular tical; disk densely punctulate. index 3:1; ocular sinus about three-fifths Male genitalia.—As in figures 441 and length of eye; postocular lobe narrow, 442; median lobe moderately broad, apex fringed; antenna (figure 445) reaching hu- expanded; ventral valve broadly triangu- merus. Pronotum bell-shaped (figure 443), lar, apical margin arcuate; armature of strongly convex, slightly depressed along internal sac with large, wishbone-shaped sides of basal lobe; lateral margins feebly sclerite in apical one-third, cluster of fine emarginate; lateral carina obsolete; cervi- denticles in middle, symmetrical, setose cal sulcus well-defined. Scutellum as long structure and circular closure valve in api- as wide, slightly tapered, bidentate. Elytra cal one-third; lateral lobes slender (figure with lateral margins gently arcuate (figure 442), apices oblique, cleft two-thirds their 443); striae parallel except 2nd deflected length. laterally at base to join 3rd stria, 4th, 5th, and 6th striae with basal denticles; in- Size.—Body length 1.4–2.7 mm; width terstices flat, finely imbricate. Metacoxa 0.9–1.7 mm. densely punctulate except fossula smooth; Type depository.—MCZC, holotype No. hind leg as in figure 444, metafemur with 25053. one small, acute denticle and three minute denticles following; metatibia with only

116 dorsomesal carina present, lateral face de- Acanthoscelides herissantitus Johnson void of carinae; mucro slender, slightly cur- (Figures 448–452) vate, about one-half as long as basitarsus, lateral and coronal denticles minute. Male Acanthoscelides herissantitus Johnson abdomen with small tuft of white setae be- 1983a:96 tween metacoxal bases; pygidium in both Color.—Body black; basal four or five an- sexes densely punctulate and setose, apex tennal segments, fore legs, and mid legs of male only slightly reflexed into ventral reddish orange, apex of metafemur usually emargination. dark red. Vestiture uniformly silvery gray; Male genitalia.—As in figures 446 and 447; elytral interstices distinct with three to median lobe moderately broad, straight- four rows of setae; pygidial vestiture lack- sided; ventral valve narrow, sharply trian- ing pattern. gular; armature of internal sac consisting Structure.—Vertex densely punctulate, of cluster of fine spicules at apical orifice, punctures on frons elongated, median line basal one-third of sac lined with blunt impunctate but not carinate; ocular index denticles, middle with two thornlike scler- 2.6:1; ocular sinus about one-half length ites and a dense patch of fine denticles, of eye; antenna (figure 450), extending to extreme apex lined with fine spicules and anterior margin of metacoxa. Pronotum closure valve as figured; lateral lobes slen- bell-shaped (figure 448), lateral margins der (figure 447), cleft nearly to base, apices sinuate; disk evenly convex, sparsely slightly expanded, setose. punctulate; lateral carina evanescent; cer- Size.—Body length 1.4–1.7 mm; width vical sulcus distinct but short. Scutellum 0.8–1.1 mm. quadrate, bidentate. Elytra together 1.1 times as long as wide (figure 448), strongly Type depository.—CNCI, holotype No. convex, greatest width at one-third from 10906. apex; striae parallel, distinct, 3rd, 4th, 5th, Type locality.—Florida, Gainesville. and sometimes 6th minutely denticulate Distribution.—FL, NC. at basal margin; interstices densely setose. Metacoxa with narrow, dense band Host plants.—Helianthemum corymbosum. of punctures, fossula smooth; hind leg as Natural enemies.—None recorded. in figure 449; metafemur with one to three minute denticles on ventral margin; metat- Immatures.—Not described. ibia with lateral, ventral, and dorsomesal Discussion.—Acanthoscelides helian-the- carinae complete; mucro short, less than mum is one of the smallest species in the one-eighth length of basitarsus, lateral North American fauna. Its size, ocellate denticle and three coronal denticles sub- elytral vestiture, minute denticles in the equal in size; 1st abdominal segment not pecten, and the slender, curved mucro modified, 5th segment deeply emarginate distinguish it. It appears to be related to in male; male pygidium reflexed into ter- A. alboscutellatus in the form of the male minal emargination; disk sparsely punctu- genitalia and ocellate elytral vestiture but late. is much smaller than that species. (See Male genitalia.—As in figures 451 and 452; discussion of A. atomus.) slender, ventral valve elongate-triangu- The host genus, Helianthemum, and the lar; armature of internal sac consisting of host genus for Acanthoscelides atomus, strands of minute denticles in basal one- Lechea, are both in the family Cistaceae, third of sac, a median, thornlike sclerite, whereas the host genus for A. alboscutel-la- two pairs of curved, serrate sclerites, a tus, Ludwigia, belongs to the Onagraceae. brushlike apical cluster, and a crescentic closure valve; lateral lobes short (figure

117 452), broad, cleft about one-half their tibia. Vestiture of fine intermixed golden length. and gray setae on pronotum, elytra with Size.—Body length 1.0–1.4 mm; width two pairs of golden discal maculae cen- 0.7–0.9 mm. tered on third interstice, three lateral, golden maculae on each elytron; venter of Type depository.—USNM. body gray. Type locality.—Mexico, Queretaro, 7 mi N Structure.—Vertex and frons densely Queretaro. punctulate, vestiture on vertex long, fine; Distribution.—TX; Mexico. Known from Big frons with impunctate median line, fronto- Bend National Park, Texas. clypeal suture lacking; ocular index 3.5:1; ocular sinus about one-half length of eye; Host plants.—Not yet associated in United antenna as in figure 455. Pronotum bell- States. In Herissantia crispa (Linnaeus) shaped (figure 453), evenly convex, lateral Brizicky and Malvastrum bicuspidatum margins gently arcuate, disk irregularly (Wats.) Rose in Mexico; however, both foveolate and punctulate; lateral carina hosts are found in the southwestern Unit- evanescent; cervical sulcus short, incon- ed States. spicuous. Scutellum transverse, densely Natural enemies.—Not recorded. setose. Elytra together 1.2 times as long as Immatures.—Not described. wide (figure 453), convex, lateral margins feebly arcuate; striae parallel, inconspicu- Discussion.—Johnson places this species ous, 3rd, 4th, and 5th with minute basal in his aequalis species group, all mem- denticles; interstices imbricate, densely bers of which attack seeds of malvaceous setose. Metacoxal face densely punctulate genera. The male genitalia are remark- except fossula smooth; hind leg as in figure ably similar to those of A. brevipes (Sharp) 454; metafemoral pecten with one long which ranges from northern Mexico to denticle followed by 0–3 minute denticles; Colombia. Externally, A. herissantitus most metatibia carinate only on ventral margin, closely resembles A. aequalis among the other carinae absent; mucro slender, about U.S. species, and genitalic characters are four-tenths as long as basitarsus. Fifth the only certain means of distinguishing abdominal sternum of male broadly emar- the two, although dimensions of A. aequa- ginate; male pygidium reflexed into termi- lis average slightly larger and its meta- nal emargination of abdomen; disk densely femur and metatibia are largely reddish punctulate. orange. Male genitalia.—As in figures 456 and 457; Acanthoscelides inquisitus (Fall) median lobe slender, elongated; ventral valve ovate, apex acute; armature of inter- (Figures 453–457) nal sac consisting of mixture of transverse Bruchus inquisitus Fall 1910:180; Pic and acute denticles in basal two-thirds, 1913a:29. two slender, spinescent bands and symmetrical, spinescent structure in apical Mylabris inquisitus: Leng 1920:305. one-third; closure valve circular; lateral Acanthoscelides inquisitus: Johnson lobes slender (figure 457), cleft about two- 1968:1267; Bottimer 1968c:1019,1038; thirds their length, apices scarcely broad- Johnson 1970:55; Johnson and King- ened, setose. solver 1982:414; Udayagiri and Wadhi 1989:49. Size.—Body length 2.3–3.0 mm; width 1.3–1.6 mm. Color.—Body black except basal four or five antennal segments, fore legs, mid legs, api- Type depository.—MCZC, holotype No. cal one-half of metafemur, and all of tibia 25054. red, metatarsi sometimes darker than

118 Type locality.—California, Deep Creek, San prominent, impunctate; ocular index 2.5:1; Bernardino Mountains, 6,500 ft. ocular sinus about three-fifths length of Distribution.—CA. eye; antenna (figure 460) reaching middle of metepisternum. Pronotum bell-shaped Host plants.—Not known. (figure 458), strongly convex, disk densely Natural enemies.—Not recorded. microfoveolate, interspaces punctulate, lateral margin not carinate; cervical sulcus Immatures.—Not described. deep, narrow. Scutellum quadrate, emar- Discussion.—Distinguishing characters for ginate, bidentate. Elytra widest near apices Acanthoscelides inquisitus are body integu- (figure 458), strongly convex; striae paral- ment black, fore legs and mid legs red, lel, shallow, slightly sinuate, 2nd, 3rd, 4th, metafemur and metatibia partly red, and and 5th with minute basal denticles; inter- vestiture of elytra and pronotum interstices punctate-imbricate. Metacoxa dense- mixed golden brown and white. The fron- ly punctulate except fossula smooth; hind toclypeal suture is absent, and the lateral, leg as in figure 459, pecten with three den- ventrolateral, and dorsomesal carinae of ticles; metatibia with carinae complete to the metatibia are lacking. Male genita- apex; mucro one-third as long as basitar- lic characters include median lobe long sus, lateral and coronal denticles small. and slender and lateral lobes slender and Male 1st abdominal sternum with median straight. pubescent spot, 5th sternum nearly divid- Johnson (1970) placed this species near ed by deep emargination; female 5th ster- A. pauperculus and A. napensis, both of num slightly emarginate; male pygidium which lack lateral and ventrolateral metati- strongly reflexed, disk in both sexes shal- bial carinae. lowly foveolate. Male genitalia.—As in figures 461 and 462. Acanthoscelides kingsolveri Johnson Ventral valve triangular, armature of in- (Figures 458–462) ternal sac with two short rows of spicules basally, a large saddle-shaped sclerite in Acanthoscelides kingsolveri Johnson middle, a cluster of small spicules within 1973:169; Johnson 1974:268; Cen- a thin, sclerotized envelope, and an apical ter and Johnson 1974:1097; Jan- sclerotized ring; lateral lobes slender (fig- zen 1975:172; Center and John- ure 462), deeply cleft. son 1976:196; Janzen 1977a:419, 1978:185; Johnson 1979b:63; Jan- Size.—Body length 1.5–2.5 mm; width zen 1980:946; Johnson 1981b:79, 1.0–1.7 mm. 1981e:1009; Johnson and Kingsolver Type depository.—USNM, holotype No. 1982:414; Johnson 1983a:106; Udaya- 71400. giri and Wadhi 1989:49 Type locality.—Mexico, Sinaloa, 29 mi Color.—Integument mostly black except north of Mazatlan. red in spot on side of head behind eye, Distribution.—AZ, TX; Mexico to Panama. basal four or five antennal segments, fore legs and mid legs, at least one-half of Host plants.—Indigofera lindheimeriana, I. meta-femur, all of metatibia and tarsus, sphaerocarpa, I. suffruticosa. Other spe- and broad median stripe on each elytron. cies of Indigofera are host to this bruchid Vestiture white, evenly distributed on body in Mexico and Central America (Johnson except faint median stripe on pronotum 1983a). and pygidium, darker maculae either side Natural enemies.—None recorded. of median line on pronotum, and spots of bronzy hairs on elytra. Immatures.—Not described. Structure.—Vertex and frons punctulate Discussion.—Distinguishing characters for and minutely granulose, frontal carina not A. kingsolveri are the black body with red

119 median stripe of variable width, fore legs lateral carina lacking; cervical sulcus elon- and mid legs red, hind legs usually all red, gate, conspicuous. Scutellum slightly lon- sometimes with base black, frontal carina ger than wide, emarginate apically. Elytra prominent, 2nd, 3rd, 4th, and 5th ely- together slightly longer than wide, convex tral striae with prominent, equally spaced but depressed at scutellum; striae paral- basal denticles, and the basal abdominal lel, deep, narrow, 2nd, 3rd, and 4th striae segment of the male with a pubescent de- denticulate basally, interstices flat, finely pression. The large, asymmetrical sclerite imbricate. Metacoxa densely micropunctate in the internal sac of the male genitalia is except fossula glabrous, lateral one-fourth unique. of coxal face densely setose; hind leg as in This species resembles A. submuticus in figure 465, pecten with one acute denticle size and host preferences. The integument and one or two minute denticles; metati- of A. submuticus is uniformly red, frontal bia nearly straight, slightly dilated, lateral, carina and prominent basal denticles are ventral, and dorsomesal carinae complete, lacking, and armature of the internal sac ventrolateral carina evanescent; mucro is composed of many small denticles. one-fourth as long as basitarsus, lateral denticle and two or three coronal denticles Acanthoscelides lobatus (Fall) subequal in length. First abdominal sternum of male with mesal concavity extend- (Figures 463–468) ing as a lobe fringed with golden setae from Bruchus lobatus Fall 1910:182; Pic the posterior margin of the segment, 5th 1913a:32. sternite nearly divided by emargination; female 1st and 5th sternites not modified; Mylabris lobatus: Leng 1920:305. pygidium of both sexes finely punctate, Acanthoscelides lobatus: Johnson that of male strongly reflexed. 1968:1267; Bottimer 1968c:1019,1038; Johnson 1970:56; Center and John- Male genitalia.—As in figures 467 and 468; son 1976:196; Faustini and Hal- median lobe moderately broad, gradually stead 1982:47; Johnson and King- widened toward apex; ventral valve broadly solver 1982:415; Udayagiri and Wadhi triangular, apex attenuate; armature of 1989:50. internal sac consisting of transverse rows of minute spicules in basal one-half, two Color.—Body and appendages entirely pockets lined with fine denticles and a black, basal antennal segments sometimes median, hooked sclerite at middle, a sym- dark red. Vestiture of fine white hairs uni- metrical, elongate, spinose structure and formly distributed on body, except ventral C-shaped closure valve at apex; lateral lobe of male 1st abdominal sternum with lobes (figure 468) fused for two-thirds their golden fringe; pygidium with small basal length, apical lobes subelliptical, setose on triangle (figure 464). apical margins and on mesal faces. Structure.—Vertex and frons densely Size.—Body length 2.1–2.6 mm; width punctulate, interspaces granulose, frons 1.2–1.7 mm. with median impunctate line; ocular index 3:1, ocular sinus six-tenths as long as eye; Type depository.—MCZC, lectotype No. postocular lobe narrow, setose; antenna 25055, by Johnson 1968. sexually dimorphic (figure 466), that of Type locality.—New Mexico, Las Vegas, Hot female reaching middle of metepisternum, Springs. that of male reaching middle of metacoxa. Pronotum subconical (figure 463), lateral Distribution.—AZ, CA, CO, ND, NM, TX, UT, margins slightly sinuate, constricted near WY. apex by cervical sulcus; disk strongly con- Host plants.—Astragalus humistratus, A. vex, basal margin with slight depressions, lentiginosus, A. mollissimus bigelovii, A. surface densely microfoveolate;

120 mollissimus marcidus, A. mollissimus moll- humerus. Pronotum bell-shaped (figure issimus, A. troglodytus. 469), strongly but evenly convex, reticu- Natural enemies.—None recorded. late-punctate; lateral carina absent; cervical sulcus distinct. Scutellum quadrate, Immatures.—Not described. densely pubescent. Elytra evenly convex Discussion.—The salient characters for A. (figure 469); striae distinct, deep, parallel, lobatus include the setiferous sternal lobe 4th with distinct basal denticle; interstices of the male, the total black coloration of flat, imbricate-punctate; metacoxa densely the body and appendages, and absence of punctulate except fossula glabrous; hind the ventrolateral carina on the metatibia. leg (figure 470) with tridenticulate pecten; Females of A. lobatus are difficult to sepa- metatibia moderately dilated toward apex, rate from those of A. fraterculus. tibial carina complete to apex, mucro long, curved, two-thirds length of basitarsus, Acanthoscelides longistilus (Horn) lateral denticles separated from mucro by deep sinus, three coronal denticles acute. (Figures 469–473) Male 1st abdominal sternum with median Bruchus longistilus Horn 1873:331; Blatch- tuft of setae, 5th segment nearly divided by ley 1910:1240; Fall 1910:171; Pic pygidial emargination; pygidium convex, 1913a:32; Zacher 1952:462. disk densely, shallowly foveolate. Bruchus longistylus: Ulke 1902:30; Brimley Male genitalia.—As in figures 472 and 473; 1938:232 (misspelling). median lobe broad at apex, ventral valve Mylabris longistilus: Leng 1920:305. subtriangular, lateral margins arcuate; Acanthoscelides longistilus: Bridwell internal sac armed with many small, acute 1935:186; Bissell 1940:846; Bradley denticles, four larger spines, and a sleeve- 1947:40; Blackwelder and Blackwelder like, thinly sclerotized structure lined with 1948:44; Bottimer 1968c:1019,1039; many fine spicules; lateral lobes flat (figure Johnson 1968:1267; Johnson and 473), spatulate, appearing slightly bowed. Kingsolver 1982:415; Udayagiri and Size.—Body length 1.8–2.3 mm; width Wadhi 1989:51. 0.9–1.3 mm. Acanthoscelides sp. near longistylus: Glick Type depository.—MCZC, lectotype No. 1957:9. 3894, by Johnson 1968. Color.—Integument black except basal two Type locality.—Tennessee (lectotype). or three antennal segments reddish brown, fore legs and mid legs dark brown to red, Distribution.—AL, CT, DC, FL, GA, IL, IN, mesal face of metafemur red, and entire KS, LA, MD, MI, MO, NC, NH, NJ, NY, OK, metatibia red. Vestiture of fine, white setae ON, PA, SC, TN, VA, TX. in two elongated patches each side on Host plants.—Desmodium illinoense; Les- pronotal disk, in broken stripe pattern on pedeza capitata, L. frutescens, L. hirta, L. elytra, each elytron with two large, round- intermedia, L. texana, L. virginica. “Said to ed maculae of dark brown setae; sides breed in mallows” (Blatchley 1910). Zacher of metathorax and abdomen with dense, (1952) listed Desmodium virginianum, but white patches, remainder of body with this apparently a misidentification. sparse white vestiture; pygidium with three Natural enemies.—Zatropis incertus; basal white triangular patches, sometimes Catolaccus sp.; Tetrastichus sp. with median white line. Immatures.—Not described. Structure.—Vertex and frons densely imbricate-punctulate, frontal carina absent; Discussion.—This species is one of four ocular ratio 2:1; ocular sinus three-fourths similar and probably related species (A. bi- as long as eye; antenna eccentric from ustulus, A. longistilus, A. pedicularius, and 5th segment (figure 471), reaching elytral A. stylifer) that have long, curved

121 metatibial mucro, maculate elytra, basal Structure.—Vertex punctulate, punctures abdominal segment of male with setose tending to coalesce into sulci on frons; dimple, and fourth stria with basal tu- frontal carina in both sexes prominent bercle. The geographical distribution of (figures 475 and 476); male eyes promi- A. longistilus (generally east of the 100th nent; ocular index 14:1 in male, 6.25:1 in meridian) usually will separate it from the female; ocular sinus one-half length of eye; southwestern U.S. distributions of the oth- male antenna strongly serrate (figure 479), er three. In areas of overlap, male genitalia reaching 1st abdominal segment, female will need to be dissected. The mucro length antenna (figure 479) moderately serrate, of A. longistilus, A. stylifer, and A. pedicu- reaching middle of metepisternum. Prono- larius is about one-half the length of the tum subconical, lateral margins feebly basitarsus, whereas that of A. biustulus is arcuate (figure 474), disk evenly convex, about one-third. Body and appendage color feebly depressed at posterior corners, of A. pedicularius is totally black, whereas shallowly sulcate on basal lobe; surface the hind legs of the other three are often densely foveolate, foveolae concealed by partly reddish orange. Male genitalia of vestiture; lateral carina evanescent; cervi- these four species are distinctive. cal sulcus distinct. Scutellum quadrate, densely setose, bidentate. Elytra together Acanthoscelides macrophthalmus slightly longer than wide (figure 474), (Schaeffer) convex laterally, depressed medially; striae shallow but distinct, 3rd and 4th each (Figures 474–481) with subbasal denticle set on common Bruchus macrophthalmus Schaeffer gibbosity, remaining striae lacking basal 1907:300; Fall 1910:168; Pic 1913a:33. denticles; interstices flat, parallel-sided, Mylabris macrophthalmus: Leng 1920:305. densely setose; metacoxa finely, densely Acanthoscelides macrophthalmus: John- punctulate except fossula glabrous; hind son 1968:1267; Bottimer 1968c:1019; leg as in figure 477; pecten with one long Pfaffenberger and Johnson 1976:26; acute denticle separated by gap from two Johnson 1979b:64; Johnson and King- smaller denticles; distal ventral face of solver 1982:415; Johnson 1983a:116, femur sulcate; metatibia basally arcuate, 1985:209; Mead 1989:2; Udayagiri and apically dilated, tibial carinae present and Wadhi 1989:51. complete, mucro acute, one-sixth as long as basitarsus, lateral denticle short, acute, Color.—Integument red, occasionally with coronal denticles inconspicuous. Male 5th diffuse piceous marginal shading on ely- abdominal sternum deeply, broadly emar- tra. Vestiture of fine dark brown, gray, and ginate; male pygidium apically reflexed, golden setae in pattern shown in figure female pygidium oblique; disk punctulate, 474, pattern usually more pronounced in densely setose. females; head and pronotum golden with little or no pattern; elytra with 1st, 2nd, Male genitalia.—As in figures 480 and 481; 4th, 7th, and 10th interstices golden, 3rd, slender, slightly expanded at apex, ventral 5th, 7th, and 9th with brown, golden, and valve subtriangular, lateral margins in- white quadrate or elongate maculae; male curved; internal sac armature consisting pygidial vestiture golden, evenly distrib- of many blunt denticles, apex of sac with uted, that of female golden with divergent symmetrical, spiny structure; lateral lobes brown maculae sometimes joined dorsally cleft three-fourths their length, apically to form inverted “U” or “V” (figure 478), spatulate (figure 481). some specimens with indistinct golden se- Size.—Body length 2.9–3.8 mm.; width tal patches; venter of body gray, vestiture 1.8–2.0 mm. evenly distributed. Type depository.—USNM, lectotype No. 42342, by Johnson 1968.

122 Type locality.—Texas, Brownsville, Esper- 1982:415; Udayagiri and Wadhi anza Ranch. 1989:51. Distribution.—FL, HI, TX; Mexico, Central Color.—Body and appendages black except America. ventral face of three basal antennal segments red. Vestiture very fine, mostly white Host plants.—Leucaena leucocephala, except coppery on dark patches at middle L. pulverulenta, L. retusa. Johnson of pronotum and elytra; venter of body (1983a:117) lists other species of Leucaena evenly gray. in Mexico and Central America. Structure.—Vertex and frons densely punct- Natural enemies.—None recorded. ulate, sparsely setose, some specimens Immatures.—Pfaffenberger and Johnson with distinct frontal carina; ocular index 1976:26 (first larval instar). 2.6:1; ocular sinus two-thirds as long as Discussion.—Johnson (1983a) redescribed eye; antenna (figure 484) reaching middle and illustrated Acanthoscelides macro- of metepisternum. Pronotum bell-shaped phthalmus. Salient characters for this (figure 482), apex constricted in dorsal species are red integument, eyes strongly aspect by cervical sulci; disk evenly convex enlarged in both sexes, narrowly divided on except posterior corners slightly depressed; midline in male, frons carinate and nar- densely foveolate, foveolae crowded, some- rowed, third and fourth striae ending in times coalescing, each with seta emerging denticles on a gibbosity, and female pygidi- from its anterior rim; lateral carina absent; um with paired, curvate maculae. Johnson cervical sulcus deep, reaching nearly to (1983a) placed A. macrophthalmus in the dorsal midline. Scutellum quadrate, emar- mexicanus group, but the male genita- ginate, sparsely setose. Elytra together lon- lia are of different construction (compare ger than wide (figure 482), broadest behind figure 480 with figure 614 ofA. speciosus, middle; striae shallow but distinct, paral- which is also in the mexicanus group). Ar- lel, 2nd to 6th with minute basal denticles; mature of the metafemur and configuration interstices flat, sparsely punctate-imbricate of the metatibial mucro resemble those and setose. Metacoxal face densely punc- found in species of the mexicanus group tate and setose except fossula glabrous; (compare figures 477 and 612) and of some pecten of metafemur (figure 483) consisting species of Merobruchus (compare figure of one short, acute denticle followed by two 773). minute denticles; metatibia slender, bent only at base, slightly dilated toward apex; Acanthoscelides macrophthalmus was re- ventral and dorsomesal carinae present, cently collected in Florida for the first time ventrolateral carina absent, lateral carina (Mead 1989) although Leucaena leucoceph- represented by an ephemeral, obtuse ridge, ala has been growing in the State for many or tibial face evenly rounded; mucro slen- years, and it was collected for the first time der, short, one-sixth as long as basitarsus, in Hawaii associated with Leu-caena leuco- lateral denticle short, acute, two coronal cephala (Samuelson 1991:2). denticles. First sternum of male abdomen This bruchid has been introduced into medially flattened with small median pit, South Africa for possible suppression of posterior margin of flattened area sinuate Leucaena leucocephala (B. Grobbelaar, per- with fringe of long setae; posterior bor- sonal communication, 1996). der of male 5th sternum nearly divided by emargination; male pygidium strongly Acanthoscelides margaretae Johnson reflexed, female vertical; disk in both sexes feebly, evenly variolate. (Figures 482–486) Male genitalia.—As in figures 485 and 486; Acanthoscelides margaretae Johnson median lobe slender, ventral valve subtri- 1970:58; Johnson and Kingsolver

123 angular, lateral margins incurved, apex 1976:196; Pfaffenberger and John- acute; internal sac with minute spicules son 1976:27; Johnson and Kingsolver and large, saddle-shaped sclerite in mid- 1982:415; Pfaffenberger 1985a:3; dle, apex with pair of slender, setose lobes; Udayagiri and Wadhi 1989:52. lateral lobes spatulate (figure 486), cleft Color.—Integument black except basal about one-half their length. two or three antennal segments, legs, Size.—Body length 1.7–2.8 mm; width and rounded spot covering most of api- 0.9–1.3 mm. cal one-half of each elytron orange, rarely extending to base. Vestiture of fine, golden Type depository.—USNM, holotype No. and white setae, colors often intermixed 69687. on pronotum and elytra, pronotum with Type locality.—California, San Mateo Co., feeble, median white stripe, each elytron 10 mi south of Half Moon Bay. with elongate white spot on 3rd interstice, Distribution.—CA. occasionally with indistinct white stripe on 7th and 9th interstices; scutellum, pygidi- Host plants.—Astragalus pycnostachys. um, and venter of body white. Natural enemies.—None recorded. Structure.—Vertex and frons densely punct- Immatures.—Not described. ulate, intervals granulose; frons sometimes with impunctate median line; ocular in- Discussion.—The integument of this spe- dex 3:1; ocular sinus two-thirds length of cies is usually entirely black except that eye; antenna not dimorphic (figure 490), the legs are sometimes piceous. The meta- reaching middle of metepi-sternum. Pro- tibial lateral carina is usually evanescent, notum bell-shaped (figure 487), lateral and an occasional specimen will have an margins arcuate; disk convex, sparsely intermittent lateroventral carina. The male foveolate; lateral carina marked by blunt 1st abdominal sternum is dimpled, de- ridge; cervical sulcus well-marked. Scutel- pressed, and fringed. The male genitalia lum quadrate, densely pubescent. Elytra closely resemble those of Acanthoscelides convex (figure 487) except feebly depressed mixtus (figures 485 and 491) and ofA. pul- around scutellum; striae inconspicuous, lus (figure 572), which led Johnson (1970) parallel, partly obscured by vestiture, 3rd to place these three species in a subgroup and 4th each with a minute basal denticle; of the aureolus group. Elytral apices of Ac- interstices flat, densely pubescent, finely anthoscelides mixtus are usually yellowish imbricate beneath vestiture. Metacoxa red and lack distinct spots, whereas those densely punctate except fossula glabrous, of A. pullus are usually gray with well-de- lateral one-half of coxa densely pubescent; fined brown spots. Vestiture ofmargaretae hind leg as in figure 489; pecten with three is easily abraded, and the pattern is often denticles; ventrolateral carina of metatibia obscured. Distribution of margaretae is absent, the others complete to apex; mucro confined to San Mateo and Marin Counties slender, about one-fourth as long as basi- in California whereas the other two species tarsus. Male 1st abdominal sternum with have a wider range. median tuft of setae set on slight concav- Acanthoscelides mixtus (Horn) ity, integumental setae surrounding tuft in circular pattern, posterior border of 1st (Figures 487–492) sternum sinuate with fringe of fine setae; Bruchus mixtus Horn 1873:340; Fall male 5th sternum broadly, deeply emargin- 1910:172,180; Pic 1913a:35. ate; pygidium subtriangular (figure 488), convex, apex of male reflexed. Mylabris mixtus: Leng 1920:305. Acanthoscelides mixtus: Johnson Male genitalia.—As in figures 491 and 492; 1968:1268; Bottimer 1968c:1019,1039; median lobe broad, ventral valve subtri- Johnson 1970:62; Center and Johnson

124 angular, apex bluntly angulate, internal 1989:53; Johnson 1990c:433; Kingsolv- sac with large, scutiform median sclerite er 1995a:170. bearing a blunt tooth, apex with paired, Bruchus atrocephalus Pic 1938:78. brushlike, slender sclerites; lateral lobes Acanthoscelides atrocephalus: Blackwelder broad (figure 492), spatulate cleft about 1946:758; Johnson 1990c:433. one-third their length. Color.—Body varying from all red with pro- Size.—Body length 1.7–2.2 mm; width notum piceous, to all piceous with broad 0.9–1.2 mm. red stripe on elytron, to all piceous; anten- Type depository.—MCZC, lectotype No. na piceous except segments 1–3 are red; 3896, by Johnson 1968. legs all red to all piceous. Vestiture fine, short, gray, everywhere dense but evenly Type locality.—Utah. distributed. Distribution.—AZ, CA, NM, OR, TX, UT, WA. Structure.—Body obovate (figure 493); head Host plants.—Eysenhardtia texana; Astrag- turbiniform, densely punctulate, frons alus allochrous, A. lentiginosus, A. loncho- vaguely carinate or with median glabrous carpus, A. praelongus, A. sabulonum, A. line; eyes prominent; ocular index 8:1 in thurberi, A. wootoni. male, 4:1 in female; ocular sinus one-half Natural enemies.—Eupelmus sp., Micro- length of eye; male antenna (figure 495) dontomerus anthonomi. reaching first abdominal segment, female antenna reaching middle of metepister- Immatures.—Pfaffenberger and Johnson num; ocular lobe absent in male, narrow 1976:27 (first larval instar). and fringed in female. Pronotum subconi- Discussion.—This species usually can be cal, lateral margins slightly sinuate; disk distinguished by the yellowish-gray ap- evenly convex, densely punctulate, with pearance with the apical one-third to one- shallow antescutellar depression; basal half of the elytra reddish yellow, the male margin strongly sinuate; lateral margin 1st abdominal segment concave with a me- represented by sinuate ridge. Prosternum dian setal tuft and posterior fringe, lateral acutely triangular. Scutellum quadrate, carina of metatibia evanescent, and ven- apical angles minutely dentate. Elytra con- trolateral carina absent. The male genitalia vex, together as long as broad, lateral mar- are similar to those of A. margaretae and A. gins gently arcuate, disk briefly depressed pullus (figures 485 and 572); see also the around scutellum; striae regular in course discussions of these species. except 2nd, 3rd, and 4th divergent laterad Nelson and Johnson (1983a,b) reported at base, 3rd and 4th ending short of basal effects of selenium on A. mixtus in three margin, 4th ending in small denticle; interspecies of Astragalus. stices flat, of equal width, finely granulose. Fore and mid leg not especially modified; Acanthoscelides modestus (Sharp) metacoxa finely, evenly punctulate except fossula glabrous; metafemur (figure (Figures 493–497) 494) not strongly swollen, pecten consist- Bruchus modestus Sharp 1885:461; Pic ing of one long and two smaller denticles; 1913a:35. metatibia with mucro and lateral denticle small, of equal size, corona with one or two Acanthoscelides modestus: Blackwelder minute denticles; lateral, dorsomesal, and 1946:760; Kingsolver and Whitehead ventral carinae complete, ventrolateral ca- 1976:2; Johnson 1979b:65; John- rina obsolete near apex. Pygidium broadly son and Kingsolver 1982:415; John- triangular, moderately convex in male, son 1983a:134; Udayagiri and Wadhi nearly flat in female.

125 Male genitalia.—As in figures 496 and 497. This species does not appear to be closely Median lobe slender (figure 496), lateral related to any other U.S. species. It shares margins constricted in middle of lobe; some characteristics with Acanthoscelides ventral valve ogival; armature of internal macrophthalmus (Schaeffer): 4th stria end- sac consisting of two parallel rows of elon- ing in basal denticle, male eyes enlarged, gate denticles set diagonally and extend- male antenna elongate and serrate; but in ing nearly one-half distance from base to macrophthalmus the lateral margins of the margin of cucullus then gradating into pronotum are arcuate, the basic integu- thornlike denticles in apical portion of sac; mental color is dark red, the male genitalia lateral lobes (figure 497) spatulate, setose, are distinctly different, and the metafemur cleft nearly to base. is broader (compare figures 477 and 494). Size.—Body length 1.4–2.5 mm; width 0.9–1.5 mm. Acanthoscelides mundulus (Sharp) Type depository.—BMNH (modestus); (Figures 498–503) MNHP (atrocephalus). Bruchus mundulus Sharp 1885:448; Pic Type locality.—Guatemala, San Gerónimo. 1913a:36. Acanthoscelides mundulus: Blackweld- Distribution.—FL (introduced). Brazil, Brit- er 1946:760; Bottimer 1961:294, ish Guiana, Colombia, Cuba, Dominica, 1968c:1021,1039; Kingsolver Grenada, Guatemala, Honduras, Jamaica, and Whitehead 1976:2; Johnson Mexico, Panama, Puerto Rico, Trinidad, 1977b:161, 1981b:79; Johnson Venezuela. and Kingsolver 1982:415; Johnson Host plants.—Aeschynomene indica, A. 1983a:136; Udayagiri and Wadhi histrix incana. The latter host is a recent 1989:53. introduction from tropical America (Isely Color.—Integument black; antenna black 1990). Johnson (1990c) lists other host except segments two and three yellowish species of Aeschynomene from tropical red; legs red. Vestiture dark brown, light America. brown, and white; pronotum mostly light Natural enemies.—None recorded. brown with feeble lateral patches of white, Immatures.—Not described. midline with elongated, triangular white stripe (figure 498); scutellum white; elytra Discussion.—A record from Texas is based mixed dark brown and light brown with on a single specimen identified by C.D. two diagonal series of white maculae; py- Johnson, but the presence of this bruchid gidium light brown with elongate, intensely in central Florida (Polk County) is puzzling. white basal stripe (figure 499), remainder Previous records of the host plant, Ae- of disk with variable white patches; venter schynomeme hystrix var. incana, are from of body mostly white with brown patches the extreme northwest part of the state. on metepisternum and abdomen, the ab- Aeschynomeme indica is widespread in the dominal patch divided by a white line of southeastern United States. setae. Johnson (1990c) places this bruchid in Structure.—Vertex and frons densely punc- the megacornis group of Acanthoscelides, tulate, sparsely setose, frontal carina which comprises Neotropical species. feeble; ocular ratio 3.5:1; ocular sinus two- Salient characters for this species are the thirds length of eye; postocular lobe fringed enlarged male eyes, strongly serrate male with long, brown setae; antenna (figure antenna, the denticle at the base of the 501) with distal seven segments strongly fourth elytral stria, denticle-like mucro, eccentric, not dimorphic. Pronotum sub- and details of the male genitalia. conical (figure 498), lateral margins

126 nearly straight, disk strongly convex, feebly Discussion.—This species is immediately impressed either side of basal lobe, surface recognizable by the brown basic body densely microfoveolate, each foveola with color with contrasting white setal stripes centrally placed seta; lateral carina brief, of the pronotal disk and pygidium and arcuate. Scutellum slightly longer than by the conspicuous white scutellum. The wide, densely pubescent. Elytra together prominent denticle at the base of the 4th slightly longer than wide (figure 498), stria and the male genitalia are diagnos- evenly convex except slightly depressed tic. Johnson (1983a) places mundulus by around scutellum; striae narrow, distinct, itself in the mundulus group, and it is the 3rd to 5th not reaching base, 4th denticu- only North American representative of the late at base; 3rd, 7th, and 9th interstices group. Other species in the group are scat- slightly wider than alternates; metacoxa tered on the South American continent. densely, evenly punctulate except fossula glabrous; pecten of metafemur (figure 500) Acanthoscelides napensis Johnson of one long and two minute denticles, ven- (Figures 504–508) tral femoral margin shallowly channeled, distally; lateral, ventral, and dorso-mesal Acanthoscelides napensis Johnson metatibial carinae distinct, complete, 1970:63; Johnson and Kingsolver ventrolateral carina arched toward base of 1982:415; Udayagiri and Wadhi mucro; mucro short, acute, less than one- 1989:53. eighth length of basitarsus; lateral denticle Color.—Integument variable in color from and two coronal denticles minute. Male 5th all black (except basal four antennal seg- abdominal segment broadly emarginate for ments) to all reddish brown (except distal apex of reflexed pygidium, female pygidium segments of antenna light brown). Vesti- oblique, pygidial disk shallowly foveolate. ture fine white and bronze setae; pronotum Male genitalia.—As in figures 502 and 503; usually white, sometimes intermixed with median lobe broad, ventral valve truncate, bronze; elytra with evanescent, rounded lateral margins deeply incurved; internal bronze spots; pygidium sparsely white; sac lined with triangular denticles, basal venter of body white. one-third with pair of lightly chitinized Structure.—Vertex and frons shallowly, sclerites with leaflike lateral appendages, coarsely foveolate, sparsely setose; ocular middle of sac with two pairs of thin, ar- index 2.5:1; ocular sinus one-half length cuate, serrate sclerites, apex of sac with of eye; antenna (figure 506) not sexually spiny, clavate sclerite; lateral lobes spatu- dimorphic, extending to middle of metepi- late (figure 503), emargination between sternum. Pronotum bell-shaped (figure lobes shallow. 504), lateral margins straight in basal one- Size.—Body length 1.7–3.0 mm; width half, apically arcuate; disk evenly convex, 1.5–1.7 mm. coarsely foveolate-imbricate, each foveola with seta emerging from anterior rim; lat- Type depository.—BMNH. eral carina lacking. Scutellum quadrate, Type locality.—Mexico, Guanajuato. sparsely setose. Elytra slightly longer than wide (figure 504), lateral margins arcuate; Distribution.—AZ; Mexico. striae parallel, shallowly impressed, strial Host plants.—Nissolia schottii, N. wislize- punctures elongate, 3rd, 4th, 5th, and nii. For Mexican host plants, see Johnson 6th minutely denticulate at basal margin; (1983a:137). interstices flat, minutely imbricate; meta- Natural enemies.—Horismenus productus; coxa shallowly foveolate, sparsely setose, Urosigalphus bruchivorus, U. neobruchi. fossula glabrous; hind leg (figure 505) moderately swollen, ventromesal margin Immatures.—Not described.

127 with one or two minute denticles; metati- solver 1975b:113; Center and John- bia lacking distinct carina; mucro slender, son 1976:196; Janzen 1977a:419, straight, one-third as long as basitarsus, 1978:185; Johnson 1979b:65; Janzen coronal denticles slender, dorsal denticle 1980:946; Johnson 1980:29–32; John- set at angle, longer than lateral denticles. son and Kingsolver 1982:415; John- First male abdominal segment shallowly son 1983a:141; Pfaffenberger 1985a:3; impressed, 5th ventral nearly divided by Udayagiri and Wadhi 1989:54. broad emargination; pygidial disk shallowly Color.—Sexually dimorphic (figures 509 foveolate-imbricate, apex of male reflexed and 510); male integument mostly dark into abdominal emargination. red, antenna and legs brown, thoracic Male genitalia.—As in figures 507 and 508; sterna piceous; female integument piceous median lobe slender, ventral valve ogival, except head dark red, antenna and legs apex narrowly produced, basal one-half of dark red. Male vestiture gray, golden, and internal sac lined with blunt, transverse dark brown; pattern subdued with small, denticles and transverse rows of minute brown pronotal patches; elytra gray with denticles, apical one-half with paired, slen- intermixed golden setae; pygidium gray der, setose structures; lateral lobes long with two faint brown stripes; female dorsal (figure 508), slender, cleft three-fourths vestiture contrasting, pronotal disk usu- their length. ally with four large, dark brown patches but sometimes coalescing, smaller lateral Size.—Body length 1.1–1.8 mm; width patches, otherwise gray with a few lateral 0.7–0.9 mm. intermixed golden setae; elytral pattern of Type depository.—USNM, holotype No. dark brown maculae on gray background 69688. with scattered golden diffusion; pygidi- Type locality.—California, Napa Co., 9 mi um gray with two divergent dark brown south of Pope Valley. stripes; venter of body in both sexes uniformly gray. Distribution.—CA, ID, OR. Structure.—Vertex and frons finely punc- Host plants.—On flowers ofMalacothrix in- tate, interspaces minutely granulose, cana and M. saxatilis implicata (Miller and frontal carina feeble; ocular index 4:1 in Davis 1986). male, 3.8:1 in female; ocular sinus two- Natural enemies.—None recorded. thirds length of eye; antenna (figure 514) not dimorphic. Pronotum bell-shaped Immatures.—Not described. (figure 509), lateral margins slightly sinu- Discussion.—Salient characters for this ate; disk convex, feebly impressed on basal species are its small size, arcuate lateral lobe and near posterior angles, surface of pronotal margins, white vestiture with disk irregularly microfoveolate, interspaces bronze spots on the pronotum and elytra, finely imbricate; lateral carina ridgelike, 1st abdominal segment of male impressed, feebly arched at middle; cervical sulcus details of the male genitalia, the lack of short, deep. Scutellum quadrate, densely metatibial carinae, and the minute meta- setose. Elytra together as long as wide femoral denticle. (figure 509); striae distinct, narrow, strial Johnson (1970) placed A. napensis near A. punctures elongate, 2nd, 3rd, and 4th pauperculus and A. inquisitus. each with minute basal denticle; 3rd, 5th, 7th, and 9th interstices slightly wider than Acanthoscelides obrienorum Johnson alternates; metacoxal face densely, finely punctate except fossula glabrous; hind leg (Figures 509–516) as in figure 513; metafemur slightly chan- Acanthoscelides obrienorum Johnson neled in distal one-third, pecten with four 1970:65; Slobodchikoff and John- or five denticles, the first longest; base of son 1973:282; Whitehead and King-

128 metatibia arcuate, apical four-fifths nearly genitalia of the two species are of distinctly straight, slightly dilated toward apex, different construction, especially in the lateral, ventral and dorsomesal carinae shape of the ventral valve and lateral lobes. prominent and complete to apex, ventrolateral carina joining ventral near base of Acanthoscelides obsoletus (Say) mucro; mucro short, subequal to lateral (Figures 517–521) denticle, about one-eighth as long as basitarsus, coronal denticles minute. Male Bruchus obsoletus Say 1831:2; Hamil- 1st abdominal sternum with oval setose ton 1892a:162; Riley and Howard pit, 5th sternum broadly, deeply emargin- 1892c:30; Pic 1913a:37. ate; pygidial disk finely punctate-imbricate, Mylabris obsoletus: Leng 1920:305. that of male nearly concealed by vestiture, Acanthoscelides obsoletus: Yip 1936:625; male reflexed apically, female oblique, with Bradley 1947:40; Bottimer 1968b:287, slight tumosity near apex. 1968c:1019,1039; Kingsolver 1968:9; Male genitalia.—As in figures 515 and 516; Johnson 1970:69; Johnson and King- median lobe moderately broad; ventral solver 1982:415; Udayagiri and Wadhi valve semicircular with flaring base; basal 1989:54. one-third of internal sac lined with poorly Bruchus obscurus: Fitch 1861:62; Hamilton defined, broad denticles, middle one-third 1892b:253 (misspelling). lined with elongate spicules, apical one- Bruchus distinguendus: Horn 1873:336 (in third saclike, lined with fine spicules, api- part); authors (in part). cal sclerite C-shaped; lateral lobes slightly broadened apically (figure 516), cleft two- Color.—Integument black, two basal an- thirds their length. tennal segments and postocular spot red, mesal surfaces of legs usually reddish Size.—Body length 2.1–2.5 mm; width brown. Vestiture white, dark brown, and 1.3–1.7 mm. brassy; elytra mottled with brassy; head Type depository.—USNM, holotype No. white, setae directed toward median fovea; 69689. pronotum with median, broad, black stripe bisected by evanescent white setal line, Type locality.—Arizona, Pima Co., Santa lateral areas of disk intermixed white and Rita Mountains., Box Canyon. brassy, white condensed into spots; scutel- Distribution.—AZ, TX; Mexico to Costa lum, pygidium, and venter of body white. Rica, Jamaica. Structure.—Vertex and frons punctulate, Host plants.—Senna armata, S. polyantha, transverse sulcus well marked; frontal S. wislizenii wislizenii. Johnson (1983a) carina represented by impunctate median lists S. wislizenii from Mexico. line; ocular index 3.5:1; ocular sinus two- Natural enemies.—Eupelmus sp.; Steno- thirds length of eye; antenna of both sexes corse bruchivora. serrate, dimorphic (figure 519). Pronotum bell-shaped (figure 517), lateral margins Immatures.—Not described. straight; disk evenly convex, uniformly Discussion.—Few species of Bruchidae are foveolate, each foveola with a seta under its sexually dimorphic in color pattern to the anterior rim; lateral carina obsolete; cervi- extent found in Acanthoscelides obrien- cal sulcus deep, well-marked. Scutellum orum. I could find no intergradation be- conspicuous, slightly longer than wide, tween the bold pattern of the female and bidentate. Elytra about as long as wide (fig- the muted pattern of the male. ure 517), widest near apex, convex; striae narrow, deep, punctures oval, setose, Johnson (1983a) placed A. obrienorum as a 3rd and 4th with prominent denticles on marginal member of the mexicanus group, basal margin; interstices regularly foveo- which includes A. speciosus, but the male late, imbricate; metacoxal face densely,

129 evenly punctulate, fossula glabrous. Hind The color pattern of A. obsoletus is one leg (figure 518) with pecten of metafemur that frequently recurs in this genus: 3rd bearing one long denticle and two or three interstice with a rectangular, pale setal minute denticles; metatibia arcuate in spot anteriorly and posteriorly limited by basal one-third, dilated apically, carinae a darker, quadrate spot, and lateral parts reaching apex of tibia, ventrolateral ca- of elytra with irregular bands composed rina approaching mucro base; mucro long, of pale setal patches. The dimorphic, all- slender, curved, less than one-half length black antennae, modified male abdomen, of basitarsus, lateral and coronal denticles and distinctive male genitalia should easily inconspicuous. First abdominal sternum separate this species. In contrast, the male of male shallowly sulcate and depressed, abdomen of A. obtectus is not modified, the partly glabrous, setae surrounding de- apical segment of the antenna and most of pression arranged in circular pattern; 5th the abdomen is red, antennae are not di- sternum deeply emarginate, segments morphic, and male genitalia are diagnostic telescoped ventrally; 5th female sternum (figure 525). shallowly emarginate; pygidium in both sexes foveolate, interspaces punc-tulate, Acanthoscelides obtectus (Say) male pygidium reflexed. (Figures 11, 15, 24, 44, 522–526) Male genitalia.—As in figures 520 and 521; Bruchus obtectus Say 1831:1; Jekel median lobe slender, ventral valve ogival, 1855:14; Horn 1873:337; Sharp slightly produced at apex; internal sac with 1885:458; Riley 1892:291; Riley and cluster of fine denticles at apical orifice, Howard 1892a:27, 1892c:165; Sling- middle of sac with numerous denticles of erland 1892; Riley 1893:171; Schilsky several sizes (figure 520), apex with sym- 1905:12; Schaeffer 1907:302; Blatch- metrical, setose structure; lateral lobes (fig- ley 1910:1238; Fall 1910:172; Cush- ure 521) suddenly expanded apically, cleft man 1911:499,507; Bridwell 1918:466, one-half their length. 1919:17. Size.—Body length 2.1–2.9 mm; width Laria obtecta: Bedel 1901:345. 1.4–1.9 mm. Mylabris obtectus: Leng 1920:305; Kannan Type depository.—Type destroyed. 1923:1; Essig 1929b:858. Type locality.—Indiana. Mylabris irresecta: Baudi 1886b:58. Distribution.—AL, AR, CT, DC, FL, GA, IL, Bruchidius (Acanthoscelides) obtectus: Reit- IN, KS, MD, MO, MS, NC, NJ, OK, PA, SC, ter 1912(IV):224. TX, VA, WI. Acanthoscelides obtectus: Skaife 1926:576; Host plants.—Tephrosia ambigua, T. spi- Bridwell 1929a:42, 1932:104; Bottimer cata, T. virginiana. 1935:129; Bridwell 1938b:4; Back 1940:7; Bridwell 1942:249; Black- Natural enemies.—Because of past confu- welder 1946:760; Bradley 1947:40; sion of the names of A. obsoletus and A. Blackwelder and Blackwelder 1948:45; obtectus, any parasitoids listed for A. obsol- Decelle 1951:173; Luk’yanovich and etus are suspect. Ter-Minassian 1957:172; Hinck- Immatures.—Not described. ley 1960:261; Bottimer 1968b:286, 1968c:1009,1019,1039,1040; Huig- Discussion.—The name Acanthoscelides nard 1968:233; Kingsolver 1968:4, obsoletus for many years was applied to 1970a:371; Johnson 1970:67; Foris- the bean bruchid, A. obtectus (Say) and ter and Johnson 1971:231; De Luca even today appears in Old World literature 1972:105; Howe 1973:571; Slobod- as the name for that species. Bottimer chikoff and Johnson 1973:282; Center (1968b) gave a thorough discussion of the and Johnson 1974:1097; Smith and confusion in usage of these two names.

130 Brower 1974:322; Rogers and Gar- Bruchus leguminarius var. melanocephalus rison 1975:242; Center and John- Blackwelder 1946:760 (error) son 1976:196; De Luca 1976:127; Because of its almost universal distribu- Pfaffenberger and Johnson 1976:29; tion, this bruchid species has a vast bank Schoonhoven 1977:691; Bell 1978:148; of literature pertaining to control, brief life Southgate 1978:221; Dobie et al. histories, and identification. I have been 1979:169; De Luca 1980:1; Stampou- selective in recording here only the most los and Huignard 1980:38; Johnson important references in which different 1981b:74; Szentesi 1981:219; John- combinations of the name appear or that son and Kingsolver 1982:415; John- contain significant contributions to biology, son 1983a:144, 1983c:10, 1985:208; physiology, illustration, or host relation- Pfaffenberger 1985a:3, 1985b:239; ships. Many references to the occurrence Bonet et al. 1987:378; Johnson and of the species in Old World countries have Kistler 1987:266; Kingsolver 1990c. been omitted since this handbook is pri- Bruchus leguminarius: Gyllenhal 1833:69 marily American in scope. (misidentification). Color.—Body mostly black except abdomen Bruchus irresectus Fahraeus 1839:18; Pic variably red, pygidium red, antenna with 1913a:38. basal four and terminal segments red, legs Bruchus melanocephalus Fahraeus red except ventral margin of each femur 1839:87 (misidentification). piceous brown. Vestiture of yellowish-gray Bruchus tetricus Gyllenhal 1839:22; Pic and dark brown slender setae, brown setae 1913a:52; Kingsolver 1979a:341. forming three vaguely defined, dark brown bands across elytra (figure 522), head, pro- Bruchus pallidipes Fahraeus 1839:91; Pic notum, and ventral areas of body uniform- 1913a:38. ly yellowish gray, pygidium with narrow Bruchus subellipticus Wollaston 1854:420; median stripe of yellowish setae, remainder Pic 1913a:38. of pygidium evenly clothed. Bruchus acanthocnemus Jekel 1855:15 Structure.—Vertex microfoveolate, frons validation of Dejean 1837 page 253 in punctate, intervals minutely granulose; synonymy with obtectus. frontal carina absent; ocular index 3:1, Bruchus fabae Fitch 1861:63; Riley ocular sinus two-thirds length of eye; 1871:52; Pic 1913a:38 (not Motschul- postocular lobe narrow but prominent, sky 1874). densely setose; antenna (figure 524) not Spermophagus incretus Motschulsky strongly dimorphic, reaching elytral hu- 1863:519. merus. Pronotum with bell-shaped lateral Bruchus breweri Crotch 1867:389; Pic margins (figure 522), disk evenly convex 1913a:38. except shallowly sulcate on basal lobe and with shallow depressions near pos- Bruchus fabi Rathvon 1870:119. terior corners; surface densely microfo- Bruchus obsoletus, of authors (not Say veolate, intervals finely punctate; lateral 1831): Horn 1873:337; Pic 1913a:37, carina evanescent; cervical sulcus short, and other authors. deep, nearly hidden by vestiture. Scutel- Bruchus gilvipes Motschulsky 1874:238. lum quadrate, densely setose, apex deeply Bruchus varicornis Motschulsky 1874:239 emarginate and bidentate. Elytra together (not Brulle 1832); Pic 1913a:38; King- slightly longer than wide (figure 522), later- solver 1979a:342. ally convex but medially depressed, striae mostly parallel, occasionally slightly sinu- Bruchus mimosae: Gemminger and Harold ate, shallow, with elongated punctures, 3rd 1873:3226 (not Fabricius 1781). and 4th striae with minute basal denticles; interstices flat, shallowly imbricate, 3rd,

131 5th, 7th and 9th interstices wider than V. subterranea, V. umbellata, V. unguicu- alternates. Metacoxa densely punctate ex- lata sesquipedalis, V. unguiculata unguicu- cept fossula glabrous; pecten of metafemur lata. Doubtful records include Albizia sp., (figure 523) with one long and two shorter Dolichos melanophthal-mus, Lupinus albus, denticles; metatibia slender, slightly dilated Zea mays, and inflorescences ofDaucus toward apex; lateral, ventral, and dor- carota. somesal carinae prominent and complete, Natural enemies.—Bruchobius laticeps; ventrolateral carina evanescent in apical Chryseida spinola bennetti; Dinarmus la- one-third of tibia; mucro short, slender; ticeps; Eupelmus cushmani, E. cyaniceps; lateral denticle short, acute, two coronal Pyemotes ventricosus; Torymus atheatus; denticles minute. First abdominal ster- Uscana semifumipennis. num not modified, 5th sternum broadly emarginate in male, shallowly emarginate Immatures.—Kannan 1923 (egg, first lar- in female; pygidium evenly convex, slightly val instar); Skaife 1926 (egg, larva); Lar- reflexed into sternal emargination in male; son and Fisher 1938 (all stages); Lepesme disk of pygidium moderately densely foveo- 1944; Pfaffenberger 1985b (larval instars), late, intervals punctate. 1991:564 (larva). Male genitalia.—As in figures 525 and 526; Discussion.—This species, along with Cal- median lobe elongate, slender; ventral losobruchus maculatus (Fabricius) and Cal- valve short and broadly triangular; arma- losobruchus chinensis (Linnaeus), are well ture of internal sac lined with fine, acute known pests of stored legumes in nearly all spicules in basal one-half, broad scalelike parts of the world. Probably originating in denticles in apical one-half, apical one- Middle America or northern South Amer- fourth with scattered denticles, closure ica, A. obtectus has spread through com- valve circular; lateral lobes deeply cleft mercial interchange of infested seed stocks (figure 526), expanded at apices. or of beans for consumption. Type depository.—Type apparently de- Following Horn’s (1873) error of confusing stroyed. Bruchus obtectus and Bruchus obsoletus, obtectus was listed as a synonym of obsol- Type locality.—Louisiana. etus in many publications including Mau- Distribution.—Cosmopolitan. AL, BC, CA, rice Pic’s bruchid fascicle in Catalogus Co- CT, DC, DE, FL, GA, HI, IA, IL, IN, KS, LA, leopterorum (1913a). The application of the MA, MB, MD, MN, MO, MS, NE, NJ, NY, name obsoletus to the common bean weevil OH, OK, ON, OR, PA, PQ, RI, SC, SD, TN, persisted until the 1960’s. See discussion TX, VA. following the description of obsoletus and Bottimer (1968b:286–288) for more com- Host plants.—Many host plants have been plete discussion of the confusion of these listed for this species in the literature. The two names. primary hosts are species and varieties of the fabaceous genus Phaseolus (common The oviposition habit of A. obtectus well beans, lima beans, etc.), but seeds of other equips it to be a storage pest. In the field, fabaceous genera are attacked as well. Re- eggs are almost always attached to bean ported host plants from which this species seeds either through partly dehisced pod has been reared include Cajanus cajan; valves or on seeds left in containers, in Cicer arietinum; Dolichos melanophthalmus; sheds, or in bean fodder. Larson and Fish- Glycine max; Lablab purpureus; Lathyrus er (1938:35), however, state that females sativus; Lens culinaris; Phaseolus acuti- will oviposit on immature pods after chew- folius latifolius, P. coccineus, P. lunatus, P. ing a hole in the pod in the valley between ritensis, P. vulgaris; Pisum sativum arvense; developing seeds. On more mature pods, Sesbania aegyptiaca, S. sesban; Vicia faba; Vigna aconitifolia, V. radiata radiata,

132 the female will chew a hole in the suture Host resistance.—Stampoulos and Huig- through which she inserts her ovipositor nard 1980; Stampoulos and Desroches and then drops eggs into the seed cav- 1981; Thiery 1982a,b; Thiery and Jarry ity. In storage, eggs are attached to naked 1985; Stampoulos 1987; Gatehouse et al. beans, sometimes several to a seed if popu- 1989. lations are dense. Eggs are also laid in old Integumental glands.—Bushnell 1936; Bié- emergence holes in seeds. Hatching and mont et al. 1989. larval penetration of the seed was reported by Thiery and Jarry (1985). Laboratory rearing techniques.—Strong et al. 1968. Other aspects of morphological and physiological studies may be found in the follow- Larval development.—Leroi and Jarry ing references. Complete information can 1981. be found in the bibliography. Individual Larval parasites.—Perez 1982; Perez and papers can be found in Fujii et al. (1990). Bonet 1984. Adult feeding.—Leroi 1978, 1981; Huig- Larval illustration.—Larson and Fisher nard and Leroi 1981; Jarry 1987. 1938; Pfaffenberger and Johnson 1976 Adult illustration.—Larson and Fisher (first instar); Pfaffenberger 1985b (all in- 1938; Lepesme 1944; Johnson 1983a. stars); Costa et al. 1988 (as A. obsoletus). Chromosomes.—Smith and Brower 1974; Lateral oviduct secretions.—Hamon et al. Garaud and Lecher 1982; Garaud 1984. 1982, 1983. Coevolution.—Huignard et al. 1990. Life history.—Hamilton 1892b; Riley 1892; Riley and Howard 1892a,b; Slingerland Diapause.—Biémont and Bonet 1981; 1892; Riley 1893; Gibson 1906; Manter Hodek et al. 1981. 1917; Daviault 1928; Menusan 1934b; Digestive tract.—Leroi et al. 1984. Herford 1935; Menusan 1936; Zachariae Disease transmission.—Heinze 1959. 1959; Filipek 1962; Howe and Currie 1964; Biémont and Bonet 1981; Bonet 1981; Enzymes.—Horler 1970; Agadzhanyan Bonet et al. 1987; Shade et al. 1987; Pich- 1984; Leroi et al. 1984; Gatehouse et al. ard et al. 1991. 1989. Male accessory glands.—Huignard 1968, Egg development.—Leroi 1981; Biémont 1975; Huignard et al. 1977; Cassier and and Bonet 1981; Loi and Fornasari 1985. Huignard 1979; Das et al. 1980; Huignard Eclosion.—Lepesme 1942; Thiery and Jarry 1983, 1984. 1985. Male genitalia.—Huignard 1968; Johnson Female genitalia.—Huignard 1968; John- 1970; Kingsolver 1970a; Johnson 1983a. son 1983a. Mating and reproduction.—Labeyrie 1968; Field infestation.—Back 1940:7; Biémont Huignard 1971, 1976; Biémont 1979a; and Bonet 1981; Jarry 1981; Menten and Huignard and Biémont 1981; Huignard Menten 1984; Jarry et al. 1985; Bonet et and Leroi 1981; Labeyrie 1981; Burkholder al. 1987. 1982; Biémont and Jarry 1983; Monge 1985. Flight.—Perttunen and Häyrinen 1969, 1970a,b; Perttunen 1972; Menten et al. Natural enemies.—See above. 1981. Oogenesis.—Bushnell and Boughton 1940; Genetics.—Garaud and Lecher 1982; Ga- Huignard 1970; Pouzat 1970; Biémont raud 1984. 1979a,b; Pouzat 1982; Biémont et al. 1981, 1987; Butare and Biémont 1987. Geographical origin.—Essig 1929b; South- gate 1975; Bonet et al. 1987.

133 Oogenesis, effect of host on.—Sandner and shaped (figure 527), lateral margins slight- Pankanin 1974; Huignard 1976; Pouzat ly arcuate to the brief apical constriction 1977, 1978, 1981; Leroi 1980; Huignard by the cervical sulci, disk convex with trace and Leroi 1981; Leroi and Jarry 1981; depressions along base, surface densely Monge 1985; Jarry et al. 1987. imbricate-punctate, lateral carina basally evanescent, apically absent; cervical sulcus Oviposition.—Kannan 1923; Menusan long, narrow. Scutellum nearly twice as 1934a; Huignard 1970; Labeyrie 1970; long as wide (figure 527), densely setose, Pouzat 1970; Sandner and Pankanin 1974; apex shallowly emarginate. Elytra together Pouzat 1975, 1976; Szentesi 1976; Pouzat about as long as wide, convex; striae deep, 1977; Szentesi 1981; Hamon et al. 1982; narrow, parallel except 3rd stria slightly Pouzat 1983; Menten and Menten 1984. arched toward 4th at base, 3rd, 4th, Parasites.—Perez 1982; Perez and Bonet 5th, and 6th striae minutely denticulate, 1984; Fujii and Khin Mar Wai 1990. arched laterad toward base; interstices Pathogens.—Mierzejewska 1982. densely, finely punctate. Metacoxal face densely punctate and setose except fos- Penetration of host seeds.—Thiery 1982a,b; sula glabrous; metafemur as in figure 528, Thiery and Jarry 1985. pecten usually with one minute denticle, Pheromones.—Hope et al. 1967; Horler occasionally with two; metatibia relatively 1970:859; Halstead 1973; Errard 1981a,b; slender, expanded gradually toward apex, Szentesi 1981. all tibial carinae lacking, mucro short, less Photoperiodism.—Stampoulos 1989. than one-sixth length of basitarsus, lateral carina and three slender coronal denticles Population density.—Desroches 1983. subequal in form. First abdominal sternum Spermatheca.—Surtees 1961. of male slightly concave with median setal tuft, 5th sternum nearly divided by api- Radiation, effects of.—MacLeod 1933. cal emargination, 1st sternum of female Acanthoscelides oregonensis Johnson not modified, 5th sternum not emarginate; pygidium finely punctate, densely setose, (Figures 527–531) apex of male strongly reflexed apically into Acanthoscelides oregonensis Johnson sternal emargination. 1970:69; Johnson and Kingsolver Male genitalia.—As in figures 530 and 531; 1982:415; Udayagiri and Wadhi median lobe slender, slightly expanded 1989:57. toward apex; ventral valve subtriangular, Color.—Body mostly black, elytra varying apex slightly produced; armature of inter- from all yellow with base black to all black; nal sac consisting of transverse rows of legs yellow to yellowish red, antenna black minute spicules in basal one-third, minute except basal three or four segments yellow. spicules and an elongate, acute sclerite in Vestiture uniformly silvery gray, without middle one-third, a pair of setose, elongate pronotal or elytral pattern. lobes, fine spicules and C-shaped closure valve in apical one-third; lateral lobes Structure.—Vertex and frons sparsely fused two-thirds their length (figure 531), punctulate, interspaces granulose; frontal apices slightly divergent, setose. carina vague in some specimens but usually appears as an impunctate median line; Size.—Body length 1.5 mm; width 0.9 mm. ocular index 3:1 in male, 5:2 in female; Specimen size is remarkably uniform. ocular sinus about two-thirds length of Type depository.—USNM, holotype No. eye; postocular lobe very narrow, sparsely 69690. fringed; male antenna as in figure 529, male antenna reaching nearly to metacoxa, female to elytral humerus. Pronotum bell-

134 Type locality.—Oregon, Malheur Co., On- piceous sutural stripe; legs reddish yellow tario. to yellow; basal four segments of antenna Distribution.—AB, ID, OR, WA, WY. reddish yellow, distal segments piceous. Vestiture everywhere dense, yellowish gray Host plants.—Dalea ornata. on pronotum, elytra, and legs, gray on re- Natural enemies.—None recorded. mainder of body. Immatures.—Not described. Structure.—Head elongate; vertex and frons sparsely punctate, interspaces finely gran- Discussion.—The small size, black body ulate, frontal carina absent but often with with yellow legs and variable yellow and impunctate median line; ocular index 3:1; black elytra, the single minute femoral ocular sinus three-fifths as long as eye; denticle, the lack of tibial carinae, and the postocular lobe with short, white fringe; three slender coronal denticles should suf- antenna as in figure 535, extending to fice to distinguish this species. Coloration middle of metepisternum. Pronotum bell- in Acanthoscelides pallidipennis is similar shaped (figure 532), lateral margins slight- but that species has well defined tibial ly sinuate, disk strongly convex, slightly carinae and three denticles in the pecten. depressed at posterior corners, surface Johnson (1970) places A. oregonensis ten- densely microfoveolate, interspaces finely tatively near A. aureolus. imbricate, sculpture hidden by dense vestiture; lateral carina evanescent; cervical Acanthoscelides pallidipennis (Motschulsky) sulcus conspicuous, deep. Scutellum 1.5 (Figures 532–537) times as long as wide, densely pubescent, lateral margins arcuate, apex bifid. Elytra Bruchus pallidipennis Motschulsky together slightly longer than wide (figure 1874:210; Pic 1913a:39; Bottimer 532), strongly convex, striae shallow, hid- 1968c:1040 (as unrecognized species). den by vestiture, parallel except 2nd bent Mylabris pallidipennis: Leng 1920:306. laterad at base; 3rd, 4th, and 5th each Acanthoscelides pallidipennis: Kingsolver with minute basal denticle. Metacoxal face 1979a:341; Wendt 1980:223, 1984:165; densely punctate and pubescent except Udayagiri and Wadhi 1989:57. fossula glabrous; hind leg as in figure 534; Bruchus collusus Fall 1910:176; Pic pecten with one short denticle followed 1913a:22 by two minute denticles; metatibia slen- Mylabris collusus: Leng 1920:305. der basally, gradually expanded api-cally, with lateral, ventral, and dorsomesal ca- Acanthoscelides collusus: Johnson rina complete, ventrolateral carina absent; 1968:1267; Bottimer 1968c:1018,1038; mucro one-seventh as long as basitarsus, Johnson 1970:44; Rogers and Garrison lateral denticle and two coronal denticles 1975:241. subequal in size. First abdominal sternum Bruchus perplexus Fall 1910:177; Pic of male with tomentose pit, 5th sternum 1913a:40. deeply emarginate; 1st and 5th sterna Mylabris perplexus: Leng 1920:305. in female not modified; pygidium in both Acanthoscelides perplexus: Johnson sexes convex (figure 533), apex of male re- 1968:1268; Bottimer 1968c:1020,1030; flexed into 5th sternal emargination. Johnson 1970:44; Pfaffenberger and Male genitalia.—As in figures 536 and Johnson 1976:25. 537; median lobe moderately broad, sides Acanthoscelides tarnawskii Borowiec straight; ventral valve triangular, lateral 1980:167; Wendt 1984:165. margins incurved, apex attenuate; arma- Color.—Body piceous with following ex- ture of internal sac consisting of many ceptions: elytra usually reddish yellow to minute denticles and one large, thornlike reddish brown, rarely piceous, often with

135 median spine in basal one-half, apical one- the body, but identification is sometimes half with pair of elongated, setose struc- problematical because of the multiplic- tures, apex lined with fine spicules, closure ity of forms (compare with A. mixtus). One valve C-shaped; lateral lobes cleft nearly series of specimens from Washington State one-half their length (figure 537), apices is nearly indistinguishable from A. floridae, rounded, flat, densely setose. even in characters of the male genitalia. Size.—Body length 1.1–2.7 mm; width Acanthoscelides pauperculus (Le Conte) 0.9–1.6 mm. Type depositories.—ZMUM (pallidipen- (Figures 538–542) nis); MCZC (collusus, holotype No. 25050; Bruchus pauperculus LeConte 1857:52 perplexus, holotype No. 25058); Borowiec, (not Philippi and Philippi 1864); Horn private collection, Wroclaw, Poland, (tar- 1873:329; Fall 1901:29; Fall and Cock- nawskii). erell 1907:201; Schaeffer 1907:299; Type localities.—”Californie” (pallidipennis); Fall 1910:172, 1912:322; Pic 1913a:40; California, S. Bernadino Mountains, Deep Zacher 1952:462. Creek, 6,500 ft. (collusus); New Mexico, Mylabris pauperculus: Leng 1920:305; Albuquerque (perplexus); Bulgaria, East Kannan 1923:24 (probable misidentifi- Rodopi (tarnawskii). cation). Distribution.—AZ, CA, CO, IA, ID, MT, NE, Acanthoscelides pauperculus: Bottimer NM, OK, TX, WA, WY; Bulgaria. 1968b:285, 1968c:1020,1039; John- son 1968:1268; Bottimer 1969b:1188; Host plants.—Amorpha californica, A. cane- Johnson 1977d:303; Johnson and scens, A. fruticosa fruticosa; Astragalus sp.; Kingsolver 1982:415; Udayagiri and Dalea sp.; Desmanthus virgatus acumina- Wadhi 1989:58. tus, D. virgatus virgatus; Errazu-rizia rotundata; Glycyrrhiza sp.; Lotus sp.; Parryella Bruchus pulicarius Motschulsky 1874:233; filifolia. Pic 1913a:43; Bottimer 1968c:1040 (as unrecognized name); Kingsolver Natural enemies.—Horismenus missou-rien- 1979a:341. sis, H. productus. Mylabris pulicarius: Leng 1920:306. Immatures.—Pfaffenberger and Johnson Bruchus rufus Motschulsky 1874:223; 1976:25 (as A. collusus; first larval instar) Pic 1913a:47; Bottimer 1968c:1040 Discussion.—Several color forms are dis- (as unrecognized name); Kingsolver cussed by Johnson (1970). Specimens from 1979a:341. northern California reared from Amorpha Bruchus simplex Motschulsky 1874:220; californica have bright golden yellow ves- Pic 1913a:50; Bottimer 1968c:1040 titure, whereas those from Errazurizia (as unrecognized name); Kingsolver rotundata from northern Arizona have 1979a:341. dense yellow vestiture. A form in Arizona, Mylabris simplex: Leng 1920:306. New Mexico, and Texas of which the type of perplexus is representative has all white Color.—Body and appendages black except vestiture. mouthparts and basal four antennal segments reddish brown, occasional speci- The male genitalia indicate that this spe- mens all reddish brown (teneral). Vestiture cies is very near A. aureolus but differ in of intermixed bronzy and gray setae on the shape of the medial spine (compare pronotum and elytra, often with vague gray figures 360 and 536). This species can maculae on elytra, sometimes with gray usually be distinguished by the reddish or postscutellar stripe, ventral areas of body yellowish elytra and legs contrasted with gray. the piceous pronotum and underside of

136 Structure.—Vertex and frons densely but Type depository.—MCZC, lectotype No. shallowly foveolate, frontal carina lacking 4469, by Bottimer (1968b) (pauperculus); but sometimes with evanescent impunctate ZMUM (pulicarius, rufus, simplex). median line; ocular index 5:2; ocular sinus Type locality.—California, San Jose and about three-fifths length of eye; antenna San Diego (pauperculus); “Californie” (puli- as in figure 540, extending to middle of carius, rufus, simplex). metepisternum. Pronotum bell-shaped (figure 538), lateral margins arcuate, disk Distribution.—AZ, BC, CA, CO, ID, OR, WA. convex, slightly depressed near posterior Host plants.—Trifolium obtusiflorum. John- corners; disk densely, shallowly foveolate, son (1977d:303–304) lists several plants each foveola with seta emerging from un- from which adults of A. pauperculus have der its anterior rim, interspaces glabrous, been swept (Achillea sp., Asclepias sp., impunctate; lateral carina evanescent; Lotus sp., Rorippa nasturtium-aquaticum, cervical sulcus conspicuous, extending Veronica anagallis-aquatica, Boisduvalia two-thirds distance to midline, visible in stricta), but specimens have been reared dorsal aspect as a constriction of apex of only from T. obtusiflorum. pronotum. Scutellum quadrate, bidentate Natural enemies.—None recorded. apically. Elytra 1.2 times as long as wide (figure 538), lateral margins arcuate, disk Immatures.—Kannan 1923 (egg). convex; striae moderately deep, parallel, Discussion.—This species can usually be not basally divergent, basal margin with distinguished from others in the genus by upturned carina dentate at each stria; the black body and appendages, the lack of interstices finely imbricate. Metacoxal face metatibial carinae (figure 539), slender mu- finely, densely foveolate, conspicuously se- cro, deep clefts between coronal denticles, tose only in lateral one-fourth; hind leg as three denticles in the pecten, and host in figure 539; pecten with three denticles; plant association. The species appears to metatibia usually lacking carinae, mucro be most closely related to Acantho-scelides slender, straight, about one-third length of inquisitus, a larger (2.3–3.0 mm.) sympatric basitarsus, lateral denticle and two coro- species also with a black body but with the nal denticles acute, subequal. First ab- legs partly red. The male genitalia of the dominal sternum in male not modified, 5th two species are similar. A. napensis is a sternum deeply emarginate in male, only small (1.1–1.8 mm), similar California spe- slightly emarginate in female; pygidium in cies without metatibial carinae and either both sexes shallowly punctate, that of male one or no denticles in the pecten. strongly reflexed into sternal emargination. Male genitalia.—As in figures 541 and 542; Acanthoscelides pectoralis (Horn) median lobe long, slender; ventral valve (Figures 543–548) subelliptical, acute; armature of basal three-fourths of internal sac consisting of Bruchus pectoralis Horn 1873:333; Blatch- fine, transverse rows of minute setae, and ley 1910:1238 (probably a misidentifi- minute, acute denticles, apex with sym- cation); Fall 1910:170; Pic 1913a:40. metrical, brushlike structure; closure valve Mylabris pectoralis: Leng 1920:305. small, C-shaped; lateral lobes long (figure Acanthoscelides pectoralis: Bradley 542), arms slender, only slightly expand- 1947:40; Blackwelder and Black- ed toward apex, cleft three-fourths their welder 1948:44; Bottimer 1968c:1020; length. Johnson 1968:1268; Johnson and Size.—Body length 1.2–2.3 mm; width Kingsolver 1982:415; Luckow and 0.8–1.3 mm.

137 Johnson 1987:49; Udayagiri and Wadhi Male genitalia.—As in figures 547 and 548; 1989:58. median lobe long (figure 547), slender; Color.—Body mostly dark red except tho- ventral valve triangular, lateral margins racic sternal areas, coxa, femur, and basal incurved, apex acute; armature of internal one-fourth of each leg, frons, clypeus, sac consisting of densely set, minute denti- humerus, and apical margin of each ely- cles entire length of sac, middle of sac with tron piceous, antenna dark red to piceous; elongate clusters of spines; closure valve pronotum and 3rd elytral interstice with circular; lateral lobes cleft about one-half variable patches of dark brown. Vestiture their length (figure 548), apices broadly mostly of grayish-yellow, short, slender expanded, densely setose. setae; pronotal and elytral patches dark Size.—Body length 2.3–3.1 mm.; width brown; small patch of white setae occa- 1.4–1.7 mm. sionally present on elytral margin. Type depository.—MCZC, holotype No. Structure.—Vertex densely punctulate, 8205. frons and clypeus coarsely punctate; fron- Type locality.—Texas. tal carina usually present, sometimes only as a punctate knob; ocular index 4:1; ocu- Distribution.—AZ, TX; Mexico. lar sinus two-thirds length of eye; postocu- Host plants.—Desmanthus sp., D. lepto-lo- lar lobe narrow, fringed; antenna (figure bus, D. obtusus, D. reticulatus, D. velutinus, 546) strongly serrate, reaching metacoxa. D. virgatus virgatus. Pronotum subconical (figure 543), lateral margins nearly straight; disk strongly Natural enemies.—None recorded. convex, basal lobe sulcate, basal margin Immatures.—Not described. slightly depressed; surface densely foveo- Discussion.—Distinguishing characteristics late, sculpture nearly concealed by vesti- for this species are body largely red with ture; lateral carina lacking; cervical sulcus black frons and vertex, thoracic sterna, short, deep. Scutellum quadrate, biden- legs, and elytral border; dorsal vestiture tate. Elytra together as long as wide (figure yellowish; and mucro short. The color 543), evenly convex, humeri prominent; pattern resembles that of Acanthoscelides striae parallel, shallow, concealed by vesti- griseolus; however, the elytral border and ture, basal denticles of 3rd and 4th striae legs of the latter species are yellow. Male set on feeble tumescences; interstices flat, genitalia of the two species are radically finely imbricate. Metacoxa densely punctu- different (figures 441 and 547). late, fossula glabrous; hind leg as in figure 545, metafemur swollen, pecten with one Acanthoscelides pedicularius (Sharp) long, acute denticle and two smaller denticles; metatibia nearly straight, slightly di- (Figures 549–554) lated with lateral, ventral, and dorsomesal Bruchus pedicularius Sharp 1885:479; Pic carinae prominent and complete, ventrolat- 1913a:40. eral carina evanescent in apical one-half; Acanthoscelides pedicularius: Blackwelder mucro short, acute, only slightly longer 1946:760; Johnson and Kingsolver than lateral denticle; coronal denticles 1982:415; Johnson 1983a:152; Udaya- three or four. First abdominal sternum not giri and Wadhi 1989:58. modified, 5th sternum broadly emarginate in male, evenly rounded in female; pygidial Bruchus pulloides Fall 1910:179; Pic disk in both sexes finely punctate, puncta- 1913a:43. tion hidden by dense vestiture (figure 544), Mylabris pulloides: Leng 1920:305. male pygidium reflexed apically into ster- Acanthoscelides pulloides: Johnson nal emargination. 1968:1268; Bottimer 1968c:1020,1039; Johnson 1970:78; Slobodchikoff and

138 Johnson 1973:282; Center and John- truncate; ventral valve ogival, apex acute; son 1976:196; Johnson 1976b:260; armature of internal sac consisting of a Johnson and Kingsolver 1982:415; large, median, thornlike spine set in midst Johnson 1983a:152; Udayagiri and Wa- of longitudinal rows of triangular denticles, dhi 1989:60. sometimes appearing as elongate, serrate Color.—Body and appendages black. Vesti- rods, a cluster of three or four slender ture on elytra white or of intermixed white spines in middle, a spinose, symmetrical and bronzy or golden setae often with dark, structure and many fine spicules in api- lateral maculae, white setae on pronotum cal one-half, closure valve circular; lateral more laterally abundant; underside of body lobes elongate (figure 554), slender, cleft gray. three-fourths their length, apices with only a few setae. Structure.—Vertex and frons shallowly foveolate; vague transverse sulcus between Size.—Body length 1.3–1.5 mm; width vertex and frons; frontal carina sometimes 0.8–0.9 mm. distinct, more often an impunctate me- Type depositories.—BMNH (pedicularius); dian line, ocular index 3:1; ocular sinus MCZC, holotype No. 25061 (pulloides). about one-half length of eye; postocular Type localities.—Guatemala, near the city lobe marked by very narrow fringe of se- (pedicularius); New Mexico (pulloides). tae; antenna as in figure 552. Pronotum bell-shaped (figure 549), lateral margins Distribution.—AZ, NM; Mexico, Guatemala. straight with slight apical restriction, disk Host plants.—Dalea carnea albida. convex, slightly depressed either side of basal lobe; surface densely foveolate, each Natural enemies.—None recorded. setose foveola deeper on anterior mar- Immatures.—Not described. gin; lateral carina absent; cervical sulcus Discussion.—This species belongs to John- moderately deep. Scutellum quadrate, son’s pertinax group characterized by small biden-tate, setose. Elytra together slightly size, lateral maculae on the elytra, an elon- longer than wide (figure 549), convex; gate, curved mucro, and slender lateral striae conspicuous, narrow, deep, paral- lobes. The single denticle at the base of 4th lel, 4th stria with minute basal denticle; stria is a good key character. Because sev- interstices minutely imbricate. Metacoxal eral other species of Acantho-scelides are face confusedly punctulate in lateral two- marked with similar lateral elytral macu- thirds, fossula glabrous; hind leg as in lae, positive identification may require dis- figure 551; pecten with one long and two section of male genitalia. short denticles; metatibia slender at base, expanded toward apex, lateral, ventral, and Acanthoscelides perforatus (Horn) dorso-mesal carinae complete, ventrolateral evanescent or absent; mucro slender, (Figures 555–560) slightly curved, three-tenths as long as Bruchus perforatus Horn 1873:335; basitarsus, base of mucro with deep sinus; Schaeffer 1907:301; Fall 1910:173,182; lateral denticle and two coronal denticles Pic 1913a:40. subequal. First abdominal sternum of male Mylabris perforatus: Leng 1920:305. flattened medially or slightly concave with setae set in circular pattern, 4th sternum Acanthoscelides perforatus: Bradley of male narrow and broadly emarginate, 1947:40; Blackwelder and Blackwelder sterna of female not modified; pygidium of 1948:45; Bottimer 1968c:1020,1039; both sexes finely punctulate, that of male Johnson 1968:1268, 1969c:54, strongly reflexed into sternal emargination. 1970:74; Center and Johnson 1976:196,198; Johnson and King- Male genitalia.—As in figures 553 and solver 1982:415; Udayagiri and Wadhi 554; median lobe moderately broad, apex 1989:58.

139 Color.—Body entirely black except ventral divided by emargination on caudal border, side of antennal segments 1–3 reddish that of female with slight emargination; brown, legs of some specimens dark red. pygidial face densely microfoveolate, inter- Body uniformly clothed with short, acicu- spaces punctulate, apex of male strongly late setae; pronotal, elytral, and pygidial reflexed into sternal emargination. pattern lacking. Male genitalia.—As in figures 559 and 560; Structure.—Vertex densely punctulate median lobe slender, elongate, apex dilat- with median area nearly bare but finely ed; ventral valve ogival, apex acute; arma- granulose, interspaces of vertex and frons ture of internal sac consisting of transverse granulose, frons with median impunctate denticles in basal one-half, longitudinal line sometimes faintly carinate; ocular rows of fine, acute denticles, a single spine index 3:1; ocular sinus two-thirds length of near a symmetrical, spinous structure; eye; postocular lobe with narrow fringe of closure valve circular; lateral lobes fused white setae; antennae sexually dimorphic two-thirds their length (figure 560), apices (figure 558), that of male reaching margin setose. of abdomen, that of female reaching meta- Size.—Body length 1.9–2.7 mm; width coxa. Pronotum bell-shaped (figure 555), 1.2–1.7 mm. lateral margins slightly sinuate, apically constricted, disk strongly convex, slightly Type depository.—MCZC, lectotype No. depressed along basal margin, surface 7145, by Johnson 1968. densely set with small, round, setate fove- Type locality.—Arizona. olae, interspaces punctulate and setate; Distribution.—AB, AZ, DC, IA, IL, KS, MD, lateral carina evanescent; cervical sulcus MI, MN, MO, NC, ND, NY, SD, TX, VA, VT, conspicuous, deep, extending dorsad and WI, WV. delimiting apical constriction. Scutellum slightly longer than wide, api-cally deeply Host plants.—Astragalus canadensis. emarginate. Elytra nearly quadrate (fig- Natural enemies.—None recorded. ure 555), widest behind humeri, convex but slightly depressed around scutellum; Immatures.—Not described. striae conspicuous, deep, parallel, strial Discussion.—No other U.S. Acanthoscelides punctures large, rounded, encroach- has the deep, wide strial punctations of A. ing into interstices, each puncture with perforatus. This character, in addition to caudally directed seta, 1st to 6th striae the all-black body and appendages, min- each ending basally in large, marginal pit ute femoral denticles, relative size, and with anterior rim. Metacoxal face densely, the concave and fringed basal abdominal very finely punctulate except fossula gla- sternum of the male should sufficiently brous, lateral two-thirds of face densely segregate the species. The female pygidium setose; hind leg as in figure 557, pecten is sharply triangular compared with the with one small denticle followed in some more rounded pygidium of A. fraterculus specimens by a minute denticle; metatibia (compare figures 427 and 556). nearly straight, slightly expanded toward Johnson (1970) redescribed and illustrated apex, lateral, ventral, and dorso-mesal the male genitalia of this species. carinae present in some specimens, evanescent in others, ventrolateral carina ab- Acanthoscelides prosopoides (Schaeffer) sent; mucro short, curved, about one-fifth length of basitarsus, lateral denticle and (Figures 561–567) one or two coronal denticles subequal in Bruchus prosopoides Schaeffer 1907:299; size. First abdominal sternum of male with Fall 1910:169; Pic 1913a:43. median concave or flattened area, posterior Mylabris prosopoides: Leng 1920:305. margin of sternum lobed and with fringe of long hairs, 5th sternum of male nearly

140 Acanthoscelides prosopoides: Johnson hind leg as in figure 563, pecten with one 1968:1268; Bottimer 1968c:1020,1039; long denticle and two smaller denticles; Johnson 1969c:284, 1970:75; Foris- metatibia slender, slightly dilated, lateral, ter and Johnson 1971:224; Johnson ventral, and dorsomesal carinae distinct and Kingsolver 1982:415; Johnson and complete, ventrolateral carina two- 1983a:160; Udayagiri and Wadhi thirds length of tibia; mucro slender, acute, 1989:59. extending at slight angle to ventral mar- Color.—Body dark red to piceous; append- gin, lateral denticle prominent and acute, ages reddish brown to piceous except basal coronal denticles three. First abdominal four antennal segments always reddish sternum of male not modified, 5th sternum brown ventrally. Vestiture predominantly broadly, deeply emarginate for pygidial gray with dark brown in elytral and prono- apex; 5th sternum of female slightly emar- tal patterns (figure 561); pattern of prono- ginate; pygidial disk densely micropunctu- tum varying from uniformly gray to a large late, vestiture dense, concealing punc- central brown patch sometimes bisected tation, male pygidium apically reflexed, by narrow gray line; elytra varying from all female oblique. gray with faintly indicated humeral, lateral, Male genitalia.—As in figures 566 and 567; and terminal brown maculae to more ex- median lobe moderately broad, elongate, tensive dark brown maculations; pygidium dorsal valve hoodlike, ventral valve ogival, usually gray with intermixed dark brown apex acute; armature of internal sac con- setae in vague pattern (figure 562); ventral sisting of cluster of fine spicules at apical areas of body uniformly gray. orifice, five thornlike spines at middle, a Structure.—Vertex minutely punctate, frons pair of setose lobes, closure valve subtri- more coarsely punctate, clypeus imbricate, angular; lateral lobes elongated (figure frontal carina absent; ocular index 3.3:1; 567), cleft nearly to base, arms slender, ocular sinus one-half length of eye, post- moderately expanded and setose at apex. ocular lobe narrow with fringe of short Size.—Body length 1.8–3.1 mm; width setae; antenna moderately serrate, sexually 1.3–1.9 mm. dimorphic (figures 564 and 565), reaching Type depository.—USNM, lectotype No. middle of mesepisternum. Pronotum sub- 42341, by Johnson 1968. conical (figure 561), lateral margins nearly straight, disk strongly convex, basal lobe Type locality.—Texas, New Braunfels. sulcate, basal margin depressed either side Distribution.—AZ, CA, NM, OK, TX; Mexico. of basal lobe, surface densely, irregularly foveolate; lateral carina threadlike, extend- Host plants.—Condalia lycioides; Ziziphus ing from posterior corner of prono-tum to obtusifolia. procoxal cavity; cervical sulcus hidden by Natural enemies.—Eupelmus cushmani; vestiture. Scutellum 1.5 times as long as Urosigalphus neobruchi. wide, attenuate, apex emarginate, biden- Immatures.—Forister and Johnson tate. Elytra together slightly longer than 1971:224 (egg and first larval instar); wide (figure 561), laterally convex from 6th Pfaffenberger and Johnson 1976:30 (first stria to margin, median one-third subde- larval instar). pressed; striae narrow, distinct, parallel except 2nd to 5th curved laterad at base, Discussion.—Salient characters for recogni- 3rd and 4th on common, bidentate basal tion of this species are the color pattern, elevation; interstices densely setose, ocel- the elongate scutellum, subconical prono- late, minutely imbricate; humeri imbricate. tum, and especially the laterally deflected Metacoxa densely punctulate, densely 3rd and 4th strial bases ending in a cari- setose in lateral one-half, fossula glabrous; nate or toothed subbasal ridge. It

141 resembles species of Algarobius in the densely setose, apically emarginate. Elytra dorsal pattern and elongate scutellum but together about as long as wide (figure 568), does not have the pygidial sulci found in moderately convex; striae usually paral- females of that genus. lel, sometimes slightly sinuate, narrow but Johnson redescribed Acanthoscelides shallow, 2nd to 6th basally denticulate; prosopoides twice (1970, 1983a). Forister interstices flat, finely punctate-imbricate. and Johnson (1971) published on its life Metacoxal face entirely punctulate except history. fossula glabrous, densely setose in lateral one-half; hind leg as in figure 570; pecten Acanthoscelides pullus (Fall) with one long and two shorter denticles; metatibia straight, slightly dilated toward (Figures 568–573) apex; lateral carina threadlike, not promi- Bruchus pullus Fall 1910:180; Fall nent, ventrolateral carina evanescent, ven- 1912:322; Pic 1913a:43. tral and dorsomesal carinae evident and complete; mucro about one-fifth as long Mylabris pullus: Leng 1920:305. as basitarsus, set at slight angle to ventral Acanthoscelides pullus: Bridwell carina; lateral denticle acute; three coronal 1923b:260; Zacher 1952:465; Bottimer denticles of uneven lengths. First abdomi- 1968b:285, 1968c:1020,1038; John- nal sternum of male concave with mesal son 1968:1268; Udayagiri and Wadhi setose pit surrounded by long hairs set in 1989:60. radiating pattern, caudal margin of ster- Bruchus brunneostictus Fall 1912:322; Pic num arcuate, 5th sternum nearly divided 1913a:19. by broad emargination with liplike carina, Mylabris brunneostictus: Leng 1920:305. 1st and 5th sterna of female not modified; Acanthoscelides brunneostictus: Johnson pygidium of both sexes convex, densely 1968:1267; Bottimer 1968c:1017,1038. punctate, that of male strongly reflexed. Color.—Body black, legs all black to black Male genitalia.—As in figures 572 and 573; with apex of femur and of tibia reddish median lobe narrowed in middle, moder- brown, mesal face of each leg sometimes ately broad at apex; ventral valve subtri- reddish brown, antenna black except basal angular, lateral margins incurved, apex three segments reddish brown. Vestiture acute; armature of internal sac consisting largely yellowish gray with dark brown of mixed acute denticles and transverse characteristic pattern on pronotum and rows of minute setae in basal one-half, elytra (figure 568), ventral areas of body an irregular, toothed sclerite in middle, a uniformly gray but 1st sternum of male pair of elongate, setose lobes near apex, with caudal fringe of golden setae. closure valve C-shaped; lateral lobes fused two-thirds their length (figure 573), apices Structure.—Vertex and frons densely im- truncate and setose. bricate-punctate except for mesal impunctate line on frons; ocular index 3:1; ocular Size.—Body length 1.5–2.7 mm; width sinus one-half length of eye; postocular 0.9–1.5 mm. lobe narrow, fringed; antenna (figure 571) Type depository.—MCZC, holotype No. serrate from 4th segment, reaching middle 25062 (pullus); holotype No. 25048 (brun- of metepisternum. Pronotum bell-shaped neostictus). (figure 568), lateral margins arcuate, api- Type locality.—California, Ojai (pullus); cally slightly constricted by cervical sulci, California, Contra Costa Co., Alhambra disk convex, slightly depressed along (brunneostictus). basal margin, surface densely microfoveolate, interspaces imbricate; lateral carina Distribution.—AZ, BC, CA, ID, NV, OR, UT. evanescent; cervical sulcus deep, prominent. Scutellum slightly longer than wide,

142 Host plants.—Astragalus allochrous, A. Color.—Body black, fore leg and mid leg asymmetricus, A. bolanderi, A. crotalaria, dark brown to black, hind leg piceous with A. douglasii, A. lentiginosus, A. oxyphysus, reddish-brown suffusion in some speci- A. praelongus, A. thurberi, A. trichopodus mens, basal four segments of antenna red- antisellii, A. trichopodus phoxus, A. wootoni. dish yellow, terminal segments piceous. Natural enemies.—None recorded. Vestiture mostly gray with bronze setae in three vague bands on elytra, ventral areas Immatures.—Not described. of body gray. Discussion.—Salient characters for A. pul- Structure.—Vertex and frons imbricate to lus are the gray color with brown spots of granulose, frontal carina weakly defined; varying size on the elytra, pronotum with ocular index 3:1; ocular sinus six-tenths a large central, black maculation usually as long as eye; postocular lobe narrow, with a median and transverse gray line of fringed; antenna as in figure 576, reach- setae forming a cross on the pronotum, ing middle of metepisternum. Pronotum and the concave 1st abdominal sternum bell-shaped (figure 574), lateral margins of the male with a radiating fringe of setae. arcuate in anterior one-half, slightly con- Johnson (1970:81–82) correlated variations stricted apically by cervical sulci; disk in color pattern with host and altitude in evenly convex, basal margin with slight California. depression; surface densely microfoveo- The male genitalia of A. pullus are indis- late, foveolae nearly concealed by vestiture; tinguishable from those of A. mixtus and lateral carina represented by blunt ridge; A. margaretae; however, in A. mixtus, the cervical sulcus deep but nearly concealed apical one-half of the elytra and at least by vestiture. Scutellum slightly longer than the fore and middle legs are yellowish red, wide, apically bidentate, densely setose. whereas in A. margaretae, the body and Elytra together slightly longer than wide legs are all black, the vestiture is sparse, (figure 574), convex except depressed and the elytral pattern indistinct. The first around scutellum; striae parallel, evenly abdominal segment is lobed similarly to spaced, deep, narrow, 2nd stria bent lat- that of A. lobatus. erad at base, 3rd, 4th, and 5th denticulate; interstices flat, finely imbricate. Metacoxal Nelson and Johnson (1983a,b) described face shallowly punctate in caudal one- the effects of selenium in Astragalus spp. half, fossula glabrous; hind leg as in figure on seed size and its effect on host prefer- 575, pecten with three small denticles, the ence of Acanthoscelides pullus, A. aureolus, first sometimes slightly larger than the and A. mixtus. others; metatibia nearly straight, slightly dilated; lateral, ventrolateral, ventral, and Acanthoscelides pusillimus (Sharp) dorso-mesal carina usually complete but (Figures 574–578) sometimes evanescent; mucro one-fourth Bruchus pusillimus Sharp 1885:479; Sharp as long as basitarsus, lateral denticle and 1886:37; Zacher 1952:462. two coronal denticles subequal. First sternum of male abdomen flattened or slightly Bruchidius pusillimus: Zacher concave, densely punctate, setae divergent 1952:464,473. from midline, 5th sternum of male nearly Acanthoscelides pusillimus: Chujo divided by broad emargination, 5th sternal 1937a:43, 1937b:193; Blackwelder margin of female even; pygi-dium in both 1946:760; Johnson 1981b:80; John- sexes convex, male reflexed into sternal son and Kingsolver 1982:415; John- emargination. son 1983a:168; Udayagiri and Wadhi Male genitalia.—As in figures 577 and 578; 1989:152. median lobe moderately broad; ventral Acanthoscelides sp. 20: Johnson 1979b:70. valve broadly triangular, lateral margins at

143 base rounded, apex right-angled; armature Acanthoscelides quadridentatus: Zacher of internal sac consisting of crownlike clus- 1952:465,470; De Luca 1965:55; Bot- ter of denticles around apical orifice, clus- timer 1968c:1020,1039; Johnson ter of acute denticles and one U-shaped 1968:1268; Bottimer 1969b:1192; sclerite in middle, apical one-third with Johnson 1970:19; Janzen 1977a:418, setose structure, closure valve angulate; 1978:185; Johnson 1979b:66; Janzen lateral lobes fused about one-half their 1980:946; Kingsolver 1980b:282; Jan- length (figure 578), arms slender, apices zen 1981:272; Johnson and Kingsolver suddenly expanded, setose. 1982:415; Johnson 1983a:172; Udaya- Size.—Body length 1.1–1.5 mm; width giri and Wadhi 1989:61; Mead 1989:2; 0.5–1.0 mm. Kassulke et al. 1990; Center and Kipker 1991; Harley et al. 1995. Type depository.—BMNH, lectotype by Acanthoscelides quadridentatum: Moreno Johnson 1983a. and Bibby 1943:23. Type locality.—Guatemala, Guatemala Color.—Body reddish brown, eyes black, City. head often with median black stripe, tho- Distribution.—TX; Mexico to Honduras. racic sterna and sometimes abdominal sterna with dusky suffusions, pronotum Host plants.—Johnson (1983a) lists sev- and pygidium usually with paired elongate eral species of Dalea in Mexico and Central piceous stripes, elytra usually with variable America, including Dalea scandens pauci- piceous markings; legs invariably red, an- folia, also a U.S. species, but the specific tenna usually uniformly red, occasionally host in Texas is not known. The species is with apical margin of each segment pice- occasionally intercepted in cut flowers from ous. Vestiture of fine straw yellow and dark south of the Rio Grande. Zacher’s (1952) brown setae in variable pattern (figure record of Trifolium pratense is question- 579), usually strongly striped on pronotum able. and pygidium (figures 580 and 581), faintly Natural enemies.—None recorded. striped on elytra, ventral areas of body uni- Immatures.— Not described. formly straw yellow. Discussion.—Salient characters include the Structure.—Vertex convex, finely, densely minute dimensions, short mucro, min- punctate, frons with evanescent median ute spines in the pecten, and the con- ridge marked at the dorsal end by a small cave, punctulate 1st sternum of the male. pit, ridge occasionally appearing as a ca- Johnson (1983a) was unable to place A. rina; ocular index 3:1; ocular sinus about pusillimus in any of the species groups two-thirds length of eye; postocular lobe he had established in Acanthoscelides. It very narrow, densely fringed; antenna as in resembles A. pertinax but the short mucro, figure 583, reaching middle of metepister- small femoral spines, and characters of the num. Pronotum bell-shaped (figure 579), male genitalia exclude it from the pertinax lateral margins slightly sinuate, slightly group. constricted apically by cervical sulci; disk evenly convex, slight depressions near Acanthoscelides quadridentatus (Schaeffer) posterior corners, surface densely microfoveolate, lateral sculpture nearly hidden (Figures 579–585) by vestiture; lateral carina represented by Bruchus quadridentatus Schaeffer short ridge; cervical sulcus conspicuous. 1907:304; Fall 1910:186; Cushman Scutellum quadrate, apically emarginate 1911:492,507; Pic 1913a:43; White and bidentate, setose. Elytra together as 1941:190; Zacher 1952:462. long as wide (figure 579), convex; striae Mylabris quadridentatus: Leng 1920:306. parallel, 2nd stria slightly bent at base,

144 2nd, 3rd, and 4th striae minutely denticu- pronotum broadly striped and pygidium late basally; interstices subequal in width, usually narrowly striped, elytral striae flat, minutely imbricate-punctate. Meta- parallel, femoral pecten with one long and coxa finely, evenly punctate, setose only four shorter denticles, mucro short, and at lateral angle, fossula glabrous; meta- male genitalia distinctive. Acanthoscelides femur (figure 582) with one long denticle tridenticulatus Bottimer and A. quadriden- and usually four smaller, slender denticles tatus were reared from the same collection set on triangular blade; metatibia slender of seed pods of Mimosa strigillosa in Texas. basally, moderately expanded apically; lat- The two species are similar in appear- eral, ventrolateral, ventral, and dorsomesal ance but with the following differences: (A. carinae conspicuous and complete; mucro tridenticulatus in brackets) antenna not less than one-sixth length of basitarsus, sexually dimorphic, segments slightly ec- lateral denticle and three coronal denticles centric (figure 583) male antenna strongly subequal. First abdominal sternum of male serrate [figure 640; 3rd elytral interstice not modified, 5th sternum deeply, broadly evenly pubescent throughout its length fig- emarginate; 5th sternum of female only ure 579) 3rd interstice interrupted by two slightly emarginate; pygidium convex in brown spots figure 637; median lobe as in both sexes, strongly reflexed into sternal figure 584 (compare with figure 641). emargination in male. Acanthoscelides quadridentatus and A. Male genitalia.—As in figures 584 and 585; puniceus Johnson, the latter from Mexico, median lobe broad, dorsal valve hoodlike, were selected for experiments in biocontrol ventral valve triangular, apex blunt; ar- of Mimosa pigra introduced into Australia mature of internal sac consisting of dense and Thailand (Harley et al. 1988; Forno setal patches lining dorsal hood and api- et al. 1989; Harley et al. 1995). The biol- cal orifice, two elongate spines, a cluster ogy of these two species in Australia was of fine denticles near apex, closure valve published by Kassulke et al. (1990), and C-shaped; lateral lobes cleft three-fourths male and female reproductive organs were their length (figure 585), broad, mesal illustrated. faces setose entire length of cleft. Center and Kipker (1991) investigated the Size.—Body length 2.1–3.3 mm; width biology of this species in Florida. 1.1–1.9 mm. Type depository.—USNM, lectotype No. Acanthoscelides rufovittatus (Schaeffer) 42347, by Johnson 1968. (Figures 586–591) Type locality.—Texas, Brownsville. Bruchus rufovittatus Schaeffer 1907:303; Distribution.—TX, FL; Mexico through Cen- Fall 1910:172; Pic 1913a:47. tral America to Brazil and Paraguay; Aus- Mylabris rufovittatus: Leng 1920:305. tralia and Thailand (introduced). Acanthoscelides rufovittatus: Johnson Host plants.—Mimosa pigra, M. pigra ber- 1968:1268; Bottimer1968c:1020, landieri, M. strigillosa. 1039; Johnson 1969a:284, 1970:82; Natural enemies.—Eupelmus sp.; Hetero- Slobodchikoff and Johnson 1973:282; spilus bruchi, H. prosopidis; Parasierola Center and Johnson 1976:196; John- distinguenda; Phanerotoma sp.; Stenocorse son 1979b:66, 1981b:79; Johnson bruchivora; Urosigalphus bruchi, U. neobru- and Kingsolver 1982:415; Johnson chi. 1983a:178; Udayagiri and Wadhi 1989:62. Immatures.—None described. Color.—Body varying from all black with Discussion.—Salient characters for identi- basal four antennal segments, fore and fication of this species are the straw yellow middle legs, and lateral spot on head red, color with dark brown dorsal markings,

145 to body and appendages all red with black armature of internal sac consisting of one median stripe on vertex of head. Vestiture slender, acute sclerite and four burrlike mostly gray with brown or bronze in in- sclerites in middle, a pair of spiny lobes distinct pattern on pronotum and elytra; near apex, and apex lined with fine setae; pygidium often with median stripe of setae closure valve C-shaped; lateral lobes (figure (figure 587). 591) slender, slightly spatulate, cleft three- Structure.—Vertex sparsely to densely fourths their length. punctulate, frons densely punctulate, Size.—Body length 1.9–2.5 mm; width frontal carina usually present, sometimes 1.3–1.7 mm. as an impunctate line; ocular index 2.8:1; Type depository.—USNM, holotype No. ocular sinus about two-thirds as long as 69685. eye; postocular lobe narrow, densely setose; antenna (figure 589) reaching Type locality.—Arizona. anterior margin of metacoxa. Pronotum Distribution.—AZ, TX; Mexico to Venezuela. bell-shaped (figure 586), lateral margins Host plants.—Galactia wrightii; Tephrosia arcuate, disk convex, without depressions, cinerea, T. purpurea, T. thurberi. Other spe- surface densely microfoveolate, nearly cies of Tephrosia were reported by Johnson concealed by vestiture; lateral carina repre- (1979b) from Mexico. sented by obsolete ridge; cervical sulcus narrow, deep, narrowly separated from an- Natural enemies.—None recorded. terior margin. Scutellum nearly quadrate, Immatures.—Not described. densely setose. Elytra 1.1 times as long as wide (figure 586), convex; striae subparal- Discussion.—Despite the implication of the lel except 2nd stria bent laterad at extreme name, most specimens of A. rufovittatus base, 3rd and 4th striae, sometimes 2nd are black, although entirely red and partly and 5th, with basal denticles; interstices red specimens are known. Key characters flat, of equal width, densely, finely -im are the distinct frontal carina, the flattened bricate. Metacoxa densely micropunctate male 1st sternum with a median pit, and except fossula glabrous; metafemur (figure the distinctive male genitalia (figure 590). 588) with one long, slightly curved denticle This species appears to have no close rela- and two smaller denticles, metatibia slen- tives in the genus. der and strongly bent basally, expanded toward apex, lateral, ventrolateral, ventral, Acanthoscelides schaefferi (Pic) and dorsomesal carinae all prominent and (Figures 592–597) complete; mucro slender, slightly curvate, Bruchus rufescens Schaeffer 1907:304 four-tenths length of basitarsus, with deep (preoccupied, Motschulsky 1874); Fall sinus at base, lateral denticle short, broad; 1910:184; Pic 1912:92. two or three small coronal denticles. First abdominal sternum in male flattened medi- Acanthoscelides rufescens: Johnson ally with rounded setose pit, 5th sternum 1968:1268. broadly emarginate; 1st sternum in female Bruchus schaefferi Pic 1912:92; Pic not modified, 5th sternum only slightly 1913a:47. emarginate; pygidium (figure 587) in both Mylabris schaefferi: Leng 1920:306. sexes slightly convex, punctulate but Acanthoscelides schaefferi: Bottimer punctures concealed by vestiture, apex in 1968c:1020,1039; Johnson 1977a:118; male reflexed into sternal emargination. Johnson and Kingsolver 1982:416; Male genitalia.—As in figures 590 and Johnson 1983a:183; Udayagiri and Wa- 591; median lobe moderately broad; dor- dhi 1989:62. sal valve rounded, densely setose, ventral valve ogival with truncate base, apex acute;

146 Color.—Body variable in color, head usu- Male genitalia.—As in figures 596 and 597; ally black but with red lateral spots above median lobe slender, elongated; dorsal eyes, pronotum usually black, sometimes valve angulate, setose, ventral valve semi- dark red; each elytron usually with a broad circular, apex slightly angled; armature of red stripe; pygidium black; antenna with internal sac consisting of two small, bur- first three segments red, remainder brown rlike sclerites near apical orifice, major to black, legs red. Vestiture gray, except portion of sac lined with minute, thornlike very dark specimens with bronzy spots on sclerites, apex with two large, thornlike elytra; some specimens with bronzy setae sclerites, closure valve circular; lateral on piceous median stripe. lobes fused one-half their length (figure Structure.—Body elongate (figure 592). 597), apical lobes divergent, rounded, se- Vertex and frons minutely punctate, frons tose. with evanescent carina in some speci- Size.—Body length 1.8–2.5 mm; width mens; ocular index 3:1; ocular sinus three- 1.1–1.6 mm. fourths length of eye, appearing in lateral Type depository.—USNM, holotype No. aspect to nearly divide the eye; postocular 42346. lobe narrow, fringed; antenna (figure 595) moderately serrate, terminal segment elon- Type locality.—Texas, Brownsville. gate-ovate. Pronotum bell-shaped (figure Distribution.—AZ, CA, TX; Mexico to Guate- 592), lateral margins arcuate, slightly con- mala. stricted at apex by cervical sulci, disk con- Host plants.—Not known. vex, without depressions, surface minutely foveolate, more coarsely imbricate toward Natural enemies.—None recorded. lateral margins, lateral carina evident only Immatures.—Not described. at posterior corners, cervical sulcus deep, conspicuous. Scutellum quadrate, densely Discussion.—Salient characters are the setose. Elytra together slightly longer than small size, narrow body, arcuate pronotal wide (figure 592), convex; striae parallel, margins, the red stripe on each elytron shallow, nearly hidden by vestiture, 2nd, (most specimens), and the large number of 3rd, and 4th striae minutely denticulate at denticles lining the internal sac in the male base, interstices flat, minutely imbricate. genitalia. Metacoxa densely, uniformly punctulate except fossula glabrous, densely setose in Acanthoscelides schrankiae (Horn) lateral one-third; hind leg as in figure 594; (Figures 598–603) pecten with one long and two smaller den- Bruchus schrankiae Horn 1873:330; Riley ticles; metatibia slender basally, gradually and Howard 1892c:165; Fall 1910:182; expanded; lateral, ventral, and dorsomesal Cushman 1911:499,508; Pic 1913a:47; carinae complete and prominent, ventro- Zacher 1952:462,471,474; Johnson lateral carina evanescent; mucro slender, 1968:1268, 1983a:184. about one-half as long as basitarsus, deep sinus at base of mucro, lateral denticle and Mylabris schrankiae: Leng 1920:305. two or three coronal denticles subequal in Acanthoscelides schrankiae: Bissell length, slender. First abdominal sternum 1940:846; Kingsolver 1965a:128; Bot- of male not modified, 5th sternum broadly timer 1968c:1020,1039, 1969b:1189; emarginate, 5th sternum of female evenly Johnson 1970:16,19,43, 1979b:66; rounded; pygidium (figure 593) in both Johnson and Kingsolver 1982:416; sexes convex, that of male reflexed into Udayagiri and Wadhi 1989:63. sternal emargination. Color.—Body reddish brown to black, elytral stripe and pygidium with reddish-

147 brown integument; legs red; antenna with presents a full description of their relation- basal four or five segments red, terminal ships. The distribution of A. schrankiae segment sometimes red. Vestiture yellow- generally follows the distribution of Schran- ish gray to white uniformly distributed over kia in the United States, but in Mexico body except condensed into white stripe on and farther south it moves to Mimosa. On pronotum, a short postscutellar line, and the other hand, A. chiricahuae, feeding in often in a narrow line on pygidium. Mimosa spp., is found only in Texas, New Structure.—Body almost identical to that of Mexico, and Arizona and follows the gen- Acanthoscelides chiricahuae but with fol- eral range of the host genus. lowing differences: vertex and frons finely Salient characters are the stubby appear- punctate, frontal carina varying from im- ance of the body, especially of the male, punctate line to raised ridge; ocular index raised basal denticles on 3rd and 4th stri- 3:1; metacoxal face punctate as in figure ae, uniform distribution of vestiture except 600, without arcuate ridge and radiating for median pronotal and pygidial stripes striae (compare figure 384). on some specimens, and the distinctive Male genitalia.—As in figures 602 and 603; male genitalia with sinuate lateral lobes median lobe slender, slightly expanded and arcuate, serrate apical sac sclerites. A. at apex; ventral valve subelliptical, apex chiricahuae has 5 to 8 denticles on each of acute, set at right angle to ventral face of the sclerites, in contrast to 9 to 13 for A. valve; armature of internal sac consisting schrankiae (compare figures 387 and 602). of dense lining of mixed acute and truncate denticles extending nearly to apex, two Acanthoscelides seminulum (Horn) curved, serrate sclerites near apex, each (Figures 604–610) with 9 to 13 teeth; lateral lobes long, slen- Bruchus seminulum Horn 1873:342; Fall der, sinuate, cleft three-fourths their length 1901:160; Cockerell 1902:379; Fall (figure 603), apices triangularly expanded, and Cockerell 1907:201; Blatchley setose. 1910:1238; Fall 1910:188; Bridwell Size.—Body length 1.7–2.3 mm; width 1935:186; Fox 1943:206. 1.1–1.3 mm. Bruchus seminulus: Pic 1913a:49 (misspell- Type depository.—MCZC, holotype No. ing). 3895 (Johnson 1968). Mylabris seminulum: Leng 1920:306. Type locality.—Missouri, near St. Louis. Abutiloneus seminulum: Bradley 1947:41; Distribution.—AR, FL, GA, KS, MO, MS, Blackwelder and Blackwelder 1948:45. ND, OK, SC, SD, TX, WY; Mexico to Ven- Megacerus seminulum: Zacher 1952:466. ezuela. Acanthoscelides seminulum: Bottimer Host plants.—Acacia amentacea; Desman- 1935:129, 1968c:1020,1039; Johnson thus virgatus acuminatus; Mimosa sp., M. 1968:1268, 1969c:55, 1970:84; Center borealis; Schrankia microphylla, S. nuttallii, and Johnson 1976:196,200; Johnson S. roemeriana, S. uncinata. Several species and Kingsolver 1982:416; Udayagiri of Mimosa south of the Rio Grande are re- and Wadhi 1989:63. corded as hosts by Johnson (1983a). Color.—Body uniformly dark red to black except basal three or four antennal seg- Natural enemies.—None recorded. ments reddish brown, mesal face of meta- Immatures.—Not described. femur and tibia reddish brown. Vestiture of Discussion.—This species and Acantho- fine, white setae uniformly distributed over scelides chiricahuae are closely related body. as indicated in the description above and that of A. chiricahuae. Johnson (1983a:60)

148 Structure.—Vertex finely imbricate-punc- thirds their length (figure 610), apical lobes tate, frons punctate except granulose rounded, densely setose. median line; ocular index 3:1; ocular sinus Size.—Body length 1.3–2.1 mm; width seven-tenths length of eye; postocular lobe 0.9–1.1 mm. very narrow, setose; antennae (figures 607 and 608) sexually dimorphic, that of male Type depository.—MCZC, lectotype No. extending to metacoxa, that of female to 8212, by Johnson 1968. middle of metepisternum. Pronotum bell- Type locality.—(Green disk) (Pennsylvania, shaped (figure 604), lateral margins nearly Nebraska, “Dacota”, California). The exact straight; disk strongly convex, slightly locality of the lectotype can not be deter- depressed along base at corners; surface mined. microfoveolate, sculpture nearly hidden Distribution.—AB, AZ, CA, CO, FL, IA, IL, by vestiture; lateral carina ridgelike; cer- IN, KS, LA, MB, MN, MO, ND, NE, NM, OK, vical sulcus conspicuous. Scutellum 1.5 OR, SD, SK, TX, WI, WY. times as long as wide, bidentate apically. Elytra slightly longer than wide (figure Host plants.—Dalea aurea, D. candida, 604), strongly convex; striae narrow, shal- D. enneandra, D. feayi, D. frutescens, D. low, well-defined, parallel except 2nd and stanfieldii, D. tenuis. Some of these names 3rd converging basally, 3rd, 4th, and 5th appear in literature under the genus Pet- basally denticulate; interstices finely im- alostemon. On dogwood flowers. bricate; metacoxal face evenly punctulate Natural enemies.—None recorded. and setose except fossula glabrous; hind leg as in figure 606; pecten usually with Immatures.—Not described. one or two minute denticles hidden by Discussion.—Salient characters for this vestiture, denticles sometimes lacking; species are small size, black body and metatibia slightly dilated, lateral, ventral, appendages (except antennal base), deep and dorsomesal carinae complete, ventro- clefts between tibial spines, 1st abdominal lateral carina absent; mucro one-third as sternum flattened with a median pit and long as basitarsus, deep sinus at base of long curved hairs, and metafemur with one mucro, lateral denticle and three coronal or two minute spines, or lacking spines. denticles slender, acute, and separated by deep sinuses. First abdominal sternum of Acanthoscelides speciosus (Schaeffer) male flattened with a median setose pit, (Figures 611–615) setae of flattened area long, curved toward midline, posterior margin of sternum sinu- Bruchus speciosus Schaeffer 1907:301; ate, 5th sternum of male nearly divided by Fall 1910:170; Pic 1913a:50. deep emargination; 1st sternum of female Mylabris speciosus: Leng 1920:305. sinuate on posterior border, 5th sternum Acanthoscelides speciosus: Johnson slightly emarginate; pygidium in both sexes 1968:1268; Bottimer 1968c:1020,1039, convex, that of male strongly reflexed into 1969b:1189,1197; Udayagiri and Wadhi ventral emargination, sculpture of pygidial 1989:52. disk obscured by vestiture (figure 605). Acanthoscelides mexicanus: Johnson Male genitalia.—As in figures 609 and 610; 1970:60; Johnson and Kingsolver median lobe moderately broad, straight- 1982:415 (not mexicanus Sharp 1885). sided; ventral valve sharply triangular; Color.—Dorsal integument mostly red with armature of internal sac consisting of a elongated, black elytral maculae; venter of lining of mixed acute and rounded den- body mostly black with variable red areas ticles, a large spine in middle, two similar, on abdominal sterna and metepisternum; elongate, spinose structures; closure valve antennae red, legs red except ventral C-shaped; lateral lobes fused about two- margin of metafemur black; mouthparts

149 usually black. Dorsal vestiture black, anal opening, this patch lacking in male; white, dark brown and golden; pronotum pygidium feebly convex, sexes similar. mostly golden with variable brown spots, Male genitalia.—As in figures 614 and 615; white lateral spots and elongate basal median lobe broad; ventral valve narrow, stripe; elytra mostly golden with brown hu- attenuate, bluntly rounded at apex; dorsal meral and basal spots, alternate interstices valve membranous, margin setose; inter- with series of brown, white, and golden nal sac with clusters of short denticles spots, lateral maculae often coalescing into at apical orifice; middle section and two large, submarginal patch; pygidium white lobes of sac densely lined with spicules to golden with denser, white patches at and denticles, apex of sac lined with fine middle of basal margin and on lateral mar- spicules; closure valve C-shaped; lateral gins, sometimes with median white stripe, lobes strongly bowed (figure 615), cleft two- usually with paired, subapical brown spots thirds their length, apices bilobed. of variable size; venter of body white to golden or golden with white spots, especial- Size.—Body length 1.5–2.6 mm; width ly along lateral margins of abdomen and on 1.0–1.7 mm. metepi-sternum. First and 2nd abdominal Type depository.—USNM, lectotype No. sterna usually with glabrous spots where 42343, by Johnson 1968. metafemora rub. Type locality.—Arizona, Huachuca Moun- Structure.—Vertex and frons densely tains. punctulate, frons with prominent, median Distribution.—AZ, TX; Mexico. boss; ocular index 3.7:1, eyes not dimorphic; ocular sinus one-half length of eye; Host plants.—Mimosa biuncifera, M. mala- antenna (figure 613) not dimorphic. Prono- cophylla, M. wherryana. tum bell-shaped (figure 611), lateral mar- Natural enemies.—None recorded. gins sinuate; disk convex, feebly impressed at posterior corners, sulcate on basal lobe; Immatures.—Not described. surface of disk sparsely microfoveolate on Discussion.—This species belongs to the middle of disk, dense and crowded laterally mexicanus group which extends from the with interspaces imbricate; lateral carina southwestern United States into South blunt, ridgelike, arched; cervical sulcus America (Johnson 1983a:131). It superfi- short, deep, nearly concealed by vestiture. cially resembles A. bisignatus in its color Scutellum quadrate, deeply emarginate, pattern but does not have the dimorphic densely setose. Elytra together as long as eyes of that species. The apical antennal wide (figure 611); striae subparallel, alter- segment of A. bisignatus is red, the pygi- nate interstices slightly wider, 3rd and 4th dium lacks subapical spots, and the male denticulate at base, both denticles on a genitalia are distinctive (figure 370). John- slight elevation. Metacoxa densely, evenly son (1970:60) synonymized A. speciosus punctate except fossula glabrous; hind leg with A. mexicanus (Sharp), but in 1983a as in figure 612; pecten with one long and (p.131) resurrected it to a valid species two shorter denticles, ventral margin fee- name. It closely resembles A. mexicanus bly channeled; metatibia arcuate, lateral, (see Johnson 1970). ventral, and dorsomesal carinae complete, ventrolateral carina joining ventral carina Acanthoscelides stylifer (Sharp) at base of mucro; mucro short, about one- (Figures 616–621) eighth as long as basitarsus, lateral and coronal denticles inconspicuous. First ab- Bruchus stylifer Sharp 1885:479; Pic dominal segment of male not modified, 5th 1913a:51. segment broadly emarginate; 5th segment Acanthoscelides stylifer: Blackwelder of female with glabrous patch each side of 1946:761; Johnson and Kingsolver

150 1982:416; Johnson 1983a:191; Udaya- two or three coronal denticles subequal in giri and Wadhi 1989:64. size. First abdominal sternum of male with Bruchus pugiunculus Fall 1910:178. small, shallow, densely setose pit, 5th sternum of male deeply, broadly emarginate, Bruchus pygionculus: Pic 1913a:43. 5th sternum of female slightly emarginate; Mylabris pugiunculus: Leng 1920:305. pygidium in both sexes sparsely foveolate, Acanthoscelides pugiunculus: Johnson densely setose, concealing foveae, pygidi- 1968:1268; Bottimer 1968c:1020,1039; um of male strongly reflexed into sternal Johnson 1970:77; Slobodchikoff and emargination. Johnson 1973:282; Center and John- Male genitalia.—As in figures 620 and 621; son 1976:196; Johnson 1976b:260; median lobe broad; ventral valve broadly Johnson and Kingsolver 1982:415; triangular, lateral margins ogival, apex Johnson 1983a:191; Udayagiri and Wa- blunt; armature of internal sac consist- dhi 1989:68. ing of cluster of minute spicules at apical Color.—Body black, legs red except basal orifice, a large V-shaped median sclerite, one-half to three-fourths of metafemur a cluster of thornlike denticles, a spinose, black, apical denticles of metatibia usually bilaterally symmetrical structure near black, antenna black except basal four seg- apex; closure valve C-shaped; lateral lobes ments dark red. Vestiture of predominantly spatulate (figure 621), cleft nearly to base, gray setae with some intermixed bronzy apices setose. or dark brown setae on pronotal disk and condensed into maculae of variable inten- Size.—Body length 1.3–1.9 mm; width sity on elytra; vestiture of ventral areas 0.9–1.1 mm. uniformly white. Type depository.—BMNH (stylifer, lectotype Structure.—Vertex and frons densely, by Johnson 1983a:193); MCZC, holotype finely punctate, frontal carina visible in No. 25060 (pugiunculus). some specimens, usually only as an im- Type locality.—Mexico, Guanajuato (styl- punctate line; ocular index 2.5:1, ocular ifer); Arizona, Chiricahua Mountains (pugi- sinus two-thirds as long as eye, postocular unculus). lobe densely fringed; antenna as in figure Distribution.—AZ, NM, TX; Mexico. 619. Pronotum nearly semicircular (figure 616), slightly constricted at apex by Host plants.—Desmodium sp., D. grahamii. cervical sulci, disk evenly convex, surface Natural enemies.—None recorded. imbricate-punctate; lateral carina absent; Immatures.—Not described. cervical sulcus conspicuous but narrow. Scutellum quadrate, densely setose. Elytra Discussion.— The small size, the elongate, together one-fifth longer than wide (figure curved mucro, black body, legs red ex- 616), convex; striae parallel, deep, narrow, cept metafemur bicolored, and distinctive bases of 3rd, 4th and 5th striae minutely male median lobe and lateral lobes should denticulate; interstices minutely imbricate- readily separate this species. Its apparent punctate. Metacoxal face densely, shallow- nearest relative is Acanthoscelides biustu- ly punctate in posterior one-half, glabrous lus which differs in its all-black metafemur anteriorly; hind leg as in figure 618, pecten and slightly shorter mucro and in details with three small denticles; meta-tibia ba- of the male genitalia (figure 375). sally slender, apically strongly expanded, lateral, ventral, and dorsomesal carinae Acanthoscelides subaequalis Johnson prominent and complete, mucro long, slen- (Figures 622–626) der, curved, one-half to six-tenths as long as basitarsus, base of mucro with deep Acanthoscelides subaequalis Johnson sinus; lateral denticle and 1970:85; Johnson and Kingsolver 1971:145; Slobodchikoff and Johnson

151 1973:282; Center and Johnson valve semicircular; armature of internal 1976:196; Johnson and Kingsolver sac consisting of two large, spinose thorn- 1982:416; Johnson 1983a:195; Udaya- like sclerites, two slender spines, two bur- giri and Wadhi 1989:65. rlike sclerites, and a sleevelike, spinose Color.—Body black; front and middle legs, structure; closure valve C-shaped; lateral apical one-fourth to three-fourths of meta- lobes widely separated nearly to base femur, metatibia, and basal four segments (figure 626), arms slender, apices slightly of antenna reddish brown, apical segments expanded, densely setose. piceous. Vestiture of pronotum and elytra Size.—Body length 1.4–1.9 mm; width gray to yellowish gray, sometimes with a 0.9–1.3 mm. golden sheen, ventral areas of body gray. Type depository.—USNM, holotype No. Structure.—Vertex and frons minutely 69940. punctate, frons with carina or impunc- Type locality.—Arizona, Santa Catalina tate line; ocular index 3:1; ocular sinus Mountains, Sabino Canyon. two-thirds length of eye; postocular lobe narrow; antenna as in figure 624, reach- Distribution.—AZ, TX; Mexico. ing middle of metepisternum. Pronotum Host plants.—Abutilon abutiloides, A. ber- nearly semicircular in outline (figure 622), landieri, A. incanum, A. trisulcatum. lateral margins arcuate, disk strongly convex, with slight depressions on basal Natural enemies.—Horismenus missouri-en- border, surface finely imbricate with scat- sis; Zatropis incertus. tered small foveolae, setae densely set; Immatures.—Not described. lateral carina reaching half the distance to Discussion.—Salient characters to dis- coxal cavity; cervical sulcus inconspicu- tinguish this species are the minute size, ous, nearly hidden by vestiture. Scutel- quadrate appearance, semicircular prono- lum quadrate, apically emarginate. Elytra tum, single denticle in the pecten, short together as long as wide (figure 622), even- mucro, slender, widely spaced lateral lobes, ly convex, striae parallel, deep, narrow, and the pattern of the armature in the me- slightly sinuate, 2nd and 3rd slightly bent dian lobe. Abutiloneus idoneus Bridwell has at base, 2nd to 6th basally denticulate; in- been reared from the same lot of seeds of terstices flat, minutely imbricate. Metacox- Abutilon berlandieri; it is similar in appear- al face densely punctulate in lateral two- ance and size but the legs and antennae thirds, mesally sparsely punctate, fossula are bright yellow. glabrous; hind leg as in figure 623, pecten with one denticle sometimes followed by Acanthoscelides submuticus (Sharp) another minute denticle; metatibia slender, apically slightly expanded, lateral, ven- (Figures 627–631) tral, and dorsomesal carinae evanescent, Bruchus exiguus Horn 1873:341 (not ventrolateral carina lacking; mucro short, Rosenhauer 1856); Riley and How- scarcely longer than lateral denticle, three ard 1892c:166; Fall 1901:160; Fall coronal denticles subequal. First sternum and Cockerell 1907:201; Blatchley of male not modified, convex; 5th broadly, 1910:1238; Fall 1910:175,184; Cush- deeply emarginate, 5th sternum of female man 1911:506; Pic 1913a:28; Bridwell perceptibly emarginate; pygidium in both 1918:479; Zacher 1952:462. sexes convex, densely, minutely punctate, Mylabris exiguus: Leng 1920:305. that of male reflexed into sternal emargination. Acanthoscelides exiguus: De Luca 1965:54; Bottimer 1968c:1019; Johnson Male genitalia.—As in figures 625 and 626; 1968:1267, 1969c:55. median lobe moderately broad; ventral Bruchus submuticus Sharp 1885:455; Pic 1913a:51.

152 Acanthoscelides submuticus: Glick hind leg as in figure 628, pecten with one 1939:36; Blackwelder 1946:761; Bot- long and three minute denticles; metatibia timer 1961:294, 1968c:1019; Johnson basally slender, apically expanded, ven- 1969c:55, 1970:87; Slobodchikoff and tral and dorsomesal carina present, lateral Johnson 1973:282; Center and John- and ventrolateral carinae usually lacking; son 1976:196,198; Johnson 1979b:67; mucro four-tenths as long as basitarsus, Johnson and Kingsolver 1982:416; slender, with moderately deep sinus be- Udayagiri and Wadhi 1989:65. tween base and lateral denticle; lateral and Bruchus horni Pic 1912:92 (as new name two coronal denticles subequal in shape for exiguus Horn, not Rosenhauer and length; basal tarsal segment without 1856). lateral carina. First abdominal sternum of male with small setose pit between coxae, Mylabris horni: Leng 1920:305; Kannan 5th sternum broadly emarginate about 1923:16. one-half its length, 5th sternum of female Acanthoscelides horni: Caffrey 1943:27; slightly emarginate; pygidium shallowly Peck 1963:956; De Luca 1965:54; Bot- foveolate, in male strongly reflexed into timer 1968c:1019,1038. sternal emargination. Color.—Body mostly red with thoracic ster- Male genitalia.—As in figures 630 and na and sometimes 1st and 2nd abdominal 631; ventral valve broadly triangular, apex sterna black, head black with red patch with short nipple; armature of internal sac above each eye, pronotum red with darker consisting of cluster of fine setae at apical red or piceous median spot, elytra red, orifice, lining of mixed acute and rounded sometimes with black suture and border, denticles in middle, a pair of spinose struc- occasionally with piceous lateral maculae, tures near apex, fine setae near C-shaped pygidium red to piceous, legs red except closure valve; lateral lobes cleft about base of metafemur and coronal denticles three-fourths their length (figure 631), usually piceous, antenna with basal four apices spatulate, densely setose. segments red, remainder piceous. Vestiture yellowish gray, maculae brown, ventral Size.—Body length 1.4–2.4 mm; width areas uniformly white. 0.8–1.5 mm. Structure.—Vertex and frons shallowly Type depository.—BMNH (submuticus); punctate, appearing granulose, frontal MCZC (exiguus, lectotype No. 8211, by carina present in some specimens, in Johnson 1968). others frons with a glabrous knob; ocu- Type locality.—Mexico, southern Sonora lar index 2.5:1; ocular sinus about one- (submuticus); Kansas (exiguus). half length of eye; antenna as in figure 629. Pronotum bell-shaped (figure 627), Distribution.—AZ, CA, CO, IA, IL, KS, MN, slightly constricted apically by cervical MO, NE, NM, OK, TX, WI, WY; Mexico. sulci, disk convex, without depressions, Host plants.—Amorpha californica, A. fruti- surface densely microfoveolate, interspaces cosa fruticosa, A. fruticosa angustifolia, A. sparsely punctulate, lateral carina lacking; fruticosa occidentalis. Some records taken cervical sulcus deep, conspicuous. Scutel- from the older literature are questionable lum slightly longer than wide, apically because of confusion with Acanthoscelides bidentate. Elytra together nearly quadrate floridae. (figure 627), convex, striae narrow, deep, Natural enemies.—Eupelmus brevicauda, parallel except 2nd and 3rd slightly bent E. bruchivorus, E. cyaniceps, E. inyoensis; at base, 3rd and 4th striae basally den- Eurytoma sp., E. tylodermatis; Heterospi- ticulate; interstices minutely imbricate. lus prosopidis; Horismenus missouriensis, Metacoxal face evenly punctulate except H. productus; Microdontomerus anthonomi; fossula glabrous, setose in lateral one-fifth; Stenocorse bruchivora; Zatropis incertus,

153 Z. orontas. Parasite records from older Elytra convex, one and one-fourth times literature may be found under Acanthos- as long as wide (figure 632); striae narrow, celides submuticus, A. horni, or A. exiguus, parallel, not denticulate; interstices flat but may also be confused with records of with random black, denuded spots; meta- Acanthoscelides floridae. coxa shallowly, indistinctly punctulate, Immatures.—Kannan 1923 (egg, first larval fossula glabrous; hind leg as in figure 633; instar). metafemur with a single, minute denticle, sometimes with two additional minute den- Discussion.—This species is similar in color ticles; metatibia with ventral and dorsome- and size to the more eastern Acanthos- sal carinae present, lateral and ventrolat- celides floridae, but the two species can eral carinae lacking; mucro slightly longer be distinguished by examining the male than lateral denticle; two coronal denticles. genitalia. The internal sac of A. submuticus Male abdomen with 1st sternum unmodi- does not have the large, acuminate median fied; 5th sternum not dimorphic; male spine of A. floridae (figure 424), and the pygidium feebly bulbous, female pygidium metatibia of A. submuticus usually does nearly flat, oblique; pygidial disk minutely not have lateral and lateroventral carinae punctate. that are always present in A. floridae (figure 422). Both A. submuticus and A. flori- Male genitalia.—As in figures 635 and 636; dae attack seeds of Amorpha fruticosa. median lobe slender. dilated toward apex; ventral valve subtriangular, apex bluntly Kannan (1923) described the oviposition rounded, lateral margins sinuate; internal habits of this species. sac lined with small denticles and short, transverse rows of fine setae; lateral lobes Acanthoscelides tenuis Bottimer elongate-spatulate (figure 636), deeply (Figures 632–636) cleft, densely setose on medial faces. Acanthoscelides tenuis Bottimer Size.—Body length 1.2–1.4 mm; width 1935:127; Bradley 1947:40; Bottimer 0.6–0.7 mm. 1968c:1021,1039, 1969a:977; John- Type depository.—USNM, holotype No. son 1968:1268; Johnson and King- 50860. solver 1982:416; Udayagiri and Wadhi Type locality.—Texas, Brazoria Co., Pearl- 1989:66. and. Bruchus tenius: Blackwelder 1939:64. Distribution.—AL, FL, LA, MI, MO, ON, TX. Color.—Integument black; antenna, fore legs, and mid legs dark brown. Vestiture Host plants.—Lythrum alatum, L. linare. of long, white setae in mottled pattern on Adults collected on flowers ofRubus sp. pronotum, elytra, and pygidium; scutellum and Crataegus sp. intensely white, venter of body uniformly Natural enemies.—None recorded. white. Immatures.—Not described. Structure.—Vertex and frons densely punctulate, frontal carina lacking, middle Discussion.—Although this species is of frons with small, glabrous boss; ocular smaller than Acanthoscelides alboscutel-la- index 5:1; ocular sinus one-half length of tus, it superficially resembles that species eye; antenna as in figure 634, reaching in the mottled pronotum, elytra, and py- middle of metepisternum. Pronotum bell- gidium, the intensely white scutellum, and shaped (figure 632), apex evenly rounded, bulbous male pygidium. The latter charac- lateral margins nearly straight; disk evenly ter is not as pronounced in A. tenuis as in convex, feebly impressed either side of A. alboscutellatus. The distribution of the basal lobe, shallowly microfoveolate, inter- host plants suggests a wider range for this spaces punctulate, lateral carina lacking; bruchid than is indicated by the known cervical sulcus well defined. Scutellum records. scutiform, densely pubescent. 154 Acanthoscelides tridenticulatus Bottimer apex broadly acute; internal sac armature consisting of mass of slender spicules at (Figures 637–642) apical orifice flanked by spherical, burrlike Acanthoscelides tridenticulatus Bottimer sclerites, middle of sac with a second mass 1969b:1193; Kingsolver 1980b:288; of slender spicules, a pair of hooked scler- Johnson and Kingsolver 1982:416; ites, a pair of thick, elongated sclerites, Udayagiri and Wadhi 1989:69. apex with symmetrical spiculate structure; Color.—Integument mostly reddish brown, lateral lobes broadly spatulate (figure 642), thoracic sterna of some specimens black; cleft to two-thirds their length. male antenna sometimes dark brown. Ves- Size.—Body length 2.3–2.7 mm; width titure mostly yellowish gray except brown 1.4–1.7 mm. on elytral spots; pronotum with two broad, bare stripes separated by evanescent me- Type depository.—CNCI, holotype No. dian line of setae; pygidium with a pair of 10907. elongate spots. Type locality.—Texas, Brownsville. Structure.—Vertex finely imbricate-punc- Distribution.—LA, TX. tate, frons with impunctate median line; Host plants.—Mimosa strigillosa. ocular index 4:1; ocular sinus one-half as long as eye; antenna dimorphic (figure Natural enemies.—None recorded. 640), that of female reaching elytral hu- Immatures.—Not described. merus, that of male reaching middle of me- Discussion.—This species closely resem- tepisternum. Pronotum bell-shaped (figure bles Acanthoscelides quadridentatus, but 637), lateral margins sinuate; disk evenly the male antenna of A. tridenticulatus is convex, densely microfoveolate in middle of strongly serrate and the male genitalia of disk, lateral foveolae smaller; lateral ca- the two species are radically different (com- rina ridgelike. Scutellum quadrate, densely pare figures 584 and 641). The 3rd elytral setose, apex emarginate. Elytra together as interstice of A. quadridentatus is uniformly long as wide (figure 637), moderately con- grayish yellow for its entire length, whereas vex, slightly depressed around scutellum; that of A. tridenticulatus is interrupted by striae narrow, parallel except 3rd and 4th two brown spots. The pygidium of female approximate at base, 2nd to 6th each with A. quadridentatus is grayish yellow strong- blunt denticle on basal margin; interstices ly marked with brown, whereas that of A. flat, finely imbricate; metacoxa densely tridenticulatus has a faint median stripe punctulate in lateral one-third, sparsely and is otherwise uniformly gray. Both spe- punctate in medial two-thirds, fossula cies have been reared from the same host glabrous; hind leg as in figure 639, pecten plant, same collection of seeds. with one long and three or four shorter denticles; metatibia arcuate at base, di- The male genitalia are unlike those of any lated toward apex, carinae complete except other species I have seen. ventrolateral feeble; mucro long, slender, one-third length of basitarsus; lateral and Genus Algarobius Bridwell coronal denticles acute. First sternum of Algarobius Bridwell 1946:54; Bottimer male abdomen not modified; 5th sternum 1968c:1021,1039; Johnson 1968:1268; deeply emarginate; pygidium arcuate, that Kingsolver 1972b:116, 1986:110; John- of male strongly reflexed, that of female son and Kingsolver 1982:416; Borow- vertical at apex; disk finely, shallowly fo- iec 1987:105; Udayagiri and Wadhi veolate. 1989:68; Johnson and Siemens 1997a. Male genitalia.—As in figures 641 and 642; Type species: Bruchus prosopis LeCon- median lobe broad; ventral valve subtri- te, by original designation. angular, lateral margins feebly arcuate,

155 Small to medium sized bruchids (2.2–5.0 Algarobius bottimeri Kingsolver mm). Two species in the United States, six (Figures 60, 643–647) in genus. Bruchus prosopis, of authors (not LeConte): Integument red to piceous, vestiture gray Fullaway 1913:24; Bridwell 1918:475, with brown to black maculations. 1919:17, 1920a:337, 1920b:403; Elongate-ovate, subfusiform. Antenna Swezey 1925:3; Bridwell 1929a:43; (figure 653) reaching metepisternum. Krauss 1945:315. Pronotum bell-shaped, convex, sometimes Algarobius prosopis: Hinckley 1960:261. gibbous, lateral carina evanescent. Scu- tellum two times as long as wide (figure Algarobius bottimeri Kingsolver 1972b:116; 648). Elytra evenly convex, with fine basal Kingsolver et al. 1977:115; Ward et denticles on 3rd and 4th striae; metafemur al. 1977:5; Johnson and Kingsolver with three subapical denticles (figure 654); 1982:416; Johnson 1983c:27; King- mucro nearly as long as width of tibial solver 1986:119; Udayagiri and Wadhi apex; male pygidium evenly convex, female 1989:68. pygidium with pair of subapical sulci. Color.—Integument dark red to nearly black; eyes, female pygidial sulci, and Internal sac of male genitalia with charac- humeri black. Vestiture yellowish gray, teristic H-shaped sclerite (figure 646). brassy, and black; dorsal elytral pattern Geographic range of genus is southwestern mottled, female more contrasting than United States to Colombia and Venezuela. male. All host associations are with Prosopis spp. Structure.—Female pygidial sulci (figure (mesquite). 645) located on either side of midline about Taxonomy: Kingsolver 1972b, 1986. one-half of the distace from lateral margin to midline and about two-thirds of the distance from base to apex of pygidium. Male genitalia.—As in figures 646 and 647; Key to Species of U.S. Algarobius median lobe with apical frame thick and massive (figure 646), ventral valve short, 1 With pair of polished sulci on pygidium (figures 645 and not keeled, membrane dorsad of apical ori- 652) (females)...... 2 fice with cluster of acute denticles, internal Without sulci on pygidium (figures 644 and 651) (males) sac armature as illustrated; lateral lobes ...... 3 flat (figure 647), cleft about one-third their length. 2(1) Pygidium slightly convex, sulci centered one-third from apex (figure 645) Size.—Body length 2.7–4.25 mm; width 1.5–2.25 mm...... bottimeri Kingsolver Pygidium strongly convex apically, sulci nearly confluent Type depository.—USNM, holotype No. 70389. with apical margin (figure 652)...... prosopis (LeConte) Type locality.—Texas, Bentsen State Park, 3(1) Median lobe of genitalia with thickened frame surround- Hidalgo Co. ing ventral valve; internal sac armature as in figure 646. Distribution.—NM, OK, TX, HI (introduced); ...... bottimeri Kingsolver Mexico. Median lobe of genitalia with membranous frame sur- Host plants.—Prosopis glandulosa glan- rounding ventral valve; internal sac armature as in figure dulosa, P. pallida (Hawaii only), P. reptans 655. cinerascens, P. reptans reptans...... prosopis (LeConte) Natural enemies.—Heterospilus prosopidis; Lariophagus texanus; Stenocorse bruchivora

156 Crawford. These parasites are listed as Algarobius prosopis: Bridwell 1946:54; attacking Algarobius prosopis but because Blackwelder and Blackwelder 1948:45; their distribution is given as Texas or Ha- Werner and Butler 1958:7; Arnett waii, the association instead is undoubt- 1962:957; Bottimer 1968c:1021; John- edly with A. bottimeri. son 1968:1268; Kingsolver 1972b:119; Immatures.—Not described. Center and Johnson 1974:1101; King- solver 1972b:119; Smith and Ueckert Discussion.—Although the males of this 1974:61; Swier 1974:5; Pfaffenberger species are externally indistinguishable and Johnson 1976:31; Kingsolver et al. from those of Algarobius prosopis, the two 1977:114; Ward et al. 1977:5; John- species belong to different species groups son and Kingsolver 1982:416; Johnson based on male genitalia and the position 1983c:27; Kingsolver 1986:113; Borow- of the female pygidial sulcus. Females are iec 1987:105; Udayagiri and Wadhi easily differentiated in the United States by 1989:69; Decelle 1990:20; Johnson and the position of the sulci (compare figures Siemens 1997a:38. 645 and 652). Bruchus uniformis LeConte 1858:77; Horn The distribution lies largely within Texas 1873:333; Fall 1901:160, 1910:174; with only scattered localities in border- Cushman 1911:508; Zacher 1952:463. ing states, plus Hawaii. Collections in Mylabris uniformis: Kannan 1923:17; Essig Hawaii indicate that A. bottimeri attacks 1929a:486 (desertorum and prosopis as only seeds of Prosopis pallida, itself an synonyms). immigrant into Hawaii. None of the Texan Algarobius uniformis: Blackwelder and species of Prosopis are currently in Hawaii Blackwelder 1948:45; Johnson according to Fosberg (1966:134). 1968:1268. Records listing Bruchus prosopis or Alga- Bruchus desertorum: LeConte 1858:77; robius prosopis from Hawaii prior to 1972 Horn 1873:328, 1894:345; Schaeffer are almost certainly A. bottimeri. Bridwell 1907:292; Fall 1910:174; Cushman (1929a:43) was the first to recognize that 1911:508; Zacher 1952:462. the Hawaiian specimens were not A. prosopis, but he never described the species. Mylabris desertorum: Leng 1920:305 Acanthoscelides desertorum: Blackwelder Prosopis velutina and P. glandulosa were 1946:759. recently inadvertently introduced into South Africa and both Algarobius bottimeri Algarobius desertorum: Blackwelder and A. prosopis have been introduced as and Blackwelder 1948:45; Johnson possible biocontrol agents. (B. Grobbelaar, 1968:1268. personal communication, 1990; Hoffmann Color.—Integument brownish yellow; eyes, et al. 1993) female pygidial sulci, and humeri black. Vestiture of yellow, white, dark brown, light Algarobius prosopis (LeConte) brown, and black fine setae in variable pattern (figures 648 and 649); venter of body (Figures 61, 648–656) mostly white with some yellow spots. Fe- Bruchus prosopis LeConte 1858:77; Horn male pygdial sulci located on apical margin 1873:331; Sharp 1885:475; Fall either side of midline. 1901:160; Schaeffer 1907:299; Cush- Male genitalia.—As in figures 655 and man 1911:497,507; Zacher 1952:462. 656. Ventral valve subtriangular, keeled; Mylabris prosopis: Leng 1920:305. dorsal hood rounded; armature of inter- Acanthoscelides prosopis: Blackwelder nal sac consisting of two curved sclerites 1946:760. connected by a bridge; an elongate, tapered spine flanked each side by a slender,

157 curved spine; two tapered, crossed spines of the pygidial sulci (compare discussion of in middle of sac; and an amorphous, thinly Algarobius bottimeri). sclerotized structure toward apex; closure Teran (1967) first illustrated the male geni- valve circular; lateral lobes flat (figure 656), talia of this species. cleft one-half their length. Algarobius prosopis is the most common Size.—Body length 2.1–4.1 mm; width bruchid attacking mesquite in its range. 1.0–2.2 mm. This species has been introduced into Type depository.—MCZC (lectotypes: deser- South Africa for possible control of Proso- torum, No. 4468; prosopis, No. 4470; uni- pis (B. Grobbelaar, personal communica- formis, No. 4471; all designated by King- tion, 1990) Scullen and Wold (1969) re- solver 1986). ported it as the prey of the hymenop-teran Type locality.—California, Colorado Desert Cerceris truncata Cameron. (desertorum, prosopis, and uniformis). Kistler (1982) reported on the effects of Distribution.—AZ, CA, NM, NV, TX, UT; temperature on this species. Mexico; South Africa, Arabian Peninsula Swier (1974) investigated the interac- (J. Decelle, personal communication, tions of four species of bruchids attacking 1990). Prosopis velutina in Arizona—Algarobius Host plants.—Prosopis chilensis, P. glan-du- prosopis, Mimosestes amicus, Mimosestes losa torreyana, P. laevigata, P. velutina, P. protractus, and Neltumius arizonensis. See pubescens, P. reptans cinerascens, P. alba also Kingsolver et al. (1977). (introduced into Arizona). Zacher’s (1952) inclusion of Hymenaea courbaril and Aca- Genus Althaeus Bridwell cia confusa as hosts of this bruchid should be discounted. Althaeus Bridwell 1946:55; Bradley 1947:40; Arnett 1962:956–957; Bot- Natural enemies.—Cerceris truncata; Chari- timer 1968c:1021,1040; Johnson topodinus terryi; Glyptocolastes texanus; and Kingsolver 1982:416; Borow- Heterospilus prosopidis; Horismenus mis- iec 1987:95; Kingsolver et al. 1989. souriensis, H. productus; Lariophagus Type species: Bruchus hibisci Olivier texanus; Stenocorse bruchivora; Urosigal- 1795:21, monotypic. phus bruchi, U. bruchivorus, U. neobruchi; Uscana semifumipennis. Bridwell separated Althaeus by the broad, serrate subapical denticle on the ventral Immatures.—Kannan 1923:17 (as Mylabris margin of the metafemur (figure 660). The uniformis, first instar); Pfaffenberger and generic characterization is here expanded: Johnson 1976:31 (first instar). Head (figure 664) with frontal carina eva- Discussion.—Specimens reared from Pro- nescent, ocular sinus one-half length of sopis pubescens (screwbean) average two- eye, antenna subserrate and moderately thirds the size of those reared from the dimorphic (figure 665), extending to middle straight bean (P. glandulosa, P. torreyana, of metepisternum; pronotum semicircular and P. velutina) and are usually uniformly (figure 663), convex, lateral margins arcu- yellowish with little or no pattern. Male ate, lacking lateral carina; elytral striae genitalia, however, are identical to those of parallel, 3rd and 4th striae sometimes the larger specimens. This species exhib- minutely denticulate basally; basitarsus its a wide range of intensity of color pat- of fore leg short (figure 667), subequal to tern with females usually, but not always, 2nd segment, that of mid leg 1.5 times as darker than males. In zones of distribution long as 2nd (figure 668); metafemur not overlap in eastern New Mexico and western strongly swollen; mucro of metatibia short, Texas, males must be dissected, but fe- one-fifth or less length of basitarsus (figure males can be distinguished by the position 660).

158 Apex of internal sac of male genitalia with numerous small denticles (figures 661, Key to Species of Althaeus 670, and 676). 1 Legs black except tarsi reddish brown; basal four Althaeus is apparently closely related to or five segments of antenna dark red, antennal club Acanthoscelides. The character combina- black; metatibial mucro subequal in size to lateral and tion of elongate, serrate femoral denticle, coronal denticles; denticle of metafemur as in figure short, broad body, ocellate elytral pubescence, and short fore tarsal segments will 675...... steineri Kingsolver separate the three species included here Some portion of legs usually red or reddish yellow; from those species now placed in Acanthos- antenna usually yellow, sometimes dusky; metatibial celides. mucro longer than lateral and coronal denticles; denticle Borowiec (1987) redescribed the genus but of metafemur as in figures 660 and 666...... 2 based his description only on Althaeus 2(1) Male genitalia with slender ventral valve, apex attenuated hibisci. He failed to note the minute basal (figure 670); lateral lobes apically rounded (figure 671) elytral tubercles, the dimorphic antennae, ...... hibisci (Olivier) and the short tarsal segments. Male genitalia with broad ventral valve, apex minutely The geographical range of A. hibisci lies pointed (figure 661); lateral lobes apically, obliquely predominantly in the Gulf Coast and East- truncate (figure 662) ern Seaboard States with some scattered ...... folkertsi Kingsolver records in the Mississippi Valley up to Il- linois. The range of A. folkertsi is predominantly midwestern from Oklahoma and Althaeus folkertsi Kingsolver Kansas eastward to Virginia and Maryland, where it is sympatric with A. hibisci. The (Figures 65, 657–662) third species, A. steineri, has been collect- Althaeus folkertsi Kingsolver, in Kingsolver ed only in eastern North Carolina. et al. 1989:67. The species included in Althaeus apparent- Color.—Similar to Althaeus hibisci but ly are obligatorily associated with species generally with a more grayish appearance in the plant family Malvaceae. than yellowish; however, the variation in Taxonomy: Kingsolver et al. 1989. vestitural color in both species makes this an unreliable character. Color of legs and antennae are also variable in both species. Structure (figures 657 and 658).—Similar to Althaeus hibisci but with distinctive differences in the shape of the ventral valve and lateral lobes of the male genitalia. Antenna (figure 659) slightly dimorphic. Male genitalia.—As in figures 661 and 662; median lobe slender (figure 661) in apical one-half, broadly rounded at base; ventral valve ogival with small apical nipple; base of internal sac finely denticulate, with coarser denticles in middle of sac and 11 thornlike denticles in apical portion, each with fine spines on its enlarged base, apex of sac with ringlike, denticulate, apical band; lateral lobes (figure 662) short, broad, obliquely truncate at apex.

159 Size.—Body length 1.9–2.5 mm; width legs largely reddish yellow, sometimes with 1.1–1.7 mm. bases of femora black, hind leg (figure 666) Type depository.—USNM. usually basally black, apically red, extent of black infusion variable, metatibia and Type locality.—Maryland, Plummers Island tarsus reddish. Vestiture of short, slender, (Potomac River). silvery gray to yellowish-gray hairs, pat- Distribution.—AR, DE, DC, IL, IN, IA, KS, tern of hairs on elytra mottled, dark spaces LA, MD, MO, MS, NJ, OH, OK, PA, TX, VA, between patches of hair each centered on WV. a minute golden hair; pronotum with hairs condensed into two median patches and Host plants.—Abutilon theophrasti; Hibiscus two subbasal patches, remaining areas on moscheutos lasiocarpos, H. militaris. Visit- disk mottled; venter of body evenly clothed ing records: Amelanchier; Crategus mollis; with gray hairs except for prominent Hibiscus; Malva; Spiraea; swamp mallow; patches on mesepisternum, caudal end of wafer ash. metepisternum, and dorsal border of ab- Natural enemies.—None recorded. dominal sterna. Immatures.—Pfaffenberger (first larval Structure.—Body stout (figure 663), about instar) and Gibb (larva and pupa) (In King- 2 times as long as wide, strongly convex solver et al. 1989). above. Head ovate (figure 664), eyes pro- Discussion.—See Althaeus hibisci for com- tuberant; vertex finely, obsoletely punc- parisons and distribution. Although this tate, frons finely rugose, clypeus rugosely species breeds in seeds of Hibiscus and punctate; ocular sinus about one-half Abutilon and adults are found in the flow- length of eye, postocular lobe nearly at ers, they also feed on pollen of several spe- right angles to side of head; antenna capi- cies of plants in other families. tate (figure 665), segments eccentrically expanded beginning with 5th segment, Gibb (in Kingsolver et al. 1989) described terminal segment elliptical, antenna reach- oviposition and the biology of Althaeus folk- ing middle of metepisternum. Pronotum ertsi on Abutilon theophrasti. nearly semicircular in dorsal aspect (figure 663), basal lobe prominent, disk evenly Althaeus hibisci (Olivier) convex, microfoveolate, each foveola with a (Figures 663–671) micro-seta inserted under its anterior rim, lateral margin subcarinate from posterior Bruchus hibisci Olivier 1795:21; LeConte corner about one-third distance to anterior 1824:170; Hubbard and Schwarz margin, propleuron finely punctate, cervi- 1878:660; Fall 1910:172 (in key only); cal sulcus short, deep; prosternum short, Cushman 1911:503; Weiss 1919:5; triangular, narrow before procoxae; me- Weiss and Dickerson 1919:49; Zacher sosternum narrowly trapezoidal, slightly 1952:162,478. emarginate apically; postmesocoxal sul- Bruchus transversus Say 1831:3. cus narrow paralleling rim of coxal cav- Bruchus transvexus: Horn 1873:338 (mis- ity. Elytra together as wide as long (figure spelling). 663); striae regular in course, free apically, Althaeus hibisci: Bridwell 1946:55; Brad- originating basally from minute pits un- ley 1947:40; Arnett 1962:957; Johnson der intermittent basal bead; interstices and Kingsolver 1982:416; Borowiec flat, finely imbricate except in intermittent 1987:95; Kingsolver et al. 1989:63; bare spots surrounding each golden seta; Udayagiri and Wadhi 1989:69. meta-sternum with shallow median sulcus; fore, middle, and hind tarsi as in figures Color.—Body black; antenna mostly red- 667, 668, and 669, respectively; metacoxa dish yellow, sometimes with club segments densely punctulate; metafemur oblong-el- partly or wholly piceous; fore legs and mid liptical (figure 666), denticle length equal

160 to tibial width at their juncture, posterior Daucus carota; Hibiscus; Kosteletzkya virgi- margin of denticle usually with two fine nica; mallow; marsh mallow; okra; swamp teeth; metatibia nearly straight, bent only mallow; thistle. at extreme base, gradually dilated toward Natural enemies.—Uscana semifumipennis. apex, lateral, ventral, and dorsomedial carinae distinct and entire, ventrolateral Immatures.—Not described. carina obsolete apically; mucro acute, only Discussion.—Weiss and Dickerson (1919) slightly longer than lateral and coronal listed several localities and host plants for denticles; metatarsus including claw seg- Althaeus hibisci in New Jersey, but since ment equal in length to metatibia. Abdo- A. folkertsi is also found in New Jersey, the men of male with 1st sternum l.5 times as records could apply to either species. Their long as remaining four sterna combined, descriptions of the life stages and life his- 1st sternum without pits but with vestiture tory of A. hibisci are likewise ambiguous. in curved pattern on either side of midline; Olivier described Bruchus hibisci in 1795 5th sternum broadly emarginate to receive from “Carol.”— undoubtedly referring to apex of pygidium; 1st sternum of female Carolina. The type in the Museum National equal in length to remaining sterna com- d’Histoire Naturelle in Paris is a badly bined, 5th sternum with caudal margin rubbed and broken female specimen. not emarginate; pygidium of male broadly Enough of the specimen remained to de- obovate, evenly convex, disk sparsely mi- termine that it is probably conspecific with crofoveolate resulting in speckled appear- specimens from Canadys, SC. This locality ance, each foveola with a fine seta set in its is outside the known range of the new spe- anterior rim, vestiture patchy with vague cies A. folkertsi, and the red coloration of median stripe; female pygidium similar to the metafemur of the hibisci type precludes male’s except less strongly convex. its representing the new species A. steineri. Male genitalia.—As in figures 670 and This is one of the most commonly collected 671; median lobe slender in apical one- bruchids in the Middle Atlantic states. It half (figure 670), broadly rounded basally; commonly develops in several species of ventral valve elongate, tapered to blunt Hibiscus but has been bred from seeds apex; internal sac minutely denticulate in of Abutilon theophrasti (butterprint or basal one-fifth, middle of sac with coarser velvetleaf), a common weed in fields and denticles, apex with 12 thornlike spines, waste places. the base of each denticulate, and a ringlike, denticulate band; lateral lobes (figure I have been unable to discover any consis- 671) expanded apically, apex of each lobe tent external differences between Althaeus rounded and setose, cleft for one-third hibisci and A. folkertsi. Females of the two their length. species are not distinguishable from one another at present. Only the male genitalia Size.—Body length 1.5–2.5 mm; width provide diagnostic characters. Both species 0.9–1.5 mm. are readily separated from A. steineri by Type depository.—MNHP. the characters in the key. Type locality.—”Carol.” (probably Carolina). Althaeus steineri Kingsolver Distribution.—AL, AR, DC, DE, FL, GA, IL, KY, LA, MD, MI, MS, MO, NJ, NY, NC, OK, (Figures 672–677) PA, SC, TN, TX, VA. Althaeus steineri Kingsolver, In Kingsolver Host plants.—Abutilon theophrasti; Hibis- et al. 1989:60. cus aculeatus, H. moscheutos lasiocarpus, Color.—Integument black, 1st to 5th anten- H. moscheutos moscheutos, H. moscheu- nal segments reddish yellow, 6th to 11th tos palustris. Floral records: Crataegus;

161 piceous, tarsi dark red to piceous, apex Fore legs with basitarsus short, mid legs of metatibia sometimes piceous. Vestiture with basitarsus slightly longer that those of of silvery gray, slender hairs; pronotum fore legs; metacoxal face with fine, densely with hairs evenly distributed on disk but placed punctures, metafemur (figure 675) with small paired patches at apical one- with ventral margin nearly straight, flat, third and on each side of basal lobe; elytral with external fringe of fine setae giving ap- vestiture evenly distributed but interstices pearance of fine carina, mesoventral mar- with small, ocellate bare spots each cen- gin with single, small, subapical denticle tered on a setose puncture (figure 672), finely serrate on caudal margin; metatibia scutellum densely setose; pygidium with (figure 675) nearly straight except at ex- mottled pattern (figure 673), vaguely dens- treme base; lateral, ventral, and dorsome- er along midline; body beneath and legs dial carinae complete, latero-ventral carina uniformly covered with gray hairs. ending one-half distance from base, mucro Structure.—Head turbiniform, eyes protu- shorter than lateral denticle, three coro- berant, ocular sinus about one-half length nal denticles subequal in length to lateral of eye, postocular lobe narrow, fringed; denticle. Abdomen with 1st sternum sub- frons broad with frontal carina obsolete, equal in length to remaining four together, or at most a denuded median line, vertex 5th sternum broadly emarginate in male, and frons finely punctate, clypeus ob- evenly arcuate in female; pygidium subtri- scurely punctate; 6th to 10th antennal angular, integument obsoletely foveolate, segments eccentric forming a loose, terete hairs arranged in obscure ocellate pattern; club, 11th conical (figure 674). Pronotum disk more strongly arcuate in male. semicircular (figure 672), about 1.5 times Male genitalia.—As in figures 676 and 677; as long as wide, lateral margins evenly median lobe (figure 676) elongate, slender arcuate, slightly flared at extreme base, in apical two-thirds, base slightly ex- disk almost evenly convex, basal lobe with panded; ventral valve broadly ogival, apex short, narrow sulcus; surface with densely, minutely pointed, basal one-half of inter- evenly placed, round-bottomed foveo- nal sac densely lined with blunt denticles, lae, each foveola with a short, white hair apical portion with about 18 spiny den- emerging from beneath its anterior rim; ticles and apical denticulate band; lateral lateral carina obsolete, margin of disk an lobes (figure 677) cleft about one-half their obtuse ridge; cervical sulcus fine, extend- length, rounded, expanded and apically ing dorsad to a point above eye; proster- setose. num triangular, short, extending about Size.—Body length 2.3–2.5 mm; width two-thirds length of coxa; mesosternum 1.5–1.6 mm. truncate; postmesocoxal sulcus narrow, following contour of coxal cavity. Scutel- Type depository.—USNM. lum quadrate, nearly covered with white Type locality.—North Carolina, Dare Co., hairs. Elytra together slightly longer than Frisco, Cape Hatteras National Seashore. wide (figure 672), margins evenly arcuate, widest at middle; striae regular in course, Distribution.—NC. only slightly deflected at extreme base, Host plants.—Kosteletzkya virginica. 2nd to 5th striae arising from basal pits Natural enemies.—None recorded. beneath marginal bead, denticulate; striae deep, lacking punctures but with regularly Immatures.—Not described. placed, short, curved, white setae, each set Discussion.—This species is distinguished in slightly widened space in stria (probably by the key characters—black legs, short an expanded strial puncture); all striae free mucro, smaller serrations on femoral apically; intervals prominently punctate, denticle, and the male genitalia. The lat- each puncture setose, surface between ter organs more closely resemble those punctures finely to obsoletely imbricate.

162 of Althaeus folkertsi than of A. hibisci, Schoenherr, not Thunberg. Type spe- whereas the external body characters seem cies: Bruchus brasiliensis Thunberg to more closely associate A. folkertsi with 1816, by automatic fixation. A. hibisci. This species might be confused Pedalophus (error), Bottimer 1968c:1041. with Acanthoscelides alboscutellatus (Horn) Pseudopachymerus Pic 1913a:10. Replace- because of the black body and appendages ment name for Pachymerus “Latreille” and vaguely ocellate pattern of setae on the of Schoenherr, not Thunberg. Type elytra, but the pronotum of A. alboscu-tel- species: Bruchus brasiliensis Thunberg latus is subconical, and the scutellum is 1816, by automatic fixation. contrastingly pure white. Falsobruchus Pic 1913b:110. Type species: Althaeus steineri is known only from collec- Bruchus cristatus Fahraeus 1839:122, tions made on the Cape Hatteras National by monotypy. Proposed as subgenus of Seashore. Although the host is known from Pachymerus Schoenherr. other localities on the mainland, the bru- Body broad, deep; elytra subquadrate chid has not been collected from there. (figures 69 and 683); vestiture dense everywhere on body. Vertex and frons densely Genus Caryedes Hummel punctulate, frontal carina prominent, Caryedes Hummel 1827:11. Type species: length of gena between eye and base of Bruchus faldermanni Mannerheim, In mandible 1.5–2 times as long as width of Hummel 1827:10; see Kingsolver and basal antennal segment, dorsal margin of Whitehead 1974:343. antennal cavity carinate; antenna gradually clavate. Pronotum subconical (figure Pachymera Berthold 1827:378. Latiniza- 69), lateral margins incurved; disk with tion of “pachymere” (Latreille 1825), a longitudinal median gibbosity, lateral sub- vernacular name. basal gibbosities prominent, separated Pachymerus Schoenherr 1833:84. Type from median gibbosity by shallow sulcus; species: Bruchus brasiliensis Thunberg discal punctation fine, hidden by vestiture. 1816:45, by original designation (Bru- Scutellum small, quadrate. Elytra together chus faldermanni Mannerheim listed as long as wide (figure 69), medially de- in synonymy) (not Pachymerus Thun- pressed; striae mostly hidden by vestiture berg 1805). In describing Pachymerus, in Caryedes helvinus, 3rd and 4th striae Schoenherr (1833) listed two groups laterally deflected to subbasal denticles under the genus. One was the group set on rounded gibbosity; interstices flat. set out by Latreille (1825) as Pachy- Pygidium subtriangular (figures 679 and mere but not formally described. The 684). Metacoxal face densely, evenly punc- other group was later expanded into the tate except fossula glabrous; metafemur subfamily Pachymerinae, an entirely strongly inflated, mesoventral margin with different bruchid group. The appelation three to four minute spines nearly hid- “Latreille” was used by by Gistel (1848) den in vestiture and one long and three or and Pic (1913a) to indicate to which four short, subapical denticles; metatibia group they were referring. slender, curved; lateroventral, ventral, and Andromisus Des Gozis 1881:cxiii. (Original dorsomesal carinae complete; mucro slen- spelling Adromisus, emended by Des der, curved, one-third to one-half as long Gozis 1885:125). Replacement name for as basitarsus. Pachymerus Latreille: of Schoenherr, Median lobe of male genitalia elongate (fig- not Thunberg. Type species: Bruchus ure 681), fractured near apex; lateral lobes brasiliensis Thunberg 1816, by auto- fused (figure 682), straplike, divided only matic fixation. at apex. Pedapholus Gistel 1848:xi. Replacement name for Pachymerus “Latreille” of

163 Two species in this Neotropical genus Caryedes helvinus (Motschulsky) [Caryedes incensus (Sharp) and C. helvi- (Figures 26, 68, 69, 678–682) nus (Motschulsky)] were recovered from field-grownCentrosema macrocarpa at Ft. Pachymerus helvinus Motschulsky Pierce, FL, in April 1989. The known geo- 1874:244. graphical ranges of both species do not Pseudopachymerus helvinus: Pic 1913a:11. normally extend north of southern Mexico. Caryedes helvina: Blackwelder 1946:758. The description applies to these two species only. For a more complete description Caryedes helvinus: Kingsolver and White- of the genus, see Kingsolver and White- head 1974:376; Udayagiri and Wadhi head (1974:345). 1989:73. Pachymerus scabricollis Chevrolat 1877: The only species in Florida with similar xcviii. morphological characteristics is Mero-bruchus lysilomae Kingsolver (figure 758), but Pseudopachymerus scabricollis: Pic M. lysilomae is more slender than either 1913a:12. species of Caryedes, the genal length is Caryedes scabricollis: Blackwelder less than the width of the 1st antennal seg- 1946:758. ment, and the lateral margins of the prono- Bruchus calderensis Sharp 1885:444. tum are arcuate, not incurved. Pseudopachymerus calderensis: Pic Gibbobruchus mimus (Say), which has 1913a:10. similar characteristics, has been collected Caryedes calderensis: Blackwelder at Gainesville, FL, from its preferred host 1946:758. plant, Cercis canadensis. Gibbobruchus is Pseudopachymerus multimaculatus var. easily separated from other North Ameri- binotatus Pic 1930a:36. can genera by the serrate lateroventral carina of the metafemur, the lateral pol- Caryedes bicoloripes var. binotata: Black- ished spots of the 1st and 2nd abdominal welder 1946:757. segments, and the polished, heart-shaped Color.—Integument dark red to black, pygidial spot of the female. everywhere concealed by dense vestiture; head black, vertex sometimes red; anten- Revision: Kingsolver and Whitehead 1974. nae red except 6th to 9th segments darker red to black; pronotum, elytra, pygidium, and venter of body usually dark red; pro- Key to Caryedes of the United States notum usually with linear spots on raised 1. Vestiture dense, short, yellowish white with brown or portions of disk; elytra with black maculae black lateral spots (figure 69); pygidium mostly yellowish at humeri and middle of lateral margin; pygidium with two subapical spots some- white with vague darker areas and two subapical brown times vaguely defined. Vestiture of yellow- or black spots (figure 679); front and middle legs yellow, ish-gray, white, and black setae in mottled hind leg yellowish white (figure 680) with vague brown pattern on body, black setae on black banding integumental spots...... helvinus (Motschulsky) Structure.—Body broad (figure 69), vertex Vestiture largely dark brown with some intermixed white finely punctate, frontal carina fine but patches, apical one-half of elytra with diagonal rows distinct (figure 678); ocular index 4:1; ocu- of white setae (figure 683); scutellum and basal dart of lar sinus one-half as long as eye; antenna pronotum contrastingly white; pygidium (figure 684) clavate. Pronotum subconical (figure 69), yellow and white with median dark brown denuded spot, lateral margins incurved, basal lobe well sometimes with brown subapical spots; hind leg dark defined, middle of disk gibbous, vaguely brown with white setal patches (figure 685) sulcate, with subbasal gibbosity either side ...... incensus (Sharp) of median gibbosity; lateral carina effaced.

164 Scutellum quadrate, densely pubescent. has not been found elsewhere in the Unit- Elytra together as long as wide (figure 69), ed States or Canada. The host plants are medially depressed between 7th striae, not found naturally in the United States. striae appearing as thin lines in dense vestiture, 2nd stria slightly divergent at base, Caryedes incensus (Sharp) 3rd and 4th striae ending subbasally in (Figures 683–687) dentate gibbosities. Metacoxal face densely, finely punctate, densely pubescent. Hind Bruchus incensus Sharp 1885:445. leg as in figure 680; pecten with one long Pseudopachymerus incensus: Pic 1913a:11. denticle and three short denticles; metati- Caryedes incensa: Blackwelder 1946:758. bia sinuate, apically expanded, lateral, ventral, and dorsomesal carina complete Caryedes incensus: Kingsolver and White- to apex, lateroventral carina aborted near head 1974:379; Mead 1989:2; Udaya- base of mucro; mucro slender, three-fifths giri and Wadhi 1989:74. as long as basitarsus. Abdomen not modi- Color.—Body and appendages dark red to fied except 5th sternum of male deeply black; antenna with 6th to 10th segments emarginate for apex of pygidium; pygidium black, remainder red. Vestiture of golden of male convex with subapical and lateral brown, black, and white setae in pattern depressions, of female nearly flat with only shown in figure 683, head mostly dark subapical depressions (figure 679). brown, vertex paler dorsally; pronotum mottled with black and brown hairs, nar- Male genitalia.—As in figures 681 and 682; row median band white, scutellum and median lobe slender (figure 681), similar prescutellar spot usually white, sometimes to that of C. incensus except internal sac yellowish; elytra largely black with brown armature sparse, apex of sac with bilobed patch at one-fourth from base, disk with structure; lateral lobes elongate (figure ill-defined diagonal bands of intermixed 682), fused nearly to rounded apical lobes. brown and white hairs, 7th interstice with Size.—Body length 2.8–3.7 mm; width small but prominent white spot; pygidium 1.9–2.4 mm. variable, usually with median and subapi- Type depository.—ZMUM. cal brown spots on white to yellowish-white background (figure 684); venter of body Type locality.—Venezuela, Santa Lucia. largely brown but with yellowish-white to Distribution.—FL (possibly established near white spots on mesepimeron, metepister- Ft. Pierce); Mexico to Brazil. num, lateral angle of metacoxal face, and lateral margins of abdominal segments; Host plants.—Centrosema macrocarpa, C. hind legs with vague yellow banding. pubescens. Structure.—Head 1.6 times as long as Natural Enemies.—None recorded. width across eyes; ocular index 3:1; ocu- Immatures.—None described. lar sinus one-half as long as eye; antenna Discussion.— This species can hardly be clavate, club segments strongly eccentric, confused with any other bruchid in this antenna reaching middle of metepister- handbook. Salient characters are the num; pronotum (figure 683) with median dense yellowish-gray appearance, de- gibbosity elongate and prominent, bisected pressed medial two-thirds of elytra, black by shallow sulcus, subbasal gibbosi- elytral humeri and lateral marginal macu- ties elongated, prominent, basal margin lae, pronotal color pattern, pygidial spots, sharply sinuate; disk densely punctulate; and long, slender mucro. lateral carina threadlike. Scutellum prominent. Elytra convex (figure 683) but with The species apparently escaped from ex- area around scutellum depressed; striae perimental plots near Ft. Pierce, FL, but

165 shallow, parallel, punctures prominent, Kingsolver 1982:413; Borowiec 3rd and 4th arising from minute denticles 1987:75; Udayagiri and Wadhi 1989:81. on prominent subbasal gibbosity; inter- Small to medium sized beetles (2.3–4.7 stices minutely imbricate. Hind leg as in mm). Three species in the United States. figure 685, mesoventral margin with three Body subdepressed, elytra subquadrate; minute median denticles, pecten with one head with prominent or evanescent frontal long and four shorter denticles; mucro carina; eye with moderately deep ocular one-third as long as basitarsus. Pygidium sinus; antenna with club segments strong- densely punctulate, apex of male distinctly ly serrate, not sexually dimorphic (figure reflexed. 692). Pronotum bell-shaped (figure 688), Male genitalia.—As in figures 686 and 687; lateral margins incurved, disk with median median lobe long, slender (figure 686); ven- and subbasal gibbosities, median gibbos- tral valve ogival with apex acute; internal ity sulcate; lateral carina evanescent or sac densely lined with fine denticles and lacking. Scutellum small, quadrate, biden- one saberlike sclerite; apex of sac bilobed; tate. Elytra with striae narrow (figure 688), lateral lobes long (figure 687), fused two- deep, 3rd and 4th deflected laterad at base thirds their length, broadly spatulate api- and ending in prominent gibbosity; lateral cally. umbones prominent. Pygidium dimorphic, Size.—Body length 2.7–3.3 mm; width pubescent in male, at least partly glabrous 1.7–2.2 mm. in female; 1st and 2nd abdominal sterna with polished lateral spots. Fore leg and Type depository.—BMNH. mid leg not modified; hind leg with femur Type locality.—Panama, Volcan de Chiriqui. dorsoventrally strongly swollen, ventrome- Distribution.—FL (possibly established near sal margin with one long denticle usually Ft. Pierce); Costa Rica to Colombia. separated by gap from three or four distal denticles; ventrolateral margin with row Host plants.—Centrosema macrocarpa of short, blunt denticles; metatibia arcu- (Florida), C. pubescens (Costa Rica) ate fitting ventral margin of femur; mucro Natural enemies.—None recorded. elongate, acute. Immatures.—Not described. Male genitalia with median lobe moderately broad, ventral valve subtriangular; internal Discussion.—Characters given in the gener- sac densely armed with fine spicules. ic discussion will separate Caryedes incensus from Gibbobruchus and Mero-bruchus. Genus ranges from United States to Brazil. No other U.S. bruchid species is likely to All three U.S. species were placed in the be confused with C. incensus. mimus group by Whitehead and Kingsolver (1975a). Genus Gibbobruchus Pic Revision: Whitehead and Kingsolver 1975a. Pachymerus subg. Gibbobruchus Pic 1913b:110. Type species: Bruchus spe- culifer Gyllenhal 1833, by subsequent designation, Bridwell 1932:105. Pseudopachymerus subg. Gibbobruchus: Pic 1913a:10. Gibbobruchus: Bridwell 1932:105; Blackwelder 1946:762; Bottimer 1968c:1022,1039; Whitehead and King- solver 1975a:169; Johnson and

166 Color.—Integument reddish brown to black, Key to Species of Gibbobruchus in more teneral specimens yellowish brown with dark red variegation; antenna with 1 Pygidial spot round or oval (figures 691 and 699); pecten basal segments yellowish, middle segments with 1st denticle separated from the remaining by distinct often infuscated; pronotum and elytra var- gap (figures 693 and 701)...... 3 iegated red; dorsal half of metafemur red, Pygidium with large, polished central spot (figures 691, ventral half black; female pygidium with 699, and 707) (females)...... 2 large, oval, glabrous spot. Vestiture white, Pygidium with at most a small, elongate median spot yellow, light brown, dark brown, and black; (figures 690, 698, and 706) (males)...... 4 head yellow; pronotum mostly yellow but somewhat variegated, posterior slope of 2(1) Pygidial spot cordate (figure 707); pecten of metafemur median gibbosity with elongate, intensely with denticles regularly spaced, no gap behind 1st den- white patch; elytra variegated light brown, ticle (figure 709) dark brown, yellow, and white with me- ...... mimus (Say) dian V-shaped spot velvety black; male Pygidial spot round or oval (figures 691 and 699); pecten pygidium yellow except paler spots on twin with 1st denticle separated from the remaining by distinct gibbosities, female pygidium with brown- gap (figures 693 and 701)...... 3 ish-yellow fringe around glabrous median spot; venter of body white except yellow 3(2) Elytra with short, V-shaped, median dark brown spot and brown spots on lateral sclerites; row (figure 688); base of pronotal gibbosity with elongate, of small, white spots on dorsal margin of pale spot...... cristicollis (Sharp) abdominal sterna; 1st, 2nd, and 3rd sterna Elytra with elongated, velvety black, median spot (figure with lateral, glabrous patches. 696); base of pronotal gibbosity lacking elongate pale Structure.—Vertex densely, finely punc- spot; basal triangle of pygidium prominent (figure 699) tate, transverse sulcus shallow, frons with ...... divaricatae Whitehead and Kingsolver linear punctures, frontal carina not prominent (figure 689); ocular index 3.3:1; ocu- 4(1) Pygidium sparsely pubescent, convex, with short median lar sinus five-eighths as long as eye; anten- sulcus (figure 698); basal triangle of pygidium prominent na moderately eccentric from 4th segment, ...... divaricatae Whitehead and Kingsolver terminal segment elliptical (figure 692); Pygidium densely, conspicuously pubescent...... 5 antenna extended to humerus. Pronotum with strongly incurved lateral margins 5(4) Pygidium with two projecting gibbosities (figures 688 (figure 688); median gibbosity with four and 690)...... cristicollis (Sharp) slight elevations, median sulcus shallow, Pygidium lacking gibbosities but with three glabrous, subbasal gibbosities prominent, ridgelike; subapical spots (figure 706) cervical sulcus hidden in vestiture. Scu- ...... mimus (Say) tellum quadrate, emarginate. Elytral disk depressed between 6th interstices (figure 688), slight ridge extending from humerus Gibbobruchus cristicollis (Sharp) diagonally to middle of elytron; striae sinuate, subparallel, discal striae divergent (Figures 688–695) in basal one-half, 3rd and 4th ending in Bruchus cristicollis Sharp 1885:442. denticulate basal gibbosity, interstices Pseudopachymerus cristicollis: Pic densely pubescent, punctulate-imbricate; 1913a:10. metacoxal face densely punctate, setose, setae directed laterad; pecten with four to Caryedes cristicollis: Blackwelder five denticles irregularly positioned, ventro- 1946:758. lateral carina denticulate (figure 693). First Gibbobruchus cristicollis: Whitehead and sternum of male with yellow patch of fine Kingsolver 1975a:190; Johnson and setae, 5th sternum broadly emarginate. Kingsolver 1982:413; Udayagiri and Wadhi 1989:81.

167 Male genitalia.—As in figures 694 and 695; with scattered white patches; elytra varie- ventral valve semicircular (figure 694), gated yellowish brown and white, elongate apex produced, internal sac with linear median patch velvety black (figure 696); rows of fine spicules in basal one-third, male pygidium with fine, white setae and apical two-thirds lined with fine denticles; scattered yellowish patches, conspicuous lateral lobes concave on mesal faces (figure white basal triangular patch (figure 698), 695), bowed, cleft two-thirds their length. female pygidium with white basal patch Size.—Body length 2.5–4.7 mm; width and scattered white patches (figure 699); 1.5–2.8 mm. venter of body mostly fine, white setae, row of white patches on sides of abdomen, yel- Type depository.—BMNH. low fringes on mesopleuron and metapleu- Type locality.—Mexico, Oaxaca, Yolos. ron. Distribution.—TX; Mexico to Costa Rica. Structure.—Vertex, frons, and basal one- half of clypeus densely microfoveolate; Host plants.—Bauhinia lunarioides (Texas); ocular index 3:1 (figure 697); ocular sinus B. coulteri, B. divaricata, B. macranthera, B. 0.7 as long as eye; antenna (figure 700) ec- pauletia (extralimital). centric from 5th segment. Pronotum bell- Natural enemies.—None recorded. shaped (figure 696), apex rounded, basal Immatures.—Not described. margin strongly sinuate, lateral margins strongly incurved; disk with prominent, Discussion.—This species is similar to Gib- sulcate median ridge, subbasal gibbosities bobruchus divaricatae in size and form but prominent; gibbosities densely punctate- differs in density of the male pygidial pu- imbricate, remainder of discal sculpture bescence, and extent and placement of the with umbilicate foveolae; lateral carina glabrous female pygidial spot. Male genita- obsolete; cervical sulcus distinct. Scutel- lia offer little assistance in distinguishing lum transverse, emarginate, sparsely pu- the two species. From the much smaller bescent. Elytral length and width subequal G. mimus, the pygidial characters offer the (figure 696); disk depressed between 6th easiest differentiating characters (compare interstices; lateral margin impressed be- figures 690, 698, and 706 for males; fig- hind humerus; striae sinuate, 3rd and 4th ures 691, 699, and 707 for females). ending at base in prominent, denticulate gibbosity; interstices variable in width, 3rd, Gibbobruchus divaricatae Whitehead 5th, and 7th with setose protuberances, and Kingsolver surface finely imbricate-punctate; meta- (Figures 696–703) coxal face densely micro-foveolate, setae directed laterad, fossula glabrous; meta- Gibbobruchus divaricatae Whitehead and femur with ventrolateral carina denticu- Kingsolver 1975a:183; Johnson and late (figure 701), ventromesal carina finely Kingsolver 1982:413; Udayagiri and denticulate, pecten consisting of one large, Wadhi 1989:81. slender denticle separated from smaller Color.—Integument mostly piceous with distal, serrate denticles by gap; metatibia some reddish variegation; antenna pale arcuate (figure 701), mucro nearly one-half yellow with 7th to 10th segments usually as long as basitarsus; 1st abdominal ster- fuscous; legs dark red, tarsi yellow, profe- num with patch of fine setae in male only, mur, mesofemur and tibia, and metatibia 5th sternum nearly divided by broad emar- banded with white. Vestiture variegated gination; male pygidium strongly reflexed black, yellowish brown, and white; head apically (figure 698), disk feebly channeled yellowish brown in broad band across one-third from base, apical one-half with transverse sulcus and median spot on ver- sulcate swelling limited laterally by rugose tex, frons yellowish brown with scattered white setae; pronotum yellowish brown

168 vertical channels, punctation of remain- Kingsolver 1975a:203; Johnson der of disk dense, variable in size; female 1977d:304; Johnson and Kingsolv- pygidium with prominent, glabrous, oval or er 1982:413; Udayagiri and Wadhi cordate swelling (figure 699), basal one- 1989:81. half of disk finely punctate-granulose, lat- Bruchus crataegi Fahraeus 1839:119; Pic eral margins of swelling rugosely punctate. 1913a:35. Male genitalia.—As in figures 702 and 703; Bruchus murinus Schoenherr 1839:132 median lobe slender (figure 702); ventral (misspelling of mimus, not murinus Bo- valve triangular; internal sac lined with heman 1829). masses of fine spicules and denticles of Bruchus borealis Schoenherr 1839:132 (as various sizes; lateral lobes flat, spatulate, new name for murinus Schoenherr). bowed (figure 703). Pseudopachymerus crataegi: Pic 1913a:10. Size.—Body length 2.6–4.1 mm; width Color.—Integument mostly black, dark red 1.8–2.6 mm. integumental patches on elytra, pronotum, Type depository.—USNM holotype No. and margins of abdomen, elytra sometimes 72803. all red; legs red except ventral margin of Type locality.—Mexico, Veracruz, 10 mi metafemur dark red to black, tarsi reddish east of Acayucan. yellow; antenna reddish yellow usually with some club segments dusky. Vestiture Distribution.—TX; Mexico to Honduras. yellow, white, orange, dark brown and Host plants.—Bauhinia divaricata. Three black; head yellow with condensed patch other species, Bauhina pauletia, B. pes- on vertex; pronotum mostly orange with caprae, and B. ungulata, are host to this black on gibbosities, white patches in mid- species in Mexico. dle of disk; scutellum orange; elytra with mottled pattern of white, orange, yellow, Natural enemies.—None recorded. and black, middle of disk with V-shaped Immatures.—Not described. velvety black spot (figure 704); male pygid- Discussion.—This species is known from ium yellow with pale basal triangle (figure the United States only at Brownsville, TX. 706), apex with two or three bare patches; Its distribution appears to be mainly in female pygidium with large, yellow basal Mexico. triangle (figure 707), white setae surrounding polished, median, cordate swelling; Infested seeds are almost entirely hollowed venter of body mostly white, mesopleura out by a single larva. The adult emerges and metapleura orange and yellow. through the side of the seed. Structure.—Vertex finely punctate-imbri- The key characters will separate it from the cate, frons and clypeus longitudinally other two U.S. species. imbricate, frontal carina prominent; ocular index 2.6:1; ocular sinus five-eighths Gibbobruchus mimus (Say) length of eye; antenna (figure 708) moder- (Figures 53, 704–711) ately eccentric from 5th segment, reaching middle of metepisternum. Pronotum Bruchus mimus Say 1831:2 (LeConte edi- subtriangular (figure 704), lateral margins tion 1883:260); Fall 1910:162; Cush- incurved, disk with median, sulcate ridge, man 1911:493; Pic 1913a:35. subbasal gibbosities obtuse, disk densely Mylabris mimus: Leng 1920:305; Knull punctulate on raised portions, foveolate 1934:211. on depressed portions; lateral carina ob- Gibbobruchus mimus: Bridwell 1938a:74; solete; cervical sulcus hidden in vestiture. Craighead 1950:279; Bottimer Scutellum transverse, emarginate. Elytra 1968c:1022,1039; Whitehead and

169 (figure 704) slightly depressed on 2nd and pubescent, female pygidium with cordate, 4th interstices, sutural interstice elevated; glabrous swelling; ventrolateral carina of striae sinuate, nearly parallel, 3rd and 4th metafemur edentate, pecten with one long ending basally in prominent, dentate gib- and three short denticles; median ridge of bosity; interstices flat, punctate-imbricate, pronotum not as high as in Gibbobruchus 2nd, 4th, 6th, and 8th with tufts of setae divaricatae and G. cristicollis. on slight elevations; metacoxal face densely Both Cercis canadensis and C. occidentalis punctulate except fossula glabrous, setae are confirmed larval hosts. Four specimens directed laterad; metafemur with ven- were reared from Bauhinia lunarioides tromesal carina finely dentate (figure 709), (=congesta) along with several hundred pecten of one long and three short den- specimens of Gibbobruchus cristicollis. ticles not separated by gap, ventrolateral carina feebly denticulate; metatibia arcu- Genus Lithraeus Bridwell ate, mucro acute, curved, one-fifth as long as basitarsus; male 1st sternum with ovate Lithraeus Bridwell 1952:125; Bottimer patch of fine, white setae, 5th sternum 1968c:1022; Borowiec 1987:116; Uday- emarginate one-half its length; male py- agiri and Wadhi 1989:83; Kingsolver gidium densely pubescent (figure 706); fe- 1990c:49. Type species: Bruchus electus male pygidium (figure 707) extremely finely Bridwell 1952, by monotypy, unneces- punctulate on swelling, densely punctate sary new name for Bruchus elegans in surrounding areas. Blanchard 1851. Male genitalia.—As in figures 710 and 711; Small size, Acanthoscelides-like, 1.4–3.8 similar to those of Gibbobruchus divari- mm. catae; lateral lobes strongly bowed (figure Color.—Vestiture in some species con- 711), cleft one-half their length. densed into lateral stripe or spot on met- Size.—Body length 2.3–3.5 mm; width episternum, stripe sometimes extended to 1.4–2.1 mm. abdomen. Type depository.—Type lost (mimus); NHRS Structure.—Vertex and frons convex, some (crataegi). species with frontal carina; eyes prominent; ocular sinus about one-half length Type locality.—”Indiana.” (mimus); “Caro- of eye; antenna (figure 713) not sexu- lina America borealis” (crataegi). ally dimorphic. Pronotum semicircular to Distribution.—AL, AR, AZ, DC, FL, GA, IA, subconical, disk convex. Elytra with striae IL, IN, KS, KY, LA, MD, MI, MO, MS, NC, regular, not distorted. Pygidium convex, NV, OH, OK, ON, PA, SC, TN, TX, VA, WI, not modified, metafemoral denticle min- WV; Mexico. ute to one-half width of metatibia; metibia Host plants.—Cercis canadensis, C. oc- lacking longitudinal carinae; mucro short, cidentalis; Bauhinia lunarioides. Adults acute, length less than apical width of have been collected on flowers of numer- tibia. ous other species of plants not the larval Discussion.—Bridwell thought that el- hosts—Fraxinus, Magnolia sp., and Magno- egans was a homonym of Sturm 1843, but lia grandiflora, for example. Sturm’s name is a nomen nudum listed in Natural enemies.—Eupelmus cyaniceps; a catalog with no description and was later Heterospilus bruchi, H. prosopidis; Horis- cited by Gemminger and Harold (1873) as menus sp.; Stenocorse bruchivora. a synonym of Bruchus chinensis Linnaeus. See Bottimer (1968c:1022) and Kingsolver Immatures.—Pfaffenberger 1986 (larva). (1990c:49) for discussion. Discussion.—Salient characters for this Species now assigned to the genus Lith- species are male pygidium almost totally raeus are generally small (1.5–3.5 mm),

170 and the metatibia lack carinae. Nine striae deep, narrow, punctures distinct, species are found in Chile, but several 3rd, 4th, 5th, and sometimes 6th each (undescribed) are known from Brazil and ending basally in pit twice as large as a Argentina, including L. atronotatus, which strial puncture; interstices flat, minutely has been recently introduced into Hawaii. imbricate; meta-coxal face densely punctulate, fossula glabrous; metafemur (figure Lithraeus atronotatus (Pic) 714) moderately slender, lacking denticles; metatibia slender, lacking carinae, mucro (Figures 712–716) minute, lateral and coronal denticles lack- Bruchus atronotatus Pic 1929:35; Bondar ing; abdomen not modified; male pygidium 1936:43; Krauss 1962:132, 1963:282; convex, apex only slightly reflexed, female Davis and Krauss 1962:245; Davis pygidium medially convex, with shallow 1967:344. sublateral sulci. Acanthoscelides atronotatus: Blackweld- Male genitalia.—As in figures 715 and er 1946:758; Udayagiri and Wadhi 716; median lobe moderately broad (figure 1989:37. 715); ventral valve broadly triangular, apex Lithraeus atronotatus: De Luca 1967a:20; slightly produced; internal sac lined for its Kingsolver 1968:322. full length with mixed flat, scalelike denti- Color.—Pronotal integument entirely dark cles and acute, thornlike denticles; closure red or orange with dark red median band; valve circular; lateral lobes fused, strap- elytra dark red with black maculae; py- like (figure 716), slightly expanded toward gidium dark red to orange-red with dark apex, set with stout apical setae. stripes; venter of body dark red, antenna Size.—Body length 2.2 mm; width 1.2 mm. mostly black; fore legs and mid legs reddish yellow with some duskiness on femo- Type depository.—MNHP. ra, metafemur mostly black, metatibia red- Type locality.—Brazil. dish yellow. Vestiture variable from white Distribution.—Brazil, Argentina, Paraguay; to grayish yellow, elytral maculae dark introduced into Hawaii. brown; head white; pronotum white to grayish yellow, often condensed laterally, Host plants.—Schinus terebinthifolius pronotum white or grayish yellow, macu- (aroeira, Brazil); Lithraea brasiliensis (Bra- lae usually black, sometimes dark brown; zil) (De Luca 1967a). pygidium grayish yellow, basal triangle Natural enemies.—Eurytoma sp., Horis- prominent; venter of body gray, usually menus sp., Zatropis sp. condensed into setal spots on metepister- Immatures.—Not described. num and dorsal margin of abdomen. Discussion.—This species was released Structure.—Vertex finely punctate, some- in Hawaii in 1954 and 1960 by Krauss times granulose, frons sometimes with (1963) for control of Schinus terebinthifolius median impunctate line; ocular index 3:1; (Christmas berry, a serious weedy range- ocular sinus three-fifths length of eye; land pest), but it did not increase to suffi- antennal segments (figure 713) slightly ec- cient numbers to effect control. It is occa- centric, extending to humerus. Prono-tum sionally recovered in collections. bell-shaped (figure 712), lateral margins gently arcuate, evenly convex with shallow Lithraeus atronotatus is easily separated basal sulcus; disk densely microfoveolate; from other species in Hawaii by the lack lateral carina absent; cervical sulcus short, of metatibial carinae and mucro, the lack deep. Scutellum quadrate, emarginate. of metafemoral denticles, and the shallow, Elytra slightly longer than wide (figure undifferentiated female pygidial sulci. The 712), convex; pygidial sulci of female Algarobius bottimeri are deep and distinctly marked and of a

171 different texture than that of the surrounding surface. Key to Species of Meibomeus Genus Meibomeus Bridwell 1 Body and appendages black except tarsal pads yellow; vestiture evenly distributed over bodydesmoportheus. Meibomeus Bridwell 1946:54; Bottimer Kingsolver 1968b:288, 1968c:1023; Johnson and Kingsolver 1982:413; Borowiec and Whitehead 1987:81. Type species: Bruchus muscu- Body black; part or all of antennae, fore legs, mid legs, lus (Say) 1831, by original designation. and metatarsi yellow...... 2 Meibomerus: Arnett 1962:957 (error). 2(1) Vestiture yellowish gray; sides of pronotum more densely Small beetles (1.1–2.7 mm). Three species pubescent than disk (figure 725); elytra with median are treated herein. spot of dense pubescence on 3rd and on 7th interstices; pygidium with basal triangle and median line (figure 728) Elytra and pronotum convex; head with musculus (Say) prominent frontal carina; eyes prominent, ocular sinus deep, nearly dividing eye; Vestiture white, thinly distributed, not condensed on antenna elongate, serrate; pronotum bell- pronotum (figure 732) (except for spot on basal lobe), shaped, dorsally convex; 4th elytral stria elytra, or pygidium; Florida only...... surrubresus (Pic) basally abbreviated, limited by fine denticle; fore legs and mid legs normally developed, procoxae contiguous; metafemur Meibomeus desmoportheus Kingsolver and moderately swollen, lacking external ven- Whitehead tral carina or denticles, inner margin with (Figures 717–724) one to three fine denticles before pecten, pecten with one large denticle followed by Meibomeus desmoportheus Kingsolver and three to five obliquely inserted, fine den- Whitehead 1976:12; Johnson and King- ticles, or with subequal fine denticles only; solver 1982:413; Udayagiri and Wadhi metatibia carinate, arcuate, mucro short 1989:85. or absent; abdominal sterna slightly tele- Color.—Body and appendages black except scoped in male, perceptibly emarginate to tarsal pads yellow. Vestiture white, evenly receive apex of pygidium; pygidium flat, or distributed over body; scutellum intensely slightly convex, pattern not sexually dimor- white. phic. Structure.—Vertex and frons microfo-ve- Male genitalia with median lobe elongated, olate; frontal carina prominent; eyes di- slender, sometimes partly fractured near morphic: ocular index 10:1 in male (figure apex; lateral lobes fused basally into flat, 718), 5:1 in female (figure 719); ocular straplike structure, apex divided into two sinus two-thirds length of eye; antenna lobes. dimorphic (figure 721), reaching nearly to This genus is distributed from Canada to apex of elytra in male, to middle of elytra Argentina. in female; penultimate segment 1.25 times as long as wide in male, as long as wide The tentative record of Meibomeus apicicor- in female. Pronotum bell-shaped (figure nis (Pic) reported by Kingsolver (1995a:170) 717), lateral margins slightly sinuate; disk was based on an aberrant specimen of evenly convex except for slight subbasal Meibomeus musculus (Say). depressions and basal median sulcus, sur- Taxonomy: Kingsolver and Whitehead face densely, finely punctate to imbricate; 1976. lateral carina effaced; cervical sulcus deep. Scutellum small, quadrate. Keys to species: Kingsolver and Whitehead 1976 (North American continent).

172 Elytra 1.5 times as long as wide (figure Meibomeus musculus: Bridwell 1946:54; 717), convex; striae narrow, slightly im- Teran 1967:313; Bottimer 1968b:288, pressed, 4th stria abbreviated at base; 1968c:1022; Kingsolver and White- interstices flat, finely imbricate. Metacoxal head 1976:16; Johnson and King- face finely punctate except fossula pol- solver 1982:413; Udayagiri and Wadhi ished; hind leg as in figure 722; metafemur 1989:85. with three to five prepectenal denticles, Bruchus erythrocerus Riley 1871:55; Bot- pecten with one long and four to six min- timer 1968b:288. ute denticles; metatibia basally arcuate, Bruchus alboguttis Motschulsky 1874:215; apically straight, lateral, lateroventral, Kingsolver 1979a:342. ventral, and dorsomesal carinae complete to apex; mucro minute, as long as lateral Color.—Body integument black; male an- denticle; abdomen not modified, sexes tenna entirely pale yellow, female partly similar, male only slightly emarginate to re- infuscated; male mesofemur yellow, female ceive pygidial apex; pygidium nearly flat in mesofemur partly infuscated; tarsi, fore female, slightly convex in male (figure 720). leg, and mesotibia yellow. Vestiture grayish white, variegated, vaguely condensed on Male genitalia.—As in figures 723 and 724; flanks of pronotum, in an irregular band median lobe fractured near apex (figure across middle of elytra, sometimes with 723); ventral valve semicircular, nearly flat condensed spots on pronotum and on 3rd in lateral aspect, apex truncate, margin and 7th interstices of elytra; and in three setose; internal sac with paired, thread- basal spots on pygidium with setae diago- like basal tendons, middle of sac lined nally oriented toward midline (figure 728). with blunt denticles, apex with fine, acute denticles; lateral lobes divided (figure 724), Structure.—Vertex finely foveolate, fron- rounded, and slightly bowed at apices. tal carina fine, short, nearly concealed by setae (figure 727); eyes not sexually di- Size.—Body length 1.9–2.7 mm; width morphic; ocular index 4.75:1; ocular si- 1.0–1.5 mm. nus seven-eighths length of eye; antenna Type depository.—USNM, holotype No. (figure 726) not dimorphic, reaching pos- 72811. terior margin of metasternum. Pronotum bell-shaped (figure 725), lateral margins Type locality.—Arizona, Cochise Co., Chir- incurved or slightly sinuate; disk strongly icahua Mountains. convex, lacking asperities; surface dense- Distribution.—AZ; Mexico to Costa Rica. ly, finely foveolate; lateral carina obtuse Host plants.—Desmodium grahamii. or absent. Scutellum small, prominent, densely pubescent. Elytra together slightly Natural enemies.—None recorded. longer than wide (figure 725); striae normal Immatures.—Not described. except 4th abbreviated and denticulate at Discussion.—This is the only species of one-sixth from base; interstices flat, finely Meibomeus in the United States with the imbricate. Metacoxal face finely, densely body and appendages entirely black. punctate except fossula polished; hind leg as in figure 729, pecten with one long den- Meibomeus musculus (Say) ticle separated by a gap from three short, slanted denticles; metatibia arcuate at (Figures 59, 725–731) base, apical three-fourths straight; lateral, Bruchus musculus Say 1831:3; Horn ventrolateral, ventral, and dorso-mesal 1873:340; Schwarz 1878:457; carinae complete; mucro short, slender, Hubbard and Schwarz 1878:660; as long as lateral denticle. Abdomen tele- Blatchley 1910:1241; Fall 1910:185; scoped, 5th sternum deeply emarginate for Pic 1913a:36. pygidial apex; male pygidium more deeply reflexed at apex (figure 728).

173 Male genitalia.—As in figures 730 and 731; Acanthoscelides surrubresus: Schoonhoven median lobe slender (figure 730), not frac- 1977:692. tured near apex; ventral valve triangular, Meibomeus surrubresus: Kingsolver and acute; internal sac with prominent, curved, Whitehead 1976:20; Janzen 1980:948; tuberculate structures on either side of Mead 1989:2; Udayagiri and Wadhi apical orifice, sac lined with fine denticles 1989:86. in basal three-fourths, apex lined with fine, Color.—Body integument and hind legs, ex- beadlike denticles; lateral lobes with broad cept tarsi, black; antennae usually bright apical emargination between rounded lobes yellow, sometimes with terminal five or six (figure 731). segments dusky, fore legs, mid legs and Size.—Body length 1.6–2.7 mm; width metatarsi yellow. Vestiture white, incon- 1.0–1.6 mm. spicuous, evenly distributed except white Type depository.—Types destroyed (muscu- tuft on basal lobe of pronotum and on lus, erythrocerus); ZMUM (alboguttis). scutellum. Type locality.—Indiana (musculus); “Amer. Structure.—Vertex and frons umbilicate- Bor.” (erythrocerus); Alabama, Mobile (al- foveolate; frontal carina prominent (figure boguttis). 733); eyes dimorphic: ocular index 10:1 in males (figure 733), 6:1 in females (fig- Distribution.—AL, AR, CT, DC, DE, FL, GA, ure 734); ocular sinus dimorphic: one-half IL, IN, IA, KS, KY, LA, MA, MD, ME, MI, length of eye in males, two-thirds length MN, MO, MS, NC, NH, NJ, NY, OH, OK, of eye in females; antenna eccentric from ON, PA, PQ, SC, TN, TX, WI, WV, VA. 5th segment (figure 735), reaching 1st Host plants.—Desmodium canescens, D. abdominal segment in males, reaching tenuifolium, D. tortuosum, D. triflorum; Les- metacoxal margin in females. Pronotum pedeza hirta. Most specimens are collected bell-shaped (figure 732), strongly convex, by general sweeping of vegetation. without asperities; lateral margins slightly sinuate, surface punctate-imbricate, basal Natural enemies.—None recorded. lobe sulcate; lateral carina lacking; cervical Immatures.— Not described. sulcus distinct. Scutellum small, quadrate. Discussion.—The two species of Meibomeus Elytra together 1.2 times as long as wide known from the eastern part of the country (figure 732), convex, slightly depressed are easily separated by the characters in around scutellum; striae normal, 3rd, the key. The vestiture of Meibomeus muscu- 4th, and 5th not reaching basal margin, lus is dense, concealing most of the surface 4th ending in denticle; interstices finely of the elytra and pygidium, whereas that punctate-imbricate. Metacoxal face finely of M. surrubresus is sparse and short, not punctate except fossula polished; hind leg concealing the surface. The geographic as in figure 736, pecten with six to seven distribution of M. desmoportheus does not oblique, acute denticles; metatibia arcu- overlap either of these two species in the ate in basal one-half; lateral, ventrolateral, United States. ventral, and dorsomesal carinae complete to apex; mucro short, acute, about as long Teran (1967) first illustrated the male geni- as lateral denticle. Abdomen normal except talia of this species. 5th sternum of male slightly emarginate for apex of pygidium; pygidium with extremely Meibomeus surrubresus (Pic) fine, lunulate punctation. (Figures 732–738) Male genitalia.—As in figures 737 and 738; Bruchus surrubresus Pic 1933:18. median lobe not fractured (figure 737); Acanthoscelides subrubrosus: Blackwelder ventral valve ogival, acute; internal sac 1946:761 (misspelling). short, with ovate bundle of spicules at extreme base, three to four prominent,

174 dark sclerites at middle, apex lined with aspect; cervical sulcus distinct, usually fine spicules; lateral lobes rounded at narrow; cervical boss prominent, bisetose; apices (figure 738), with U-shaped median prosternum Y-shaped, acutely triangu- cleft. lar between coxae, sometimes separating Size.—Body length 1.3–2.1 mm; width coxal apices. Elytron with 3rd and 4th 0.7–1.2 mm. striae usually arising from denticles on summit of basal gibbosity (except major). Type depository.—MNHP. Pygidium of male usually strongly reflexed Type locality.—Costa Rica, Puntarenas apically to fit sternal emargination. Fore Prov., Surubres. leg and mid leg not especially modified; metafemur strongly dorsoventrally swollen; Distribution.—FL; Mexico to Panama. ventral margin basad of pecten carinate, Host plants.—Aeschynomene americana. lacking fine denticles; pecten with three to Natural enemies.—None recorded. five denticles, basal denticle usually larger than others and separated from them by Immatures.—Not described. gap of varying width; metatibia usually Discussion.—Specimens of this species straight in apical three-fourths but bent were collected in the field near Ft. Pierce, sharply at base, in some species slightly Florida. Whether it constitutes a breeding arcuate; tibial carinae usually complete to population remains to be seen. Characters apex; mucro of varying length. Fifth ab- given in the key will easily separate M. sur- dominal sternum usually carinate either rubresus from M. musculus. The distribu- side of caudal emargination, often more tion of M. surrubresus and M. desmo-porth- prominent in females. eus is sympatric in Central America but Male genitalia with ventral valve nearly not in the United States. as broad as apical width of median lobe; internal sac usually with wishbone-shaped Genus Merobruchus Bridwell median sclerite or modification thereof, Merobruchus Bridwell 1946:54; Brad- sclerites characteristic for each species. ley 1947:41; Arnett 1962:955; John- Lateral lobes spatulate, arms usually son 1967:264, 1968:1269; Bottimer bowed; ventral strut flat, not carinate. 1968c:1023; Johnson 1969c:55, Biology: Center and Johnson 1974; John- 1969d:676; Kingsolver 1970a:374; son and Siemens 1997b. Janzen 1975:180, 1980:948; Johnson Host associations: Johnson and Siemens 1981e:999; Janzen 1982:1274; John- 1997b. son and Kingsolver 1982:418; Borowiec 1987:86; Kingsolver 1988:4; Udaya- Revision: Kingsolver 1988. giri and Wadhi 1989:87; Johnson and Keys to North American species: Kingsolver Siemens 1997b. Type species: Bruchus 1988. julianus Horn 1894:410, monotypic. Small to medium-sized beetles (1.8-8.8 mm). Body dorsally subdepressed or with elytra slightly concave. Head with frontal carina usually prominent (figure 747); ocular sinus about one-half length of eye; antenna gradually clavate from segment four, individual segments ovate or elliptical (figure 739). Lateral carina of pronotum with distinct sinuation at middle in lateral

175 Key to Species of Merobruchus 5(4) Pygidium uniformly pubescent with contrasting basal triangle...... 6 1 Elytra with 3rd, 4th, 5th, and 6th striae each with a flat- Pygidium irregularly pubescent...... 7 tened, scalelike, triangular basal denticle (figure 763); subapical metafemoral denticle large, triangular, with 6(5) Ventral valve of median lobe truncate or broadly emargin- five to six serrations on posterior margin, large denticle ate (figure 761); Florida and West Indies...... lysilomae longer than metatibial width at their intersection (figure Kingsolver 767); body length 5.2-8.8 mm Ventral valve of median lobe rounded apically (figure ...... major (Fall) 774); Kansas, Oklahoma, Texas, Arizona through Central Third and 4th striae each with subbasal denticle usually America set on gibbosity, sometimes with denticles fused into ...... placidus (Horn) transverse carina; subapical metafemoral denticle as in 7(5) Third elytral interstice with elongate, median golden figure 743 or 755; body length 1.8-5.5 mm...... 2 stripe, and short, subapical spot distinct from predomi- 2(1) Pygidium dark brown with uninterrupted, attenuate, nantly gray discal vestiture; basal margin of elytra with median white line of setae (figures 741 and 742); lateral patches of golden setae; pronotal vestiture predominantly margins of elytra irregularly darker than middle (figure gray with golden median stripe; male genitalia as in 740); body length 2.9-3.3 mm...... insolitus (Sharp) figures 750 and 751 Pygidium with short basal triangle or short line of setae, ...... julianus (Horn) or with median line not strongly differentiated ...... 3 Third interstice with median stripe distinct, evanescent, or absent; if present, is part of irregular, transverse golden 3(2) Pygidium nearly uniformly pale yellow with small, irregu- band of short stripes or spots on lateral interstices; larly rounded, median spot divided by narrow median line pronotal vestiture golden yellow or grayish yellow...... 8 of setae (figure 753); apical two-fifths of elytra usually clouded with dark brown (figure 752); body length 3.2- 8(7) Pronotal vestiture dense and conspicuous, especially on 4.1 mm...... male knulli (White) lateral portions (figure 776); mesepimeron, metepister- Pygidial and elytral patterns otherwise...... 4 num, metacoxal face, and 5th sternum with dense patches of golden setae; male genitalia as in figures 779 and 780. 4(3) Pygidium with heart-shaped, median, dark brown, bare ...... terani Kingsolver area with transverse basal band and short median dart Pronotal vestiture sparse and inconspicuous, more or pale yellow (figure 754); body length 3.2-4.1 mm less evenly distributed over disk (figure 781); mese- ...... female knulli (White) pimeron and posterior angle of metepisternum with Pygidium uniformly pubescent with contrasting basal inconspicuous patch of yellow setae; male genitalia as in triangle (figure 759) or irregularly pubescent with vaguely figures 784 and 785...... vacillator (Sharp) defined basal triangle (figure 771)...... 5

Merobruchus insolitus (Sharp) Kingsolver 1988:21; Udayagiri and Wadhi 1989:88. (Figures 740–745) Merobruchus sp.: Gibson 1972:148. Bruchus insolitus Sharp 1885:476; Pic 1913a:29. Color.—Integument dark red to black, head dark red; pronotum dark red tending to- Acanthoscelides insolitus: Blackwelder ward black; elytra red except humeri and 1946:759. lateral margins clouded with black, usu- Merobruchus insolitus: Johnson 1979a:122; ally with dark spots on 3rd interstice and Janzen 1980:948; Johnson 1981a:251; apex, lateral clouding sometimes extended Johnson and Kingsolver 1982:418; mesad as dark maculation; pygidium

176 piceous with dark red median streak; Type depository.—BMNH. antenna reddish yellow, usually with 8th, Type locality.—Guatemala, near Guatemala 9th, and 10th segments dusky; fore legs City. and mid legs reddish yellow, hind legs dark red. Vestiture yellowish white, white, Distribution.—AZ, TX; Mexico to Costa and brown; dorsal clothing mostly yellow- Rica. ish white to white, dark spots on dorsum Host plants.—Pithecellobium pallens; Lysi- brown; venter of body white, pygidium loma microphylla thornberi. Fourteen ad- intermixed brown and white patches with ditional hosts from Mexico southward are median, attenuate white dart reaching listed by Kingsolver (1988:35). See also nearly to apex (figures 741 and 742), some- Johnson and Siemens (1997b). times “pinched” in middle but not inter- Natural enemies.—Chryseida spinola ben- rupted. netti; Horismenus missouriensis; Urosigal- Structure.—Vertex above transverse sulcus phus bruchi, U. neobruchi. densely punctate-foveolate, frontal punc- Discussion.—The salient characters that tures longitudinal, frontal carina distinct; mark this species are the dark red body, ocular index 4:1; ocular sinus one-half elytral margins clouded with black, white length of eye; antenna clavate, extending basal pygidial triangle elongated into even- to humerus. Pronotum subconical, lateral ly tapered dart, and, in the male genitalia, margins nearly straight (figure 740); disk the apical nipple on the ventral valve, three evenly convex, surface densely umbilicate- subequal median sclerites, and the basal foveolate, foveolae sometimes coalescing; pair of small, thornlike sclerites. lateral carina ridgelike, sinuate; cervical sulcus deep. Scutellum transverse, apically Merobruchus julianus (Horn) emarginate; postmesal sulcus angulate. (Figures 49, 64, 74, 746–751) Elytral disk depressed (figure 740); striae narrow, shallow, parallel except 2nd, 3rd, Bruchus julianus Horn 1894:410; Fall and 4th slightly deflected laterad at base 1910:186, 1912:320; Wenzel 1912:140. with 3rd and 4th ending in small, adjacent Mylabris julianus: Leng 1920:304; Boeving denticles on low gibbosity; strial punc- 1927:141. tures fine, elongate; metacoxal face, except Acanthoscelides julianus: Blackwelder fossula, densely punctate; hind leg as in 1946:759. figure 743; metafemur with one long and Merobruchus julianus: Bridwell 1946:54; three short denticles in pecten; metatibia Blackwelder and Blackwelder with four prominent carina, mucro short, 1948:44; Bottimer 1968c:1024; John- angulate. Pygidium slightly convex, densely son 1967:264, 1968:1269; Bottimer punctulate. 1969b:1187; Johnson 1969c:55, Male genitalia.—As in figures 744 and 745, 1969d:676; Forister and Johnson median lobe broad at apex, ventral valve 1970:84; Kingsolver 1970a:374; John- semicircular with distinct apical nipple (fig- son 1979a:122; Johnson and Kingsolv- ure 744); internal sac with cluster of min- er 1982:418; Borowiec 1987:86; King- ute denticles and two thornlike denticles solver 1988:11; Udayagiri and Wadhi near base, middle of sac with three acute, 1989:88. subequal spines, apex partly lined with Bruchus ochreolineatus Fall 1910:186; minute denticles; closure valve circular; Cushman 1911:492; Johnson lateral lobes flat, cleft one-half their length 1968:1269; Johnson and Kingsolver (figure 745), apices spatulate, mesal faces 1982:418; Kingsolver 1988:11; Udaya- setose. giri and Wadhi 1989:88. Size.—Body length 2.9-3.3 mm; width 1.0- Bruchus ochrolineatus: Pic 1913a:38 (mis- 2.0 mm. spelling).

177 Color.—Integument dark red to black; dis- pygidium densely foveolate, crowded, co- tal antennal segments usually darker than alescent, middle of disk with shallow me- proximal segments. Vestiture gray, cop- dian depression (figure 748). pery, brown, and yellow; head gray, lateral Male genitalia.—As in figures 750 and slopes of pronotum gray with yellow patch- 751, median lobe broad (figure 750); ven- es, disk brown with white patches and tral valve nearly semicircular; internal sac median yellow stripe, elytra with variably lined with minute spicules and denticles, intermixed brown and gray, 3rd interstice middle of sac with one large, broad-based, with elongate stripe and subapical spot yel- ogival sclerite flanked either side by ta- low, evanescent gray or yellow bands two- pered, elongate spine, apex of sac sparsely fifths from base and one-fifth from apex; denticulate; closure valve circular; lateral pygidium yellow with evanescent basal lobes cleft three-fourths their length (figure triangle, disk with variable brown patches; 751), arms moderately broad, apices di- venter of body mostly gray, three yellow agonally truncated, mesal faces setose for patches on metepisternum and metacoxa, entire length. yellow spots sometimes evident on lateral margin of abdomen; legs gray. Size.—Body length 3.2-5.5 mm; width 1.9- 3.1 mm. Structure.—Vertex and frons densely punctulate, transverse sulcus distinct, Type depository.—CASC (julianus, lec- frontal carina prominent (figure 747); totype No. 285 D, 10474, by Johnson ocular index 3:1; ocular sinus one-half as 1969d); MCZC (ochreolineatus, holotype long as eye; antenna eccentric from 5th No. 25057). segment, extending to middle of metepi- Type locality.—Texas, San Ignacio (lecto- sternum; Pronotum subconical (figure type) (julianus); Arizona, Jerome (ochreolin- 746), lateral margins nearly straight; disk eatus). slightly depressed, posterolateral corners Distribution.—TX; Mexico. depressed; sculpture microfoveolate, each foveola setate beneath its anterior rim; lat- Host plants.—Acacia berlandieri, A. greggi, eral carina represented by intermittent row A. rigidula, A. roemeriana, A. wrightii; Des- of denticles hidden in vestiture. Scutellum manthus illinoensis (questionable); Mimosa transverse, apical margin deeply emar- borealis (questionable); Pithecel-lobium eba- ginate. Elytra slightly depressed between no (all in the United States). Acacia coulteri 5th interstices (figure 746); striae parallel and A. juncifolia are listed from Mexico. except 3rd, 4th, and 5th deflected laterad According to Rico-Arce (1991), the name at base, 3rd and 4th each ending in basal “ebano” is a senior synonym of “flexicaulis” denticle on prominent gibbosity, strial (Benth.) Coult. punctures foveolate, encroaching on inter- Natural enemies.—Eupelmus cushmani, E. stitial margins; interstices finely imbricate; cyaniceps; Stenocorse bruchivora; Urosi- metacoxa densely punctulate and setose galphus bruchi, U. bruchivorus, and three except fossula glabrous; hind leg as in unidentified species ofEurytoma. figure 749; ventral face of metafemur flattened, ventromesal margin carinate, pecten Immatures.—Not described. with four to five denticles, the 1st denticle Discussion.—Johnson (1968) designated longer than others; metatibial carinae a lectotype from Texas in the Academy of prominent, complete; mucro one-fourth as Natural Sciences in Philadelphia but later long as basitarsus; lateral denticle one- rescinded that action in favor of selection fourth as long as mucro. Margin of 5th from a series of specimens in CASC from abdominal sternum finely carinate in both which he (1969d) designated a lectotype sexes, not developed into processes; that corresponded to the data given in the original description. Kingsolver (1988)

178 erroneously listed MCZC as type depository spots) yellow; venter of body gray with of M. julianus following Johnson’s 1968 yellow patches on metepisternum; male paper. The bruchid types in the Academy elytra yellow with gray spots, female elytra were later transferred to the Museum of gray, female pygidial patch with fine brown Comparative Zoology at Harvard. setae. Kingsolver (1988:12) included some of Structure.—Vertex and frons densely punct- Bottimer’s field notes in which he indi- ulate with different sizes; frontal carina cated the differences in feeding habits of weak, impunctate with irregular margins; M. julianus in two host seed pods. A larva ocular index 3.5:1; ocular sinus one-half attacking Acacia berlandieri, because of its length of eye; antenna clavate, 5th to 10th size relative to that of the seed, will glue segments eccentric. Pronotum subconi- several seeds to the inside of the pod valve cal (figure 752), lateral margins straight; to provide sufficient seed material to com- disk convex, faintly sulcate on basal lobe, plete its development. After eclosion, the posterolateral corners impressed; surface adult then will cut through the valve to densely, irregularly foveolate-punctate, escape (Johnson 1967). Although seeds of lateral carina arcuate at middle one-third; this host normally fall to the ground when cervical sulcus deep, narrow. Scutellum the pod dehisces, those fastened to the transverse, emarginate. Elytra broadly valve remain attached. In contrast, M. julia- depressed between 5th striae (figure 752); nus, in attacking the larger-seeded Acacia striae shallow, narrow, 3rd and 4th end- greggi, does not glue the seeds to the pod ing at base in triangular denticles on low valve since each larva requires only one gibbosity, 5th and 6th sometimes denticu- seed to complete its development, and the late, interstices flat, densely imbricate; adult will cut through only the seed coat, metacoxal face densely, evenly punctulate not the valve, to escape. See also Forister except fossula glabrous, polished. Hind leg and Johnson (1970:84) for further descrip- as in figure 755; pecten with long denticle tion of the bionomics of M. julianus. separated by small gap from three smaller, distal denticles; metatibia slightly arcu- Merobruchus knulli (White) ate, dilated toward apex; tibial carinae complete to apex; mucro one-sixth to one- (Figures 752–757) eighth as long as basitarsus, coronal den- Bruchus knulli White 1941:189. ticles three. Pygidial disk finely imbricate. Mylabris knulli: Blackwelder and Black- Male genitalia.—As in figures 756 and 757; welder 1948:44. median lobe broad (figure 756), apically Merobruchus knulli: Bottimer 1968c:1024; expanded; ventral valve truncate, internal Johnson 1968:1269, 1979a:123, sac with large, V-shaped, skewed sclerite 1981a:251; Johnson and Kingsolver near base, paired cluster of fine spicules in 1982:418; Kingsolver 1988:18; Udaya- middle of sac; lateral lobes slender (figure giri and Wadhi 1989:88. 757), spatulate, deeply cleft, densely, finely Color.—Integument reddish brown; api- setose on mesal faces. cal one-third of elytra, especially in male, Size.—Body length 3.2-4.1 mm; width 1.9- clouded with dark brown to black (figure 2.6 mm. 752); male pygidium with small, irregularly rounded spot divided by medial line Type depository.—OSUC. of setae (figure 753); female pygidium Type locality.—Arizona, Huachuca Moun- with large, brown, cordate patch (figure tains. 754); basal seven antennal segments yel- Distribution.—AZ; Mexico to Honduras. low, terminal four dark brown. Vestiture gray, yellow, dark brown, and black; head, pronotum, and pygidium (except median

179 Host plants.—Lysiloma microphylla thorn- parallel, slightly deflected laterad at base, beri in the United States; L. watsoni and L. 3rd and 4th minutely denticulate on basal acapulcensis extralimital. margin, strial punctures distinct; inter- Natural enemies.—None recorded. stices flat, imbricate; metacoxal face finely, evenly punctate except fossula glabrous; Immatures.—Not described. hind leg as in figure 760; pecten with Discussion.—Nothing is known of the life three or four denticles gradually decreas- history of this species except the host as- ing in length; metatibia abruptly bent at sociations. It is most similar to Merobru- base, gradually dilated toward apex; cari- chus xanthopygus from southern Mexico nae complete except ventrolateral obsolete and Central America, but it can be dis- apically; mucro short, only slightly longer tinguished from other U.S. species by than lateral denticle, coronal denticles its pygidial pattern and distinctive male three. Pygidial disk finely strigose, strigae genitalia. unevenly distributed; ventral emargination of 5th sternum carinate. Merobruchus lysilomae Kingsolver Male genitalia.—As in figures 761 and (Figures 758–762) 762; median lobe broad (figure 761); ventral valve as broad as median lobe, apical Merobruchus lysilomae Kingsolver 1988:25; margin shallowly emarginate; internal sac Kingsolver 1992b. with folds near base lined with fine spic- Color.—Integument reddish brown to dark ules, middle of sac with inverted Y-shaped brown, elytra mottled with dark brown sclerite (wishbone), and two thornlike spots and evanescent bands (figure 758), sclerites, middle one-third of sac lined with antenna, fore legs and mid legs, metatarsi fine denticles; apex with C-shaped closure yellowish red; pronotum often with broad, valve; lateral lobes as in figure 762, cleft red, median stripe. Vestiture short, yellow- nearly to base. ish gray except dark brown on elytral spots Size.—Body length 2.4-3.1 mm; width 1.4- and occasionally on pygidium, pronotal 1.7 mm. flanks with dense yellow clothing; pygidium with basal triangular pad of setae Type depository.—CNCI. (figure 759). Type locality.—Florida, Royal Palm Park. Structure.—Vertex transversely punctate, Distribution.—FL; West Indies. transverse sulcus feeble, frons densely punc-tate, sometimes with triangular Host plants.—Acacia richei, A. simplex (A. impunctate boss, frontal carina feeble or simplicifolia of authors); Albizia lebbeck, A. absent; ocular index 4:1; ocular sinus one- polyphylla; Lysiloma latisiliqua. In Cuba, half as long as eye; antenna gradually clav- this bruchid is known from Lysiloma sa- ate, segments eccentric from 5th segment, bicu Bentham. extending to humerus. Pronotum sub- Natural enemies.—None recorded. conical (figure 758), lateral margins nearly Immatures.—Not described. straight, apex rounded; disk convex, basal lobe feebly sulcate, surface densely but Discussion.—Salient characters for this discretely microfoveolate, each foveola with species are the sparse body clothing, even- median seta, interspaces rugosely punc- ly convex pronotal disk, minute basal strial tate; lateral carina intermittent, arched denticles, and male genitalia with Y-shaped at middle; cervical sulcus deep, narrow. sclerite and two mesal, thornlike sclerites. Scutellum transverse, broadly emarginate. It is the only species of Mero-bruchus in the Elytra together slightly longer than broad eastern part of the United States. (figure 758); disk depressed, especially around scutellum; striae shallow,

180 Merobruchus major (Fall) lateral margins incurved with slight bulge at middle; disk convex, feebly sulcate but (Figures 72, 763–769) deeper on basal lobe, with subbasal gib- Bruchus major Fall 1912:320; White bosities either side opposite bases of 4th 1941:189. interstices; lateral carina arched at middle. Mylabris major: Leng 1920:304. Scutellum transverse, broadly emarginate. Elytra broadly depressed between 5th Merobruchus major: Bradley 1946:41; striae (figure 763); striae narrow, parallel, Blackwelder and Blackwelder slightly deflected laterad in basal one-half, 1948:44; Johnson 1967:264; Bottimer bases of 4th, 5th, 6th, and sometimes 3rd 1968c:1023; Johnson and Kingsolver armed with large, flat denticles; interstices 1982:418; Kingsolver 1988:7; Udayagiri mostly flat, minutely imbricate; metacoxal and Wadhi 1989:88. face densely punctulate in lateral one-half, Bruchus flexicaulis Schaeffer 1904:229 less dense in proximal one-half, polished, (manuscript name). impunctate either side of fossula; hind leg Acanthoscelides flexicaulis: U.S. De- as in figure 767; pecten with four to five partment of Agriculture 1940:36, denticles gradually decreasing in length; 1942:7,24, 1944:6,24, 1945:7,24; basal one-fourth of metatibia arcuate, Zacher 1952:465. gradually dilated toward apex; lateral, ven- Merobruchus flexicaulis: Wheeler et al. trolateral, ventral, and dorsomesal carinae 1950:23,37. complete to apex; mucro acute, slender, one-fourth length of basitarsus, coronal Acanthoscelides flexicaule: Zacher denticles three. Pygidium (figures 765 and 1952:470 (lapsus). 766) broadly sulcate along midline, middle Bruchus julianus: Schaeffer 1904:229 of disk with inverted V-shaped depression, (misidentification); Cushman 1911:491 punctation concealed by vestiture. (misidentification). Male genitalia.—As in figures 768 and 769; Mylabris julianus: Boeving 1927:133 (mis- median lobe broad (figure 768), ventral identification). valve cordate, apex rounded; internal sac Color.—Integument dark red to black; head densely lined with fine spicules, devoid of dark red, antenna red, usually with 8th to large sclerites; lateral lobes short (figure 10th segments dusky; pronotum, elytra, 769), bowed, flat, cleft two-thirds their abdominal sterna, and hind legs mottled; length, mesal faces densely setose. pygidium mostly uniformly dark red, occasionally with darker median spot. Vesti- Size.—Body length 5.2-8.8 mm; width 2.8- ture white, dark brown, and yellow; ver- 4.7 mm. tex, frons, and median stripe of pronotum Type depository.—MCZC (major, holotype yellow, flanks of pronotum and legs mostly No. 25056); USNM (flexicaulis, lectotype white; pygidium white to yellowish brown No. 42446). with white basal triangle (figures 765 and Type locality.—Texas, Brownsville (major); 766). Texas, Esperanza Ranch (flexicaulis). Structure.—Vertex finely, densely punctu- Distribution.—TX; Mexico. late, hidden by vestiture, frons more coarsely punctate, transverse sulcus and Host plants.—Pithecellobium ebano. Both frontal carina minutely granulose (figure Merobruchus major and Stator beali are 764); ocular index 3.5:1; ocular sinus one- seed predators on this host; however, S. third as long as eye; postocular lobe elon- beali also attacks other species of Pithecel- gated, lunate; antennal segments eccentric lobium whereas M. major is an obligate from 5th segment, reaching elytral humer- predator of P. ebano. According to us. Pronotum bell-shaped (figure 763),

181 Rico-Arce (1991), the name “ebano” pre- Merobruchus placidus: Johnson 1967:264; dates “flexicaulis” and must be used for Bottimer 1968c:1024; Johnson this species. 1968:1269, 1969c:55, 1979a:123, Natural enemies.—None recorded. 1981a:251; Johnson and Kingsolver 1982:418; Kingsolver 1988:27; Udaya- Immatures.—Not described. giri and Wadhi 1989:89. Discussion.—This species is the largest in Bruchus limpidus Sharp 1885:456; Cush- this genus. It is sometimes confused with man 1911:498; Pic 1913a:31. Merobruchus julianus but size overlaps only Acanthoscelides limpidus: Blackwelder slightly, elytral 3rd, 4th, 5th, and 6th strial 1946:760. bases of M. major are set with prominent, Merobruchus limpidus: Johnson and King- flattened denticles, and male genitalia are solver 1977:154. devoid of large sclerites. Color.—Integument dark red to dark brown Schaeffer (1904) discussed, under the with some sternal areas black; antennae, name “julianus,” a “gigantic Bruchus on fore legs, and mid legs yellowish red, hind Acacia flexicaulis, in the large seed pods legs dark red. Vestiture short, grayish yel- of which it undoubtedly breeds.” He dis- low, white, and brown, the latter on elytral tributed specimens under the manuscript maculations; venter of body white; prono- name “flexicaulis” but never formally tal vestiture varying from nearly uniformly described the species. The name was first yellowish gray to a pattern of median and listed as a synonym of Bruchus julianus lateral lines; pattern of elytra varying from and subsequently as an unavailable name. nearly unicolorous to pattern of lateral and According to the 1985 International Code apical dark brown maculae with elongate, of Zoological Nomenclature, a name may yellowish spot on 3rd interstice; pygidium be made available if it was used as the yellowish with variable triangular basal name of a taxon prior to 1961. In 1940, the spot (figures 771 and 772). U.S. Department of Agriculture List of In- tercepted Plant Pests lists Acantho-scelides Structure.—Vertex densely, finely punc- flexicaulis, and subsequent issues used tate, frons more coarsely punctate; trans- this combination (U.S. Department of Ag- verse sulcus feeble; frontal carina vaguely riculture 1940, 1942, 1943, 1944, 1945). delimited; ocular index 3:1; ocular sinus Wheeler et al. (1950), listed Merobruchus five-eighths length of eye; antenna gradu- flexicaulis (Schaeffer), and in 1952 Zacher ally clavate, segments eccentric from 5th used Acanthoscelides flexicaulis. Although segment, extending to humerus. Pronotum flexicaulis predates major, I prefer to use bell-shaped (figure 770), lateral margins major in the interest of stability since it slightly sinuate, apex gently rounded; disk has been used in several taxonomic pub- mostly convex, sulcate on median lobe, lications. Application to the International impressed on posterior corners; surface Congress should be made to suppress the densely but irregularly foveolate, each fo- name “flexicaulis.” veola with median seta; interspaces densely punctate; lateral carina discernible only Merobruchus placidus (Horn) at middle of margin; cervical sulcus short, deep. Scutellum transverse, broadly emar- (Figures 770–775) ginate. Elytra about as long as wide (fig- Bruchus placidus Horn 1873:341, ure 770), convex except depressed around 1894:345; Fall 1910:184; Cushman scutellum; striae parallel, only slightly 1911:498. deflected laterad near base, 3rd and 4th Mylabris placidus: Leng 1920:306. ending in basal denticles on slight protuberance; striae shallow, narrow; Acanthoscelides placidus: Blackwelder 1946:760.

182 interstices flat, imbricate; metacoxal face the genus on the North American conti- discretely punctate in lateral one-half, nent. This variation is reflected in the sev- punctures crowded at mesal one-half, fos- eral variants in form of the male genitalia sula glabrous; hind leg as in figure 773; (see Kingsolver 1988). Additional and sys- pecten with four denticles gradually de- tematic collecting is needed to determine creasing in size; metatibia strongly bent at the true status of the species—whether it base, dilated toward apex, all tibial carinae is a widely varying species or a cluster of complete to apex, or with ventrolateral closely related sibling species. Most of the carina abbreviated at apex; mucro short, host records are of Acacia angustissima angulate, lateral and coronal denticles and its varieties. minute. Fifth sterna not carinate in either The habitus of this bruchid closely sex; pygidium convex; surface minutely approaches that of some species of Acan- imbricate. thoscelides, but the basal strial denticles, Male genitalia.—As in figures 774 and 775; enlarged hind leg, and form of male genita- median lobe broad (figure 774), ventral lia are diagnostic. valve arcuate, variable in outline; armature of internal sac consisting of paired, elon- Merobruchus terani Kingsolver gate sclerites near base, middle with sub- (Figures 776–780) triangular sclerite, and apex with paired thornlike sclerites, sac lined with spicules Merobruchus species 4: Johnson and denticles of various sizes; closure valve 1979a:123. C-shaped; lateral lobes short, broad (figure Merobruchus terani Kingsolver 1980a:256; 775), cleft two-thirds their length. Janzen 1980:936,948; Johnson and Size.—Body length 1.8-2.8 mm; width 1.1- Kingsolver 1982:418; Kingsolver 1.8 mm. 1988:20; Udayagiri and Wadhi 1989:90. Color.—Integument deep red to piceous, Type depository.—MCZC (placidus, lecto- antennae yellowish red with 7th to 10th type No. 8210, by Johnson 1968); BMNH segments dusky; fore tarsi and mid tarsi (limpidus). yellowish red. Vestiture yellowish orange, Type locality.—”Texas” (placidus); Guate- gray, and dark brown; head, pronotum, mala, Capetillo (limpidus). scutellum, and elytra orangish yellow, Distribution.—AZ, OK, KS, TX; Mexico to darker spots in pattern with dark brown Costa Rica. setae; pygidium dense yellow; venter and pleuron of body gray except yellow patches Host plants.—Acacia angustissima an- on mesepimeron, metepisternum, and gus-tissima, Acacia angustissima hirta, A. sides of 5th sternum. angustissima suffrutescens, A. angustissima texensis; Astragalus sp. The variety A. Structure.—Vertex finely, densely punc- angustissima angustissima is found only tate, frons more coarsely punctate; trans- in Mexico and Central America. Kingsolver verse sulcus well-marked; frontal carina (1988) erroneously listed Acacia suffrute- an impunctate median line; ocular index scens as A. suffruticosa. See Kingsolver 3.3:1; ocular sinus six-tenths as long as (1988) for additional host species of Acacia eye; antenna clavate, extending to hu- in Mexico and Central America. merus, 5th to 10th segments eccentric, terminal segment elliptical. Pronotum bell- Natural enemies.—None recorded. shaped (figure 776), lateral margins nearly Immatures.—Not described. straight; disk convex but sulcate on basal Discussion.—Merobruchus placidus is the lobe, slightly impressed near lateral cor- most variable and widespread species in ners; surface with densely crowded umbilicate foveolae, interspaces punctulate; lateral carina medially arcuate but obsolete

183 near forecoxal cavity. Scutellum trans- vacillator are longer than wide and the verse, broadly emarginate; cervical sulcus pronotal margins are straight or slightly short, deep. Elytra as long as wide (figure bowed. Male genitalia are distinctive (com- 776); disk depressed between 5th striae; pare figures 779 and 784). striae parallel, 3rd and 4th each ending at base in flat denticle on low tumescence, Merobruchus vacillator (Sharp) striae shallow but distinct; interstices flat, (Figures 71, 781–785) minutely imbricate; postmesocoxal sulcus strongly angulate; metacoxal face densely Bruchus vacillator Sharp 1885:457; Pic punctate except fossula glabrous; hind leg 1913a:54. as in figure 778; pecten with one long and Acanthoscelides vacillator: Blackwelder three short denticles. 1946:761. Male genitalia.—As in figures 779 and 780; Merobruchus vacillator: Bottimer median lobe (figure 779) suddenly expand- 1968c:1024; Johnson and Kingsolver ed near apex, lateral margins embracing 1982:418; Kingsolver 1988:15; Udaya- base of ventral valve heavily sclerotized; giri and Wadhi 1989:90. ventral valve transverse, lunate; internal Color.—Integument reddish brown to deep sac lined with fine denticles and setae, purplish red with pale reddish variegation; middle of sac with inverted Y-shaped scler- basal seven antennal segments reddish ite; apex of sac denticulate; closure valve yellow, 8th to 11th dark red; fore legs and circular; lateral lobes moderately broad mid legs usually paler red than hind leg. (figure 780), slightly bowed, apices expand- Dorsal vestiture grayish yellow in vague ed mesad, cleft three-fourths their length. pattern (figure 781), pronotum with trans- Size.—Body length 2.9-4.5 mm; width 1.9- verse band and two spots; venter of body 2.5 mm. gray with yellow patches on metepisternum and sides of abdomen; pygidium (fig- Type depository.—USNM, holotype No. ure 782) yellow with median bare spot. 72818. Structure.—Vertex finely punctate, trans- Type locality.—Costa Rica, Guanacaste verse sulcus feeble, frons more coarsely Prov., Santa Rosa N.P. punctate, frontal carina absent or inter- Distribution.—TX; Mexico to Costa Rica. mittent; ocular index 3:1; ocular sinus This species was first reported from South- six-tenths as long as eye; antenna gradu- most in extreme southern Texas. ally clavate, segments eccentric from 5th, Host plants.—No host records are known extending to humerus. Pronotum bell- in the United States, but two of the host shaped (figure 781), lateral margins nearly plants listed by Kingsolver (1988) from straight, apex evenly rounded; disk convex, northern Mexico, Acacia berlandieri and A. basal lobe feebly sulcate, basal margin angustissima, also occur in Texas. The type impressed either side of basal lobe; surface series was reared from A. tenuifolia Willd. densely foveolate, each foveola umbilicate and setose; lateral carina ridgelike, arched. Natural enemies.—None recorded. Scutellum quadrate, emarginate, densely Immatures.—Not described. pubescent. Elytra slightly longer than wide (figure 781); striae parallel, 1st, 2nd, and Discussion.—Merobruchus terani is most 3rd laterally deflected toward base, 3rd similar to M. vacillator among the U.S. spe- and 4th each ending in basal denticle on cies, but the elytra of M. terani are nearly common tumescence, striae narrow, deep; quadrate and the pronotal margins are interstices flat, coarsely imbricate; meta- concave, whereas the elytra of M. coxal face densely, uniformly punctate except fossula glabrous; hind leg as in figure 783; pecten with one long, sharp

184 denticle, separated by gap from two small- Johnson 1978; Johnson 1983b; Borow- er distal denticles; metatibia bent at base, iec 1987:86; Udayagiri and Wadhi slightly dilated toward apex; all carinae ex- 1989:90; Johnson and Siemens 1995b. cept ventrolateral complete to apex; mucro Type species: Bruchus sallaei Sharp short, scarcely longer than lateral denticle. 1885, by original designation (=nubig- Female 5th sternum with carinate projec- ens Motschulsky). tions either side of anal notch, projections Cercidiestes Bridwell 1946:55. Type spe- lacking in male; pygidial disk densely mi- cies: Bruchus ulkei Horn 1873, by origi- crofoveolate, each foveola bearing seta from nal designation. its anterior rim. Small to medium sized beetles (1.6–6.0 Male genitalia.—As in figures 784 and 785; mm). Seven species. median lobe broad (figure 784), expanded Elongate-ovate, subfusiform. Integument at apex; ventral valve transverse, arcuate; dark red to black; vestiture variable ac- armature of internal sac consisting of fine cording to species. Head with frons usually denticles at apical orifice, an inverted V- glabrous, impunctate frontal line expanded shaped sclerite near base of sac, two large, into Y-shaped or diamond-shaped boss thornlike denticles in middle, an amor- above and between eyes (figure 57); ocular phous, sclerotized structure near apex, sinus less than one-half length of eye; pos- and fine denticles lining extreme apex; terior margin of eye protruding from side closure valve circular; lateral lobes slender of head or merging with lateral surface of (figure 785), slightly bowed, cleft nearly to head; antenna moderately eccentric (as in base, densely setose on mesal faces. figure 818), usually extending to humerus, Size.—Body length 2.7-3.3 mm; width 1.6- not sexually dimorphic. Pronotum bell- 1.9 mm. shaped, dorsally convex, sometimes with Type depository.—BMNH. slight impressions along basal margin, lateral carina varying from weakly inter- Type locality.—Mexico, Guanajuato. mittent to strong and denticulate; pro- Distribution.—TX; Mexico. coxae contiguous or narrowly separated by prosternal process; cervical sulcus usually Host plants.—No records for the United distinct, deep. Scutellum small, usually States. Lysiloma divaricatum in Mexico. quadrate. Elytra longer than broad, evenly Natural enemies.—None recorded. convex; 3rd and 4th striae, sometimes Immatures.—Not described. 5th and 6th, with basal strial denticles prominent, recumbent, striae shallow but Discussion.—This species, like M. terani, is distinct. Abdomen unmodified (exceptM. represented by one collection in southern protractus). Metafemur moderately swollen, Texas, the northern extension of a Mexico- channeled on ventral face, male often with Central American distribution. It most setose ventral pocket; pecten composed of closely resembles M. terani, but in M. terani one long denticle followed by one to four the elytra are quadrate, the dorsal vesti- smaller denticles (except M. ulkei with one ture is bright yellow, and the male genitalia spine); metatibial carina prominent, ven- are distinctive (see discussion of M. terani). trolateral abbreviated near apex; mucro as long as or shorter than lateral denticle. Genus Mimosestes Bridwell Male genitalia with ventral valve not articu- Mimosestes Bridwell 1946:54; Bradley lated; membranous dorsal valve hoodlike. 1947:39; Blackwelder and Blackwelder Revisions: Kingsolver and Johnson 1978. 1948:45; Bottimer 1968c:1024; John- son 1968:1269; Kingsolver and Keys: Kingsolver and Johnson 1978.

185 Key to U.S. Species of Mimosestes 4(3) Anterolateral corner of pronotum with denticulate shoulder (figure 808); posterodorsal margin of eye subangulate 1 Elytra with white, attenuated, median stripe extending (figure 807) from base to apex bordered by broad, lateral black stripe ...... nubigens (Motschulsky) (figure 818); pygidium white with variable black spots either side of midline (figure 819); metafemur with single Anterolateral corner of pronotum gently rounded, lack- denticle (figure 820) and with shallow ventral channel; ing denticles; posterolateral margin of eye not angulate lateral carina of pronotum denticulate ...... ulkei (Horn) (figure 802)...... 5 Elytra at most with evanescent maculae or stripes; 5(4) Body length 1.6–2.7 mm; pygidium uniformly yellowish pygidium variously colored but not white; metafemur with brown; southwestern United States and Mexico multiple denticles...... 2 ...... acaciestes Kingsolver and Johnson 2(1) Posterior margin of eye merging with side of head behind Body length 2.3–4.4 mm; pygidium vaguely maculate eye, not protruding (figure 791); color brown to bluish (figure 797); Florida and Hawaii gray, sometimes with reddish suffusion... amicus (Horn) ...... 6 Margin of eye protruding from side of head; color various . 6(5) Eye slightly flattened on dorsal margin (figure 802); ...... 3 pygidium with median stripe and two lateral white spots; one record from southern Florida (also West Indies, 3(2) Pygidium with median, hourglass-shaped, dark mark Mexico, Central America)...... mimosae (Fabricius) (figure 813); male metafemur with shallow ventral sulcus; 2nd to 5th ventral sternites of abdomen with broad con- Eye evenly rounded dorsally (figure 796); pygidium with cavity (figure 814)...... protractus (Horn) basal triangle of white setae and median, dark brown macula; Hawaii (also Colombia and West Indies) Pygidium with vague median stripe and lateral spots or ...... insularis Kingsolver and Johnson without pattern; male metafemur with deep, setose ventral sulcus; abdomen lacking ventral concavity...... 4

Mimosestes acaciestes Kingsolver Structure.—Vertex and frons finely punc- and Johnson tate; frontal boss broadly Y-shaped; ocular index 3:1; ocular sinus three-fifths length (Figures 786–789) of eye; antenna extending to humerus; pro- Mimosestes new species: Center and John- notal disk microfoveolate; lateral carina as son 1976:196. denticulate ridge extending to coxal cavity. Mimosestes acaciestes Kingsolver and Scutellum transverse, narrowly emargin- Johnson 1978:12; Johnson and King- ate. Basal denticles on 3rd, 4th, and 5th solver 1982:417; Johnson 1987:348; elytral striae (figure 786); striae nearly hid- Udayagiri and Wadhi 1989:90. den in vestiture; metacoxal face with dense setal patch in lateral one-third; metafemo- Color.—Integument mostly black with fol- ral pocket deep, pecten as in figure 787; lowing areas red: labrum; four basal an- mucro not as long as lateral denticle; apex tennal segments, but in some specimens of male 5th sternum slightly emarginate. all segments red; broad stripe in center of each elytron; pygidium; legs. Vestiture Male genitalia.—As in figures 788 and 789; uniformly yellowish brown in some speci- ventral valve arcuate (figure 788), evenly mens, usually gray and brown in indistinct rounded; dorsal valve semicircular; inter- pattern on elytra and pygidium, pronotum and pygidium usually with two white spots; scutellum white.

186 nal sac trilobed, densely lined with fine Center and Johnson 1976:196; Pfaffen- denticles; closure valve C-shaped; lateral berger and Johnson 1976:34; Mitchell lobes cleft nearly to base (figure 789), arms 1977:644; Johnson and Kingsolver slender, bowed, apical margin diagonal, 1982:417; Kistler 1982; Pfaffenberger mesal faces densely setose. 1985a:4; Johnson 1987:346; Udayagiri Size.—Body length 1.6–2.7 mm; width and Wadhi 1989:90; Kingsolver 1995a. 0.9–1.7 mm. Color.—Integument reddish orange to black; basal four antennal segments red, Type depository.—USNM, holotype No. apical segments usually black, sometimes 72778. reddish; pronotum usually black with red- Type locality.—Arizona, Yavapai Co., 7 1/2 dish basal suffusion; elytra usually black mi southeast of Camp Verde. with reddish-orange stripe, but may be all Distribution.—AZ, NM, TX; Mexico. black or all reddish orange; legs black with apices of mesofemur and metafemur red- Host plants.—Acacia amentacea, A. ber- dish. Vestiture white, golden, light brown, landieri, A. constricta, A. neovernicosa, A. and dark brown intermixed but usually rigidula, A. vernicosa. without discernible pattern, occasionally Natural enemies.—Eupelmus sp., E. am- with vague white lines on pronotum, elytra, icus; Heterospilus prosopidis; Horismenus and pygidium; venter of body white. missouriensis; Stenocorse bruchivora; Urosi- Structure.—Vertex and frons densely mi- galphus bruchivorus, U. neobruchi; Zatropis cropunctulate, frons with subtriangular, incertus. granulose, frontal boss (figure 57); ocular Immatures.—Not described. index 2.5:1; ocular sinus one-half length of eye; eye not protruding from side of head, Discussion.—This species can be distin- blending smoothly into contour of head guished by its small relative size, black (figure 791); pronotum bell-shaped (figure head, vaguely maculate elytra, and host 790), lateral margins gently curved; disk plant preference. The apical margin of the evenly convex, basal lobe feebly sulcate, ventral valve is arcuate, unlike the tongue- basal margin impressed near posterior like forms in other U.S. species. corners; lateral carina visible as short ridge Mimosestes amicus (Horn) in middle of margin; cervical sulcus concealed by vestiture. Scutellum quadrate, (Figures 27, 57, 58, 790–794) emarginate. Elytra slightly longer than Bruchus amicus Horn 1873:337; Riley wide (figure 790), evenly convex except and Howard 1892c:165; Ashmead slightly depressed around scutellum; striae 1894:342; Horn 1894:345; Townsend shallow, narrow, strial punctures elongate, 1895:277; Cockerell 1902:379; Fall 3rd, 4th, and 5th denticulate on basal and Cockerell 1907:201; Schaeffer margin, 6th sometimes denticulate; inter- 1907:292; Fall 1910:164; Cushman stices flat, finely imbricate; metacoxal face 1911:495; Zacher 1952:47. densely punctate except fossula glabrous; hind leg as in figure 792; male femur with Acanthoscelides amicus: Blackwelder deep, setose pocket; pecten with one large 1946:758. denticle and two or three small denticles; Mylabris amicus: Leng 1920:305; Kannan metatibia with all carinae complete except 1923:20. ventrolateral carina apically obsolete; mu- Mimosestes amicus: Muesebeck et al. cro shorter than lateral denticle; coronal 1951:467; Hinckley 1960:261; Bibby denticles three. Male 5th sternum slightly 1961:325; Peck 1963:956; De Luca emarginate; male pygidium moder- 1965:67; Bottimer 1968c:1024,1039; Johnson 1968:1269, 1969c:55; Smith and Ueckert 1974:63; Swier 1974;

187 ately reflexed at apex, female pygidium Mimosestes species and from nearly all oblique; disk densely punctulate with scat- other U.S. bruchids by the character of tered microfoveolae. the posterior margin of the eye merging Male genitalia.—As in figures 793 and 794; smoothly with the side of the head. It prob- median lobe broad (figure 793); dorsal ably breeds most successfully in seeds of valve arcuate with apex slightly produced; Parkinsonia species but is abundant in ventral valve liguliform, truncate; internal Prosopis seeds as well. This bruchid ovi- sac densely lined with fine spicules; middle posits only on green pods. Its exit holes of sac with flat, crescentic sclerite; closure are used by other species of bruchids for valve circular; lateral lobes bowed (figure oviposition sites. 794), cleft nearly to base, arms narrow, Kannan (1923) described oviposition by M. apices strongly expanded mesad; mesal amicus. faces setose. Wheeler and Longino (1988) found M. am- Size.—Body length 2.1–3.8 mm; width icus as an inquiline in galls of Dishol-cas- 1.4–2.5 mm. pis cinerosa Bassett (Cynipidae) in Texas. Type depository.—MCZC, lectotype No. Scullen and Wold (1969) collected M. am- 8208 (by Johnson 1968). icus as prey of the wasp Cerceris truncata Cameron. Type locality.—Texas (lectotype). Distribution.—AZ, CA, CO, FL, HI, KS, NV, Mimosestes insularis Kingsolver NM, TX; Mexico to Costa Rica. and Johnson Host plants.—Rearing records: Acacia (Figures 795–800) constricta, A. farnesiana, A. glauca (Ha- Mimosestes insularis Kingsolver and John- waii); Leucaena leucocephala; Parkinsonia son 1978:35; Johnson and Kingsolver aculeata, P. florida, P. microphylla, P. texana 1982:418; Johnson 1987:348; Udayagiri macra, P. texana texana; Prosopis glandu- and Wadhi 1989:92; Kingsolver 1992b. losa glandulosa, P. glandulosa torreyana, P. velutina, P. pallida, P. pubescens; Sesba- Color.—Integument mostly reddish orange; nia sesban (Hawaii), S. aegyptiaca. Proso- antenna reddish, sometimes with api- pis julianus records in the United States cal seven segments brown; pygidium with should be referred to P. velutina in Arizona, median or apical dark brown maculation; California, and New Mexico and to P. glan- legs reddish orange. Vestiture intermixed dulosa glandulosa in Texas. Records in the white, golden, and brown; pronotum with literature that require verification areCae - lateral patches and median line of white salpinia coriaria and Cassia occidentalis. setae, usually with two small white spots Found in flowers ofLippia wrightii Gray. on disk; elytra white with lateral patches of darker setae; pygidium with basal tri- Natural enemies.—Cerceris truncata; Eu- angular patch of white sometimes extend- pelmus amicus, E. cyaniceps; Hetero-spilus ing as a median line to apex, dark brown prosopidis; Horismenus productus; Steno- integumental maculation with brown setae, corse bruchivora; Urosigalphus bruchi, U. marginal setae white; venter of body white. neobruchi; Uscana semifumipennis. Structure.—Vertex and frons densely Immatures.—Kannan 1923 (egg); Pfaffen- punctulate, punctures crowded and co- berger and Johnson 1976:34 (first larval alescent; frontal boss broadly V-shaped, instar). granulose; ocular index 3.5:1; ocular sinus Discussion.—Mimosestes amicus is one three-fifths length of eye; posterior margin of the most abundant and widespread of eye evenly rounded (figure 796); antenna bruchids in the southwestern United slender, extending to middle of metepi- States. It is easily separated from other sternum. Pronotum bell-shaped (figure

188 795), lateral margins arcuate; disk convex, Immatures.—Not described. feebly sulcate on meson, lightly impressed Discussion.—Among the species treated either side of meson and along basal marin this handbook, Merobruchus insularis gin, surface densely but irregularly foveo- would most easily confused with M. nu- late; lateral carina ridgelike, finely den- bigens. Since they both occur in Hawaii, ticulate; cervical sulcus deep, constricting they can easily be separated by the shape neck of pronotum. Scutellum transverse, of the posterior margin of the eye—that of emarginate. Elytra slightly longer than M. insularis is evenly rounded, and that wide (figure 795), evenly convex; striae of M. nubigens is somewhat angulate. The lightly impressed, punctures distinct, 3rd, form of the “shoulder” of the pronotum of 4th, 5th, and 6th striae denticulate on M. nubigens is protruding and denticulate, basal margin; interstices flat, finely imbri- whereas that of M. insularis is rounded cate, punctate in basal one-half; metacoxal behind the cervical sulcus but not denticu- face densely punctulate, fossula glabrous; late. Male genitalia are diagnostic (compare hind leg as in figure 798, pecten with one figures 799 and 810). The only other spe- long and two or three shorter denticles; cies of Mimosestes in Hawaii is M. amicus, femoral pocket of male more than one-half which is readily separated by the flattened length of femur, ventromesal margin of posterior margin of the eye and gray ap- pocket not visible in lateral aspect; metati- pearance. bial carina distinct but with ventrolateral carina extending only one-half distance to Mimosestes insularis was most likely in- apex; mucro short, only as long as lateral troduced into Hawaii from the West Indies, denticle, coronal denticles three or four. since both of its host plant species occur in Fifth sternum of male slightly emargin- both areas. ate; pygidium of both sexes oblique, male feebly reflexed at apex; discal surface finely Mimosestes mimosae (Fabricius) granulose with scattered fine foveolae, (Figures 801–805) densely setose. Only the references to original descriptions Male genitalia.—As in figures 799 and 800; are given here. For a complete listing of median lobe moderately slender (figure synonyms and combinations, see King- 799); dorsal valve with slight apical an- solver and Johnson (1978:42). gulation; ventral valve liguliform, evenly Bruchus mimosae Fabricius 1781:76. rounded; internal sac densely lined with minute denticles; apical one-half with Bruchus dominicanus Jekel 1855:12. hourglass-shaped, flat sclerite; closure Bruchus breweri Crotch 1867:389. valve circular; lateral lobes slightly bowed Bruchus inornatus Horn 1873:333. (figure 800), cleft nearly to base, arms Bruchus subrufus Motschulsky 1874:225. slender, expanded mesad apically. Bruchus strigatus Motschulsky 1874:237. Size.—Body length 2.3–4.4 mm; width Bruchus immunis Sharp 1885:474. 1.5–2.8 mm. Bruchus innotatus Pic 1912:92 (proposed Type depository.—USNM, holotype No. for inornatus Horn, not Küster 1850). 72779. Acanthoscelides breweri: Blackwelder Type locality.—Hawaii, Oahu, Waikiki. 1946:760. Distribution.—HI; West Indies, Colombia. Acanthoscelides dominicanus: Blackwelder Host plants.—Acacia farnesiana; Prosopis 1946:759. glandulosa glandulosa, P. juliflora, P. pal- Acanthoscelides immunis: Blackwelder lida (last two in Hawaii). 1946:759. Natural enemies.—None recorded. Acanthoscelides innotatus: Bradley 1947:40.

189 Acanthoscelides mimosae: Blackwelder of elytra; metacoxal face finely, discretely 1946:760. punctate except fossula glabrous, lateral Acanthoscelides strigatus: Blackwelder one-fourth densely pubescent; hind leg (fig- 1946:761. ure 803) with ventromesal carina denticulate, pecten with one long and two smaller Acanthoscelides subrufus: Blackwelder denticles, male femoral pocket about one- 1946:761. half length of femur, ventro-mesal border Mimosestes dominicanus: Johnson and of pocket visible in lateral aspect; metatibia Kingsolver 1977:154. slender, all carinae present except ventro- Mimosestes immunis: Kingsolver 1975a:60. lateral abbreviated at apex; mucro only as Mimosestes innotatus: Bottimer long as lateral denticle, coronal denticles 1968c:1024. three, none as long as lateral denticle. Mimosestes inornatus: Johnson 1968:1269. Fifth ventral segment of abdomen broadly, shallowly emarginate; pygidium in both Mimosestes mimosae: Decelle 1975:138. sexes oblique, male slightly reflexed; disk Mimosestes strigatus: Johnson and King- densely, finely punctate, with scattered, solver 1977:154. small foveolae. Mylabris immunis: Leng 1920:305. Male genitalia.—As in figures 804 and 805; Mylabris innotatus: Leng 1920:305. median lobe moderately broad (figure 804); Color.—Head black to red-orange, V-shaped dorsal valve semicircular, membranous; frontal boss black; antenna varying from ventral valve liguliform, broad, truncate; all red-orange to having basal four seg- internal sac finely denticulate, middle of ments red-orange and apical segments sac with two arcuate clusters of coarse dark brown. Body usually red-orange vary- denticles, median sclerite oblong, rectan- ing to dark reddish brown; thoracic sterna gular, apex of sac densely, finely denticu- sometimes black; pronotum and elytra late; closure valve circular; lateral lobes usually darker than remainder of body. strongly bowed (figure 805), cleft to base, arms narrow, only slightly expanded at Structure.—Vertex and frons minutely im- apices. bricate-punctate; frontal boss Y-shaped, granulose; ocular index 3.2:1, ocular sinus Size.—Body length 2.0–4.3 mm; width one-half as long as eye; dorsal margin of 1.3–2.5 mm. eye arcuate, not evenly rounded (figure Type depository.—UZMC (mimosae); BMNH 802); antenna normal for genus, extend- (breweri, dominicanus, immunis); MCZC ing to middle of metepisternum. Pronotum (inornatus); ZMUM (strigatus, subrufus). bell-shaped (figure 801), lateral margins arcuate; disk convex, feebly sulcate on Type locality.—”Americae meridionalis” basal lobe, basal margin impressed; sur- (mimosae); Santa Domingo (dominicanus); face densely foveolate, foveolae discrete, Azores (breweri); “Middle States” (inornat- each with anteriorly positioned seta, inter- us); Colombia (subrufus); Mexico (strigatus); spaces punctulate; lateral carina ridgelike, Panama (immunis). finely denticulate; cervical sulcus short, Distribution.—FL; Mexico, West Indies, nearly hidden by vestiture. Scutellum, South America; introduced into Europe. small, quadrate, bidentate. Elytra as long Host plants.—In Florida: Caesalpinia co- as wide (figure 801), convex but feebly riaria. Extralimital: Acacia sp., A. cochli- depressed along suture; striae parallel, acantha, A. cymbispina, A. farnes-iana, A. slightly divergent toward base, 3rd, 4th, globulifera, A. hindsii, A. macracantha, A. 5th, and 6th prominently denticulate on pennatula; Caesalpinia spp., C. coriaria, C. basal margin; interstices flat, finely imbri- sclerocarpa; Ceratonia siliqua; Cordia sp. cate, distinctly foveolate in basal one-half (questionable); Lotus sp.

190 Natural enemies.—None recorded from and Johnson 1976:196; Pfaffenberger United States. See Hetz and Johnson and Johnson 1976:36. (1988) for extralimital parasites. Color.—Integument reddish orange to Immatures.—Not described. black; frons often with black maculation; four basal antennal segments red, club Discussion.—This species is included here segments black; ventral areas usually on the basis of a single collection at Key darker than pronotum and elytra. Vestiture West, Florida. It is closely similar to M. white, yellow, and dark brown; dorsal sur- insularis and M. nubigens in color pattern face with variable pattern; pronotum usu- and body form. The denticulate pronotal ally with two sublateral white spots; elytra “shoulder” and subangulate postocular variably mottled, usually in linear pattern, lobe of M. nubigens will separate it from medial sublateral and apical maculae usu- the other two species. If M. insularis should ally evident; pygidium usually with median be collected in Florida, the key characters white line of pubescence and pair of sub- and male genitalia will separate it from lateral white spots, disk sometimes uni- both M. nubigens and M. mimosae. formly clothed. Mimosestes nubigens (Motschulsky) Structure.—Vertex and frons imbricate- punctulate, frontal boss broadly V-shaped, (Figures 806–811) granulose; ocular index 2.8:1; ocular sinus Bruchus nubigens Motschulsky 1874:237. one-half as long as eye; postocular lobe Acanthoscelides nubigens: Blackwelder subangulate (figure 807); antenna extend- 1946:760. ing to humerus. Pronotum trapezoidal, lateral margins convergent to apex, con- Mimosestes nubigens: Johnson and King- stricted by cervical sulcus, anterolateral solver 1977:154; Kingsolver and John- corner swollen, strongly denticulate (figure son 1978:49; Johnson and Kingsolver 808); disk convex, basal lobe feebly sul- 1982:418; Borowiec 1987:86; John- cate, basal margin impressed, surface of son 1987:348; Udayagiri and Wadhi disk finely foveolate, interspaces minutely 1989:93; Johnson and Seeno 1993. punctate; lateral carina ridgelike, denticu- Bruchus sallaei Sharp 1885:475; Schaef- late; cervical sulcus short, deep. Scutellum fer 1907:292; Fall 1910:169; Cush- small, transverse, bidentate. Elytra slightly man 1911:491; Bridwell 1919:16, longer than wide (figure 806), striae shal- 1920a:337, 1920b:405, 1921b:465; low, narrow, parallel, 1st to 6th slightly Swezey 1925:3. divergent near base, 3rd to 6th denticulate Bruchus sallei: Pic 1913a:47 (misspelling). on basal margin; interstices flat, punctate- Mylabris sallei: Leng 1920:305 (misspell- granulose; metacoxal face finely, densely ing). punctate, fossula glabrous, lateral one- Mylabris sallaei: Kannan 1923:22. fourth of face densely pubescent; hind leg as in figure 809, femoral pocket of male Acanthoscelides sallaei: Blackwelder about one-half length of femur, mesal 1946:761. margin of pocket visible in lateral aspect, Mimosestes sallaei: Bridwell 1946:54; ventromesal carina denticulate basad of Blackwelder and Blackwelder 1948:45; pecten, pecten with one long and two short Zacher 1952:462; Hinckley 1960:261, denticles; metatibia slender, carinae dis- 1961:526; Peck 1963:956; De Luca tinct except ventrolateral abbreviated near 1965:67; Bottimer1968c:1025,1039, apex; mucro small, shorter than lateral 1041; Kingsolver 1970a:383; Foris- denticle, coronal denticles three, all small- ter and Johnson 1971:232; Bottimer er than lateral denticle. Fifth ventral seg- 1973:549; Howe 1973:575; Decelle ment of male abdomen broadly emarginate; 1975:137; Janzen 1975:157; Center pygidium oblique, apex of male

191 slightly reflexed; discal face finely imbricate Discussion.—This species of Mimosestes with scattered minute foveolae. is easily recognizable by the unique, pro- Male genitalia.—As in figures 810 and 811; nounced denticulate “shoulders” of the median lobe broad (figure 810), dorsal pronotum. Other characters are the sub- valve hoodlike, apex subangulate, ventral angulate postocular lobe of the eye and the valve broadly liguliform, truncate or slight- large, transverse plate in the internal sac. ly emarginate; internal sac lined with fine The preferred host for M. nubigens is denticles in basal one-half and in apex, probably Acacia farnesiana both on the middle of sac with transverse, sclerotized mainland and in Hawaii. Large numbers plate; closure valve circular; lateral lobes of beetles emerge from samples taken cleft nearly to base (figure 811), arms flat, anywhere within its host range. Swezey gradually expanded to apices, mesal faces (1936) and Hinckley (1960) report rearing densely setose. M. nubigens from both A. farnesiana and Size.—Body length 2.2–4.1 mm; width Prosopis pallida (as Prosopis chilensis) in 1.3–2.7 mm. Hawaii. Fosberg (1966) detailed usage of names applied to the Hawaiian species of Type depository.—ZMUM (nubigens); BMNH Prosopis and concluded that P. pallida, but (sallaei). neither P. chilensis nor P. juliflora, is the Type locality.—”Bresil” (nubigens); Mexico, correct name. Guanajuato (sallaei). Kannan (1923) described oviposition of this Distribution.—AZ, CA, FL, HI, TX; Mexico to species. Brazil; introduced into New Caledonia and the Philippines. Mimosestes protractus (Horn) Host plants.—Rearing records: Acacia cor- (Figures 812–817) nigera, A. farnesiana, A. schaffneri, A. tor- Bruchus protractus Horn 1873:332; Sharp tuosa; Lotus sp.; Prosopis glandulosa glan- 1885:476; Horn 1886:xi, 1894:345; dulosa, P. glandulosa torreyana, P. pallida Schaeffer 1907:291; Fall 1910:169; (Hawaii). Records needing confirmation: Cushman 1911:507. Acacia amentacea; Caesalpinia coriaria; Mylabris protractus: Leng 1920:305. Ceratonia siliqua; Cordia sp.; Gleditsia triacanthos; Prosopis chilensis, P. juliflora. The Acanthoscelides protractus: Blackwelder common name “klu” is applied to Acacia 1946:760; Zacher 1952:465,471. farnesiana in Hawaii. Literature records Mimosestes protractus: Johnson for Prosopis chilensis in Hawaii should be 1968:1270; Bottimer 1968c:1025,1039; referred to P. pallida (Fosberg 1966). See Swier 1974; Center and Johnson appendix I. 1976:196; Pfaffenberger and Johnson Natural enemies.—Argiope sp.; Bracon sp.; 1976:35; Kingsolver et al. 1977; King- Catolaccus sp.; Cephalonomia hyalini-pen- solver and Johnson 1978:58; Johnson nis; Eupelmus bruchivorus, E. cushmani, and Kingsolver 1982:418; Pfaffenberger E. cyaniceps; Eurytoma sp., E. tyloderma- 1985a:4; Udayagiri and Wadhi 1989:94. tis; Heterospilus prosopidis; Horismenus Bruchus longiventris Sharp 1885:476; Horn bruchophagus; Lariophagus texanus; 1894:345. Monomorium sp.; Parasierola distinguenda; Mimosestes longiventris: Johnson and Pyemotes boylei; Stenocorse bruchivora; Kingsolver 1977:154. Urosigalphus bruchi, U. neobruchi; Uscana Color.—Integument of head usually black, semifumipennis; Zelus renardii. body and appendages reddish orange, Immatures.—Hinckley 1960 (as M. sallaei; elytra usually with darker lateral maculae, egg); Pfaffenberger and Johnson 1976:36 (as M. sallaei; first instar).

192 pygidium with median, hourglass-shaped lined with blunt denticles, apical part of maculation; metafemur with dark brown sac with slender spicules; closure valve maculation. Vestiture white, golden, and small, circular; lateral lobes divergent (fig- brown; head white; pronotum mostly white ure 817), outline as figured. intermixed with brown and with brown lat- Size.—Body length 2.0–3.5 mm; width eral patches; elytra white with dark brown 1.2–2.1 mm. lateral and apical maculae (figure 812); pygidium white with median maculation Type depository.—MCZC (protractus, holo- dark brown constricted by white patches at type No. 8200); BMNH (longiventris). midpoint; venter of body white. Type locality.—Lower California (protractus); Arizona (longiventris). Structure.—Body elongate, lateral margins subparallel. Vertex and frons densely Distribution.—AZ, CA, NV, TX, UT; Mexico. punctulate; frontal boss broadly V-shaped, Host plants.—Prosopis glandulosa glandulo- granulose; ocular index 3.2:1; ocular sinus sa, P. glandulosa torreyana, P. laevigata, P. four-tenths length of eye; antenna extend- velutina; Parkinsonia microphylla, P. texana ing to middle of metepisternum. Pronotum texana. Names of Prosopis species have bell-shaped (figure 812), lateral margins been confused in the literature. I followed sinuate, apex evenly rounded; disk con- Johnston (1962) in application of names, vex, impressed along basal margin; surface but where species or varieties overlap, densely foveolate, each foveola bearing especially in western Texas, literature re- seta on anterior rim, interspaces punctu- cords are unreliable. late; lateral carina ridgelike, denticulate, Natural enemies.—Cerceris truncata Cam- densely pubescent; cervical sulcus distinct, eron (Scullen and Wold 1969). deep. Scutellum quadrate, densely setose. Elytra 1.2 times as long as wide (figure Immatures.—Pfaffenberger and Johnson 812); moderately convex; striae shallow, 1976:35 (first instar). sometimes evident only by a line of punc- Discussion.—The elongate, parallel-sided tures, subparallel, 2nd to 6th divergent form, reddish-orange appendages, and laterad in basal one-half, 3rd to 6th den- depression of the 5th sternum should eas- ticulate on basal margin; interstices flat, ily separate this species from other North imbricate-granulose; metacoxal face evenly American Mimosestes. punctulate, fossula glabrous; hind leg as in figure 815, ventral margin of male meta-fe- This bruchid has been found as an inqui- mur shallowly sulcate in basal one-fourth; line in galls of Disholcaspis cinerosa (Bas- pecten with one long and two small den- sett) (Cynipidae) by Wheeler and Longino ticles; metatibial carinae present except (1988). ventrolateral evanescent in apical one-half; mucro not as long as lateral carina, three Mimosestes ulkei (Horn) coronal denticles minute. Male 5th abdom- (Figures 818–822) inal sternum with deep, round depression Bruchus ulkei Horn 1873:324; Riley and (figure 814), female 5th sternum shallowly Howard 1892c:165; Fall 1910:164; sulcate or slightly impressed; pygidial Cushman 1911:494,508. disk densely foveolate as on pronotal disk, interspaces punctulate, sculpture nearly Bruchus (Cercidiestes) ulkei: Zacher hidden by vestiture. 1952:463. Cercidiestes ulkei: Bridwell 1946:55 (type Male genitalia.—As in figures 816 and 817; species); Blackwelder and Blackwelder median lobe slender (figure 816); dorsal 1948:45; Zacher 1952:463,472; Bot- valve evenly rounded; ventral valve trian- timer 1968c:1021,1039; Johnson gular, elongate; internal sac lined with fine 1968:1269, 1969a:54; Center and spicules at apical orifice, middle one-half Johnson 1976:196.

193 Mimosestes ulkei: Kingsolver and John- concealed by vestiture; mucro short, son 1978:62; Johnson and Kingsolver one-half as long as lateral denticle; three 1982:413; Johnson 1987:348; Udaya- coronal denticles minute. Pygidium feebly giri and Wadhi 1989:94. convex, disk densely pubescent; minutely Mylabris ulkei: Leng 1920:305; Kannan punctate with scattered foveolae. 1923:19. Male genitalia.—As in figures 821 and 822; Color.—Integument reddish orange to median lobe moderately slender (figure black; head black, antenna reddish or- 821); dorsal valve evenly rounded, densely ange; elytra black; fore legs and mid legs setose; ventral valve broadly liguliform, reddish brown to orange, hind legs black, apex broadly angulate; internal sac lined often with mesal face brown; abdomen of- with blunt denticles in basal one-half, with ten with orange suffusion. Vestiture white clusters of spicules in apical one-half; clo- covering entire body except dark brown to sure valve circular; lateral lobes with arms black on lateral one-half of each elytron, strongly bowed (figure 822), densely setose median, wedge-shaped elytral stripe gray at apices. (figure 818), two black spots of variable Size.—Body length 2.5–4.9 mm; width size and shape on pygidium (figure 819), 1.9–2.9 mm. and a black spot on 5th sternum each side of meson. Type depository.—MCZC, holotype No. 1873. Structure.—Vertex and frons densely punctulate; frontal boss broadly triangu- Type locality.—Arizona. lar, granulose; ocular index 4:1; ocular Distribution.—AZ; Mexico. sinus eight-tenths length of eye; posterior Host plants.—Parkinsonia aculeata, P. margin of eye subangulate; antenna sub- florida, P. texana texana. serrate, extending to humerus. Pronotum bell-shaped (figure 818), strongly convex, Natural enemies.—None recorded. lateral margins gently arcuate, basal lobe Immatures.—Kannan 1923 (egg). sulcate, basal margin impressed, apical margin evenly rounded; surface sparsely Discussion.—This strikingly marked bru- foveolate and punctulate, nearly concealed chid is probably the most easily recognized by dense vestiture; lateral carina ridge- in North America. It was designated the like, denticulate, densely pubescent, form- type species of the genus Cercidiestes by ing sharp “shoulder” behind eye; cervical Bridwell in 1946, but Kingsolver and John- sulcus nearly hidden by vestiture. Scutel- son (1978) moved it to Mimosestes based lum quadrate, bidentate, pubescent. Elytra on form of male genitalia, male metafemur slightly longer than wide (figure 818); con- with ventral pocket, and head with promi- vex, subdepressed around scutellum; stri- nent boss. This species, while not rare in ae distinct in gray mesal area but difficult collections, is less abundant than other to distinguish within black lateral stripes, species of Mimosestes. subparallel, slightly divergent toward base, Essig (1929a) gave the common name 3-5 denticulate on basal margin; interstic- “retama weevil” to this bruchid after a local es flat, imbricate-granulose; metacoxal face name for Parkinsonia aculeata. densely punctulate and pubescent except fossula glabrous; hind leg as in figure 820; Genus Neltumius Bridwell male with small, shallow pocket on ventral Neltumius Bridwell 1946:54; Bradley face of femur near apex of trochanter; pect- 1947:35; Kingsolver 1964a:105; Bot- en with one long, acute denticle; metatibia timer 1968c:1025,1039,1041; Johnson with all carinae present except ventrolater- 1968:1270; Johnson and Kingsolver al abbreviated at midpoint, carinae nearly 1982:416; Borowiec 1987:97; Udayagiri and Wadhi 1989:95.

194 Type species: Bruchus arizonensis New World. Structure of the metatibia Schaeffer, by monotypy. resembles that of Old World Bruchidius, Integument black; vestitural pattern dis- but because of the close association of two tinctive for each species. of the three species with the plant genus Prosopis, itself highly developed in the New Frons carinate, sparsely pubescent; anten- World, Neltumius is likely a relict group na serrate, short, extending to humerus, originating in this hemisphere. Bradley not dimorphic. Pronotum strongly convex, (1947) thought that Neltumius was a close gibbous; lateral carina lacking; cervical relative of Gibbobruchus because of the sulcus narrow, hidden by pubescence. gibbous pronotum, but other characters Elytra together slightly longer than wide; of Gibbobruchus (pecten with several den- striae deep, narrow, lacking basal denticles, strong basal elytral denticles, and ticles. Pygidium dimorphic in color pattern form of the male genitalia) negate this as- and convexity. Metafemur with single, sub- sociation. apical denticle (figure 832); metatibia with mucro shorter. Revision: Kingsolver 1964a. Neltumius does not seem to be closely related to any of the other genera in the

Key to Species of Neltumius elongated white patches separated by dark brown spots 1 Metatibia uniformly gray; dorsal pattern mostly gray- (figures 823 and 838); humerus carinate or granulose.....2 ish white with prominent patches of dark brown and 2(1) Pronotum with prominent, paired gibbosities separated yellowish setae; pronotum with single median gibbosity by shallow mesal and transverse channels (figure 823), near apex (figure 830); middle of 3rd elytral interstice posterior gibbosities more prominent than anterior pair with elongate, white spot (figure 829); humerus with fine, (figure 824); humerus with fine, transverse carina transverse carina ...... arizonensis (Schaeffer) ...... (Schaeffer) gibbithorax Pronotum with gibbosities and channel only slightly Metatibia with brown band at middle of dorsal margin; developed (figures 838 and 839); humerus granulose dorsal pattern of strongly contrasting dark brown and ...... texanus (Schaeffer) gray elongated patches; 3rd interstice with two or more

Neltumius arizonensis (Schaeffer) Johnson 1983c:28; Udayagiri and Wa- dhi 1989:95. (Figures 823–828) Color.—Vestiture black, white, and reddish Bruchus arizonensis Schaeffer 1904:229; brown in distinctive pattern (figure 823) Fall 1910:162. on pronotum and elytra; pygidial pattern Mylabris arizonensis: Leng 1920:305; Kan- dimorphic (figures 825 and 826); venter of nan 1923:21. body gray except brown spots on metepi- Neltumius arizonensis: Bridwell 1946:54; sternum, lateral margins of abdominal Bradley 1947:36; Bibby 1961:325 (as sterna, and on each femur and tibia. griseolus); Kingsolver 1964a:108; Bot- Structure.—Head densely punctulate and timer 1968c:1025,1039,1041; John- pubescent, nearly concealing surface, son 1968:1270; Forister 1970:68; frontal carina distinct; ocular index 5:1; Swier 1974; Pfaffenberger and Johnson ocular sinus three-fifths as long as eye. 1976:37; Kingsolver et al. 1977:115; Pronotum strongly gibbous with median, ohnson and Kingsolver 1982:416; longitudinal sulcus separating the two

195 pairs of gibbosities, sulcus broader basally Immatures.—Kannan 1923:21 (egg, first in- (figure 823). Elytral striae deep, narrow, star); Pfaffenberger and Johnson 1976:37. 6th and 7th striae connected subbasally by Discussion.—The pronotum of N. arizo-ne- short, dentate carina; alternate interstices nsis is more deeply sulcate than that of N. broader and more elevated, surface finely gibbithorax and bears two subequal pairs imbricate; metacoxal face densely punc-tu- of gibbosities, whereas the prominent, late, pubescent in lateral angle; ventrolat- anteriorly placed gibbosity of N. gibbitho- eral carina of metatibia absent, the other rax is not sulcate. The pronotal gibbosity three nearly concealed by vestiture; mucro of N. texanus is much less prominent and smaller than lateral denticle, coronal den- is placed nearer the base of the pronotum ticles minute. First abdominal sternum of than in the other two species. Pygidial pat- male with small tuft of hair; male pygidium terns of the three species, size differences moderately convex, strongly reflexed into between N. texanus and the other two sternal emargination, female pygidium species, and details of male genitalia are convex in basal one-half, strongly swollen salient characters for discrimination. apically. Male genitalia.—As in figures 827 and 828; Neltumius gibbithorax (Schaeffer) median lobe slender (figure 827); ventral (Figures 829–837) valve semicircular with small membranous area at apex; armature of internal sac con- Bruchus gibbithorax Schaeffer 1904:230; sisting of two burrlike sclerites near base, Fall 1910:162; Pic 1913a:27. a mass of spicules in radiating pattern in Neltumius gibbithorax: Bradley 1947:36; middle of sac, and two thornlike sclerites Bottimer 1968c:1025,1039; Johnson near apex; closure valve small, circular; 1968:1270; Kingsolver et al. 1977:115 lateral lobes as figured (figure 828), cleft et seq.; Johnson and Kingsolver three-fourths to base, arms bowed and 1982:416; Johnson 1983c:28; Udaya- setose on mesal faces. giri and Wadhi 1989:95. Size.—Body length 2.5–4.0 mm; width Neltumius gibbothorax: Kingsolver 1.4–2.1 mm. 1964a:106 (misspelling). Mylabris gibbithorax: Leng 1920:305. Type depository.—USNM, holotype No. 42285. Color.—Vestiture of gray intermixed with orange and brown (pattern, figure 829); Type locality.—Arizona, Pinal Mountains. pygidial pattern as in figures 833 and 834; Distribution.—AZ, CA, NV, TX; Mexico. venter of body gray intermixed with orange. Host plants.—Prosopis velutina, P. glandu- Structure.—Punctation of vertex and frons losa glandulosa, P. glandulosa torreyana. nearly concealed by dense vestiture; frontal Records of Prosopis chilensis and P. juliflora carina prominent, sometimes with pit at are probably the result of misidentification dorsal end; ocular index 4:1; ocular si- of the plants since neither species is within nus three-fifths length of eye; antenna not the range of N. arizo-nensis according to dimorphic (figure 831). Pronotum gibbous Johnston (1962). Records of Chilopsis in anterior one-half of disk (figure 830), not linearis (Cav.) Sweet and Larrea tridentata channeled (figure 829); vestiture dense, (Sesse and Mocino) Cov. are undoubtedly concealing surface; lateral carina absent; accidental. cervical sulcus concealed. Scutellum small, Natural enemies.—Urosigalphus arizonensis triangular. Elytra together as long as wide Crawford, U. bruchivorus Crawford. This (figure 829); strial punctures elongate, bruchid is also listed as the prey of Cerce- striae slightly impressed; interstices ris truncata Cameron (Hymenoptera) (Scul- len and Wold 1969).

196 densely pubescent; 3rd, 5th, and 7th inter- Neltumius texanus (Schaeffer) stices wider than 4th, 6th, and 8th; hu- (Figures 67, 838–844) merus with transverse carina, sometimes dentate; metacoxa densely punctate, punc- Bruchus texanus Schaeffer 1904:231; Fall tation concealed by dense pubescence ex- 1910:162. cept fossula narrowly glabrous; ventrolat- Mylabris texanus: Leng 1920:305. eral carina of metatibia absent, remaining Neltumius texanus: Bradley 1947:36; carinae concealed by dense pubescence. Kingsolver 1964a:110; Bottimer First abdominal sternum of male with tuft 1968c:1025,1039; Johnson 1968:1270; of setae; male pygidium strongly reflexed at Kingsolver et al. 1977:115; John- apex, female pygidium bulbous; punctation son 1978:432; Johnson and King- concealed by dense pubescence. solver 1982:416; Udayagiri and Wadhi Male genitalia.—As in figures 835, 836, 1989:95; Johnson 1995a:42. and 837; median lobe relatively broad (fig- Color.—Vestiture white and reddish brown ure 835); ventral valve nearly semicircular, arranged in pattern in figure 838; head apex slightly produced and membranous; with mixed reddish brown and gray; prono- internal sac with two large, burrlike scler- tum gray with paired reddish-brown spots ites occupying basal one-third, a dense, near apex, another pair either side of white fanlike cluster of long, slender spicules in pubescent wedge on basal lobe; humeri middle of sac, and two flat, lunate sclerites black, elytra otherwise gray with reddish- in apical one-half; extreme apex of sac with brown maculae, 3rd interstice with alter- sleevelike, spinous structure; lateral lobes nating gray and reddish-brown elongate (figure 837) relatively flat, deeply cleft, spots; pygidium gray with evanescent hori- arms broad, apex with transverse, setose zontal maculae (figures 840 and 841); ven- lobe, mesal faces of lobes densely setose. ter of body gray with two or three reddish- Size.—Body length 2.6–4.2 mm; width brown spots on metepisternum; abdominal 1.5–2.0 mm. sterna mixed reddish brown and gray; legs faintly reddish brown with distinct brown Type depository.—USNM, lectotype No. spot on dorsal face of metatibia. 42286, by Johnson 1968. Structure.—Vertex finely punctate; frontal Type locality.—Arizona, Pinal Mountains carina distinct, sharp; ocular index 4:1; (lectotype). ocular sinus one-half length of eye; an- Distribution.—AZ, CA, NV, UT; Mexico. tenna not dimorphic, extending past 1st Host plants.—Prosopis glandulosa glandu- abdominal segment. Pronotum bell-shaped losa, P. glandulosa torreyana, P. pubescens. (figure 838), strongly convex, briefly sul- A record of Pluchea sericea is probably ac- cate on prominent basal lobe; lateral mar- cidental. gins feebly incurved; lateral carina lacking; cervical sulcus hidden by vestiture. Scutel- Natural enemies.—None recorded. lum small, densely pubescent. Elytra one Immatures.—Not described. and one-tenth times as long as wide (figure 838), evenly convex, without asperities, Discussion.—This species is easily distin- humeri granulose; striae parallel, 3rd and guished by the rounded protuberance in 4th congruent basally, strial punctures the apical one-half of the pronotal disk, the deep, elongate, inconspicuous; interstices body size, and the feebly patterned pygid- flat, minutely imbricate; metacoxal face ium in both sexes. It is subequal in size to finely, densely punctate except fossula gla- N. arizonensis, but larger than N. texanus. brous; metafemur with single, small denticle; metatibia with lateral, ventral, and dorsomesal carinae complete to apex,

197 ventrolateral carina absent; mucro sub- Condalia spp., a genus in the Rhamnaceae, equal in length to lateral denticle, apex whereas host records of the other two spe- with three minute coronal denticles. First cies of Neltumius are with Prosopis spp. in abdominal sternum without tuft of hairs the Mimosoideae. Johnson (1995a) record- similar to those found in N. arizonensis ed additional host and distributional data and N. gibbithorax. for N. texanus. Male genitalia.—As in figures 842, 843, and 844; median lobe slender (figures 842 Genus Sennius Bridwell and 843); ventral valve triangular, apex Sennius Bridwell 1946:55; Brad- bent ventrad; armature of internal sac ley 1947:37,39; Blackwelder and consisting of rodlike cluster of fine spicules Black-welder 1948:45; Bottimer and denticles at apical orifice, apex of sac 1968c:1025,1039,1041; Johnson densely lined with fine denticles, middle of 1968:1270; Center and Johnson 1973; sac with cluster of coarse spines, a cluster Johnson and Kingsolver 1973:1; John- of minute denticles, and a terminal cluster son and Slobodchikoff 1979; Johnson of coarse spines; lateral lobes with arms and Kingsolver 1982:419; Johnson slender (figure 844), cleft nearly to base, 1984a; Borowiec 1987:101; Udaya- apices broadly expanded and with coarse giri and Wadhi 1989:99. Type species: spines on ventral faces. Bruchus cruentatus Horn, by original Size.—Body length 2.0–2.5 mm; width designation. 1.1–1.2 mm. Small beetles (1.8–2.8 mm). Type depository.—USNM, holotype No. Vertex densely punctulate, frons with im- 42287. punctate line, obtuse carina, or sharp cari- Type locality.—Texas, Esperanza Ranch, na; eyes protuberant; antenna eccentric Brownsville. from 5th segment except apical segment ovate, not sexually dimorphic, uniform in Distribution.—AZ, CA, TX; Mexico. outline, except elongated in S. whitei (fig- Host plants.—Condalia correllii, C. globosa ure 904). Pronotum bell-shaped, evenly globosa, C. globosa pubescens, C. hookeri convex, without asperities; disk densely, hookeri, C. obovata, C. spathulata, C. war- uniformly punctulate; lateral carina ob- nockii kearneyana. Zanthoxylum clava-her- tuse or lacking; cervical sulcus narrow culis is probably accidental. but distinct; prosternum triangular, pro- Natural enemies.—Horismenus missou-rien- coxae contiguous. Mesepimeron reduced sis; Urosigalphus bruchivorus. to triangular sclerite near elytral humerus. Scutellum quadrate, apical margin emar- Immatures.—Johnson 1978 (egg). ginate. Elytra convex, disk in some species Discussion.—This species, although obvi- slightly depressed, lateral margins and ously belonging to Neltumius, is distinctive apex arcuate; striae regular, narrow and in details of the male genitalia, granulate shallow, without basal denticles or gibbosi- humeri, lack of setal tuft on 1st abdominal ties in U.S. species, strial punctures fine; sternum, relative size, less protuberant interstices flat, subequal in width, imbri- pronotal gibbosities, and pygidial pattern. cate. Fore legs and mid legs not modified; In pronotal and elytral pattern, it more metacoxal face densely, uniformly punctu- closely resembles N. arizonensis than it late except fossula impunctate, polished; does N. gibbithorax. The stout, thornlike metafemur slightly swollen, ventromesal processes at the apices of the lateral lobes margin with one subapical denticle, ven- are unique in the Bruchidae. trolateral margin rounded, not carinate; Johnson (1978) described the life history metatibia straight with lateral, ventrolat- of this species. All host records are with eral (most species), ventral, and dorsomesal carinae present; mucro relatively short, scarcely longer than 198 coronal denticles. Abdomen unmodified except male 5th sternum emarginate to Key to Species of Sennius receive apex of pygidium; male pygidium more strongly reflexed at apex than female, Some species of Sennius vary greatly in the extent of red elytral macula- disk densely punctulate. tion. Some species that normally are adorned with a large macula may vary Male genitalia with curved sclerite on ei- to an entirely black form (for example, S. cruentatus, S. simulans) ther side of apical orifice (hinge sclerites); whereas others are consistent in their color patterns. Because of these internal sac usually armed with fine denti- variables, construction of an unambiguous key to species is difficult. In cles, some species with clusters of coarser questionable cases, male genitalia should be extracted and compared. denticles; lateral lobes spatulate, fringed Use elytral pattern outlines in conjunction with key because the red with sensitive setae. integumental color does not register well in photographs. Sennius is undoubtedly a valid genus, but 1 Elytra usually with distinct, red maculae, or marginal or separating its members from some spe- medial stripes (figures 846, 852, and 868); legs variously cies and species groups now placed in Acanthoscelides can be difficult.Acantho- colored...... 2 scelides is wide-ranging in its variation and Elytra lacking red maculae or stripes, or with maculae is difficult to characterize as a taxon, mak- indistinct; legs red with base of metafemur black...... 12 ing its discrimination from certain other 2(1) Pygidium red, or reddish brown, sometimes with darker genera difficult. The senniine characters markings; elytral maculae yellowish red, chevron shaped of hinge sclerites, short mucro, and single metafemoral denticle will differentiate (figure 868); basal margin of each elytron usually black, Sennius from the bulk of Acanthoscelides apical one-half of elytra brown to black species. Resemblance in color pattern of ...... lebasi (Fahraeus) some species of Sennius to those in Sta- Pygidium black; elytral maculae of different tor is striking, but structural characters of configuration...... 3 Sennius, particularly the lack of a lateral pronotal carina, short metatibial mucro, 3(2) Elytral maculae red, confined to basal one-half, and presence of a cervical sulcus, lack of a extending from lateral margin to 2nd stria but not reach- ventrolateral carina on the metafemur, and ing basal margin nor humerus (figure 902); metatibial presence of hinge sclerites separate Sen- denticle minute (figure 905); male frontal carina dis-tinct nius species. (figure 903); antenna elongated Irwin and Barneby (1982) revised the legu- (figure 904)...... whitei Johnson and Kingsolver minous tribe Cassiinae—including Cassia, Elytral macula of different configuration...... 4 Senna, and Chamaecrista—and transferred 4(3) Elytral macula medial, extending from lateral margin to Senna most of the species hitherto placed in Cassia. Host names herein in- toward but not reaching suture, basal margin, or apex cluded have been updated in accordance (figures 846, 881, and 892)...... 5 with the 1982 paper. Elytral macula elongate or confined to apical one-half Johnson and Slobodchikoff (1979) includ- (figures 852, 862, and 897)...... 7 ed 34 species of bruchids in their investi- 5(4) Hind leg red, or red with basal one-fourth black; elytral gations of the coevolution of Cassia (Senna) pattern as in figure 881 and Bruchidae. medialis (Sharp) Hind leg entirely black...... 6 Revision: Johnson and Kingsolver (1973) 6(5) Elytral maculation as in figure 846 (North America)...... abbreviatus (Say) Elytral maculation as in figure 892 ...... obesulus (Sharp)

199 Bruchus abbreviatus Horn, listed in Dohrn 7(4) Elytral macula confined to apical one-half (figures 892 1879:187 as synonum of bivulneratus and 897)...... 8 Horn 1873. Elytral macula elongate, extending full length of elytron Mylabris abbreviatus Dohrn: Leng (figures 852 and 862), sometimes in the form of a median 1920:305 (Leng erroneously attributed stripe on each elytron the name to Dohrn)...... 10 Sennius abbreviatus: Bottimer 1968c:1025; 8(7) Elytral macula occupying most of apical one-half (figure Johnson 1968:1270, 1969c:55; John- 897)...... simulans (Schaeffer) son and Kingsolver 1973:17, 1982:419; Udayagiri and Wadhi 1989:99. Elytral macula confined to small subapical patch...... 9 Bruchus bivulneratus Horn 1873:325; 9(8) Male genitalia as in figure 899 Dohrn 1879:187; Riley and Howard ...... simulans (Schaeffer) 1892c:165; Schaeffer 1907:293; Blatch- Male genitalia as in figure 894 ley 1910:1237; Fall 1910:165; Cush- ...... obesulus (Sharp) man 1911:494; Pic 1913a:19; Zacher 1952:461. 10(7) Red macula extending from near humerus diagonally Mylabris bivulneratus: Leng 1920:305. toward, but not reaching, suture (figure 852); legs red except in all-black form (see couplet 12); male genitalia Sennius bivulneratus: Bradley 1947:39; Blackwelder and Blackwelder 1948:45; as in figure 854; eastern United States De Luca 1965:68...... cruentatus (Horn) Color.—Integument black; basal four an- Red stripe of each elytron variable in width, sometimes tennal segments reddish orange, terminal evanescent, sometimes occupying entire width; elytron segments reddish orange to black; elytral usually with indistinct lines of white vestiture (figure macula reddish orange extending from lat- 862); Georgia and Florida...... fallax (Boheman) eral margin to 1st or 2nd stria, basal mar- Red stripe usually limited by black marginal and sutural gin narrowly black (figure 846); legs dark borders, lacking vestitural lines; southwestern United brown to black. Vestiture white, gold, and States...... 11 brown, sparse on pronotal meson, elytra with sparse, intermixed gold and brown se- 11(10) Dorsal vestiture yellowish gray; male genitalia as in figure tae; venter of body dense white; pygidium 859...... discolor (Sharp) white condensed into basal triangle, some- Dorsal vestiture white; male genitalia as in figure times extending as thin line to apex. 889 ...... morosus (Sharp) Structure.—Vertex and frons densely re- 12(1) Elytral vestiture in cruciform white pattern ticulate-foveolate; frons with impunctate (figure 875); male genitalia as in figure 878 boss sometimes extended into brief carina; ocular index 3:1; ocular sinus two-thirds ...... leucostauros Johnson and Kingsolver length of eye. Pronotum subconical (fig- Elytra without pattern; male genitalia as in figure 854 ure 845), apex rounded, lateral margins ...... melanistic form of cruentatus (Horn) straight to slightly arcuate, disk evenly convex, feebly sulcate on basal lobe; lateral Sennius abbreviatus (Say) carina evident only at posterior corner of pronotum; cervical sulcus short, shallow. (Figures 63, 845–850) Elytra together about as long as wide (fig- Bruchus abbreviatus Melsheimer 1806:30 ure 845); striae parallel, deep, strial punc- (catalogue name). tures ovate; interstices coarsely imbricate. Curculio abbreviatus Say 1824:307,308. Metafemur with small denticle (figure 847); metatibial carinae complete except ventro- Bruchus abbreviatus Say 1824:307,308 lateral evanescent toward apex, [validation of name by comparing it in discussion with Bruchus now Mega-cerus] discoidus Say.]

200 mucro short, less than one-eighth as long Sennius cruentatus (Horn) as basitarsus. (Figures 851–855) Male genitalia.—As in figures 848, 849, and 850; median lobe moderately slender Bruchus cruentatus Horn 1873:325; (figure 848), slightly expanded at apex; Sharp 1885:469; Riley and Howard ventral valve semicircular, apex produced; 1892c:165; Schaeffer 1907:294,296; armature of internal sac consisting of two Blatchley 1910:1237; Fall 1910:165; linear clusters of setae at apical orifice, Cushman 1911:498,506; Pic 1913a:23; paired, crescentic hinge sclerites, a dense Zacher 1952:461. cluster of fine spicules and denticles, a Mylabris cruentatus: Leng 1920:305. pair of clusters of coarse, curved spines, Acanthoscelides cruentatus: Blackwelder two lateral sacs lined with fine spicules, 1946:759. extreme apex lined with denticles, closure Sennius cruentatus: Bridwell 1946:55; valve circular; lateral lobes with arms Bradley 1947:39; Blackwelder and slender (figure 850), slightly bowed, apices Blackwelder 1948:45; Bottimer expanded mesad, mesal faces setose. 1968c:1026,1039,1041; Johnson Size.—Body length 2.1–3.0 mm; width 1968:1270; Kingsolver 1968:319; 1.5–2.1 mm. Johnson 1969c:55; Johnson and Kingsolver 1973:38, 1982:419; Borow- Type depository.—Type destroyed (abbre- iec 1987:101; Udayagiri and Wadhi viatus); MCZC (bivulneratus, lectotype No. 1989:101. 8194, by Johnson 1968). Bruchus depressus Fall 1912:321; Pic Type locality.—Unknown (abbreviatus); 1913a:23. “Southern and western states” (bivulneratus). Sennius depressus: Johnson 1968:1270; Bottimer 1968c:1026. Distribution.—AR, DC, GA, IL, IN, KS, KY, Bruchus nictitans Motschulsky 1874:241; LA, MD, MS, MO, NY, NC, OH, PA, TN, TX, Pic 1913a:37; Zacher 1952:462. VA. Mylabris nictitans: Leng 1920:306. Host plants.—Cassia sp.; Senna marilandica. Also collected in flowers ofAsclepias Sennius nictitans: Bottimer 1968c:1025. syriaca; Cicuta maculata; Cryptotaenia Bruchus nigrinus Horn 1873:327; Schaef- canadensis; Eupatorium sp.; Taenidia fer 1907:298; Blatchley 1910:1237; integerrima. Baskin and Baskin (1977) Fall 1910:166; Pic 1913a:37; Zacher found 22 to 91.1 percent seed predation 1952:462. of S. marilandica in Tennessee. This host Sennius nigrinus: Bradley 1947:39; Black- also appears in literature as Cassia mari- welder and Blackwelder 1948:45; Bot- landica. timer 1968c:1026; Johnson 1968:1270, Natural enemies.—Horismenus sp. 1969c:55. Immatures.—Not described. Mylabris nigrinus: Leng 1920:305. Color.—Integument mostly black; each ely- Discussion.—The color pattern of Sennius tron entirely black or black with a lateral, abbreviatus most closely resembles that elongate, red macula sometimes extending of S. medialis, but the reddish-yellow legs from humerus nearly to apex but not ex- of S. medialis, larger femoral denticle, and tending mesad of fourth stria (figure 852); male genitalia separate them easily. Other fore leg and mid leg usually reddish brown; species with partly or wholly red elytra can hind leg reddish brown to black; antenna be separated by characters in the key and with basal four segments yellow, apical by the male genitalia. segments black. Vestiture white, sparse on dorsum, slightly more dense on venter.

201 Structure.—Vertex and frons strongly con- Type locality.—Texas (cruentatus, restricted vex, densely punctulate, frontal carina by lectotype); Florida, Orlando? (depres- evanescent, usually only an impunctate sus); “Am. Bor.” (nictitans); “Middle States” line; ocular index 3.25:1; ocular sinus one- (nigrinus). half as long as eye; antenna not modified, Distribution.—AL, AR, CT, DC, FL, GA, IL, extending to middle of metepisternum. IN, IA, KS, LA, MD, MA, MN, MO, MS, NE, Pronotum nearly semicircular (figure 851), NJ, NY, NC, OH, OK, PA, SC, SD, TN, VA; strongly convex; basal lobe sulcate; surface Mexico. densely foveolate, interspaces punctulate, lateral portions of disk more densely sculp- Host plants.—Chamaecrista fasciculata, C. tured than middle; lateral carina ridgelike, nictitans nictitans; Parkinsonia sp. often cervical sulcus lacking. Scutellum quad- collected. Records in literature of Cassia rate, emarginate. Elytra together about as chamaecrista and Chamaecrista nictitans long as wide (figure 851); disk flattened nictitans are referable to Chamaecrista fas- between 5th striae; striae parallel, shal- ciculata. Often collected in inflorescences low but distinct, lacking basal denticles, of other plant species and in spanish moss interstices flat, punctate-imbricate in basal (Tillandsia usneoides) as a hibernaculum one-half, imbricate toward apex; metacoxal (Rosenfield 1911). face densely, evenly punctate, fossula gla- Natural enemies.—None recorded. brous; hind leg as in figure 853; meta-fem- Immatures.—Not described. oral denticle longer than width of tibia at base; metatibial carinae, with exception of Discussion.—The shape of the elytral abbreviated ventrolateral carina, complete maculation will usually separate S. cruen- to apex; mucro shorter than lateral den- tatus from all the U.S. species except S. ticle, coronal denticles subequal to lateral simulans. The maculation of S. cruentatus denticles; abdomen not modified; pygidium usually extends basad to the humerus, convex, discal surface densely microfoveo- whereas the maculation of S. simulans is late. confined to the apical one-third of the each elytron. Male genitalia of the two species Male genitalia.—As in figures 854 and 855; are similar except that the apex of the me- median lobe slender (figure 854), apically dian lobe is truncate in S. cruentatus but slightly expanded; ventral valve semicir- acute in S. simulans. Little overlap occurs cular, apex produced, truncate; armature in the distribution of the two species, S. of internal sac consisting of two small cruentatus being found east of the 100th clusters of denticles at apical orifice, hinge meridian whereas S. simulans is known sclerites short, remainder of internal sac only from Arizona. The occasional totally sparsely lined with denticles; apex with black individual of S. cruentatus must be circular closure valve; lateral lobes bowed separated by male genitalia. Other Sennius (figure 855), cleft nearly to base, arms species can be separated by color pattern slender, apices expanded toward meson, and male genitalia. mesal faces densely setose. Size.—Body length 1.6–2.9 mm; width Sennius discolor (Horn) 1.2–2.2 mm. (Figures 856–860) Type depository.—MCZC (cruentatus, lectotype No. 3886, by Johnson 1968; depres- Bruchus discolor Horn 1873:326; sus, lectotype No. 25051, by Johnson Fall 1901:165; Fall and Cockerell 1968; nigrinus, lectotype No. 126, by John- 1907:200; Schaeffer 1907:292; Cush- son 1968); ZMUM (nictitans, holotype). man 1911:506; Pic 1913a:24; Zacher 1952:462. Sennius discolor: Bottimer 1968c:1026; Johnson 1968:1270, 1969c:54,55;

202 Johnson and Kingsolver 1973:43, consisting of two lateral rows of closely 1982:419; Udayagiri and Wadhi packed, elongate spicules extending two- 1989:102. fifths from base, middle of sac with two Mylabris discolor: Leng 1920:305. dense rows of minute denticles and numerous, randomly placed denticles, apex simi- Bruchus discopterus Fall 1910:167; Fall larly armed; closure valve circular; lateral 1912:322; Pic 1913a:24. lobes bowed (figure 860), cleft nearly to Sennius discopterus: Bottimer 1968c:1027; base, arms slender, moderately expanded Johnson 1968:1270. apically. Mylabris discopterus: Leng 1920:305. Size.—Body length 1.3–2.3 mm; width Bruchus infirmus Sharp 1885:481. 0.8–1.6 mm. Acanthoscelides infirmus: Blackwelder Type depository.—ICCM (discolor, Ulke Col- 1946:759. lection); MCZC (discopterus, holotype No. 25052); BMNH (infirmus); MNHP (mana- Bruchus managuanus Pic 1935:66. guanus). Acanthoscelides managuanus: Blackwelder Type locality.—Texas (discolor); Southern 1946:760. California, Elsinore (discopterus); Guate- Color.—Integument mostly black but each mala (infirmus); Nicaragua (managuanus). elytron largely reddish orange with margin- Distribution.—AZ, AR, CA, NM, TX; Mexico al, basal, and sutural borders black (figure to Nicaragua. 857); legs reddish orange, antenna reddish brown to brown, pygidium black. Dorsal Host plants.—Senna lindheimeriana, S. vestiture of dense, intermixed white and roemeriana, S. wislizeni wislizeni; “Cassia golden setae; venter of body with dense sp.” Cushman (1911) reported this spe- white pubescence; pygidium white with cies from Prosopis sp. and Parkinsonia sp. evanescent basal triangle. but these records have not been verified by subsequent rearings. Structure.—Vertex and frons strongly convex, densely punctulate, impunctate Natural enemies.—None recorded. frontal line; ocular index 3:1; ocular sinus Immatures.—Not described. two-thirds length of eye; pronotum subconical (figure 856), lateral margins nearly Discussion.—This species most closely straight; disk strongly convex and densely resembles S. morosus among U.S. species punctulate; cervical sulcus short, deep. of Sennius. The metafemoral denticle of Scutellum quadrate, emarginate. Each ely- S. morosus is usually as long as or longer tron convex but slightly depressed around than width of the tibia at base, but this is scutellum (figure 856); striae deep, narrow; difficult to measure. Male genitalic charac- interstices minutely imbricate; metafemo- ters are the most reliable (compare figures ral denticle acute (figure 858), length equal 859 and 889). No U.S. species other than to tibial width at base; metatibia with these two has the red elytral patch cover- ventrolateral carina lacking; mucro, lateral ing nearly the entire surface. denticle, and coronal denticles subequal in length. Sennius fallax (Boheman) Male genitalia.—As in figures 859 and 860; (Figures 861–866) median lobe slender (figure 859), slightly Bruchus fallax Boheman 1839:59; Pic expanded apically, dorsal valve semicir- 1913a:25; Zacher 1952:462. cular, ventral valve subtriangular, lateral Acanthoscelides fallax: Blackwelder margins incurved, apex blunt; apical orifice 1946:759. lined with fine denticles; hinge sclerites strongly arcuate, internal sac armature

203 Sennius fallax: Johnson and Kingsolver ventral valve triangular, lateral margins 1973:54, 1982:419; Udayagiri and Wa- not arcuate, apex slightly produced; hinge dhi 1989:102. sclerites massive, sinuate; middle of sac Bruchus xanthopus Suffrian 1870:156; Pic with paired clusters of fine, elongate spic- 1913a:57. ules, middle of sac and lateral pockets with scattered fine denticles; closure valve cir- Acanthoscelides xanthopus: Blackwelder cular; lateral lobes cleft nearly to base (fig- 1946:759; Zacher 1952:465. ure 866), arms slender, apices spatulate, Sennius xanthopus: Bottimer 1961:295, with mesal expansion, mesal faces setose. 1968c:1027,1039. Size.—Body length 1.3–2.5 mm; width Bruchus probus Sharp 1886:481; Pic 0.8–1.7 mm. 1913a:43. Type depository.—NHRS (fallax, californi- Acanthoscelides probus: Blackwelder cus); Cuba, Havana, Academia de Ciencias 1946:760. de Cuba (xanthopus); BMNH (probus). Bruchus californicus Boheman 1859:114; Pic 1913a:19; Kingsolver 1979a:342. Type locality.—Jamaica (fallax); Cuba (xanthopus); Guatemala (probus); California Acanthoscelides californicus: Bottimer (californicus). 1968c:1017. Color.—Integument dark red to black, each Distribution.—FL, GA; Mexico to Panama, elytron usually with some red integument West Indies. varying from faintly red in basal areas to Host plants.—Senna bicapsularis bicap-su- the entire elytron red to reddish orange laris, S. biflora, S. obtusifolia, S. occiden- (figure 862); head and pronotum black; talis. Johnson and Kingsolver (1973) and antenna and legs red; pygidium dark red Johnson (1977b) record eight additional to black. Vestiture dense, intermixed white hosts from Mexico and Central America. and brown setae on pronotum; elytra faint- Natural enemies.—None recorded. ly striped in most specimens; pygidium with three basal white patches (figure 863); Immatures.—Not described. venter of body white. Discussion.—Sennius fallax is also wide- Structure.—Vertex and frons densely re- spread in the Neotropics and is variable in ticulate-punctate, frons with median line; integument color. Specimens found from ocular index 3:1; ocular sinus one-half Mexico southward on the mainland are length of eye; antenna reaching middle usually smaller with the pronotum and of metepisternum. Pronotum bell-shaped elytra entirely black, whereas those speci- (figure 861), lateral margins straight in mens from the West Indies and southern basal one-half, apex evenly arcuate; disk United States are larger with reddish- evenly convex and densely microfoveolate; brown pronotum and elytra usually with a basal lobe sulcate; lateral carina not vis- red stripe of varying width on each elytron. ible; cervical sulcus distinct, deep. Scutel- Male genitalia of the two forms are identi- lum transverse, densely pubescent. Elytral cal. margins convex (figure 861), striae parallel, Bottimer (1961) discussed the life history deep, narrow, strial punctures fine; inter- of this species under the name Sennius stices flat, sparsely punctate and minutely xanthopus. imbricate. Metafemoral denticle minute (figure 864); ventrolateral carina of metatibia evanescent; pygidial disk foveolate- punctulate. Male genitalia.—As in figures 865 and 866; median lobe moderately broad (figure 865);

204 Sennius lebasi (Fahraeus) deep, continuous ventrally. Scutellum quadrate, emarginate, densely pubescent. (Figures 867–873) Elytral striae shallow (figure 867), relatively Bruchus lebasi Fahraeus 1839:25; Pic broad, strial punctures slightly encroach- 1913a:30. ing on interstitial borders; interstices flat, Acanthoscelides lebasi: Blackwelder finely imbricate; metafemoral denticle 1946:760. about one-third width of tibial base (figure Sennius lebasi: Kingsolver 1979a:342; 871); ventrolateral carina of metatibia lack- Johnson and Kingsolver 1982; Udaya- ing; pygidium shallowly micropunctate. giri and Wadhi 1989:104. Male genitalia.—As in figures 872 and 873; Bruchus celatus Sharp 1885:499; Pic moderately broad; ventral valve broad with 1913a:20. lateral margins sinuate (figure 872); hinge sclerites strongly arcuate, basal one-half of Acanthoscelides celatus: Blackwelder internal sac densely lined with long spic- 1946:759. ules, rows of slender spines in apical one- Sennius celatus: Bottimer 1961:294, half; closure valve circular; lateral lobes 1968c:1026,1039; Moldenke 1971:108. bowed (figure 873), cleft nearly to base, Bruchus rufescens Motschulsky 1874:222; expanded mesad in apical one-half, mesal Blackwelder 1946:761. faces densely setose. Sennius rufescens: Kingsolver 1979a:342. Size.—Body length 1.5–2.9 mm; width Acanthoscelides rufescens: Blackwelder 0.9–1.9 mm. 1946:761. Type depository.—NHRS (lebasi); BMNH Color.—Body reddish yellow to black, most (celatus); ZMUM (rufescens). often dark red; head black, pronotum red Type locality.—Colombia, “Carthagena” (le- to black, often with paler red median line; basi); Panama, Bugaba (celatus); “Colum- elytra reddish yellow to dark brown, each bia” (rufescens). elytron with broad, oblique, paler band usually extending from side to side, sepa- Distribution.—TX; Mexico to Colombia, Bra- rated from base by narrow black border zil, Trinidad. extending briefly along suture, apical one- Host plants.—Senna bicapsularis bicap-su- half dark brown with light brown median laris. Six additional host plants are record- macula, apical border usually black (fig- ed from Mexico south (Waterworth 1986). ure 868); antennae and legs red, sometimes with base of metafemur dark brown; Natural enemies.—None recorded. sternal areas dark brown to black except Immatures.—Not described. terminal abdominal segments reddish; Discussion.—The pygidium of this species pygidium varying from uniformly reddish is red or reddish brown, usually with dark- yellow to a slightly darker form with paired er, arcuate maculae. The elytral pattern is dark brown maculae. Vestiture inconspicu- variable. In some specimens, the color is ous, sparse yellow on pronotum, elytra, nearly all yellowish red; but variation rang- and abdomen, white on thoracic sterna es from that to dark brown or even black, and scutellum; pygidium yellow, vestiture darker specimens with a chevron-shaped condensed in basal one-half and in median transverse band across the elytra slightly stripe. above the middle. The pronotum is usually Structure.—Vertex and frons strongly con- much darker than the elytra. vex, reticulate-punctate; frontal carina distinct; ocular index 3.25:1; ocular sinus six-tenths as long as eye; pronotum strongly convex (figure 867); disk densely foveolate-punctate; cervical sulcus narrow,

205 Bottimer (1961) briefly discussed the life strongly curved; basal one-half of internal history of this species in Texas under the sac with massive cluster of elongate spic- name Sennius celatus. Johnson (1977b) ules, lateral pockets of apical one-half lined listed host plants also under the name with spicules, remainder of sac with scat- Sennius rufescens. tered spicules. Lateral lobes cleft nearly to base (figure 879), arms slender, apices Sennius leucostauros Johnson and slightly expanded mesad; mesal faces Kingsolver densely setose. (Figures 874–879) Size.—Body length 1.8–2.3 mm; width 1.1–1.7 mm. Sennius leucostauros Johnson and Kingsolver 1973:75; Johnson Type depository.—USNM, holotype No. 1977b:128,130; Johnson and King- 71397. solver 1982:419; Udayagiri and Wadhi Type locality.—Mexico, Oaxaca, Temascal. 1989:419. Distribution.—TX; Mexico to Guatemala. Color.—Integument black; antenna reddish brown, fore legs and mid legs reddish Host plants.—Senna bicapsularis bicapsu- brown, metafemoral base black, apical laris. one-third to one-half red. Vestiture white Natural enemies.—None recorded. and dark brown; pronotum with median Immatures.—Not described. stripe and lateral patches white; scutellum white; elytra black, dorsal setal pattern Discussion.—No other U.S. species has a of brief basal patches, sutural stripe, and white, cross-shaped pattern on a black ely- transverse median band white, remainder tral integument. Johnson and King-solver of each elytron with dark brown vestiture (1973) placed Sennius leucostauros in the (figure 875); pygidium with median stripe abbreviatus species group, but its appear- and lateral patches white (figure 876); ven- ance is more similar to Sennius bondari ter of body uniformly white. (Pic) from South America than to other members of the abbreviatus group. Structure.—Vertex and frons microfoveate, frons with impunctate median line; Sennius medialis (Sharp) ocular index 3:1; ocular sinus three-fifths (Figures 880–884) as long as eye; antenna short, extending only to humeri. Pronotum bell-shaped Bruchus medialis Sharp 1885:470; Pic (figure 874), evenly convex, disk densely 1913a:34. punctulate with scattered foveae; cervical Acanthoscelides medialis: Blackwelder sulcus short, deep, not continuous across 1946:760. pro-sternum. Scutellum transverse, apex Sennius medialis: Johnson and King- emarginate. Elytra striae consisting of solver 1973:78; Center and Johnson lightly impressed rows of elongate punc- 1973:670; Johnson and Kingsolver tures at base, gradually becoming deep 1982:419; Udayagiri and Wadhi and narrow toward apex; interstices finely, 1989:105. densely imbricate. Metafemoral denticle Bruchus auctus Fall 1910:166. equal to width of tibia at base (figure 877); ventrolateral carina of metatibia absent; Mylabris auctus: Leng 1920:305. mucro as long as lateral denticle. Pygidial Sennius auctus: Johnson 1968:1270; Bot- disk microfoveate, foveae elliptical, crowd- timer 1968c:1026. ed, sometimes coalescing. Color.—Integument mostly black; elytra Male genitalia.—As in figures 878 and 879; with broad, reddish-orange, transverse median lobe moderately broad (figure 878); ventral valve broadly triangular, lateral margins straight; hinge sclerites large,

206 band continuous from margin to margin Type depository.—BMNH (medialis); MCZC and centered about one-third from base, (auctus, lectotype No. 325046, by Johnson sometimes with narrow, black, sutural 1968). stripe (figure 881); antennae usually red- Type locality.—Mexico, Guanajuato (me- dish orange or reddish brown; legs red dialis); Santa Rita Mountains, 8,000 feet except base of each metafemur sometimes (auctus). black. Vestiture yellow, dark brown, and white; pronotum mostly yellow with indis- Distribution.—AZ; Mexico to Guatemala. tinct white basal patches. Scutellum white. Host plants.—Senna hirsuta leptocarpa. Each elytron with irregular, white band Johnson and Kingsolver (1976) list also along basal margin, more conspicuous on Cassia tomentosa (now Senna hirsuta hir- 5th and 6th interstices; black elytral por- suta) from Mexico. tions with dark brown setae, orange band Natural enemies.—Urosigalphus bruchi- with yellow setae; pygidium and ventral vorus; Horismenus missouriensis. areas white. Immatures.—Center and Johnson Structure.—Vertex and frons with coarse, 1973:670 (egg). elongate punctures, setae directed mesad, frons with broad, impunctate ridge; ocular Discussion.—Center and Johnson (1973) index 3:1; ocular sinus one-half length of described oviposition and larval feeding in eye; antenna extending to humerus; pro- Senna hirsuta leptocarpa. Eggs have two notum nearly semicircular (figure 880), elongate anchoring strands at each end. disk densely punctulate and foveolate, Larvae develop one to a seed, devouring it foveae sparse along midline; cervical sulci almost completely, and pupation occurs continuous across prosternum. Scutellum inside the intact seed coat. It does not quadrate, emarginate. Elytral striae shal- form a cocoon, and the adult does not bore low, punctures deep (figure 880); inter- through the pod valve but escapes when stices flat, coarsely imbricate; metafemoral the valves dehisce. denticle about two-thirds as long as width This species most closely resembles Sen- of tibial base (figure 882), sometimes ser- nius abbreviatus among the U.S. species, rate on distal margin; ventrolateral carina but the hind legs of S. abbreviatus are of metatibia absent, mucro and lateral black, not red. denticle subequal in length. Pygidial disk densely set with shallow, round varioles. Sennius morosus (Sharp) Male genitalia.—As in figures 883 and 884; (Figures 885–890) median lobe moderately broad (figure 883); ventral valve subtriangular, lateral margins Bruchus morosus Sharp 1885:467; Pic slightly sinuate; hinge sclerites slender, 1913a:36. moderately arcuate; basal one-half of sac Acanthoscelides morosus: Blackwelder with small, paired clusters; apical one-half 1946:760. with scattered rows and small clusters of Sennius morosus: Johnson and King- minute denticles; closure valve large, cir- solver 1973:84; Center and Johnson cular; lateral lobes cleft three-fourths their 1976:669; Pfaffenberger and John- length (figure 884), arms long, slender, api- son 1976:38; Johnson and King- ces expanded mesad, mesal faces setose. solver 1982:419; Udayagiri and Wadhi Size.—Body length 1.6–2.4 mm; width 1989:105. 1.2–1.7 mm. Bruchus discolor: Fall and Cockerell 1907:200 (not Horn 1873). Color.—Integument mostly black, each elytron with large, subelliptical reddish-

207 orange macula extending from humerus Size.—Body length 1.3–2.7 mm; width nearly to apex and mesad to 2nd stria, 0.9–1.7 mm. bordered basally, laterally, and apically Type depository.—BMNH. with black (figure 886); basal four segments of antenna yellow, apical segments Type locality.—Mexico, Jalapa. brown to black; fore legs and mid legs Distribution.—AZ, NM, TX; Mexico to Pana- reddish brown, metafemur red with basal ma. one-fourth black, tibia and tarsus red. Host plants.—Senna bauhinioides, S. cove- Vestiture white, sparse on dorsum, dense sii, S. durangensis durangensis, S. hirsuta on sides of abdomen and on pygidium; 3rd leptocarpa, S. obtusifolia, S. roemeriana. and 5th striae with elongate, white, basal Senna occidentalis is reported from Mexico. patches. Natural enemies.—Horismenus missourien- Structure.—Vertex reticulate-foveolate, sis. foveae coarser on frons bordering broad, impunctate ridge; ocular index 2.5:1; Immatures.—Pfaffenberger and Johnson ocular sinus three-fifths as long as eye; 1976:38 (first larval instar); Center and antenna extending to humerus. Pronotum Johnson 1973:669 (egg). bell-shaped (figure 885), lateral margins Discussion.—Sennius morosus is similar to straight; disk convex, densely, evenly fo- S. discolor but can be separated from it by veolate, intervals micropunctate; cervical the larger metafemoral denticle and details sulcus nearly hidden by vestiture, continu- of the male genitalia (compare figures 859 ous across prosternum. Scutellum quad- and 889). rate, densely pubescent. Elytra convex (figure 885); strial punctures deep, round, Center and Johnson (1973) discussed the slightly encroaching on interstitial margins life history of this species in Arizona. Eggs in basal one-third but becoming more nar- are anchored by several strands to the row toward apex; interstices densely, finely substrate along the line of pod dehiscence. imbricate; 3rd, 4th, and 5th striae some- Larvae burrow into a seed and, after de- times with small basal denticles; length vouring the contents, move to the adjacent of femoral denticle equal to width of tibial seed in the pod. As many as eight seeds base, sometimes minutely serrate on distal may by eaten by a larva, but three or four margin (figure 888); ventrolateral carina are usual. Excavated seeds are glued to- of metatibia apparent only in basal one- gether by the larva, and the resultant clus- fourth; mucro subequal to lateral denticle. ter of seeds remains attached to the pod Pygidial sculpture of intermixed foveolae valve even after the pod has dehisced. Pu- and minute punctures. pation occurs within the cluster of seeds, but a cocoon is not spun. This species is Male genitalia.—As in figures 889 and 890; apparently bivoltine, according to Center median lobe moderately broad (figure 889); and Johnson (1976). lateral margins of ventral valve basally incurved, then angulate to apex; hinge scler- Sennius obesulus (Sharp) ites strongly arcuate, middle of sac with elongate mass of small, flat, ovate sclerites (Figures 891–895) flanked by masses of small denticles, api- Bruchus obesulus Sharp 1885:468; Pic cal one-third of sac lined with small den- 1913a:37. ticles; closure valve large, circular; lateral Acanthoscelides obesulus: Blackwelder lobes cleft four-fifths to base (figure 890), 1946:760. arms elongate, slender, apices expanded toward meson, setose on mesal faces. Sennius obesulus: Johnson and Kingsolver 1973:88; Johnson 1977b:130; Udaya- giri and Wadhi 1989:106.

208 Color.—Integument mostly black; each Distribution.— AZ; Guatemala, El Salvador, elytron with red hemispherical maculation Costa Rica. not attaining humerus, sometimes appear- Host plants.—Chamaecrista serpens ing as a subapical lateral ovate spot (figure wrightii. 892); basal four or five antennal segments reddish orange, apical segments dark Natural enemies.—None recorded. brown to black; fore leg and middle tibia Immatures.—Not described. reddish brown, middle femur and hind leg Discussion.—Inclusion of Sennius obesulus black. Vestiture white, everywhere dense, in this handbook is based on Johnson’s sometimes concealing sculpture. record (1977b) from Arizona, Santa Cruz Structure.—Vertex and frons convex, dense- County, near the border with Mexico. ly, evenly punctate, with short median This species is closely similar to Sennius impunctate line dorsad of fronto-clypeal simulans (Schaeffer) (see below), differ- suture; ocular index 3:1; ocular sinus one- ing only in the following (characters of S. half length of eye; antenna short, extending simulans in brackets): elytral maculation to humerus, club segments not markedly red, hemispherical, not attaining humer- eccentric. Pronotum nearly semicircular us, sometimes appearing as a subapical, in outline, moderately convex, disk evenly lateral, ovate spot (figure 892) red elytral punctulate, each puncture bearing short spot apical, nearly spanning width of apex, white seta; lateral pronotal carina vaguely occasionally only as a small red, apical traceable near posterior angle; cervical spot figure 897; armature of internal sac sulcus absent; prosternum short, triangu- densely lined with blunt denticles (figure lar, procoxal apices contiguous. Scutellum 894) internal sac with fewer blunt denticles rectangular, slightly wider than long, bi- figure 899. dentate. Elytra (figure 891) together as long as wide, broadest behind humeri; slightly Further study is needed to assess the cor- depressed behind scutellum; striae regular rect application of these two names. If they in course, shallow, strial punctures evenly prove to be synonymous, S. obesulus has spaced; interstices flat, densely punctate. priority. Hind leg as in figure 893; tibial coronal denticles fine; lateral carina complete, ven- Sennius simulans (Schaeffer) trolateral carina extending one-half dis- (Figures 896–900) tance to apex. Pygidium apically reflexed in male; disk finely, shallowly foveolate. Bruchus simulans Schaeffer 1907:296; Fall 1910:164; Pic 1913a:50. Male genitalia.—As in figures 894 and 895; median lobe slender (figure 894); ventral Mylabris simulans: Leng 1920:305. valve ogival, lateral margins sinuate, apex Sennius simulans: Johnson 1968:1270; acutely produced; hinge sclerites moder- Bottimer 1968c:1027; Johnson and ately long, slightly arcuate; internal sac Kingsolver 1973:93, 1982:419; Udaya- densely lined with mixed blunt and acute giri and Wadhi 1989:107. denticles; lateral lobes elongate (figure Color.—Integument mostly black; each 895), slender, slightly bowed, cleft about elytron with reddish-orange apical macula- two-thirds their length. tion bordered with black and not extending Size.—Body length 1.7–2.6 mm; width anteriad beyond middle of elytron (figure 1.1–1.8 mm. 897); basal four antennal segments yellow, apical segments black. Fore legs brown, Type depository.—BMNH. mid legs dark brown to black, hind legs Type locality.—Guatemala, “near the city always black. Vestiture white, everywhere Capetillo.” dense, sometimes concealing sculpture.

209 Structure.—Vertex punctulate, punctures spot on each elytron and totally black hind separated by three diameters of a punc- legs, but see the discussion of S. obesulus. ture, frons more coarsely punctate flank- Center and Johnson (1973) briefly dis- ing impunctate median ridge; ocular index cussed the aspects of the life history of S. 3:1; ocular sinus three-fourths length of simulans. Eggs are glued to the pod wall eye; antenna short, extending to humerus. as usual but with anchoring strands simi- Pronotum nearly semicircular (figure 896), lar to those of Sennius morosus. Larvae lateral margins arcuate; disk convex; sur- consume several seeds within a pod and face sculpture intermixed fine punctures pupate within the feeding cavity. Seeds and elliptical foveolae; lateral carina ridge- are apparently not glued together as in like; cervical sulcus absent, proster-nal the case of S. morosus, and no cocoon sulcus complete between cervical bosses. is formed. Pods attacked by S. simulans Scutellum quadrate, apically emarginate. dehisce only slightly, not widely as with Elytra (figure 896) slightly depressed along normal pods. Adults escape through the suture; striae shallow, strial punctures dehiscing gap but do not bore through the round, variolate, encroaching on margins pod wall. of interstices; interstices punctate-imbricate. Length of metafemoral denticle about Sennius whitei Johnson and Kingsolver two-thirds length of tibial width at base (figure 898); ventrolateral carina of metati- (Figures 901–907) bia obsolete in apical one-third. Pygidium Sennius whitei Johnson and Kingsolver densely, finely foveolate. 1973:99; Johnson and Kingsolver Male genitalia.—As in figures 899 and 1982:420; Udayagiri and Wadhi 900; median lobe of uniform width (figure 1989:107. 899); lateral margins of ventral valve nearly Color.—Body mostly black; each elytron straight, apex narrowly produced; hinge with subbasal, marginal, red macula ex- sclerites lunate, massive; internal sac rela- tending mesad to 3rd stria, apicad to tively short, without lateral apical pockets, middle of elytron (figure 902); fore leg, mid sac lined with blunt denticles; lateral lobes leg, and basal three antennal segments cleft two-thirds their length (figure 900), reddish yellow, remaining segments black; arms slender, expanded mesad, mesal hind legs black. Vestiture black, white, and faces setose. yellow; head, lateral portions of pronotum, Size.—Body length 1.7–2.6 mm; width sutural stripe, pygidium, and venter of 1.1–1.8 mm. body white, elytral vestiture yellow from 2nd stria to margin; middle one-third of Type depository.—USNM, holotype pronotum with inconspicuous black setae. No.42339. Structure.—Vertex and frons densely foveo- Type locality.—Arizona, Huachuca Moun- late, foveolae discrete, tending to be elon- tains. gate either side of distinct frontal carina Distribution.—AZ, CA; Mexico. (figure 903); ocular index 4:1; ocular sinus Host plants.—Chamaecrista nictitans men- three-fourths length of eye; antenna (figure salis (as Cassia leptadenia in literature). 904) atypical for genus, subserrate from 3rd segment, extending to middle of me- Natural enemies.—None recorded. tepisternum. Pronotum bell-shaped (figure Immatures.—Center and Johnson 1973 901), lateral margins nearly straight; disk (egg). convex except flattened on basal lobe; surface densely set with intermixed elliptical Discussion.—Adults of S. simulans usu- ovarioles and fine punctures; cervical sulci ally can be recognized by the subapical red connected across prosternum. Scutellum quadrate, apex

210 shallowly emarginate. Elytra (figure 901) Genus Stator Bridwell depressed along suture, striae shallow, Stator Bridwell 1946:55; Bradley strial punctures deep, elongate, encroach- 1947:39; Blackwelder and Black- ing on margins of interstices, basal foveae welder 1948:45; Johnson 1963:860; rounded; interstices flat or slightly convex, Teran 1967:308,316; Bottimer densely, finely imbricate; metafemoral den- 1968c:1027,1039,1041; Johnson ticle minute (figure 905); metatibial carina 1968:1270; Kingsolver 1972a:219; evanescent; mucro shorter than lateral Johnson and Kingsolver 1973:1,5, denticle. Pygidium densely, finely variolate, 1976:2; Borowiec 1987:99; John- interspaces finely punctate. son et al. 1989:7; Udayagiri and Wa- Male genitalia.—As in figures 906 and 907; dhi 1989:108; Johnson and Siemens median lobe slender (figure 906); lateral 1995a. Type species: Bruchus pruininus margins of ventral valve sinuate, apex Horn, by original designation. produced slightly; hinge sclerites elongate, Frons carinate or with finely punctate line; moderately arcuate, internal sac lined antenna not dimorphic, club segments with blunt denticles in basal one-half, with slightly eccentric, subserrate from 5th seg- acute denticles in apical one-half, closure ment, terminal segment elliptical, extend- valve large, circular, lateral pockets ab- ing to humerus; eyes protruding laterally, sent; lateral lobes cleft about one-half their separated from side of head (except Stator length (figure 907), apices bowed mesad, subaeneus); antenna not sexually dimor- setose on mesal faces. phic, terminal segment ovate. Prothorax Size.—Body length 2.5–2.9 mm; width semicircular in dorsal aspect (figure 908), 1.7–1.9 mm. convex, lacking asperities; lateral margin Type depository.—USNM, holotype No. sharply carinate, carina extending from 71391. posterolateral corner of pronotum nearly to coxal cavity; procoxae contiguous api- Type locality.—Mexico, Nayarit, 23 mi cally (except Stator beali); cervical sulcus south of Tepic. absent; cervical boss bisetose. Scutellum Distribution.—AZ; Mexico. quadrate, slightly concave; mesepi-meron reduced to triangular sclerite. Elytra sub- Host plants.—Unknown. quadrate (figure 908), slightly depressed Natural enemies.—None recorded. on disk; striae not distorted, subparallel, Immatures.—Not described. lacking basal asperities, strial punctures distinct; interstices of uniform width, flat, Discussion.—This species resembles S. cru- finely punctate-imbricate. Metacoxae en- entatus and was placed in the cruen-tatus tirely punctate (figure 956) or with median group by Johnson and Kingsolver (1973). one-half smooth (figure 949); metafemur The red elytral spot of S. whitei, however, with ventral face channeled, ventrolateral is confined to the anterior one-half (figure and ventromesal margins carinate, ven- 902), the frontal carina is sharp and dis- tromesal carina with acuminate, subapical tinct (figure 903), the median lobe is more denticle; ventrolateral carina slightly sinu- slender, lateral lobes are more shallowly ate, or emarginate subapically with blunt cleft, and the metafemoral subapical denti- angulation; meta-tibia gradually dilated to- cle is much smaller than for S. cruentatus. ward apex; mucro usually short, subequal Nothing is known of the life history of this to lateral denticle; ventrolateral carina species. sometimes evanescent apically, lateral, ventral, and dorsomesal carina strong and complete; some species with longitudinal dorsal fossa (figure 911), coronal denticles two or three. Abdominal sterna unmodified except male 5th sternum

211 usually deeply emarginate; pygidium con- Biology.—Johnson 1967, 1984b. vex, lacking asperities. Keys.—Johnson 1963; Johnson and King- Median lobe of male genitalia with one to solver 1976 (North America); Pfaffenberger several spines or spicules; lateral lobes 1981 (first larval instar); Johnson et al. deeply cleft, spatulate apically. 1989 (South America).

Key to Species of Stator of United States 1 Metacoxal face entirely punctate (figure 956)...... 2 6(5) Metafemur and antenna all red...... bottimeri Kingsolver Metacoxal face with most of medial one-half smooth (fig- Metafemur and antenna partly black ure 949); lateral one-half and small patch near trochan- ...... vachelliae Bottimer teral fossa punctate...... 7 7(1) Metafemur mostly black with no more than apical one- 2(1) Elytra entirely red, or black with red maculae...... 3 tenth reddish; elytra with intermixed white and brown Elytra entirely black...... 4 setae, without denser patches of white setae ...... pygidialis (Schaeffer) 3(2) Vestiture of pronotum and elytra uniformly yellow; black maculation of elytra extending from margin to 5th or 6th Metafemur with apical two-tenths or more dark red; elytra stria; prosternum separating procoxae for entire length with intermixed reddish-brown and brown setae, without ...... beali Johnson denser patches or rows of white setae...... 8 Vestiture of pronotum and elytra uniformly white; black 8(7) Male genitalia with large, curved, saddle-shaped spine maculation, if present, extending only to 7th stria; procox- (figure 951); apex of ventral valve arcuate ae contiguous at apices...... limbatus (Horn) ...... sordidus (Horn) 4(2) Eye flattened, nearly contiguous with lateral margin of Male genitalia either with large, curved spine or small, head...... subaeneus (Schaeffer) tapered spine (figures 912 and 932); apex of ventral valve emarginate...... 9 Eye not flattened, posterior margin protruding from side of head...... 5 9(8) Male genitalia with large spine; apex of ventral valve narrowly emarginate (figure 924)...... 5(4) Metafemur black and red, usually with apex red, or all red...... chihuahua Johnson and Kingsolver ...... 6 Male genitalia with small, tapered spine (figure 926); Metafemur entirely black...... pruininus (Horn) ventral valve broadly emargi-nate ...... coconino Johnson and Kingsolver

Stator beali Johnson segments orange to brown; fore leg and mid leg reddish yellow, hind leg usually (Figures 908–914) black, sometimes with tibial apex dark red. Stator beali Johnson 1963:861; Bottimer Vestiture yellow, sparsely distributed on 1968c:1027,1039; Johnson 1968:1270; dorsum of body except pronotum with me- Johnson and Kingsolver 1976:16, dian stripe and lateral patches; scutellum 1982:420; Johnson et al. 1989:9; Uday- white; pygidium with median basal triangle agiri and Wadhi 1989:109; Nilsson and and lateral patches yellow, more prominent Johnson 1993b; Fox and Mousseau in female (figures 909 and 910); ventral 1995. vestiture white. Color.—Body black except each elytron Structure.—Frontal carina usually a gla- mostly red with black basal and sutural brous ridge; ocular index 3:1; ocular sinus borders and lateral macula extended medi- one-half length of eye. Pronotal disk finely ally to 5th or 6th stria; antenna with basal punctate (figure 908). Metacoxal face five segments yellow or orange, apical 212 uniformly punctate; ventrolateral meta- Stator bottimeri Kingsolver femoral carina subapically sinuate (fig- (Figures 915–920) ure 911); metatibial carinae all complete, dorsal face with elongate fossa, sometimes Stator bottimeri Kingsolver 1972a:225; difficult to see. Johnson and Kingsolver 1976:17, 1982:420; Johnson et al. 1989:9; Male genitalia.—As in figures 912, 913, Udayagiri and Wadhi 1989:109; Alvarez and 914; ventral valve triangular (figure Marin and Kingsolver 1997:220. 912); internal sac with one slender median sclerite, and one apical, helmet-shaped Color.—Body black with bronzy sheen; an- sclerite; lateral lobes with arms slender at tenna, fore leg, and mid leg reddish brown base (figure 913), apices expanded toward to yellow, hind leg red except trochanter meson, cleft two-thirds their length; spicu- black. Vestitural pattern of pronotum and lum gastrale as in figure 914. elytra with black, intermixed brown and white, and white setae (figure 915); pygid- Size.—Body length 2.1–2.8 mm; width ial pattern of male intermixed brown and 1.6–2.1 mm. white setae with pair of slightly depressed, Type depository.—USNM, holotype No. black, median patches (figure 916), that 67419. of female black with lateral basal white Type locality.—Texas, Cameron Co., patches and scattered apical white setae Brownsville. (figure 917); venter of body densely white. Distribution.—TX; Mexico. Structure.—Frontal carina evanescent; ocular index 4:1; ocular sinus two-thirds Host plants.—Pithecellobium ebano (com- length of eye. Metacoxal face uniformly mon name, Texas ebony). punctate; metafemur with ventrolateral Natural enemies.—None recorded. carina sinuate (figure 918); ventrolateral carina of metatibia lacking, dorsal sulcus Immatures.—Not described. well-defined. Discussion.—This species closely resembles Male genitalia.—As in figures 919 and 920; Stator limbatus but differs in the yellow ventral valve with lateral margins arcuate dorsal vestiture, prosternum separating (figure 919), apex bluntly rounded; internal the procoxae, the lateral elytral macula sac armed with 18–20 broad, flat, bladelike reaching 5th or 6th stria, and shape of the spines; lateral lobes as in figure 920. spiculum gastrale (compare figures 914 and 934). (See S. limbatus discussion.) Size.—Body length 2.1–2.2 mm; width 1.5–1.7 mm. Bridwell, in an unpublished notebook from 1936, states, “While this species resembles Type depository.—USNM, holotype No. Stator limbatus, it affects only the seed of 70396. Texas ebony. Its eggs are laid upon the ripe Type locality.—Cuba, near Santiago. seeds within the pod, the ovipositor being inserted between the pod and seed, and as Distribution.—FL; West Indies. many as 25 eggs may be deposited at one Host plants.—Acacia farnesiana, A. pineto- time by a female on a single seed. Each rum. seed can furnish food for that many more Natural enemies.—None recorded. larvae if it is full sized.” Immatures.—None described. Nilsson and Johnson (1993b) discussed oviposition habits of S. beali, compared to Discussion.—This species is most similar to those of S. limbatus, and their attempts at Stator subaeneus among the U.S. species hybridization of these species. but is easily separated from it. The posterior margin of the eye is separated from the

213 head (partly contiguous in S. subaeneus), Type locality.—Arizona, Pima Co., Kitt the hind leg is entirely red (entirely black Peak. in S. subaeneus), and armature of the Distribution.—AZ; Mexico. internal sac is distinctive (compare figures 919 and 958). Host plants.—Acacia angustissima angustissima, A. constricta; Calliandra eriophylla, Stator chihuahua Johnson and Kingsolver C. humilis humilis; Lysiloma microphylla thornberi; Mimosa biuncifera, M. wherry- (Figures 921–925) ana. Stator chihuahua Johnson and Kingsolver Natural enemies.—Lariophagus texanus. 1976:23; Pfaffenberger 1981:255; John- son and Kingsolver 1982:420; Johnson Immatures.—Pfaffenberger 1981:255 (first et al. 1989:10; Udayagiri and Wadhi larval instar). 1989:110. Discussion.—This species is similar to Sta- Color.—Body black, legs dark red, basal five tor sordidus, S. coconino, S. pygidialis, and antennal segments reddish yellow, six api- S. vachelliae, differing principally in struc- cal segments black. Pronotum and elytra tures of the male genitalia. with intermixed coppery and white setae, evanescent white patch of setae at middle Stator coconino Johnson and Kingsolver of each 7th interstice (figure 921); male py- (Figures 926–927) gidial vestiture nearly uniformly white with scattered coppery setae near apex, basal Stator coconino Johnson and King- margin with three patches of white setae solver 1976:25, 1982:420; Johnson (figure 922); female pygidium black with et al. 1989:10; Udayagiri and Wadhi lateral basal white patches (figure 923); 1989:109. ventral vestiture mostly white with scat- Color.—Similar to Stator chihuahua but tered coppery setae on lateral portions of with base of metafemur black, the 7th abdominal sterna. elytral interstice lacking a white patch, and Structure.—Vertex and frons uniformly the apical one-half of the male pygidium punctulate, frontal carina absent; ocular nearly glabrous. index 4:1 in male, 2.4:1 in female; ocu- Structure.—Identical to Stator chihuahua lar sinus one-half length of eye; antenna except for male genitalia. reaching middle of metepisternum; medial Male genitalia.—Characters of the male one-half of metacoxal face impunctate and genitalia are distinctive (figures 926 and polished, lateral one-half punctulate; ven- 927); ventral valve is apically broadly trolateral carina of metafemur with obtuse emarginate, and the internal sac is armed angle opposite subapical denticle; metatib- only with a small median sclerite (figure ia lacking ventrolateral carina; dorsal fossa 926); lateral lobes broadly spatulate (figure absent. 927), mesally expanded, cleft two-thirds to Male genitalia.—As in figures 924 and 925; base, mesal faces finely setose. ventral valve with incurved lateral margins Size.—Similar to Stator chihuahua. Body and emarginate apex (figure 924); internal length 2.2 mm; width 1.6 mm. sac armed with large, saddle-shaped median sclerite, apical sclerite absent; lateral Type depository.—USNM, holotype No. lobes broadly spatulate (figure 925), cleft 71403. nearly to base; mesal faces finely setose. Type locality.—Arizona, Coconino Co., 8 mi Size.—Body length 1.6–2.2 mm; width north of Sedona. 1.1–1.6 mm. Distribution.—AZ. Type depository.—USNM, holotype No. Host plants.—Not known. 71402.

214 Natural enemies.—None recorded. Color.—Body black except each elytron Immatures.—Not described. usually red with black lateral macula, each macula with lobe sometimes extended Discussion.—See discussion of Stator chi- toward suture as far as 7th stria, varying huahua. from all red to a pattern of small basal and apical red maculae on black background; Stator limbatus (Horn) antennal color varying from four or five (Figures 928–934) basal segments yellow and remaining segments dark red, to all segments yel- Bruchus limbatus Horn 1873:326; low; fore leg and mid legs yellow. Vestiture Sharp 1885:454; Horn 1894:344; white, uniformly distributed over body, Fall 1901:29,160; Schaeffer pygidium sometimes with evanescent basal 1907:292,294,303; Fall 1910:165; patches. Cushman 1911:498,506; Bridwell 1920b:406,409, 1920c:332, 1921a:458, Structure.—Frontal carina short, some- 1921b:465, 1923c:262; Krauss times appearing as only an impunctate 1944:11. line; ocular index 3.75:1; ocular sinus Mylabris limbatus: Leng 1920:305; Kannan one-half length of eye; antenna extending 1923:25. to elytral humerus. Pronotum, scutellum, and elytra as for genus (figure 928). Meta- Acanthoscelides limbatus: Moreno and coxal face uniformly punctate; ventrolat- Bibby 1943:23; Blackwelder 1946:760; eral carina of metafemur sinuate; all four Zacher 1952:465,470,471. metatibial carina complete. Female pygid- Stator limbatus: Leech 1954:85 Swezey ium more elongated than in male (figures 1954; Leech 1959:60; Hinckley 929 and 930). 1960:261; Bibby 1961:325; John- Male genitalia.—As in figures 932, 933, son 1963:861; Bottimer 1968b:284, and 934; ventral valve triangular (figure 1968c:1027,1039; Johnson 1967:267, 932); internal sac with thornlike median 1968:1270, 1969c:55; Forister and sclerite and helmet-shaped, dentate api- Johnson 1970:84, 1971:231; Kingsolver cal sclerite; lateral lobes with arms slender 1972a:228; Johnson and Kingsolver (figure 933), apices mesally strongly ex- 1976:32; Pfaffenberger and John- panded, cleft two-thirds to base. Spiculum son 1976:39; Mitchell 1977:645,648 gastrale as in figure 934. (life history); Johnson and King- solver 1982:420; Kistler 1982:266; Size.—Body length 1.6–2.7 mm; width Stein 1983a,b; Johnson et al. 1989:8; 1.1–2.0 mm. Udayagiri and Wadhi 1989:110; Nils- Type depository.—MCZC (limbatus, lecto- son and Johnson 1993b:385; Johnson type No. 8196; BMNH (interruptus). 1995b:319. Type locality.—Lower California (limbatus); Bruchus interruptus Sharp 1885:470; Mexico and Guatemala (interruptus). Schaeffer 1907:297. Acanthoscelides interruptus: Blackwelder Distribution.—AZ, CA, TX, HI; Mexico to Ec- 1946:759. uador. Generally distributed in the Hawai- ian Islands (Stein 1983a). Bruchus cearanus Pic 1930c:12 Host plants.—Acacia acatlensis, A. angus- Acanthoscelides cearanus: Blackwelder tissima angustissima, A. baileyana, A. 1946:759 berlandieri, A. confusa, A. cultriformis, A. Stator cearanus: Kingsolver 1972a:225; farnesiana, A. greggi, A. koa, A. melano-xy- Johnson and Kingsolver 1976:38, lon, A. millefolia, A. retinodes, A. richei, A. 1982:420; Johnson et al. 1989:27; roemeriana, A. wrightii; Albizia lebbeck, A. Kingsolver and Silva 1991:412; John- saponaria; Arachis hypogaea; Caesalpinia son 1995b:319.

215 pulcherrima; Calliandra eriophylla, C. hu- Kistler (1982) studied the effects of temper- milis humilis, C. humilis reticulata; Cassia ature on biological processes of S. limbatus fistula, C. grandis, C. javanica indochinen- in Arizona. sis, C. javanica javanica; Delonix regia(?); Essig (1929a) gave the common name “lim- Desmanthus bicornutus; Erythrina sandwi- bate weevil” to this species. censis; Glycine max; Leucaena leucocephala, L. pulverulenta; Lysiloma microphylla Stator pruininus (Horn) thornberi; Neptunia plena; Parkinsonia (Figures 54, 935–939) aculeata, P. florida, P. micro-phylla, P. texana macra, P. texana texana; Pithecellobium Bruchus pruininus Horn 1873:327; Sharp brevifolium, P. dulce, P. ebano, P. pallens, P. 1885:453,472; Riley and Howard saman, P. unguis-cati; Prosopis glandulosa 1892c:165; Fall 1901:29,160; Fall glandulosa, P. glandulosa torreyana, P. juli- and Cockerell 1907:200; Schaeffer flora (exp.). 1907:292–297; Fall 1910:162–166; Natural enemies.—Microdontomerus an- Cushman 1911:507; Payne 1913:40; thonomi; Stenocorse bruchivora; Urosigal- Bridwell 1918:465–471, 1919:18, phus bruchivorus, U. neobruchi; Uscana 1920a:408; Swezey 1921:521; Bridwell semifumipennis. 1923a:79, 1923b:261; Swezey 1925:3, 1936:201; Bridwell 1938a:71; Swezey Immatures.—Kannan 1923 (egg and first 1954:80. larval instar); Pfaffenberger and Johnson Bruchus (Stator) pruininus: Zacher 1976 (first larval instar); Pfaffenberger 1952:462. 1981 (first instar). Mylabris pruininus: Leng 1920:305; Kan- Discussion.—This species has one of the nan 1923:25. most extensive host lists among the Bru- chidae. Waterworth (1986) lists 36 hosts Bruchidius pruininus: Herford 1935:15. taken from the literature. Acanthoscelides pruininus: Blackwelder Johnson, in his paper on bruchid guilds 1946:760. (1981a), places this species in his mature Stator pruininus: Bridwell 1946:55; Bradley seed guild, in which species oviposit only 1947:39; Zacher 1952:462,480; Hinck- on mature seeds on the plant. Mitchell ley 1960:261; Bibby 1961:325; Johnson (1977) reported that Stator limbatus en- 1963:862; Peck 1963:956; Bottimer tered seed pods of Cercidium floridum 1968b:1027,1039,1041, 1968c:285; through exit holes of another bruchid, Johnson 1967:267, 1968:1270; Bot- Mimosestes amicus (Horn). timer 1969b:1189,1191; Janzen 1969:14,19; Johnson 1969c:55; Jan- Mitchell detailed the oviposition strategy zen 1972:976; Kingsolver 1972a:221; and life history of S. limbatus, and Bridwell Center and Johnson 1976:196; John- (1923c) described oviposition through open son and Kingsolver 1976:44; Johnson valves of the seed pod. Nilsson and John- 1979a:124, 1981c:251; Pfaffenberger son (1993b) experimented with hybridiza- 1981:264; Johnson and Kingsolver tion of S. limbatus and S. beali. 1982:420; Kistler 1982:266; Johnson Among the species included in this hand- 1984b:88; Borowiec 1987:99; Luckow book, Stator limbatus and S. beali are the and John-son 1987:49; Johnson et only species with red elytra marked with al. 1989:10; Udayagiri and Wadhi black maculae. Other differentiating char- 1989:111. acters can be found in the key to species. Bruchus cognatus Sharp 1885:472. Acanthoscelides cognatus: Blackwelder 1946:759. Bruchus piger Sharp 1885:473.

216 Acanthoscelides piger: Blackwelder Desmodium uncinatum (exp.); Erythrina 1946:760. sandwicensis; Glycine max (exp.); Indigo- Color.—Body and appendages black except fera anil, I. suffruticosa; Leucaena leuco- 3rd to 5th basal antennal segments, fore cephala; Mimosa biuncifera, M. dysocarpa, leg and mid leg reddish yellow. Vestiture M. grahamii, M. laxiflora, M. wherryana; sparse, white, and uniformly distributed Neptunia plena; Olneya tesota; Parkinsonia over body. aculeata, P. texana macra, P. texana texana; Pithecellobium ebano, P. pallens, P. saman; Structure.—Frons with impunctate median Prosopis glandulosa glandulosa, P. juliflora line; ocular index 3:1; ocular sinus two- (exp.), P. velutina; Robinia pseudoacacia; thirds length of eye; antenna extending to Scaevola taccada sericea; Senna siamea elytral humerus. Pronotum, scutellum, and (exp.); Sesbania emerus, S. exaltata, S. elytra as for genus (figure 935). Metacoxal macrocarpa, S. sesban; Sophora chryso- face densely, uniformly punctate; ventro- phylla. lateral carina of metafemur sinuate; all four carinae of metatibia complete to apex, For Bridwell’s experimental rearings dorsal tibial fossa present. in Hawaii, see Johnson and Kingsolver (1976:50–51). For other host plants from Male genitalia.—As in figures 938 and 939; Mexico southwards, consult Johnson ventral valve obtusely angulate (figure and Kingsolver (1976:47–51), Johnson 938); internal sac armed with many spines (1984b:88), and Luckow and Johnson and sclerites of various sizes; lateral lobes (1987). broadly spatulate (figure 939), bowed, cleft three-fourths to base; mesal faces densely Natural enemies.—Anisopteromalus ca- setose. landrae; Charitopodinus swezeyi, C. terryi; Heterospilus prosopidis; Horismenus Size.—Body length 1.4–2.7 mm; width depressus, H. missouriensis; Lariophagus 0.9–2.0 mm. texanus; Pteromalus sp.; Uscana semifumi- Type depository.—MCZC (pruininus, lecto- pennis; Trichogrammatidae. type No. 3888); BMNH (cognatus, piger). Immatures.—Bridwell 1918:470 (eggs); Type locality.—Arizona (pruininus); Mexico Kannan 1923 (egg); Pfaffenberger and (cognatus, piger). Johnson 1976:40 (first instar); Pfaffenberg- Distribution.—AZ, CA, HI, NM, NV, OR, TX, er 1981:264 (first instar). UT; Mexico to Venezuela. The species is Discussion.—Stator pruininus is one of the generally distributed throughout the Ha- most easily recognized species in the genus waiian Islands (Stein 1983a). with its uniformly gray appearance and Host plants.—Hosts marked (exp.) are ex- reddish-yellow fore legs and mid legs. perimental host trials. Acacia berlandieri, This species evidently is adaptable to a A. californica, A. confusa, A. constricta, A. wide range of host plant situations. John- dealbata, A. decurrens (exp.), A. greggi, A. son (1981a) placed S. pruininus with S. koa, A. mearnsii, A. melanoxylon, A. neover- limbatus in his mature seed guild, a group nicosa, A. richei, A. rigidula, A. roemeriana, that oviposit on seeds in partly dehiscent A. schotti, A. vernicosa, A. wrightii; Albizia pods. Waterworth (1986) lists 38 host saponaria (exp.); Arachis hypogaea (exp.); plants for S. pruininus. Caesalpinia pulcherrima (exp.); Calliandra Kistler (1982) studied the effects of temper- eriophylla, C. humilis humilis, C. humilis ature on S. pruininus, and Johnson (1967) reticulata; Cassia fistula(exp.), C. grandis, published notes on its bionomics.Fullaway C. javanica indochi-nensis, C. javanica ja- and Krauss (1945) stated that females vanica; Coursetia microphylla; Desmanthus chew holes in koa pods and ovi- bicornutus, D. cooleyi, D. covillei, D. fruticosa, D. leptolobus, D. virgatus virgatus;

217 posit directly on the seeds. Stein (1983b) for genus (figure 940); lateral one-half of briefly detailed the biology of this bruchid. metacoxal face finely punctate, median Essig (1929a) gave the common name one-half glabrous and polished; ventro- “pruinose weevil” to this species. lateral carina of metatibia absent, dorsal coronal denticle nearly as long as mucro. Stator pygidialis (Schaeffer) Male genitalia.—As in figures 944 and 945; apex of ventral valve slightly truncated (Figures 940–945) (figure 944); internal sac armed with one Bruchus pygidialis Schaeffer 1907:297; large, saddle-shaped sclerite; lateral lobes Fall 1910:164,166. as illustrated (figure 945). Mylabris pygidialis: Leng 1920:305. Size.—Body length 1.9–2.7 mm; width Stator pygidialis: Johnson 1967:267; Bot- 1.3–1.9 mm. timer 1968c:1027,1039; Johnson Type depository.—USNM, lectotype No. 1968:1270; Bottimer 1973:545; Center 42340, by Johnson 1968. and Johnson 1976:197; Johnson and Kingsolver 1976:52; Pfaffenberger and Type locality.—Arizona, Huachuca, Moun- Johnson 1976:41; Johnson 1981a:251, tains. 1981d:241; Pfaffenberger 1981:264; Distribution.—AZ, NM, TX; Mexico to Ven- Johnson and Kingsolver 1982:420; ezuela. Johnson et al. 1989:10; Udayagiri and Wadhi 1989:112. Host plants.—Calliandra humilis humilis, C. humilis reticulata. Bruchus pythonicus Pic 1913a:43 (unnecessary new name for pygidialis Schaef- Natural enemies.—None recorded. fer). Immatures.—Pfaffenberger and Johnson Mylabris pythonicus: Leng 1920:305. 1976:41 (first larval instar); Pfaffenberger Stator pythonicus: Johnson 1963:864. 1981:264 (first larval instar). Acanthoscelides pythonicus: Johnson Discussion.—Johnson (1981d) investigated 1963:864. the bionomics of this species in Arizona. Color.—Body black, fore leg and mid leg Johnson and Kingsolver (1976) placed S. varying from dark red to black; hind leg pygidialis in the sordidus group of Stator. usually black, sometimes dark red with Characters to separate S. pygidialis from base of femur black; basal three or four other Stator species can be found in the antennal segments dark red to black, api- key. cal segments always black. Vestiture of Bottimer (1968c:1027) explained the rea- intermixed coppery and white setae on son for rejecting Bruchus pythonicus Pic pronotum, elytra, and lateral areas of ab- as a replacement name for pygidialis domen, remaining ventral areas sparsely Schaeffer. In his catalogue Pic (1913a) white; male pygidium with three white changed the spelling of Kytorhinus pygi- basal patches (figure 941), apex with mixed dalis Motschulsky to pygidialis and then coppery and white setae; female pygidium transferred the name to Bruchus, making similar to male except with two white basal Bruchus pygidialis Schaeffer a secondary patches (figure 942). homonym. Bottimer rejected Pic’s usage as Structure.—Vertex and frons densely an incorrect subsequent spelling thereby punctulate, frontal carina absent; ocu- freeing pygidialis Schaeffer to become a lar index 3:1; ocular sinus three-fourths valid name. Johnson (1963) used the name length of eye; antenna extending to middle “pythonicus” for this species. of metepisternum; pronotum and elytra as

218 Stator sordidus (Horn) smooth and polished; ventrolateral carina of metafemur angulate (figure 950); ven- (Figures 946–952) trolateral carina of metatibia complete to Bruchus sordidus Horn 1873:319; Horn apex; ventrolateral carina evanescent or 1894:344; Fall 1901:29,160, 1910:162– absent; dorsal coronal denticle nearly as 166; Schaeffer 1907:292,294,298. long as mucro. Mylabris sordidus: Leng 1920:305. Male genitalia.—As in figures 951 and 952; Acanthoscelides sordidus: Blackwelder ventral valve evenly rounded (figure 951); 1946:761. internal sac with large, saddle-shaped Stator sordidus: Johnson 1963:863; Botti- sclerite; lateral lobes as illustrated (figure mer 1968b:285, 1968c:1027,1039; 952). Johnson 1968:1270, 1969c:55; Molden- Size.—Body length 1.7–3.2 mm; width ke 1971:108; Bottimer 1973:545; John- 1.2–2.2 mm. son and Kingsolver 1976:54; Johnson Type depository.—MCZC (sordidus, ho- 1981a:251; Pfaffen-berger 1981:260; lotype No. 8193); BMNH (semicolon, usti- Johnson and Kingsolver 1982:420; color). Kistler 1982:266; Johnson 1984b:88; Johnson et al. 1989:24; Udayagiri and Type locality.—Lower California (sordidus); Wadhi 1989:112. Guatemala (semicolon); Mexico (usticolor). Bruchus semicolon Sharp 1885:472; Distribution.—AZ, TX; Mexico to Colombia. Schaeffer 1907:297. Host plants.—Acacia wrightii; Calliandra Acanthoscelides semicolon: Blackwelder humilis humilis. Johnson (1979a, 1984b) 1946:761 lists five additional hosts from Mexico and Stator semicolon: Bottimer 1973:545. Honduras. Bruchus usticolor Sharp 1885:467. Natural enemies.—None recorded. Acanthoscelides usticolor: Blackwelder Immatures.—Pfaffenberger 1981 (first larval 1946:761. instar). Stator usticolor: Bottimer 1973:545. Discussion.—Stator sordidus is placed in Color.—Body black; fore leg, mid leg, and the sordidus group with S. chihuahua, S. basal four or five antennal segments red- coconino, S. pygidialis, and S. vachelliae dish yellow, hind leg mostly dark red, (Johnson and Kingsolver 1976). Characters sometimes with base black. Vestiture of to distinguish S. sordidus from others in intermixed yellow-brown and coppery setae the group may be found in the key to spe- on pronotum and elytra; male pygidium cies. mostly yellowish brown with three slightly denser basal patches (figure 947), female Kistler (1982) investigated the effects of pygidium brown with white patch at each temperature on this species. dorsal angle (figure 948); venter of body white but with intermixed coppery setae on Stator subaeneus (Schaeffer) sides of abdomen. (Figures 953–960) Structure.—Vertex and frons densely Bruchus subaeneus Schaeffer 1907:298; punctulate, frontal carina absent; ocular Fall 1910:164. index 3:1; ocular sinus two-thirds length Mylabris subaeneus: Leng 1920:305. of eye; antenna extending to middle of Stator subaeneus: Johnson 1963:865; metepisternum. Pronotum and elytra as Janzen 1967:351,363; Bottimer for genus (figure 946). Metacoxal face with 1968c:1027,1039; Johnson 1968:1271; lateral one-half punctate, medial one-half Janzen 1969:11; Kingsolver 1972a:224; Bottimer 1973:549; Johnson and

219 Kingsolver 1976:60, 1982:420; Johnson the size and shape of the internal sac sc- et al. 1989:9; Udayagiri and Wadhi lerites (compare figures 919 and 959). 1989:113. See discussion of Stator sordidus. Color.—Body and hind leg black; fore leg, Stator subaeneus was reared from Acacia mid leg, and antennae reddish orange; farnesiana seeds (not pods) beneath the dorsum of body with metallic sheen. Vesti- trees (Bottimer 1973). ture coppery, and white setae in irregular bands and patches on pronotum and elytra Stator vachelliae Bottimer (figure 953), in evanescent basal patches and apical lines on pygidium (figures 954 (Figures 961–967) and 955); scutellum white; venter of body Stator vachelliae Bottimer 1973:546; John- sparsely white. son and Kingsolver 1976:65, 1982:420; Structure.—Vertex and frons densely Johnson et al. 1989:9; Udayagiri and punctulate, frontal carina short but dis- Wadhi 1989:114. tinct; ocular index 3:1; ocular sinus one- Bruchus sordidus: Schaeffer 1907:294 half length of eye; dorsal and caudal mar- (misidentification). gins of eye contiguous with vertex and side Stator sordidus: Johnson 1963:863 (mis- of head; antenna extending to middle of identification). metepisternum; pronotum and elytra as for genus (figure 953). Metacoxal face uni- Color.—Body black except basal four anten- formly punctulate (figure 956); ventrolat- nal segments, fore leg and mid leg, apex of eral carina of metafemur obtusely angulate metafemur, and all of metatibia dark red (figure 957); ventrolateral carina of metati- to reddish orange. Vestiture dimorphic: in bia lacking, dorsal fossa distinct. males yellowish brown uniformly distributed except slightly more dense on lateral Male genitalia.—As in figures 958, 959, portions of pronotal disk, pygidium with and 960; ventral valve subtriangular with slightly denser median and lateral patches arcuate lateral margins (figure 958); inter- (figure 963); in females fine, reddish-brown nal sac armed with 20-22 bladelike and setae uniformly distributed except py- thornlike sclerites (figure 959); lateral lobes gidium usually with three elongate, basal as illustrated (figure 960). patches of white setae (figure 964), these Size.—Body length 2.4–2.8 mm; width sometimes evanescent. Scutellum white in 1.8–2.0 mm. both sexes. Type depository.—USNM, holotype No. Structure.—Vertex and frons densely punct- 65933. ulate, frons with median, impunctate line; ocular index 3.5:1; ocular sinus one-half Type locality.—Texas, Brownsville. length of eye; antenna extending to middle Distribution.—TX; Mexico. of metepisternum; pronotum and elytra as Host plants.—Acacia cornigera; A. farnesi- for genus (figure 961); metacoxal face uni- ana. formly punctulate; ventrolateral carina of metafemur angulate opposite ventromesal Natural enemies.—None recorded. denticle (figure 965); ventrolateral carina of Immatures.—Not described. metatibia absent; dorsal fossa lacking. Discussion.—This species is easily sepa- Male genitalia.—As in figures 966 and 967; rated from other U.S. Stator by the eyes ventral valve subtriangular with incurved being contiguous with the vertex and sides lateral margins (figure 966); internal sac of the head. The male genitalia are similar armed with elongate, forked sclerite; lateral to those of S. bottimeri but are distinct in lobes as illustrated (figure 967).

220 Size.—Body length 2.0-2.8 mm; width 1.4- Stylantheus macrocerus (Horn) 1.9 mm. (Figures 13, 70, 968–975) Type depository.—CNCI, holotype. Bruchus macrocerus Horn 1873:342; Type locality.—Texas, Brownsville. Schaeffer 1907:305; Blatchley Distribution.—TX; Mexico to Venezuela. 1910:1238; Fall 1910:187; Carr 1920:7. Host plants.—Acacia farnesiana. Mylabris macrocerus: Leng 1920:306. Acanthoscelides macrocerus: Bridwell Natural enemies.—None recorded. 1935:186. Immatures.—Pfaffenberger 1981 (larva). Stylantheus macrocerus: Bridwell 1946:54; Discussion.—Although S. vachelliae is Bradley 1947:40; Blackwelder and placed in the sordidus group, it differs Blackwelder 1948:44; Bottimer from the other four species in the group by 1968c:1028; Johnson 1968:1271, its uniformly punctate metacoxal face and 1969c:55, 1976b:254; Johnson and by its sexually dimorphic vestiture. Bottim- Kingsolver 1982:413; Udayagiri and er (1973) determined that females of this Wadhi 1989:114. species oviposit on seeds on the ground Acanthoscelides mucrofer Bottimer but not on pods on the tree. The extensive 1969a:975, Ne w Sy n o n y m y . geographical distribution is no doubt due Color.—Integument everywhere black, all to its association with its widespread host, appendages black. Teneral specimens may Acacia farnesiana. be brownish black. Vestiture white, sparse Johnson and Kingsolver (1976) reported on head and pronotum; elytra with short, that S. vachelliae was the prey of a robber oblong patches separated by glabrous fly,Efferia argyrosoma (Hine). spaces, each with central seta, giving ocel- Johnson (1988a) found S. vachelliae in late effect (figures 13 and 968); pygidium Venezuela ovipositing on Parkinsonia acu- patchy (figure 970). leata seeds mixed on the ground with its Structure.—Body elongate. Vertex and normal local host seeds of Acacia flexuosa. frons finely punctate, punctures elongated, Johnson regards this as a possible exam- interspaces granulose; frontal carina ab- ple of the mechanism of host transfer. sent; ocular index 4:1; ocular sinus four- fifths as long as eye; male antenna (figure 971) serrate, eccentric from 3rd segment, Genus Stylantheus Bridwell extending beyond elytral apices; female Stylantheus Bridwell 1946:54; Bradley antenna (figure 971) not as strongly ser- 1947:40; Blackwelder and Blackwelder rate, not extending beyond apices. Prono- 1948:48; Bottimer 1968c:1028; John- tum bell-shaped (figure 968), sides straight son 1968:1271, 1970:28, 1976b:254; anteriad to cervical sulcus then abruptly Johnson and Kingsolver 1982:413; constricted; disk convex, only slightly im- Borowiec 1987:93; Udayagiri and Wadhi pressed on basal margin; surface densely 1989:114. Type-species: Bruchus mac- foveolate, foveolae elliptical, setose; in- rocerus Horn 1873:343, by monotypy. terspaces punctate, setose; lateral carina obsolete; cervical sulcus deep. Scutellum One species. Because this at present is a slightly longer than wide, emarginate. monotypic genus, the generic and specific Elytra slightly longer than wide (figure diagnoses are combined. Johnson (1976b) 968), strongly convex; striae parallel, dis- presented a more detailed description. tinct, 2nd, 3rd, and 4th ending in trans- Revision: Johnson 1976b. verse, denticulate basal carina; interstices flat, finely imbricate; metacoxal face finely, densely punctate, punctures crowded;

221 fossula glabrous; hind leg as in figures Natural enemies.—Urosigalphus bruchi- 70 and 972; pecten with three small den- vorus. ticles; metatibia slender, lateral, ventral, Immatures.—Not described. and dorsomesal carinae present; mucro elongate, curved, one-half length of basi- Discussion.—Johnson (1976b) concluded tarsus (figure 972), lateral denticle and two that Stylantheus, although closely related coronal denticles subequal. Fifth abdomi- to Acanthoscelides, is a valid genus differ- nal sternum of male broadly emarginate; ing in its deep ocular sinus, much male pygidium strongly reflexed; female elongated and broadened antennae, and pygidium vertical; discal surface irregularly uniform size and shape of the pecten den- punctate. ticles. The single species of Stylantheus closely resembles Acanthoscelides compres- Male genitalia.—As in figures 973, 974, sicornis (Schaeffer) in the long tibial mucro and 975; median lobe slender, elongated and elongated antennae but differs in the (figures 973 and 974); ventral valve nar- ocellate elytral pattern and distinctive male rowly triangular, apex acute; internal sac genitalia. lined with minute denticles, large sclerites absent; closure valve circular; lateral lobes I could find no differences between the two elongate (figure 975), arms slender, cleft type specimens of Acanthoscelides mucro- three-fourths their length, apices moder- fer Bottimer and normal Stylantheus mac- ately expanded, mesal faces setose. rocerus; therefore, I here synonymize A. mucrofer with S. macrocerus. Size.—Body length.1.7–2.2 mm; width.1.0– 1.3 mm. Type locality.—District of Columbia (macrocerus, lectotype); Texas, Kenedy County (mucrofer). Type depository.—MCZC, lectotype No. 3897, by Johnson 1968 (macrocerus); CNCI (mucrofer). Distribution.—AL, AR, DC, FL, GA, IL, IN, LA, MD, MS, MO, NC, NJ, OH, OK, SC, TN, TX, VA. Host plants.—Stylosanthes biflora.

222 Appendix I The Hawaiian species of Prosopis have been variously listed as chilensis, juliflora, Hawaiian Bruchidae and pallida. Fosberg (1966) expressed the opinion that the correct name is Prosopis All but one of the 16 species of Bruchidae pallida. Two forma of this species occur known to occur in the Hawaiian Islands in Hawaii—P. pallida pallida is spine- have been accidently introduced either less whereas P. pallida armata is spined. from the southwestern United States- Wagner et al. (1990), however, list both Mexico region or from southeastern Asia P. pallida and juliflora and state that the or through commerce. The exception is a first naturalized collections ofjuliflora deliberate introduction of Lithraeus atro- were made in 1978 on Oahu. Algarobius notatus from South America for biological bottimeri, the sole representative of the control of Christmas berry (Schinus terebin- genus in Hawaii, is found on the main- thifolius Raddi). L. atronotatus is appar- land primarily within the borders of Texas ently well established in the southern part with only a few records in adjoining states of the island of Hawaii, but it has had little (Kingsolver 1986). It attacks seeds of Pro- effect in controlling its host (B. Kumashiro, sopis glandulosa glandulosa and P. reptans personal communication, 1988). cinerascens. Yet this bruchid was somehow transported to the Islands and became the Some of these bruchids were undoubtedly principal seed pest of another species of introduced in the early part of the 20th Prosopis. Fullaway (1913:25) describes the century during establishment of a cattle- attempts to import hymen-opterous para- feed industry using protein-rich Prosopis sites to combat bruchids. The Department seeds. This venture died out within a few of Agriculture brought in from Texas con- years due to the unchecked depredations signments of bruchid-infested seeds from of three introduced bruchids (Algarobius which Heterospilus prosopidis Viereck and bottimeri, Mimosestes nubigens, and Carye- Uscana semifumipennis Girault were bred don serratus) and one stored-grain moth and released. This is probably the source (Plodia interpunctella). Bridwell (1920a) of entry of Algarobius bottimeri into Hawaii. described the effects of these four spe- The preferred host of Mimosestes nubigens cies on the industry, the bruchids being on the mainland and in the Islands is Aca- identified by him asBruchus prosopis (now cia farnesiana, but it has never been found Algarobius bottimeri), Bruchus sallaei (now in mainland Prosopis. Its damage level in Mimosestes nubigens), and Caryoborus Prosopis in Hawaii apparently was never as gonagra (now Caryedon serratus). The first high as that of Algarobius. two species primarily destroyed seeds in the field, whereasC. serratus began feed- The preferred host of Caryedon serratus ing in the field and continued into storage. throughout its tropicopolitan range is Tam- Parasites from the southwestern U.S. were arindus indica, but its host list is extensive. introduced in an attempt to control these The full-grown larva is too large to allow it bruchids, but these efforts were ineffective. to fit inside a seed ofProsopis, but Bridwell (1920b) indicates that oviposition is on ripe Bianchi (1940:381) listed the self-intro- pods and the principal feeding occurs in duced parasites of the Bruchidae of the seed storage. Since it spins a cocoon be- Hawaiian Islands. All of the bruchids fore pupating, serratus does not need the have been transferred to other genera or protection of an excavated seed as a pupal synonymized since then, but most of the shell as other species do. parasites remain in the genera in which he placed them. Six parasites were deliber- Bridwell’s pioneering papers (1918, 1919, ately introduced. 1920a) were in great part reports of experi

223 mental transfers of species of bruchids to known at that time, although several spe- native and introduced hosts. cific names have changed since his papers Hinckley (1960 and 1961) thoroughly were published. covered the history of 12 of the 16 species

Key to Hawaiian Bruchidae

The following key applies only to the Hawaiian Islands.

1 Lateral pronotal margin with prominent tooth or denticle ous (figure 712); antenna not strongly modified (figure (figure 46); ventrolateral margin of metafemur with tooth 713); body red, elytra with white patchy pattern, with or angular emargination (figure 47); elytral striae lacking lateral and apical bare spots; length 2.25 mm basal denticles: Bruchus...... 2 ...... Lithraeus atronotatus (Pic) Lateral pronotal margin smooth, lacking prominent tooth 6(4) Metafemur with 9–12 denticles on mesoventral margin; or denticle; metafemur variously armed; elytral striae with metatibia strongly arcuate (figure 1); reddish brown with or without basal denticles...... 3 scattered brown spots; length 6 mm 2(1) Ventrolateral tibial tooth long, acute, lying outside tibia ...... Caryedon serratus (Olivier) when leg is closed (figure 47); pygidium with large, Metafemur with no more than four denticles on mesoven- subapical maculae (figure 267) tral margin; metatibia nearly straight (except at extreme ...... Bruchus pisorum (Linnaeus) base), or slightly arcuate...... 7 Ventrolateral tibial tooth angular but not elongated (figure 7(6) Scutellum elongate, nearly 2 times as long as wide (figure 274); pygidial maculae small (figure 275) 643); yellowish and brown mottled; female with pair of ...... Bruchus rufimanus Boheman sulci on pygidium (figure 645); length 4–4.5 mm 3(1) Metatibial apex with two movable apical spurs (calcaria) ...... Algarobius bottimeri Kingsolver (figure 43); lateral margin of pronotum with sharp carina; Scutellum quadrate or transverse...... 8 ventrolateral margin of metafemur carinate, sinuate; body 8(7) Lateral margin of pronotum with distinct, shining carina; black with transverse white bands on elytra (female) ventrolateral margin of metafemur with strong carina (fig- (figure 191) or with yellowish mottling (male) (figure 190) ure 931), sometimes angulate or thickened; ventromesal ...... Zabrotes subfasciatus (Boheman) margin with single denticle: Stator...... 9 Metatibial apex without movable apical spurs but some- Lateral margin of pronotum lacking distinct, shining times with fixed apical spines and denticles; margin of carina; ventrolateral margin of metafemur carinate or not, metafemur varied; color varied...... 4 ventromesal margin with one or more denticles (figures 4(3) Metafemur with one or more denticles on ventromesal 44 and 792)...... 10 margin (figures 78, 523 and 937)...... 6 9(8) Body black with black-bordered red or orange elytra; Metafemur without denticles on ventromesal margin pygidium with three indistinct white patches along basal (figure 215, 714)...... 5 margin; length 2.5–3.0 mm...... Stator limbatus (Horn) 5(4) Frons with prominent median carina; strial punctures Body and elytra uniformly black; pygidium lacking of elytra deep and rounded (figure 243); male antenna patches; length 2.25–3.0 mm pectinate (as in figure 230), female antenna serrate (as ...... Stator pruininus (Horn) in figure 231); red and white alternating stripes on elytra; 10(8) Frons with V- or Y-shaped, bare, shining, raised boss length 3.5 mm (figure 57); male with deep cavity on ventral margin of ...... Megacerus leucospilus (Sharp) metafemur: Mimosestes...... 11 Frons without median carina; strial punctures inconspicu-

224 Frons with strong or faint vertical median carina; male Elytral striae extending to basal margin, lacking prominent denticles metafemur lacking ventral cavity...... 13 or gibbosities (figure 305); basal lobe of pronotum with glossy coating on setae...... 16 11(10) Body black with some reddish areas, vestiture entirely gray; eyes flat against side of head (figure 791) 15(14) Pattern of pronotum and elytra as in figure 327; pygidium ...... Mimosestes amicus (Horn) in both sexes only slightly convex, nearly immaculate Body dark red, vestiture yellowish; eyes protruding from except for subapical, paired spots; antennae as in figures side of head...... 12 328 and 329; lateral margins of 2nd to 4th abdominal sterna with uniform pubescence ...... 12(11) Anterolateral corner of pronotum sharply angulate and ...... Callosobruchus pulcher Pic finely tuberculate on shoulder (figures 807 and 808); Pattern of pronotum and elytra as in figure 297; pygidium posterodorsal corner of eye subangulate (figure 807) in both sexes convex, male with vague patches at middle ...... Mimosestes nubigens (Motschulsky) and at apex, female with more intense patches sometimes Anterolateral corner of pronotum rounded, not tubercu- connected into paired crescents; antennae as in figures late; posterodorsal corner of eye rounded (figure 796) 299 and 300; lateral margins of 2nd to 4th abdominal ...... Mimosestes insularis Kingsolver & Johnson sterna with intensely white setal patches (figure 298) 13(10) Metafemur with toothed carina on ventrolateral margin ...... Callosobruchus chinensis (Linnaeus) and single denticle on ventro-mesal margin (figure 310): 16(14) Pronotum yellowish with brown or black longitudinal Callosobruchus...... 14 lines (figure 321); elytral pattern as in figure 321; eye with Metafemur without carina on ventrolateral margin but with narrow posterior fringe ...... one long and two or three shorter denticles (figure 44); ...... Callosobruchus phaseoli (Gyllenhal) terminal segment of antenna and abdomen red Pronotum dark red to black, lacking longitudinal lines ...... Acanthoscelides obtectus (Say) (figure 305); elytral pattern variable (figures 305–307); 14(13) Elytral striae three and four each with prominent subbasal eye with posterior margin lobed denticles on slight gibbosity (figure 298)...... 15 ...... Callosobruchus maculatus (Fabricius)

Host Associations of Hawaiian First reported by Bridwell (1918), but Bruchidae specimens were collected by Van Dine in 1904 (specimen label data). This list was compiled from various sources, and plant names were updated as far Algarobius bottimeri.—Mainland distribu- as possible. Bridwell (1918) conducted a tion is almost entirely within Texas. Pri- series of experimental rearings of several mary host in Texas is Prosopis glandulosa. species of bruchids, but it is not certain Hawaiian host is Prosopis pallida. First that all of the plants he used now grow in reported by Fullaway (1913) as Bruchus Hawaii. prosopis. Refer to the main body of this handbook Bruchus pisorum.—Origin in Old World, but for illustrations and for further information species is cosmopolitan. Primary host is and to the bibliography for complete cita- Pisum sativum. First reported by Bridwell tions. (1918) but probably not established in Hawaii. Acanthoscelides obtectus.—Origin in the Neotropics but now cosmopolitan. Primary Bruchus rufimanus.—Origin in Old World hosts are various species of Phaseolus, but but nearly cosmopolitan. Primary host records of other legume genera are known. is Vicia faba. First reported by Bridwell

225 (1918) but probably not established in Cercidium, Parkinsonia, and Prosopis. In Hawaii. Hawaii, it attacks Acacia farnesiana, Leu- Callosobruchus chinensis.—Origin in Asia caena glauca, Prosopis pallida, and Ses- or Africa but now cosmopolitan. Primary bania sesban. First reported by Swezey hosts are in genera Phaseolus and Vigna, (1925). but host range is broad. First reported by Mimosestes insularis.—Distribution West Swezey (1912). Indies, Colombia, and Hawaii. In the West Callosobruchus maculatus.—Generally Indies, attacks Acacia farnesiana and Pro- same as C. chinensis. Many hosts have sopis juliflora. In Hawaii, attacks Prosopis been reported. First detected by Bridwell pallida. Described in 1978 by Kingsolver (1918). and Johnson from Hawaii and West Indies. Mimosestes nubigens.—Native to United Callosobruchus phaseoli.—Origin probably States south to Brazil. Primary host prob- in Asia but now tropicopolitan. Primary ably Acacia farnesiana and perhaps other host Lablab purpureus, but other hosts are species of Acacia. Hawaiian hosts are Aca- recorded. First reported by Bridwell (1918), cia farnesiana and Prosopis pallida. First but specimens were collected by Swezey in reported by Bridwell (1919). Listed by him 1908 (specimen label data). and subsequent authors as Bruchus or Callosobruchus pulcher.—Hosts: Cajanus, Mimosestes sallaei. Cicer, Phaseolus, and Pisum in native Phil- Stator limbatus.—Native to United States ippine Islands. In Hawaii, first collected south to Panama in Acacia spp., Cercidium by Krauss in 1965 at Haiku, Maui; subse- spp., Lysiloma spp., and Pithecellobium quently reared from Cajanus cajan at Ewa spp. In Hawaii, it attacks Acacia confusa, by Funisaki (1973), and collected at light A. koa, Albizia lebbeck, Leucaena glauca, at Manoa. Pithecellobium dulce, and Samanea saman. Caryedon serratus.—Tropicopolitan species First reported by Bridwell (1920b) on col- with Asian or African origin. Primary host lections made in 1919. probably Tamarindus indica but in Hawaii Stator pruininus.—Native to United States attacks Acacia farnesiana, Bauhinia to- south to Venezuela in Acacia spp., Mimosa men-tosa, Caesalpinia pulcherrima, Cassia spp., Pithecellobium spp., and others. In fistula, Cassia grandis, and Prosopis pal- Hawaii, it attacks Acacia confusa, A. koa, lida as well as Tamarindus. First reported Albizia saponaria, Cassia nodosa, Des- by Fullaway (1913) and Swezey (1912) as man-thus virgatus, Erythrina sandwicensis, Caryoborus gonagra. Indigofera anil, I. suffruticosa, Leucaena Lithraeus atronotatus.—Native host in Bra- leucocephala, and Sesbania sesban. First zil is Schinus terebinthifolius. Introduced reported by Bridwell (1918) from collec- into Hawaii to control the same plant. First tions made in 1917 on L. leucocephala and released in 1954 by Krauss (1963) but suc- S. sesban. cessful control not evident. Zabrotes subfasciatus.—Probably native to Megacerus leucospilus.—Native to Texas Central America but now tropicopolitan. through Central America in Ipomoea pes- Breeds principally in Phaseolus but also caprae. Found in I. carnea fistulosa, I. in Vigna spp., Vicia faba, Lablab purpu- pes-caprae, I. tuberculata, and Merremia reus, Cajanus cajan, and Cicer arietinum. aegyptia in Hawaii. Described as Mega-ce- Bridwell (1918) reared it from Phaseolus rus alternatus by Bridwell (1929b). vulgaris, P. lunatus, P. articulatus, P. acutifolius, Vigna unguiculatus, Cajanus cajan, Mimosestes amicus.—Native from south- Glycine max, Cicer arietinum, and Pisum western United States to Costa Rica in sativum. First observations by Cowan but species of Acacia, reported by Bridwell (1918) as Spermopha- gus pectoralis.

226 Appendix II Synonymical List of the Bruchidae of the United States and Canada Names are arranged in alphabetical order. Full references and dates for each name can be found in the main text, or in the bibliography.

Extant Species Bruchus rufus: Motschulsky Mylabris aureolus: Leng Abutiloneus Bridwell Mylabris pauperculus: Leng (not LeConte) Abutiloneus idoneus Bridwell Acanthoscelides baboquivari Johnson Abutiloneus flavicornis:Kingsolver Acanthoscelides bisignatus (Horn) Acanthoscelides flavicornis:Blackwelder Acanthoscelides bisignatus: Glick Althaeus idoneus: De Luca Bruchus bisignatus Horn Bruchus (Abutiloneus) idoneus: Zacher Mylabris bisignatus: Leng Bruchus flavicornis Sharp Acanthoscelides Schilsky Acanthoscelides biustulus (Fall) Acanthocelides (sic) Schilsky Acanthoscelides biustulus: Johnson Acanthoscelides aequalis (Sharp) Bruchus biustulus Fall Acanthoscelides aequalis: Moreno and Mylabris biustulus: Leng Bibby Acanthoscelides calvus (Horn) Bruchus aequalis Sharp Acanthoscelides calvus: Bridwell Mylabris aequalis: Leng Bruchus calvus Horn Acanthoscelides alboscutellatus (Horn) Mylabris calvus: Leng Acanthoselides abutilonis: Kingsolver Acanthoscelides chiricahuae (Fall) Acanthoscelides alboscutellatus: Bottimer Acanthoscelides chiricahuae: Johnson Acanthoscelides conspersus: Kingsolver Bruchus chiracahuae: Pic (error) Bruchus abutilonis Motschulsky Bruchus chiricahuae Fall Bruchus albiscutellaris: Ashmead (error) Mylabris chiricahuae: Leng Bruchus alboscutellaris: Zacher (error) Acanthoscelides compressicornis (Schaeffer) Bruchus alboscutellatus Horn Acanthoscelides compressicornis: Bruchus conspersus Motschulsky Muesebeck et al. Mylabris alboscutellatus: Leng Acanthoscelides subserripes: Johnson Acanthoscelides atomus (Fall) Bruchus compressicornis Schaeffer Abutiloneus atomus: Bradley Bruchus subserripes Fall Acanthoscelides atomus: Bottimer Mylabris compressicornis: Leng Bruchus atomus Fall Acanthoscelides comstocki Johnson Mylabris atomus: Leng Acanthoscelides daleae Johnson Acanthoscelides aureolus (Horn) (complex) Acanthoscelides desmanthi Johnson Acanthoscelides aureolus: Blackwelder Acanthoscelides distinguendus (Horn) Acanthoscelides pauperculus: Bottimer Acanthoscelides distinguendus: Bissell Bruchus aureolus Horn Bruchus distinguendus Horn Bruchus fraterculus: Fall (not Horn) Mylabris distinguendus: Leng Bruchus pulicarius: Motschulsky Acanthoscelides flavescens(Fahraeus) Bruchus pauperculus: Schaeffer (not Acanthoscelides flavescens:Blackwelder LeConte) Acanthoscelides ochraceicolor: Wolcott

227 Bruchus flavescens Fahraeus Acanthoscelides macrophthalmus (Schaeffer) Bruchus ochraceus Schaeffer Acanthoscelides macrophthalmus: Johnson Bruchus ochraceicolor Pic Bruchus macrophthalmus Schaeffer Mylabris ochraceicolor: Leng Mylabris macrophthalmus: Leng Acanthoscelides floridae(Horn) Acanthoscelides margaretae Johnson Acanthoscelides floridae:Brett Acanthoscelides mixtus (Horn) Bruchus floridaeHorn Acanthoscelides mixtus: Johnson Mylabris floridae:Leng Bruchus mixtus Horn Acanthoscelides fraterculus (Horn) Mylabris mixtus: Leng Acanthoscelides fraterculus: Trelease and Acanthoscelides modestus (Sharp) Trelease Acanthoscelides atrocephalus: Blackwelder Bruchus aureolus: Fall (misident.) Acanthoscelides modestus: Blackwelder Bruchus fraterculus Horn Bruchus atrocephalus Pic Mylabris fraterculus: Leng Bruchus modestus Sharp Acanthoscelides fumatus (Schaeffer) Acanthoscelides mundulus (Sharp) Acanthoscelides fumatus: Bottimer Acanthoscelides mundulus: Blackwelder Acanthoscelides fumatus: Johnson Bruchus mundulus Sharp Bruchus fumatus Schaeffer Acanthoscelides napensis Johnson Mylabris fumatus: Leng Acanthoscelides obrienorum Johnson Acanthoscelides griseolus (Fall) Acanthoscelides obsoletus (Say) Acanthoscelides griseolus: Bridwell Acanthoscelides obsoletus: Yip Acanthoscelides kiotoensis: Morimoto Bruchus distinguendus: Horn (in part)(error) Bruchus griseolus Fall Bruchus obscurus: Fitch (error) Bruchidius kiotoensis: Chujo Bruchus obsoletus Say Bruchus kiotoensis Pic Mylabris obsoletus: Leng Bruchus kiotensis: Udayagiri and Wadhi Acanthoscelides obtectus (Say) Mylabris griseolus: Leng Acanthoscelides obtectus: Skaife Acanthoscelides helianthemum Bottimer Bruchidius (Acanthoscelides) obtectus: Reitter Bruchus compressicornis: Blatchley Bruchus acanthocnemus Jekel Acanthoscelides herissantitus Johnson Bruchus breweri Crotch Acanthoscelides inquisitus (Fall) Bruchus fabae Fitch (not Motschulsky) Acanthoscelides inquisitus: Johnson Bruchus fabi Rathvon Bruchus inquisitus Fall Bruchus gilvipes Motschulsky Mylabris inquisitus: Leng Bruchus irresectus Fahraeus Acanthoscelides kingsolveri Johnson Bruchus leguminarius: Gyllenhal Acanthoscelides lobatus (Fall) Bruchus leguminarius var. melanocephalus Acanthoscelides lobatus: Johnson Blackwelder (error) Bruchus lobatus Fall Bruchus melanocephalus Fahraeus (misiden- Mylabris lobatus: Leng tification) Acanthoscelides longistilus (Horn) Bruchus mimosae: Gemminger and Harold (not Fabricius) Acanthoscelides longistilus: Bridwell Bruchus obsoletus: of authors (not Say) Acanthoscelides sp. near longistylus: Glick Bruchus obtectus Say Bruchus longistilus Horn Bruchus pallidipes Fahraeus Bruchus longistylus: Ulke (error) Bruchus subellipticus Wollaston Mylabris longistilus: Leng

228 Bruchus tetricus Gyllenhal Acanthoscelides pullus (Fall) Bruchus varicornis Motschulsky Acanthoscelides brunneostictus: Johnson Laria obtecta: Bedel Acanthoscelides pullus: Bridwell Mylabris irresecta: Baudi Bruchus brunneostictus Fall Mylabris obtectus: Leng Bruchus pullus Fall Spermophagus incretus Motschulsky Mylabris brunneostictus: Leng Acanthoscelides oregonensis Johnson Mylabris pullus: Leng Acanthoscelides pallidipennis (Motschulsky) Acanthoscelides pusillimus (Sharp) Acanthoscelides collusus: Johnson Acanthoscelides pusillimus: Chujo Acanthoscelides pallidipennis: Kingsolver Acanthoscelides sp. 20: Johnson Acanthoscelides perplexus: Johnson Bruchidius pusillimus: Zacher Acanthoscelides tarnawski Borowiec Bruchus pusillimus Sharp Bruchus collusus Fall Acanthoscelides quadridentatus (Schaeffer) Bruchus pallidipennis Motschulsky Acanthoscelides quadridentatum: Moreno Bruchus perplexus Fall and Bibby Mylabris collusus: Leng Acanthoscelides quadridentatus: Zacher Mylabris pallidipennis: Leng Bruchus quadridentatus Schaeffer Mylabris perplexus: Leng Mylabris quadridentatus: Leng Acanthoscelides pauperculus (LeConte) Acanthoscelides rufovittatus (Schaeffer) Acanthoscelides pauperculus: Bottimer Acanthoscelides rufovittatus: Johnson Bruchus pauperculus LeConte Bruchus rufovittatus Schaeffer Bruchus pulicarius Motschulsky Mylabris rufovittatus: Leng Bruchus rufus Motschulsky Acanthoscelides schaefferi (Pic) Bruchus simplex Motschulsky Acanthoscelides rufescens: Johnson Mylabris pauperculus: Leng Acanthoscelides schaefferi: Bottimer Mylabris pulicarius: Leng Bruchus rufescens Schaeffer (preocc.) Mylabris simplex: Leng Bruchus schaefferi Pic Acanthoscelides pectoralis (Horn) Mylabris schaefferi: Leng Acanthoscelides pectoralis: Bradley Acanthoscelides schrankiae (Horn) Bruchus pectoralis Horn Acanthoscelides schrankiae: Bissell Mylabris pectoralis: Leng Bruchus schrankiae Horn Acanthoscelides pedicularius (Sharp) Mylabris schrankiae: Leng Acanthoscelides pedicularius: Blackwelder Acanthoscelides seminulum (Horn) Acanthoscelides pulloides: Johnson Abutiloneus seminulum: Bradley Bruchus pedicularis Sharp Acanthoscelides seminulum: Bottimer Bruchus pulloides Fall Bruchus seminulum Horn Mylabris pulloides: Leng Bruchus seminulus: Pic (missp.) Acanthoscelides perforatus (Horn) Megacerus seminulum: Zacher (error) Acanthoscelides perforatus: Bradley Mylabris seminulum: Leng Bruchus perforatus Horn Acanthoscelides speciosus (Schaeffer) Mylabris perforatus: Leng Acanthoscelides mexicanus: Johnson (not Sharp) Acanthoscelides prosopoides (Schaeffer) Acanthoscelides speciosus: Johnson Acanthoscelides prosopoides: Johnson Bruchus speciosus Schaeffer Bruchus prosopoides Schaeffer Mylabris speciosus: Leng Mylabris prosopoides: Leng

229 Acanthoscelides stylifer (Sharp) Althaeus steineri Kingsolver Acanthoscelides pugiunculus: Johnson Amblycerus Thunberg Acanthoscelides stylifer: Blackwelder Anthotribus Gistel Bruchus pugiunculus Fall Spermophagus, of authors Bruchus pygionculus: Pic (error) Amblycerus eustrophoides (Schaeffer) Bruchus stylifer Sharp Amblycerus eustrophoides: Johnson Mylabris pugiunculus: Leng Spermophagus eustrophoides Schaeffer Acanthoscelides subaequalis Johnson Amblycerus ireriae Romero, Johnson and Acanthoscelides submuticus (Sharp) Kingsolver Acanthoscelides exiguus: De Luca Amblycerus nigromarginatus (Motschulsky) Acanthoscelides horni: Caffrey Amblycerus nigromarginatus: Bridwell Acanthoscelides submuticus: Glick Spermophagus nigromarginatus Motschulsky Bruchus exiguus Horn Amblycerus obscurus (Sharp) Bruchus horni Pic Amblycerus obscurus: Bridwell Bruchus submuticus Sharp Spermophagus obscurus Sharp Mylabris exiguus: Leng Amblycerus robiniae (Fabricius) Mylabris horni: Leng Amblycerus robiniae: Bridwell Acanthoscelides tenuis Bottimer Bruchus robiniae Fabricius Bruchus tenuis: Blackwelder Chrysomela gleditsiae Castiglioni Acanthoscelides tridenticulatus Bottimer Spermophagus hoffmanseggi or hoffmannseggi, of authors Algarobius Bridwell Spermophagus robiniae: Gyllenhal Algarobius bottimeri Kingsolver Amblycerus schwarzi Kingsolver Algarobius prosopis: Hinckley Amblycerus vitis (Schaeffer) Bruchus prosopis: of authors (not LeConte) Amblycerus vitis: Bottimer Algarobius prosopis (LeConte) Spermophagus vitis Schaeffer Acanthoscelides desertorum: Blackwelder Borowiecius Anton Acanthoscelides prosopis: Blackwelder Borowiecius ademptus (Sharp) Algarobius desertorum: Blackwelder and Bruchus ademptus Sharp Blackwelder Borowiecius ademptus: Anton Algarobius prosopis: Bridwell Callosobruchus ademptus: Chujo Algarobius uniformis: Blackwelder and Blackwelder Bruchidius Schilsky Bruchus desertorum LeConte Sparteus Bridwell Bruchus prosopis LeConte Spermophagus: Arnett (not Schoenherr) Bruchus uniformis LeConte Bruchidius cisti (Fabricius) Mylabris desertorum: Leng Bruchidius cisti: Southgate Mylabris prosopis: Leng Bruchidius unicolor: Bottimer Mylabris uniformis: Kannan Bruchidius unicolor var. debilis: Schilsky Althaeus Bridwell Bruchidius villosus var. pubescens: Althaeus folkertsi Kingsolver Luk’yanovich and Ter-Minassian Althaeus hibisci (Olivier) Bruchus ater Curtis Althaeus hibisci: Bridwell Bruchus ater var. pubescens: Jacquet Bruchus hibisci Olivier Bruchus autumnalis Motschulsky Bruchus transversus Say Bruchus canus Germar Bruchus transvexus: Horn (error)

230 Bruchus cisti Fabricius Bruchus granarius: of authors (not Bruchus debilis Gyllenhal Linnaeus) Bruchus nugax Motschulsky Bruchus rufimanus var. diversipubensPic Bruchus pulverulentus Motschulsky Bruchus rufimanus var. velutinusMulsant and Rey Bruchus rostratus Motschulsky Bruchus rufimanus var. rufimanus Bruchus tibiellus Stephens Luk’yanovich and Ter-Minassian Bruchus unicolor Olivier Laria rufimana:Chittenden Bruchus villosus var. pubescens: Pic Mylabris rufimana:Baudi Mylabris villosa var. pubescens: Baudi Mylabris rufimanus: Leng Spermophagus cisti: Schilsky Mylabris rufimana var. velutina:Baudi Bruchidius villosus (Fabricius) Callosobruchus Pic Bruchidius ater: Southgate Bruchus (Callosobruchus) Pic Bruchidius villosus: Zacher Callosobruchus chinensis (Linnaeus) Bruchus ater Marsham Bruchus biguttatus Fabricius Bruchus cisti sensu Paykull (not Fabricius) Bruchus biguttellus Schoenherr Bruchus fasciatus, of authors (not Olivier) Bruchus bistriatus Fabricius Bruchus villosus Fabricius Bruchus bistriotus: Bridwell (error) Mylabris villosus: Baudi Bruchus (Callosobruchus) chinensis: Pic Sparteus villosus: Bridwell Bruchus (Callosobruchus) chinensis: Zacher Spermophagus fasciatus (Olivier) of Arnett Bruchus chinensis: Cushman Bruchus brachialis Fahraeus Bruchus elegans: Sturm, Gemminger and Bruchus brachialis Fahraeus Harold Bruchus pallidicornis Mulsant and Rey Bruchus pectinicornis Linnaeus Laria brachialis: Bedel Bruchus rufobrunneus: Wollaston Mylabris brachialis: Baudi Bruchus rufus De Geer Bruchus Linnaeus Bruchus scutellaris Fabricius Acanthobruchus Ramos Callosobruchus chinensis: Bridwell Bruchoides Ramos Curculio chinensis Linnaeus Laria Scopoli (of authors) Mylabris chinensis: Harold Mylabris Müller (of authors) Pachymerus chinensis: Schilsky Bruchus pisorum (Linnaeus) Callosobruchus maculatus (Fabricius) Bruchus fabae Brulle Acanthoscelides sinuatus: Blackwelder Bruchus intermedius Motschulsky Bruchus ambiguus Gyllenhal Bruchus lunarius Rey Bruchus arachidis Fahraeus Bruchus pisi Linnaeus Bruchus litteratus Schoenherr Bruchus pisorum: Abeille Bruchus maculatus Fabricius Bruchus pisorum var. unifasciatus Rey Bruchus ornatus Boheman Bruchus sparsus Fabricius Bruchus quadrimaculatus Fabricius Dermestes pisorum Linnaeus Bruchus sinuatus Fahraeus Laria salicis Scopoli Bruchus (Acanthoscelides) trabuti Caillol Mylabris cruciger Geoffroy Bruchus vicinus Gyllenhal Mylabris pisorum: Baudi Callosobruchus maculatus: Bridwell Mylabris pisorum: Leng Callosobruchus ornatus: Hoffmann Bruchus rufimanusBoheman Callosobruchus quadrimaculatus: Shomar Bruchus fabae Motschulsky Callosobruchus sinuatus: Zacher

231 Laria quadrimaculata: Bedel Caryedon Schoenherr Mylabris quadrimaculatus: Baudi Caryedon serratus (Olivier) Pachymerus ornatus: Schilsky Bruchus fuscus Bedel, and authors Pachymerus quadrimaculatus: Schilsky Bruchus gonagra Fabricius Callosobruchus phaseoli (Gyllenhal) Bruchus serratus Olivier Bruchus conicicollis Fairmaire Caryedon gonagra: Herford Bruchus figuratusGyllenhal Caryedon serratus: Schoenherr Bruchus phaseoli Gyllenhal Caryoborus gonagra: (Fabricius) Swezey Callosobruchus phaseoli: Herford Caryoborus gonagra: Gyllenhal Mylabris phaseoli: Baudi Pachymerus acaciae Lepesme Pachymerus phaseoli: Schilsky Pachymerus gonagra: Pic Callosobruchus pulcher (Pic) Pachymerus gonager: Chujo Callosobruchus albocallosus (Pic) (misident.) Pachymerus longus Pic Caryedes Hummel Pachymerus sibutensis Pic Adromisus Des Gozis Caryobruchus Bridwell Andromisus Des Gozis Caryobruchus gleditsiae (Linnaeus) Falsobruchus Pic Bruchus arthriticus Fabricius Pachymera Berthold Bruchus fuscus Goeze Pachymere Latreille Bruchus gleditsiae: Linnaeus Pachymerus Schoenherr Caryoborus arthriticus: Schoenherr Pedalophus Bottimer Caryobruchus arthriticus: Zacher Pedapholus Gistel Caryobruchus gleditsiae: Bridwell Pseudopachymerus Pic Dermestes gleditsiae: Johansson and Lin- Caryedes helvinus (Motschulsky) naeus Bruchus calderensis Sharp Pachymerus gleditsiae: Pic Gibbobruchus Pic Caryedes bicoloripes var. binotata: Black- welder Gibbobruchus: Bridwell Caryedes calderensis: Blackwelder Pachymerus subg. Gibbobruchus Pic Caryedes helvina: Blackwelder Pseudopachymerus subg. Caryedes helvinus: Kingsolver and White- Gibbobruchus Pic head Gibbobruchus cristicollis (Sharp) Caryedes scabricollis: Blackwelder Bruchus cristicollis Sharp Pachymerus helvinus Motschulsky Caryedes cristicollis: Blackwelder Pachymerus scabricollis Chevrolat Gibbobruchus cristicollis: Whitehead and Pseudopachymerus calderensis: Pic Kingsolver Pseudopachymerus helvinus: Pic Pseudopachymerus cristicollis: Pic Pseudopachymerus multimaculatus var. Gibbobruchus divaricatae Whitehead and King- binotatus Pic solver Pseudopachymerus scabricollis: Pic Gibbobruchus mimus (Say) Caryedes incensus (Sharp) Bruchus borealis Schoenherr Bruchus incensus Sharp Bruchus crataegi Fahraeus Caryedes incensa: Blackwelder Bruchus mimus Say Caryedes incensus: Kingsolver and White- Bruchus murinus Schoenherr (error) head Gibbobruchus mimus: Bridwell Pseudopachymerus incensus: Pic

232 Mylabris mimus: Leng Mylabris leucosomus: Leng Pseudopachymerus crataegi: Pic Megacerus discoidus (Say) Kytorhinus Fischer von Waldheim Bruchus (Pachybruchus) discoideus: Pic Cytorhinus Agassiz Bruchus discoidens: Dury Kytorhinus Fischer von Bruchus discoideus: Melsheimer and au- Waldheim thors Kytorhinus Motschulsky Bruchus discoidus Say Kytorhynus Motschulsky Kytorhinus discoidii: Motschulsky Kytorrhinus Baudi Megacerus discoideus: Bridwell Pygobruchus Sharp Megacerus discoidus: Bottimer Kytorhinus prolixus (Fall) Mylabris discoideus: Leng Kytorhinus prolixus: Bridwell Megacerus impiger (Horn) Kytorrhinus prolixus: Teran Bruchus impiger Horn Mylabris (Bruchus) prolixus Fall Bruchus (Pachybruchus) impiger: Pic Lithraeus Bridwell Bruchus ramicornis Boheman (not Erichson) Lithraeus atronotatus (Pic) Megacerus eugenie Bottimer Acanthoscelides atronotatus: Blackwelder Megacerus impiger: Bridwell Bruchus atronotatus Pic Megacerus (Megacerus) impiger: Teran and Lithraeus atronotatus: De Luca Kingsolver Megacerus Fahraeus Megacerus ramicornis: Blackwelder Pachybruchus Pic Mylabris impiger: Leng Serratibruchus Teran and Megacerus leucospilus (Sharp) Kingsolver Bruchus (Pachybruchus) coryphae var. lin- Megacerus coryphae (Olivier) eatipennis Pic Bruchus coryphae Olivier Bruchus leucospilus Sharp Bruchus (Pachybruchus) coryphae: Pic Bruchus (Pachybruchus) leucospilus: Pic Megacerus coryphae: Bridwell Megacerus alternatus Bridwell Megacerus (Pachybruchus) coryphae: Teran Megacerus leucorpilus: Bridwell (error) and Kingsolver Megacerus leucospilus: Bridwell Mylabris coryphae: Leng Megacerus maculiventris (Fahraeus) Megacerus cubiculus (Casey) Acanthoscelides mactatus: Blackwelder Bruchus cubiculus Casey Bruchus impiger var.: Sharp Megacerus cubiculus: Bottimer Bruchus kytorrhinensis Pic Megacerus (Megacerus) cubiculus: Teran and Bruchus (Pachybruchus) kytorrhinensis: Pic Kingsolver Bruchus (Pachybruchus) maculiventris: Pic Mylabris impiger var. cubiculus: Leng Bruchus mactatus Pic Megacerus cubicus (Motschulsky) Bruchus maculiventris Fahraeus Bruchus cubicus: Sharp Bruchus serratifemur Schaeffer Bruchus (Pachybruchus) cubicus: Pic Bruchus (Pachybruchus) triangularifer Pic Bruchus leucosomus Sharp Bruchus (Pachybruchus) triangulifer var. Bruchus (Pachybruchus) leucosomus: Pic multisparsus Pic Kytorhinus cubicus Motschulsky Kytorrhinus quadratus Motschulsky Megacerus cubicus: Blackwelder Megacerus kytorrhinensis: Blackwelder Megacerus (Megacerus) cubicus: Teran and Megacerus maculiventris: Blackwelder Kingsolver Megacerus (Serratibruchus) maculiventris: Megacerus leucosomus: Blackwelder Teran and Kingsolver

233 Mylabris serratifemur: Leng Merobruchus julianus: Bridwell Pachybruchus triangulifer: Pic Mylabris julianus: Leng Pachybruchus triangulifer var. multisparsus: Merobruchus knulli (White) Pic Bruchus knulli White Megacerus ripiphorus (Fahraeus) Merobruchus knulli: Johnson Acanthoscelides ripiphorus: Blackwelder Mylabris knulli: Blackwelder and Blackweld- Bruchus ripiphorus Fahraeus er Megacerus ripiphorus: Teran and Kingsolver Merobruchus lysilomae Kingsolver Megacerus (Megacerus) ripiphorus: Teran Merobruchus major (Fall) and Kingsolver Acanthoscelides flexicaule:Zacher Megacerus wheelocki: Bottimer Acanthoscelides flexicaulis:U.S. Depart- Mylabris wheelocki Blatchley ment of Agriculture Megacerus schaefferianus Bridwell Bruchus flexicaulisSchaeffer Bruchus crenatus Schaeffer Bruchus julianus: Schaeffer (error) Megacerus crenatus: Johnson Bruchus major Fall Megacerus schaefferianus Bridwell Merobruchus flexicaulis: Wheeler et al. Megacerus (Serratibruchus) schaefferianus: Merobruchus major: Bradley Teran and Kingsolver Mylabris major: Leng Meibomeus Bridwell Mylabris julianus: Boeving Meibomerus: Arnett (error) Merobruchus placidus (Horn) Meibomeus desmoportheus Kingsolver and Acanthoscelides limpidus: Blackwelder Whitehead Acanthoscelides placidus: Blackwelder Meibomeus musculus (Say) Bruchus limpidus Sharp Bruchus alboguttis Motschulsky Bruchus placidus Horn Bruchus erythrocerus Riley Merobruchus limpidus: Johnson and King- Bruchus musculus Say solver Meibomeus musculus: Bridwell Merobruchus placidus: Johnson Meibomeus surrubresus (Pic) Mylabris placidus: Leng Acanthoscelides subrubrosus: Blackwelder Merobruchus terani Kingsolver (error) Merobruchus sp. 4: Johnson Acanthoscelides surrubresus: Schoonhoven (error) Merobruchus vacillator (Sharp) Bruchus surrubresus Pic Acanthoscelides vacillator: Blackwelder Meibomeus surrubresus: Kingsolver and Bruchus vacillator Sharp Whitehead Merobruchus vacillator: Bottimer Merobruchus Bridwell Mimosestes Bridwell Merobruchus insolitus (Sharp) Cercidiestes Bridwell Acanthoscelides insolitus: Blackwelder Mimosestes acaciestes Kingsolver and Johnson Bruchus insolitus Sharp Mimosestes new species: Center and John- Merobruchus insolitus: Mumford son Merobruchus sp.: Gibson Mimosestes amicus (Horn) Merobruchus julianus (Horn) Acanthoscelides amicus: Blackwelder Acanthoscelides julianus: Blackwelder Bruchus amicus Horn Bruchus julianus Horn Mimosestes amicus: Muesebeck et al. Bruchus ochreolineatus Fall Mylabris amicus: Leng Bruchus ochrolineatus: Pic (error) Mimosestes insularis Kingsolver and Johnson

234 Mimosestes mimosae (Fabricius) Mimosestes ulkei (Horn) Acanthoscelides breweri: Blackwelder Bruchus ulkei Horn Acanthoscelides dominicanus: Blackwelder Bruchus (Cercidiestes) ulkei: Zacher Acanthoscelides immunis: Blackwelder Cercidiestes ulkei: Bridwell Acanthoscelides innotatus: Bradley Mimosestes ulkei: Kingsolver and Johnson Acanthoscelides mimosae: Blackwelder Mylabris ulkei: Leng Acanthoscelides strigatus: Blackwelder Neltumius Bridwell Acanthoscelides subrufus: Blackwelder Neltumius arizonensis (Schaeffer) Bruchus breweri Crotch Bruchus arizonensis Schaeffer Bruchus dominicanus Jekel Mylabris arizonensis: Leng Bruchus immunis Sharp Neltumius arizonensis: Bridwell Bruchus innotatus Pic Neltumius gibbithorax (Schaeffer) Bruchus inornatus Horn Bruchus gibbithorax Schaeffer Bruchus mimosae Fabricius Mylabris gibbithorax: Leng Bruchus strigatus Motschulsky Neltumius gibbithorax: Bradley Bruchus subrufus Motschulsky Neltumius gibbothorax: Kingsolver (error) Mimosestes dominicanus: Johnson and Neltumius texanus (Schaeffer) Kingsolver Bruchus texanus Schaeffer Mimosestes mimosae: Decelle Mylabris texanus: Leng Mimosestes immunis: Kingsolver Neltumius texanus: Bradley Mimosestes innotatus: Bottimer Sennius Bridwell Mimosestes inornatus: Johnson Sennius abbreviatus (Say) Mimosestes strigatus: Johnson and King- Bruchus abbreviatus Horn (validation) solver Bruchus abbreviatus Melsheimer (catalog Mylabris immunis: Leng name) Mylabris innotatus: Leng Bruchus abbreviatus Say Mimosestes nubigens (Motschulsky) Bruchus bivulneratus Horn Acanthoscelides nubigens: Blackwelder Curculio abbreviatus Say Acanthoscelides sallaei: Blackwelder Mylabris abbreviatus Dohrn: Leng (error) Bruchus nubigens Motschulsky Mylabris bivulneratus: Leng Bruchus sallaei Sharp Sennius abbreviatus: Bottimer Bruchus sallei: Pic Sennius bivulneratus: Bradley Mimosestes nubigens: Johnson and King- Sennius cruentatus (Horn) solver Acanthoscelides cruentatus: Blackwelder Mimosestes sallaei: Bridwell Bruchus cruentatus Horn Mylabris sallaei: Kannan Bruchus depressus Fall Mylabris sallei: Leng (error) Bruchus nictitans Motschulsky Mimosestes protractus (Horn) Bruchus nigrinus Horn Acanthoscelides protractus: Blackwelder Mylabris cruentatus: Leng Bruchus longiventris Sharp Mylabris nictitans: Leng Bruchus protractus Horn Mylabris nigrinus: Leng Mimosestes longiventris: Johnson and King- Sennius cruentatus: Bridwell solver Sennius depressus: Johnson Mimosestes protractus: Johnson Sennius nictitans: Bottimer Mylabris protractus: Leng Sennius nigrinus: Bradley

235 Sennius discolor (Horn) Sennius morosus: Johnson and Kingsolver Acanthoscelides infirmus:Blackwelder Sennius obesulus (Sharp) Acanthoscelides managuanus: Blackwelder Acanthoscelides obesulus: Blackwelder Bruchus discolor Horn Bruchus obesulus Sharp Bruchus discopterus Fall Sennius obesulus: Johnson and Kingsolver Bruchus infirmusSharp Sennius simulans (Schaeffer) Bruchus managuanus Pic Bruchus simulans Schaeffer Mylabris discolor: Leng Mylabris simulans: Leng Mylabris discopterus: Leng Sennius simulans: Johnson Sennius discolor: Bottimer Sennius whitei Johnson and Kingsolver Sennius discopterus: Johnson Stator Bridwell Sennius fallax (Boheman) Stator beali Johnson Acanthoscelides californicus: Bottimer Stator bottimeri Kingsolver Acanthoscelides fallax: Blackwelder Str chihuahua Johnson and Kingsolver Acanthoscelides probus: Blackwelder Stator coconino Johnson and Kingsolver Acanthoscelides xanthopus: Blackwelder Stator limbatus (Horn) Bruchus californicus Boheman Acanthoscelides cearanus: Blackwelder Bruchus fallax Boheman Acanthoscelides interruptus: Blackwelder Bruchus probus Sharp Acanthoscelides limbatus: Moreno and Bruchus xanthopus Suffrian Bibby Bruchus cearanus Sennius fallax: Johnson and Kingsolver Pic Bruchus interruptus Sennius xanthopus: Bottimer Sharp Bruchus limbatus Sennius lebasi (Fahraeus) Horn Mylabris limbatus: Acanthoscelides celatus: Blackwelder Leng Stator cearanus: Acanthoscelides lebasi: Blackwelder Kingsolver Stator limbatus: Acanthoscelides rufescens: Blackwelder Leech Stator pruininus Bruchus celatus Sharp (Horn) Acanthoscelides cognatus: Bruchus lebasi Fahraeus Blackwelder Acanthoscelides piger: Bruchus rufescens Motschulsky Blackwelder Acanthoscelides pruininus: Sennius celatus: Bottimer Blackwelder Bruchidius pruininus: Sennius lebasi: Kingsolver Herford Bruchus cognatus Sennius rufescens: Kingsolver Sharp Bruchus piger Sennius leucostauros Johnson and Kingsolver Sharp Bruchus pruininus Sennius medialis (Sharp) Horn Bruchus (Stator) pruininus: Acanthoscelides medialis: Blackwelder Zacher Mylabris pruininus: Bruchus auctus Fall Leng Stator pruininus: Bruchus medialis Sharp Bridwell Stator pygidialis Mylabris auctus: Leng (Schaeffer) Acanthoscelides pythonicus: Sennius auctus: Johnson Johnson Bruchus pygidialis Sennius medialis: Johnson and Kingsolver Schaeffer Bruchus pythonicus Sennius morosus (Sharp) Pic Mylabris pygidialis: Acanthoscelides morosus: Blackwelder Leng Mylabris pythonicus: Bruchus discolor: Fall and Cockerell (not Leng Horn) Stator pygidialis: Johnson Bruchus morosus Sharp Stator pythonicus: Johnson

236 Stator sordidus (Horn) Zabrotes humboldti Kingsolver Acanthoscelides semicolon: Blackwelder Zabrotes obliteratus Horn Acanthoscelides sordidus: Blackwelder Spermophagus obliteratus: Pic Acanthoscelides usticolor: Blackwelder Zabrotes planifrons Horn Bruchus semicolon Sharp Spermophagus planifrons: Pic Bruchus sordidus Horn Zabrotes spectabilis Horn Bruchus usticolor Sharp Spermophagus spectabilis: Pic Mylabris sordidus: Leng Zabrotes stephani Kingsolver Stator semicolon: Bottimer Zabrotes subfasciatus (Boheman) Stator sordidus: Johnson Amblycerus semifasciatus: Blackwelder Stator usticolor: Bottimer Bruchus cingulatus Suffrian Stator subaeneus (Schaeffer) Bruchus leucogaster Sharp Bruchus subaeneus Schaeffer Bruchus nesapius Fahraeus Mylabris subaeneus: Leng Spermophagus basicornis Pic Stator subaeneus: Johnson Spermophagus dorsopictus Lepesme Stator vachelliae Bottimer Spermophagus minusculus Pic (error) Bruchus sordidus: Johnson (misident.) Spermophagus musculus Boheman Stator sordidus: Johnson (misident.) Spermophagus pectoralis: Back and Duckett Stylantheus Bridwell Spermophagus (Zabrotes) pectoralis: Sharp Stylantheus macrocerus (Horn) Spermophagus semicinctus Horn Acanthoscelides macrocerus: Bridwell Spermophagus semifasciatus Boheman Acanthoscelides mucrofer Bottimer Spermophagus subfasciatus Boheman Bruchus macrocerus Horn Zabrotes obtectus: Kingsolver (error) Mylabris macrocerus: Leng Zabrotes pectoralis: Blackwelder Stylantheus macrocerus: Bridwell Zabrotes semicinctus: Bottimer Zabrotes Horn Zabrotes semifasciatus: Bottimer Zabrotes amplissimus Kingsolver Zabrotes subfasciatus: Zacher Zabrotes arenarius (Wolcott) Zabrotes subnitens Horn Bruchus arenarius Wolcott Spermophagus subnitens: Pic Megacerus arenarius: Bradley (error) Zabrotes sylvestris Romero and Johnson Mylabris arenarius: Leng Zabrotes victoriensis Kingsolver Zabrotes arenarius: Bottimer Zabrotes bexarensis Kingsolver Fossil Species Zabrotes sp.1, Johnson The geologic formation in which fossil speci- Zabrotes chandleri Kingsolver mens were found is included. Zabrotes chavesi Kingsolver Bruchus Linnaeus Zabrotes vandykei Kingsolver aboriginalis Wickham 1914:482. Florissant. Zabrotes cruciger Horn anilis Scudder 1876:82. White River. Spermophagus cruciger: Pic antaeus Wickham 1917:470. Florissant. Zabrotes cruciatus: Kingsolver (error) bowditchi Wickham 1912:30. Florissant. Zabrotes cynthiae Kingsolver carpophiloides Wickham 1914:482. Floris- sant. Zabrotes densus Horn dormescens Wickham 1913a:297. Floris- Spermophagus densus: Pic sant. Zabrotes eldenensis Kingsolver exhumatus Wickham 1912:29. Florissant. Zabrotes eldenae: Kingsolver (error)

237 henshawi Wickham 1912:28. Florissant. osborni Wickham 1912:32. Florissant. succinctus Wickham 1913b:19. Florissant. Oligobruchus Kingsolver florissantensis (Wickham) 1912:30.Floris - sant. haywardi Wickham 1912:30. Florissant. primoticus Wickham 1914:480. Florissant. scudderi Wickham 1912:31. Florissant. submersus Wickham 1914:481. Florissant. wilsoni Wickham 1913b:19. Florissant. Spermophagus Schoenherr pluto Wickham 1914:480. Florissant. vivificatus Scudder 1876:82.White River.

238 Appendix III Acanthoscelides biustulus (Fall) Desmodium spp., D. batocaulon, D. cinerascens, D. grahamii, D. neomexicanus Hosts of Bruchidae of the United Acanthoscelides calvus (Horn) States and Canada, by Bruchid Aronia sp. (floral);Helianthemum canadense Records marked with (?) are questionable Acanthoscelides chiricahuae (Fall) although they may have been listed in the Mimosa biuncifera, M. borealis, M. dysocar- literature. For more extensive host associa- pa, M. grahamii, M. laxiflora, M. wherryana; tions, especially for Latin America, see the Acacia greggi many papers by Johnson and his students Acanthoscelides compressicornis (Schaeffer) (including Center and Johnson 1973, Desmanthus sp., D. cooleyi, D. covillei, D. 1974, 1976; Forister and Johnson 1970, illinoensis, D. leptolobus, D. leptophyllus, D. 1971; Johnson and Kistler 1987; Johnson obtusus, D. subulatus, D. velutinus, D. vir- and Siemans 1995a,b, 1997a,b; Nilsson gatus acuminatus, D. virgatus depressus, D. and Johnson, 1990, 1993a,b). virgatus glandulosus, D. virgatus virgatus; For author names of host plants, see Hoffmanseggia densiflora, H. drepanocarpa, H. glauca, H. tenella; Mimosa dysocarpa, M. appendix IV. grahamii Abutiloneus idoneus Bridwell Acanthoscelides comstocki Johnson Abutilon abutiloides, A. berlandieri; Allowis- Eysenhardtia texana sadula holosericea Acanthoscelides daleae Johnson Acanthoscelides aequalis (Sharp) Dalea sp.; Psorothamnus fremontii, P. schot- Abutilon berlandieri; Allowissadula holoseri- ti, P. spinosa cea, A. lozani Acanthoscelides desmanthi Johnson Acanthoscelides alboscutellatus (Horn) Desmanthus sp., D. bicornutus, D. covillei, Daucus carota (floral record);Glycyrrhiza D. leptophyllus, D. subulatus, D. virgatus lepidota; Ludwigia alternifolia, L. palustris depressus, D. virgatus virgatus Acanthoscelides atomus (Fall) Acanthoscelides distinguendus (Horn) Hudsonia ericoides (floral record);Lechea Mimosa strigillosa; Rhynchosia americana, racemulosa, L. tenuifolia R. difformis, R. erecta, R. galactoides, R. Acanthoscelides aureolus (Horn) complex intermedia, R. latifolia, R. tomentosa; Teph- Astragalus allochrous, A. amphioxys, A. rosia virginiana asymmetricus, A. bisulcatus, A. calycosus, Acanthoscelides flavescens (Fahraeus) A. crotalariae, A. douglasii, A. drummon- Abutilon hypoleucum; Eriosema violaceum; dii, A. fucatus, A. grayi, A. humistratus, A. Galactia striata; Rhynchosia longeracemosa, lancearius, A. lentiginosus, A. mollissimus R. minima; Vicia sp. mollissimus, A. oxyphysus, A. pattersonii, A. praelongus, A. lonchocarpus, A. pterocarpus, Acanthoscelides floridae (Horn) A. racemosus, A. thurberi, A. trichopodus Amorpha fruticosa fruticosa antisellii, A. trichopodus phoxus, A. trichopo- Acanthoscelides fraterculus (Horn) dus trichopodus, A. wootoni; Glycyrrhiza Abrus precatorius, Astragalus bisulcatus, lepidota; Lotus purshianus, L. scoparius; A. cibarius, A. crassicarpus crassicarpus, Oxytropis lambertii, O. sericea A. crassicarpus paysonii, A. drummondii, A. Acanthoscelides baboquivari Johnson falcatus, A. hyalinus, A. missouriensis, A. Indigofera platycarpa, I. sphaerocarpa mollissimus bigelovii, A. mollissimus mol- Acanthoscelides bisignatus (Horn) lissimus, A. pattersonii, A. pectinatus, A. pterocarpus, A. racemosus, A. utahensis; Baptisia sp.; Desmanthus sp., D. illinoen- Glycyrrhiza lepidota; Hedysarum boreale; sis, D. leptolobus, D. velutinus, D. virgatus Lotus crassifolius, L. mearnsii, L. rigidus, L. virgatus scoparius

239 Acanthoscelides fumatus (Schaeffer) Albizia sp.; Cajanus cajan; Cicer arietinum; Rhynchosia texana Daucus carota (floral); Dolichos melanophthalmus (doubtful); Glycine max; Lablab Acanthoscelides griseolus (Fall) purpureus; Lens culinaris; Lathyrus sativus; Sesbania emerus, S. exaltata Lupinus albus; Phaseolus acutifolius latifo- Acanthoscelides helianthemum Bottimer lius, P. coccineus, P. lunatus, P. ritensis, P. Helianthemum corymbosum vulgaris; Pisum sativum arvense; Sesbania aegyptiaca, S. sesban; Vicia faba; Vigna Acanthoscelides herissantitus Johnson aconitifolia, V. radiata radiata, V. subterra- Herissantia crispa; Malvastrum bicuspida- nea, V. umbellata, V. unguiculata sesquipe- tum dalis, V. unguiculata unguiculata; Zea mays Acanthoscelides inquisitus (Fall) (doubtful) Unknown Acanhoscelides oregonensis Johnson Acanthoscelides kingsolveri Johnson Dalea ornata Indigofera lindheimeriana, I. sphaerocarpa, Acanthoscelides pallidipennis (Motschulsky) I. suffruticosa Amorpha californica, A. canescens, A. fru- Acanthoscelides lobatus (Fall) ticosa fruticosa; Astragalus sp.; Dalea sp.; Astragalus humistratus, A. lentiginosus, A. Desmanthus virgatus acuminatus, D. virga- mollissimus bigelovii, A. mollissimus marci- tus virgatus; Errazurizia rotundata; Glycyr- dus, A. mollissimus mollissimus, A. troglody- rhiza sp.; Lotus sp.; Parryella filifolia tus Acanthoscelides pauperculus (LeConte) Acanthoscelides longistilus (Horn) Trifolium obtusiflorum Desmodium illinoense; Lespedeza capitata, Acanthoscelides pectoralis (Horn) L. frutescens, L. hirta, L. intermedia, L. Desmanthus sp., D. leptolobus, D. obtusus, texana, L. virginica D. reticulatus, D. velutinus, D. virgatus Acanthoscelides macrophthalmus (Schaeffer) virgatus Leucaena leucocephala, L. pulverulenta, L. Acanthoscelides pedicularius (Sharp) retusa Dalea carnea albida Acanthoscelides margaretae Johnson Acanthoscelides perforatus (Horn) Astragalus pycnostachys Astragalus canadensis Acanthoscelides mixtus (Horn) Acanthoscelides prosopoides (Schaeffer) Astragalus allochrous, A. lentiginosus, A. Condalia lycioides; Ziziphus obtusifolia praelongus, A. lonchocarpus, A. sabulonum, Acanthoscelides pullus (Fall) A. thurberi, A. wootoni; Eysenhardtia texana Astragalus allochrous, A. asymmetricus, Acanthoscelides modestus (Sharp) A. bolanderi, A. crotalariae, A. douglasii, Aeschynomene histrix incana, A. indica. In A. lentiginosus, A. oxyphysus, A. praelon- other species of Aeschynomene in Mexico gus, A. thurberi, A. trichopodus antisellii, A. and Central America trichopodus phoxus, A. wootoni Acanthoscelides mundulus (Sharp) Acanthoscelides pusillimus (Sharp) Nissolia schottii, N. wislizenii Dalea spp., Dalea scandens paucifolia; Acanthoscelides napensis Johnson Trifolium pratense (Trifolium records are Malacothrix incana, M. saxatilis implicata suspect) (all floral records) Acanthoscelides quadridentatus (Schaeffer) Acanthoscelides obrienorum Johnson Mimosa pigra, Mimosa pigra berlandieri, M. Senna armata, S. polyantha, S. wislizenii strigillosa wislizenii Acanthoscelides rufovittatus (Schaeffer) Acanthoscelides obsoletus (Say) Galactia wrightii; Tephrosia cinerea, T. Tephrosia ambigua, T. spicata, T. virginiana purpurea, T. thurberi Acanthoscelides obtectus (Say) Acanthoscelides schaefferi (Pic) Unknown

240 Acanthoscelides schrankiae (Horn) Senna corymbosa. Outside the United Acacia amentacea; Desmanthus virgatus States: Senna alata, S. bicapsularis bicapsu- acuminatus; Mimosa borealis; Schrankia laris, S. occidentalis, S. splendida, S. uni- microphylla, S. nuttallii, S. roemeriana, S. flora; Caesalpinia spp.; Cassia spp. uncinata Amblycerus obscurus (Sharp) Acanthoscelides seminulum (Horn) U.S. host unknown. Associated with Senna Dalea aurea, D. candida, D. enneandra, D. alata, S. bicapsularis bicapsularis, S. hirsuta feayi, D. frutescens, D. stanfieldii, D. tenuis leptocarpa, S. obtusifolia, S. pendula advena, S. pendula ovalifolia, and S. uniflora Acanthoscelides speciosus (Schaeffer) south of the Rio Grande Mimosa biuncifera, M. malacophylla, M. Amblycerus robiniae (Fabricius) wherryana Acacia farnesiana(?); Gleditsia aquatica, G. Acanthoscelides stylifer (Sharp) triacanthos; Robinia pseudoacacia (prob- Desmodium sp. D. grahamii able misidentification);Tillandsia usneoides Acanthoscelides subaequalis Johnson (hibernaculum) Abutilon abutiloides, A. berlandieri, A. inca- Amblycerus schwarzi Kingsolver num, A. trisulcatum U.S.A.: Hippomane mancinella. West In- Acanthoscelides submuticus (Sharp) dies: Guettarda sp.; Hippomane mancinella; Amorpha californica, A. fruticosa angustifo- Tectona grandis; Terminalia catappa; Ricinis lia, A. fruticosa fruticosa, A. fruticosa occi- communis dentalis Amblycerus vitis (Schaeffer) Acanthoscelides tenuis Bottimer Vitis arizonica Lythrum alatum, L. lineare Borowiecius ademptus (Sharp) Acanthoscelides tridenticulatus Bottimer Pueraria montana lobata Mimosa strigillosa Bruchidius cisti (Fabricius) Algarobius bottimeri Kingsolver Cynoglossum sp.; Onobrychis viciifolia; Prosopis glandulosa glandulosa, P. pallida, Spartium junceum P. reptans reptans, P. reptans cinerascens Bruchidius villosus (Fabricius) Algarobius prosopis (LeConte) Cytisus scoparius; Laburnum alpinum, L. Prosopis alba (introduced), P. chilensis, P. anagyroides; Petteria ramentacea; Spartium glandulosa torreyana, P. laevigata, P. pubes- junceum cens, P. reptans cinerascens, P. velutina Bruchus brachialis Fahraeus Althaeus folkertsi Kingsolver Daucus carota (floral record); Lathyrus Abutilon theophrasti; Hibiscus militaris, H. sativus; Vicia benghalensis, V. caroliniana, moscheutos lasiocarpus V. cracca, V. grandiflora, V. pannonica, V. sativa nigra, V. sativa sativa, V. sepium, V. Althaeus hibisci (Olivier) villosa dasycarpa, V. villosa villosa Abutilon theophrasti; Daucus carota (floral Bruchus pisorum (Linnaeus) record); Hibiscus aculeatus, H. moscheutos moscheutos, H. moscheutos lasiocarpus, H. Lathyrus sativus; Pisum elatius, P. sativum moscheutos palustris; Kosteletzkya virginica arvense; Vicia faba(?), Vicia sp.; Vigna radiata radiata Althaeus steineri Kingsolver Bruchus rufimanus Boheman Kosteletzkya virginica Vicia faba, V. sativa sativa Amblycerus eustrophoides (Schaeffer) Callosobruchus chinensis (Linnaeus) Drypetes lateriflora Cajanus cajan; Cicer arietinum; Cyamopsis Amblycerus ireriae Romero, Johnson, and Kingsolver tetragonoloba; Glycine max; Lablab purpureus; Lathyrus sativus; Lens culinaris; Host unknown Nelumbo nucifera; Phaseolus lunatus, P. Amblycerus nigromarginatus (Motschulsky) vulgaris; Pisum sativum arvense; Psophoc- arpus tetragonolobus; Vicia faba; Vigna

241 aconitifolia, V. angularis, V. mungo, V. Gibbobruchus mimus (Say) radiata radiata, V. unguiculata unguiculata Bauhinia lunarioides; Cercis canadensis, C. Callosobruchus maculatus (Fabricius) occidentalis; Fraxinus sp. (floral);Magnolia ßCajanus cajan; Cicer arietinum; Glycine sp. (floral),M. grandiflora (floral record) max; Lablab purpureus; Lathyrus aphaca, Kytorhinus prolixus (Fall) L. clymenum, L. sativus; Lens culinaris; Hedysarum boreale, H. alpinum americanum Phaseolus acutifolius, P. lunatus, P. vulgaris; Lithraeus atronotatus (Pic) Pisum sativum arvense; Vicia faba, V. lutea, V. sativa sativa; Vigna mungo, V. radiata Lithraea brasiliensis; Schinus radiata, V. unguiculata unguiculata terebinthifolius Callosobruchus phaseoli (Gyllenhal) Megacerus coryphae (Olivier) Cajanus cajan; Cicer arietinum; Crotalaria Ipomoea pes-caprae, I. sagittata spectabilis; Lablab purpureus; Phaseolus Megacerus cubiculus (Casey) lunatus, P. vulgaris; Pisum sativum arvense; Baccharis sarothroides (floral record); Sesbania sp.; Vicia faba; Vigna angularis, V. Convolvulus sp.; Ipomoea batatas, I. lacuno- radiata radiata sa, I. purpurea, I. trifida; Merremia quinque- Callosobruchus pulcher Pic folia; Senecio salignus (floral record) Cajanus cajan; Cicer arietinum; Phaseolus Megacerus cubicus (Motschulsky) vulgaris; Vigna angularis Ipomoea sp. Extralimital: Argyreia nervosa; Caryedes helvinus (Motschulsky) Calystegia sepium;, Ipomoea arborescens, I. Centrosema macrocarpa, C. pubescens carnea fistulosa, I. crassicaulis, I. hederifolia, I. leptophylla, I. meyeri, I. nil. Caryedes incensus (Sharp) Megacerus discoidus (Say) Centrosema macrocarpa, C. pubescens Calystegia sepium, C. sepium repens, C. Caryedon serratus (Olivier) macounii; Convolvulus arvensis; Ipomoea Acacia farnesiana, A. seyal, A. spirocarpa, leptophylla, I. pandurata, I. tiliacea; Hibis- A. tortilis; Adenanthera pavonina; Arachis cus sp. and Daucus carota are floral records hypogaea; Bauhinia malabarica, B. monan- Megacerus impiger (Horn) dra, B. racemosa, B. reticulata, B. rufescens, B. thonningi, B. tomentosa; Caesalpinia Calystegia atriplicifolia, C. occidentalis ful- pulcherrima; Cajanus cajan; Cassia afrofis- crata, C. occidentalis occidentalis, C. lon- tula, C. arereh, C. fistula, C. grandis, C. gipes, C. macrostegia arida, C. macrostegia javanica javanica, C. javanica indochinensis, cyclostegia, C. macrostegia macrostegia, C. C. sieberiana; Phaseolus vulgaris; Prosopis malacophylla deltoidea, C. malacophylla glandulosa glandulosa, P. pallida; Senna malacophylla, C. malacophylla pedicellata, obtusifolia; Tamarindus indica C. piersonii, C. purpurata, C. soldanella, C. subacaulis, C. stebbensii; Convolvulus Caryobruchus gleditsiae (Linnaeus) arvensis; Ipomoea sp. Cocothrinax argentata, C. martii; Livingstona Megacerus leucospilus (Sharp) chinensis; Phoenix sylvestris; Sabal etoni, S. glaucescens, S. mauritiaeformis, S. mexi- Ipomoea biloba(?), I. cairica, I. carnea fistu- cana, S. minor, S. palmetto, S. parviflora, S. losa, I. crassicaulis, I. leptophylla, I. pes-cap- uresana, S. yapa; Serenoa repens; Tilland- rae; Merremia aegyptia (Hawaii) sia usneoides (hibernaculum); Washingto- Megacerus maculiventris (Fahraeus) nia filifera Ipomoea aristolochiaefolia, I. marginisepala, Gibbobruchus cristicollis (Sharp) I. nil, I. pes-caprae, I. purpurea, I. rubriflora Bauhinia lunarioides (Texas), B. coulteri (all extralimital) (Mexico), B. divaricata, B. macranthera, B. Megacerus ripiphorus (Fahraeus) pauletia (Mexico) Ipomoea alba Gibbobruchus divaricatae Whitehead and King- Megacerus schaefferianus Bridwell solver Ipomoea longifolia Bauhinia divaricata (Texas), B. pauletia, B. pes- Meibomeus desmoportheus Kingsolver and caprae, B. ungulata (Mexico) Whitehead

242 Desmodium grahamii Acacia farnesiana; Prosopis glandulosa Meibomeus musculus (Say) glandulosa, P. juliflora, P. pallida Desmodium canescens, D. tenuifolium, D. Mimosestes mimosae (Fabricius) tortuosum, D. triflorum; Lespedeza hirta Florida: Caesalpinia coriaria. Extralimital: Meibomeus surrubresus (Pic) Acacia sp., A. cochliacantha, A. cymbispina, A. farnesiana, A. globulifera, A. hindsii, A. Aeschynomene americana macracantha, A. pennatula; Caesalpinia Merobruchus insolitus (Sharp) spp. C. coriaria, C. sclerocarpa; Ceratonia Lysiloma microphylla thornberi; Pithecello- siliqua; Cordia sp. (questionable); Lotus sp. bium pallens Mimosestes nubigens (Motschulsky) Merobruchus julianus (Horn) Rearing records: Acacia cornigera, A. farne- Acacia berlandieri, A. coulteri (extralimital), siana, A. schaffneri, A. tortuosa; Prosopis A. greggi, A. juncifolia, A. rigidula, A. roeme- pallida (Hawaii). Records needing confirma- riana, A. wrightii; Desmanthus illinoensis tion: Acacia amentacea; Caesalpinia cori- (questionable); Mimosa borealis (question- aria; Ceratonia siliqua; Cordia sp.; Gleditsia able); Pithecellobium ebano triacanthos; Lotus sp.; Prosopis chilensis, P. Merobruchus knulli (White) glandulosa glandulosa, P. glandulosa torreyana, P. juliflora Lysiloma microphylla thornberi; L. acapulcensis and L. watsoni (extralimital) Mimosestes protractus (Horn) Merobruchus lysilomae Kingsolver Parkinsonia microphylla, P. texana texana; Prosopis glandulosa glandulosa, P. glandu- Acacia richei, A. simplicifolia; Albizia leb- losa torreyana, P. laevigata, P. velutina beck, A. polyphylla; Lysiloma latisiliqua, L. sabicu (extralimital) Mimosestes ulkei (Horn) Merobruchus major (Fall) Parkinsonia aculeata, P. florida, P. texana texana Pithecellobium ebano Neltumius arizonensis (Schaeffer) Merobruchus placidus (Horn) Chilopsis linearis (floral record);Larrea Acacia angustissima angustissima, A. an- tridentata (floral record);Prosopis glandu- gustissima hirta, A. angustissima suffrutes- losa glandulosa, P. glandulosa torreyana, P. cens, A. angustissima texensis; Astragalus velutina sp. Neltumius gibbithorax (Schaeffer) Merobruchus terani Kingsolver Pluchea sericea (probably floral);Prosopis glan- Host in United States unknown; Acacia spp. dulosa glandulosa, P. glandulosa torreyana, south of Rio Grande P. pubescens Merobruchus vacillator (Sharp) Neltumius texanus (Schaeffer) Host in United States unknown; Lysiloma Condalia correllii, C. globosa globosa, C. divaricatum in Mexico globosa pubescens, C. hookeri hookeri, C. Mimosestes acaciestes Kingsolver and Johnson obovata, C. spathulata, C. warnockii kear- Acacia amentacea, A. berlandieri, A. con- neyana; Zanthoxylum clava-herculis (floral stricta, A. neovernicosa, A. rigidula, A. verni- record) cosa Sennius abbreviatus (Say) Mimosestes amicus (Horn) Cassia sp.; Senna marilandica. Floral re- Acacia constricta, A. farnesiana, A. glauca; cords: Asclepias syriaca; Cicuta maculata; Leucaena leucocephala; Parkinsonia acu- Cryptotaenia canadensis; Eupatorium sp.; leata, P. florida, P. microphylla, P. texana Taenidia integerrima macra, P. texana texana; Prosopis glandu- Sennius cruentatus (Horn) losa glandulosa, P. glandulosa torreyana, P. Chamaecrista fasciculata, C. nictitans pallida, P. pubescens, P. velutina; Sesbania nictitans; Parkinsonia sp.; Tillandsia us- aegyptiaca, S. sesban; Lippia wrightii (floral record) neoides (hibernaculum) Mimosestes insularis Kingsolver and Johnson

243 Sennius discolor (Horn) Glycine max; Leucaena leucocephala, L. “Cassia sp.”; Parkinsonia sp.(?); Prosopis pulverulenta; Lysiloma microphylla thorn- sp.(?); Senna lindheimeriana, S. roemeriana, beri; Neptunia plena; Parkinsonia aculeata, S. wislizenii wislizenii P. florida, P. microphylla, P. texana macra, P. texana texana; Pithecellobium brevifolium, Sennius fallax (Boheman) P. dulce, P. ebano, P. pallens, P. saman, P. Senna bicapsularis bicapsularis, S. biflora, unguis-cati; Prosopis glandulosa glandulosa, S. obtusifolia, S. occidentalis P. glandulosa torreyana, P. juliflora Sennius lebasi (Fahraeus) Stator pruininus (Horn) Senna bicapsularis bicapsularis Acacia berlandieri, A. california, A. confusa, Sennius leucostauros Johnson and Kingsolver A. constricta, A. dealbata, A. decurrens, A. Senna bicapsularis bicapsularis greggi, A. koa, A. mearnsii, A. melanoxylon, A. neovernicosa, A. richei, A. rigidula, A. roe- Sennius medialis (Sharp) meriana, A. schotti, A. vernicosa, A. wrightii; Senna hirsuta leptocarpa, S. hirsuta hirsuta Albizia saponaria; Arachis hypogeae; Cae- (Mexico) salpinia pulcherrima; Calliandra eriophylla, Sennius morosus (Sharp) C. humilis humilis, C. humilis reticulata; Senna bauhinioides, S. covesii, S. durangen- Cassia fistula, C. grandis, C. javanica indo- sis durangensis, S. hirsuta leptocarpa, S. chinensis, C. javanica javanica; Coursetia occidentalis (extralimital), S. obtusifolia, S. microphylla; Desmanthus bicornutus, D. roemeriana cooleyi, D. covillei, D. fruticosa, D. leptolobus, D. virgatus virgatus; Desmodium uncina- Sennius obesulus (Sharp) tum; Erythrina sandwicensis; Glycine max; Chamaecrista serpens wrightii Indigofera anil, I. suffruticosa; Leucaena leu- Sennius simulans (Schaeffer) cocephala; Mimosa biuncifera, M. dysocarpa, Chamaecrista nictitans mensalis M. grahamii, M. laxiflora, M. wherryana; Neptunia plena; Olneya tesota; Parkinsonia Sennius whitei Johnson and Kingsolver aculeata, P. texana macra, P. texana texana; Host plant unknown Pithecellobium ebano, P. pallens, P. saman; Stator beali Johnson Prosopis glandulosa glandulosa, P. juliflora, Pithecellobium ebano P. velutina; Robinia pseudoacacia; Scaevola taccada sericea; Senna siamea; Sesbania Stator bottimeri Kingsolver emerus, S. exaltata, S. macrocarpa, S. ses- Acacia farnesiana, A. pinetorum ban; Sophora chrysophylla Stator chihuahua Johnson and Kingsolver Stator pygidialis (Schaeffer) Acacia angustissima angustissima, A. Calliandra humilis humilis, C. humilis reticu- constricta; Calliandra eriophylla, C. humilis lata humilis; Lysiloma microphylla thornberi; Stator sordidus (Horn) Mimosa biuncifera, M. wherryana Acacia wrightii; Calliandra humilis humilis Stator coconino Johnson and Kingsolver Stator subaeneus (Schaeffer) Host plant unknown Acacia cornigera, A. farnesiana Stator limbatus (Horn) Stator vachelliae Bottimer Acacia acatlensis, A. angustissima angustissima, A. baileyana, A. berlandieri, A. confu- Acacia farnesiana sa, A. cultriformis, A. farnesiana, A. greggi, Stylantheus macrocerus (Horn) A. koa, A. melanoxylon, A. millefolia, A. retin- Stylosanthes biflora odes, A. richei, A. roemeriana, A. wrightii; Zabrotes amplissimus Kingsolver Albizia lebbeck, A. saponaria; Arachis hypogaea (exp.) Calliandra eriophylla, C. humilis Phaseolus ritensis humilis, C. humilis reticulata; Cassia fistula, Zabrotes arenarius (Wolcott) C. grandis, C. javanica indochinensis, C. Chamaecrista fasciculata (sweep record) javanica javanica; Caesalpinia pulcherrima; Delonix regia(?); Desmanthus bicornutus; Erythrina sandwicensis;

244 Zabrotes bexarensis Kingsolver Vicia sp., Vicia leavenworthii Zabrotes chandleri Kingsolver Host plant unknown Zabrotes chavesi Kingsolver Senna hirsuta leptocarpa, S. spectabilis (Venezuela) Zabrotes cruciger Horn Host plant unknown Zabrotes cynthiae Kingsolver Host plant unknown. Asclepias sp. (floral record); Cupressus macnabiana (floral record); Eremocarpus sp. (sweep record); Lotus sp. (floral record);Quercus sp. (floral record) Zabrotes densus Horn Host plant unknown Zabrotes eldenensis Kingsolver Host plant unknown Zabrotes humboldtae Kingsolver Host plant unknown Zabrotes obliteratus Horn Rhynchosia sp. Zabrotes planifrons Horn Chamaecrista nictitans mensalis Zabrotes spectabilis Horn Senna bauhinioides, S. covesii, S. durangensis durangensis, S. lindheimeriana, S. roemeriana Zabrotes stephani Kingsolver Host plant unknown Zabrotes subfasciatus (Boheman) Cajanus cajan; Cicer arietinum; Dipogon lignosus; Glycine max; Lablab purpureus; Phaseolus acutifolius, P. angularis angularis, P. coccineus, P. lunatus, P. vulgaris; Pisum sativum arvense (experimental); Vicia faba; Vigna angularis, V. mungo, V. subterranea, V. unguiculata unguiculata Zabrotes subnitens Horn Host plant unknown Zabrotes sylvestris Romero and Johnson Phaseolus vulgaris Zabrotes victoriensis Kingsolver Type specimen collected on Zanthoxylum clava-herculis, but this is an unlikely host plant.

245 Appendix IV Cistaceae Helianthemum canadense (Linnaeus) Michx. Hosts of Bruchidae of the United Acanthoscelides calvus States and Canada, by Host Plant Helianthemum corymbosum Michx. Acanthoscelides helianthemum Host names were updated according to Hudsonia ericoides Linnaeus the National List of Scientific Plant Names (U.S. Department of Agriculture 1982). Acanthoscelides atomus (floral record) Other sources of plant names consulted Combretaceae were Burkart (1952), Irwin and Barneby Terminalia catappa Linnaeus (1982), Isely (1973, 1975, 1990), Shetler and Skog (1978), Terrell et al. (1986), and Amblycerus schwarzi (extralimital) Wiersema et al. (1990). Extralimital host Commelinaceae plants are included for the cosmopolitan Lechea racemulosa Michx. Bruchidae in the genera Acanthoscelides, Bruchidius, Bruchus, Callosobruchus, Car- Acanthoscelides atomus eydon, and Zabrotes. For author names of Lechea tenuifolia Michx bruchid species, see appendix III. Acanthoscelides atomus Anacardiaceae Compositae Lithraea brasiliensis March. Baccharis sarothroides Gray Lithraeus atronotatus Megacerus cubiculus (floral record) Schinus terebinthifolius Raddi Eupatorium sp. Lithraeus atronotatus Sennius abbreviatus (floral record) Asclepediaceae Malacothrix incana (Nutt.) Torr. and Gray Asclepias sp. Acanthoscelides napensis (sweep record) Zabrotes cynthiae (floral record) Malacothrix saxatilis implicata (Eastw.) Hall Asclepias syriaca Linnaeus Acanthoscelides napensis (sweep record) Sennius abbreviatus (floral record) Pluchea sericea (Nutt.) Coville Neltumius gibbithorax (floral record) Apiaceae Senecio salignus DC. Cicuta maculata Linnaeus Megacerus cubiculus (floral record) Sennius abbreviatus (floral record) Cryptotaenia canadensis (Linnaeus) DC. Convolvulaceae Sennius abbreviatus (floral record) Argyreia nervosa (Burm.) Bojer Megacerus cubicus (extralimital) Bignoniaceae Calystegia atriplicifolia H. Hallier Chilopsis linearis (Cav.) Sweet Megacerus impiger Neltumius arizonensis (floral record) Calystegia longipes (S. Wats.) Brumm. Boraginaceae Megacerus impiger Cynoglossum sp. Calystegia macounii (Greene) Brumm. Bruchidius cisti Megacerus discoidus Bromeliaceae Calystegia macrostegia arida (Greene) Brumm. Tillandsia usneoides (Linnaeus) Linnaeus Megacerus impiger (hibernaculum) Calystegia macrostegia macrostegia (Greene) Amblycerus robiniae; Caryobruchus gledit- Brumm. siae; Sennius cruentatus Megacerus impiger

246 Calystegia macrostegia cyclostegia (House) Ipomoea biloba Forsk. Brumm. Megacerus leucospilus (questionable) Megacerus impiger Ipomoea cairica (Linnaeus) Sweet Calystegia malacophylla deltoidea (Greene) Megacerus leucospilus Megacerus impiger Ipomoea carnea fistulosaK. Martin Calystegia malacophylla malacophylla (Greene) Megacerus cubicus (extralimital), Megacerus Munz leucospilus Megacerus impiger Ipomoea crassicaulis (Benth.) B. Robins. Calystegia malacophylla pedicellata (Jeps.) Megacerus cubicus (extralimital), M. Munz leucospilus Megacerus impiger Ipomoea hederifolia Linnaeus Calystegia occidentalis fulcrata (Gray) Brumm. Megacerus cubicus (extralimital) Megacerus impiger Ipomoea lacunosa Linnaeus Calystegia occidentalis occidentalis (Gray) Megacerus cubiculus Brumm. Ipomoea leptophylla J. Torrey Megacerus impiger Megacerus cubicus (extralimital), M. Calystegia piersonii (Abrams) Brumm. discoidus, M. leucospilus Megacerus impiger Ipomoea longifolia Benth. Calystegia purpurata (Greene) Brumm Megacerus schaefferianus Megacerus impiger Ipomoea marginisepala O’Donnell Calystegia sepium (Linnaeus) R. Brown Megacerus maculiventris Megacerus cubicus (extralimital), M. Ipomoea meyeri (Sprengler) Dow discoidus Megacerus cubicus (extralimital) Calystegia sepium repens (Linnaeus) Gray Ipomoea nil (Linnaeus) Roth Megacerus discoidus Megacerus cubicus (extralimital), M. Calystegia soldanella (Linnaeus) R. Brown maculiventris (extralimital) Megacerus impiger Ipomoea pandurata (Linnaeus) G. Meyer Calystegia stebbensii Brumm. Megacerus discoidus Megacerus impiger Ipomoea pes-caprae (Linnaeus) R. Brown. Calystegia subacaulis Hook. and Arn. Megacerus coryphae, M. leucospilus, M. Megacerus impiger maculiventris (extralimital) Convolvulus sp. Ipomoea purpurea (Linnaeus) Roth Megacerus cubiculus Megacerus cubiculus, M. maculiventris Convolvulus arvensis Linnaeus (extralimital) Megacerus discoidus, M. impiger Ipomoea rubrifloraO’Donnell Ipomoea sp. Megacerus maculiventris Megacerus cubicus, M. impiger Ipomoea sagittata Poir. Ipomoea alba Linnaeus Megacerus coryphae Megacerus ripiphorus Ipomoea tiliacea (Willd.) Choisy Ipomoea arborescens (Sweet) Don. Megacerus discoidus Megacerus cubicus (Mexico) Ipomoea trifida(H.B.K.) G. Don Ipomoea aristolochiaefolia (H.B.K.) G. Don Megacerus cubiculus Megacerus maculiventris (extralimital) Merremia aegyptia Linnaeus Ipomoea batatas (Linnaeus) Lam. Megacerus leucospilus Megacerus cubiculus Merremia quinquefolia Linnaeus Megacerus cubiculus

247 Cupressaceae Acacia angustissima suffrutescens (Rose) Isely Cupressus macnabiana MacMurray Merobruchus placidus Zabrotes cynthiae (floral record) Acacia angustissima texensis (Torr. and Gray) Isely Ehretiaceae Merobruchus placidus Cordia sp. Acacia baileyana F. Mueller Mimosestes mimosae (questionable), M. Stator limbatus nubigens Acacia berlandieri Benth. Euphorbiaceae Merobruchus julianus; Mimosestes Drypetes lateriflora(Swartz) Krug and Urban acaciestes; Stator limbatus, S. pruininus Amblycerus eustrophoides Acacia californica T.S. Brand. Eremocarpus sp. Stator pruininus Zabrotes cynthiae (sweep record) Acacia cochliacantha Humb. and Bonpl. ex Hippomane mancinella Linnaeus Willd. Amblycerus schwarzi Mimosestes mimosae Ricinis communis Linnaeus Acacia confusa Merr. Amblycerus schwarzi (extralimital) Stator limbatus, S. pruininus Fagaceae Acacia constricta Benth. Quercus sp. Mimosestes acaciestes, M. amicus; Stator chihuahua, S. pruininus Zabrotes cynthiae (floral record) Acacia cornigera (Linnaeus) Willd. Goodeniaceae Mimosestes nubigens; Stator subaeneus Scaevola taccada sericea (Vahl) H. St. John Acacia coulteri Benth. Stator pruininus Merobruchus julianus Graminae Acacia cultriformis A. Cunningham ex G. Don Zea mays Linnaeus Stator limbatus Acanthoscelides obtectus Acacia cymbispina Sprague and Riley Leguminosae Mimosestes mimosae Abrus precatorius Linnaeus Acacia dealbata Link Acanthoscelides fraterculus Stator pruininus Acacia spp. Acacia decurrens Willd. Merobruchus terani (extralimital); Stator pruininus (experimental) Mimosestes mimosae (extralimital) Acacia farnesiana (Linnaeus) Willd. Acacia acatlensis Benth. Amblycerus robiniae (questionable); Caryedon serratus; Mimosestes amicus, Stator limbatus M. insularis, M. mimosae, M. nubigens; Acacia amentacea DC. Stator bottimeri, S. limbatus, S. subaeneus, Acanthoscelides schrankiae; Mimosestes S. vachelliae acaciestes, M. nubigens Acacia glauca (Linnaeus) Moench Acacia angustissima angustissima (Mill.) Mimosestes amicus Kuntze Acacia globulifera Safford Merobruchus placidus; Stator chihuahua, S. Mimosestes mimosae limbatus Acacia greggi Gray Acacia angustissima hirta (Nutt.) B. Rob. Acanthoscelides chiricahuae; Merobruchus Merobruchus placidus julianus; Stator limbatus, S. pruininus Acacia hindsii Benth. Mimosestes mimosae

248 Acacia juncifolia Benth. Acacia wrightii Benth. Merobruchus julianus Merobruchus julianus; Stator limbatus, S. Acacia koa Gray pruininus, S. sordidus Stator limbatus, S. pruininus Adenanthera pavonina Linnaeus Acacia macracantha Humb. and Bonpl. Caryedon serratus Mimosestes mimosae Aeschynomene spp. Acacia mearnsii Willd. Acanthoscelides modestus Stator pruininus Aeschynomene americana Linnaeus Acacia melanoxylon R. Brown Meibomeus surrubresus Stator limbatus, S. pruininus Aeschynomene histrix incana (Vogel) Benth. Acacia millefolia S. Watson Acanthoscelides modestus Stator limbatus Aeschynomene indica Linnaeus Acacia neovernicosa Isely Acanthoscelides modestus Mimosestes acaciestes; Stator pruininus Albizia sp. Acacia pennatula (Schlect. and Cam.) Benth. Acanthoscelides obtectus Mimosestes mimosae Albizia lebbeck (Linnaeus) Benth. Acacia pinetorum F.J. Hermann Merobruchus lysilomae; Stator limbatus Stator bottimeri Albizia polyphylla Fourn. Acacia retinodes Schlect. Merobruchus lysilomae Stator limbatus Albizia saponaria (Lour.) Blume Acacia richei Gray Stator limbatus, S. pruininus Merobruchus lysilomae; Stator limbatus, S. Amorpha californica Nuttall pruininus Acanthoscelides pallidipennis, A. Acacia rigidula Benth. submuticus Merobruchus julianus; Mimosestes Amorpha canescens Pursh. acaciestes; Stator pruininus Acanthoscelides pallidipennis Acacia roemeriana Scheele Amorpha fruticosa fruticosa Linnaeus Merobruchus julianus; Stator limbatus, S. Acanthoscelides floridae, A. pallidipennis, A. pruininus submuticus Acacia schaffneri (S. Wats.) F.J. Herm. Amorpha fruticosa angustifolia Pursh. Mimosestes nubigens Acanthoscelides submuticus Acacia schotti Torr. Amorpha fruticosa occidentalis (Abrams) Kearn. Stator pruininus and Peebl. Acacia seyal Delile Acanthoscelides submuticus Caryedon serratus Arachis hypogaea Linnaeus Acacia simplicifolia (Linnaeus) Druce Caryedon serratus; Stator limbatus, S. pruininus (experimental) Merobruchus lysilomae Astragalus sp. Acacia spirocarpa Hochst. ex A. Rich. Acanthoscelides pallidipennis; Merobruchus Caryedon serratus placidus Acacia tortilis (Forsskal) Hayne Astragalus allochrous Gray Caryedon serratus Acanthoscelides aureolus complex, A. Acacia tortuosa (Linnaeus) Willd. mixtus, A. pullus Mimosestes nubigens Astragalus amphioxys Gray Acacia vernicosa Standley Acanthoscelides aureolus complex Mimosestes acaciestes; Stator pruininus Astragalus asymmetricus Sheldon Acanthoscelides aureolus complex, A. pullus 249 Astragalus bisulcatus (Hook.) Gray Astragalus mollissimus marcidus (Greene ex Acanthoscelides aureolus complex, A. Rydb.) Barneby fraterculus Acanthoscelides lobatus Astragalus bolanderi Gray Astragalus mollissimus mollissimus Torrey Acanthoscelides pullus Acanthoscelides aureolus complex, A. Astragalus calycosus J. Torr. fraterculus, A. lobatus Acanthoscelides aureolus complex Astragalus oxyphysus Gray Astragalus canadensis Linnaeus Acanthoscelides aureolus complex, A. pullus Acanthoscelides perforatus Astragalus pattersonii Gray Astragalus cibarius Sheldon Acanthoscelides aureolus complex, A. fraterculus Acanthoscelides fraterculus Astragalus pectinatus (Hook.) Dougl. ex G. Don Astragalus crassicarpus crassicarpus Nuttall Acanthoscelides fraterculus Acanthoscelides fraterculus Astragalus praelongus Sheldon Astragalus crassicarpus paysonii (E.H. Kelso) Barneby Acanthoscelides aureolus complex, A. mixtus, A. pullus Acanthoscelides fraterculus Astragalus pterocarpus S. Wats. Astragalus crotalariae (Benth.) Gray Acanthoscelides aureolus complex, A. Acanthoscelides aureolus complex, A. pullus fraterculus Astragalus douglasii (Torr. and Gray) Gray Astragalus pycnostachys Gray Acanthoscelides aureolus complex, A. pullus Acanthoscelides margaretae Astragalus drummondii Douglas ex Hook Astragalus racemosus Pursh. Acanthoscelides aureolus complex, A. frater- Acanthoscelides aureolus complex, A. culus fraterculus Astragalus falcatus Lam. Astragalus sabulonum Gray Acanthoscelides fraterculus Acanthoscelides mixtus Astragalus fucatus Barneby Astragalus thurberi Gray Acanthoscelides aureolus complex Acanthoscelides aureolus complex, A. Astragalus grayi Parry ex S. Wats. mixtus, A. pullus Acanthoscelides aureolus complex Astragalus trichopodus antisellii (Gray) Jeps. Astragalus humistratus Gray Acanthoscelides aureolus complex, A. pullus Acanthoscelides aureolus complex, A. Astragalus trichopodus phoxus (M.E. Jones) lobatus Barneby Astragalus hyalinus M.E. Jones Acanthoscelides aureolus complex, A. pullus Acanthoscelides fraterculus Astragalus trichopodus trichopodus (Nutt.) Gray Astragalus lancearius Gray Acanthoscelides aureolus complex Acanthoscelides aureolus complex Astragalus troglodytus S. Wats. Astragalus lentiginosus Douglas ex. Hook. Acanthoscelides lobatus Acanthoscelides aureolus complex, A. Astragalus utahensis (Torr.) Torr. and Gray lobatus, A. mixtus, A. pullus Acanthoscelides fraterculus Astragalus lonchocarpus Barneby Astragalus wootoni Sheldon Acanthoscelides aureolus complex, A. Acanthoscelides aureolus complex, A. mixtus mixtus, A. pullus Astragalus missouriensis Nuttall Baptisia sp. Acanthoscelides fraterculus Acanthoscelides bisignatus Astragalus mollissimus bigelovii (Gray) Barneby Bauhinia coulteri MacBr. (Mex.) Acanthoscelides fraterculus, A. lobatus Gibbobruchus cristicollis

250 Bauhinia divaricata Linnaeus Calliandra humilis reticulata (Gray) L. Benson Gibbobruchus cristicollis, G. divaricatae Stator limbatus, S. pruininus, S. pygidialis Bauhinia lunarioides Gray ex Wats. Cassia spp. Gibbobruchus cristicollis, G. mimus Amblycerus nigromarginatus; Sennius Bauhinia macranthera Oliver abbreviatus, S. discolor Gibbobruchus cristicollis Cassia afrofistula Brenan Bauhinia malabarica Roxb. Caryedon serratus Caryedon serratus Cassia arereh Delile Bauhinia monandra Kurz Caryedon serratus Caryedon serratus Cassia fistulaLinnaeus Bauhinia pauletia Pers. Caryedon serratus; Stator limbatus, S. pruininus (experimental) Gibbobruchus cristicollis, G. divaricatae Cassia grandis Linnaeus Bauhinia pes-caprae Cav. Caryedon serratus; Stator limbatus, S. Gibbobruchus divaricatae pruininus Bauhinia racemosa Lam. Cassia javanica indochinensis (Irwin and Barn.) Caryedon serratus Caryedon serratus; Stator limbatus, S. Bauhinia reticulata (DC.) Hochst. pruininus Caryedon serratus Cassia javanica javanica Linnaeus Bauhinia rufescens Lam. Caryedon serratus; Stator limbatus, S. Caryedon serratus pruininus Bauhinia thonningi (Schum.) Milne-Redh. Cassia sieberiana DC. Caryedon serratus Caryedon serratus Bauhinia tomentosa Linnaeus Centrosema macrocarpa Benth. Caryedon serratus Caryedes helvinus, C. incensus Bauhinia ungulata Linnaeus Centrosema pubescens Benth. Gibbobruchus divaricatae Caryedes helvinus, C. incensus Caesalpinia spp. Ceratonia siliqua Linnaeus Amblycerus nigromarginatus Mimosestes mimosae, M. nubigens Caesalpinia coriaria (Jacq.) Willd. Cercidium spp. See Parkinsonia Mimosestes mimosae, M. nubigens Cercis canadensis Linnaeus Caesalpinia pulcherrima (Linnaeus) Swartz Gibbobruchus mimus Caryedon serratus; Stator limbatus, S. Cercis occidentalis Torr. ex Gray pruininus (experimental) Gibbobruchus mimus Caesalpinia sclerocarpa Standley Chamaecrista fasciculata (Michx.) Irwin and Mimosestes mimosae (extralimital) Barn. Cajanus cajan (Linnaeus) Huth Sennius cruentatus; Zabrotes arenarius Acanthoscelides obtectus; Callosobruchus Chamaecrista nictitans mensalis (Greene) Irwin chinensis, C. maculatus, C. phaseoli, C. and Barn. pulcher; Caryedon serratus; Zabrotes sub- Sennius simulans; Zabrotes planifrons fasciatus Chamaecrista nictitans nictitans (Linnaeus) Calliandra eriophylla Benth. Irwin and Barn. Stator chihuahua, S. limbatus, S. pruininus Sennius cruentatus Calliandra humilis humilis Benth. Chamaecrista serpens wrightii Irwin and Barn. Stator chihuahua, S. limbatus, S. pruininus, Sennius obesulus S. pygidialis, S. sordidus

251 Cicer arietinum Linnaeus Desmanthus covillei (Britt. and Rose) Wigg. ex Acanthoscelides obtectus; Callosobruchus B. Turner chinensis, C. maculatus, C. phaseoli, C. Acanthoscelides compressicornis, A. pulcher; Zabrotes subfasciatus desmanthi; Stator pruininus Coursetia microphylla Gray Desmanthus illinoensis (Michx.) MacMil. ex B. Stator pruininus Robin. and Fern. Crotalaria spectabilis Roth. Acanthoscelides bisignatus, A. compressicornis; Merobruchus julianus (question- Callosobruchus phaseoli able) Cyamopsis tetragonoloba (Linnaeus) Taubert Desmanthus fruticosa N.E. Rosa Callosobruchus chinensis Stator pruininus Cytisus scoparius (Linnaeus) Link Desmanthus leptolobus Torr. and Gray Bruchidius villosus Acanthoscelides bisignatus, A. compres- Dalea spp. sicornis, A. pectoralis; Stator pruininus Acanthoscelides daleae, A. pallidipennis, A. Desmanthus leptophyllus H.B.K. pusillimus Acanthoscelides compressicornis, A. Dalea aurea Nutt. and Pursh. desmanthi Acanthoscelides seminulum Desmanthus obtusus S. Wats. Dalea candida Willd. Acanthoscelides compressicornis, A. Acanthoscelides seminulum pectoralis Dalea carnea albida (Torr. and Gray) Barn. Desmanthus reticulatus Benth. Acanthoscelides pedicularius Acanthoscelides pectoralis Dalea enneandra Nutt. Desmanthus subulatus (Britt. and Rose) Acanthoscelides seminulum Acanthoscelides compressicornis, A. Dalea feayi (Chapm.) Barn. desmanthi Acanthoscelides seminulum Desmanthus velutinus Scheele Dalea frutescens Gray Acanthoscelides bisignatus, A. compressicornis, A. pectoralis Acanthoscelides seminulum Desmanthus virgatus acuminatus (Benth.) Isely Dalea ornata (Dougl. and Hook.) A. Eat. and Wright Acanthoscelides compressicornis, A. pallidipennis, A. schrankiae Acanthoscelides oregonensis Desmanthus virgatus depressus (Humb. and Dalea scandens paucifolia (Coult.) Barneby Bonpl. ex Willd.) B. Turner Acanthoscelides pusillimus Acanthoscelides compressicornis, A. Dalea stanfieldii (Small) Shinners desmanthi Acanthoscelides seminulum Desmanthus virgatus glandulosa B. Turner Dalea tenuis (Coultr.) Shinn. Acanthoscelides compressicornis Acanthoscelides seminulum Desmanthus virgatus virgatus (Linnaeus) Willd. Delonix regia (Bojer and Hook.) Raf. Acanthoscelides bisignatus, A. compres- Stator limbatus (questionable) sicornis, A. desmanthi, A. pallidipennis, A. Desmanthus sp. pectoralis; Stator pruininus Acanthoscelides bisignatus, A. compres- Desmodium spp. sicornis, A. desmanthi, A. pectoralis Acanthoscelides biustulus, A. stylifer Desmanthus bicornutus S. Watson Desmodium batocaulon Gray Acanthoscelides desmanthi; Stator limbatus, Acanthoscelides biustulus S. pruininus Desmodium canescens (Linnaeus) DC. Desmanthus cooleyi (A. Eaton) Trel. Meibomeus musculus Acanthoscelides compressicornis; Stator pruininus 252 Desmodium cinerascens Gray Hedysarum alpinum americanum Michx. Acanthoscelides biustulus Kytorhinus prolixus Desmodium grahamii Gray Hedysarum boreale Nutt. Acanthoscelides biustulus, A. stylifer; Acanthoscelides fraterculus; Kytorhinus Meibomeus desmoportheus prolixus Desmodium illinoense Gray Hoffmanseggia densiflora Bentham ex Gray Acanthoscelides longistilus Acanthoscelides compressicornis Desmodium neomexicanus Gray Hoffmanseggia drepanocarpa Gray Acanthoscelides biustulus Acanthoscelides compressicornis Desmodium tenuifolium Torr. and Gray Hoffmanseggia glauca (Ortega) Eifert Meibomeus musculus Acanthoscelides compressicornis Desmodium tortuosum DC. ex G. Don Hoffmanseggia tenella Thorp and Williams Meibomeus musculus Acanthoscelides compressicornis Desmodium triflorum (Linnaeus) DC. Indigofera anil Linnaeus (exp.) Meibomeus musculus Stator pruininus Desmodium uncinatum (Jacq.) DC. Indigofera lindheimeriana Scheele Stator pruininus (experimental) Acanthoscelides kingsolveri Dipogon lignosus (Linnaeus) Verde. Indigofera platycarpa Rose Zabrotes subfasciatus Acanthoscelides baboquivari Dolichos melanophthalmus DC. Indigofera sphaerocarpa Gray Acanthoscelides obtectus Acanthoscelides baboquivari, A. kingsolveri Eriosema violaceum (Aublet) G. Don Indigofera suffruticosa Miller Acanthoscelides flavescens (extralimital) Acanthoscelides kingsolveri; Stator pruininus Errazurizia rotundata (Wooton) Barn. Lablab purpureus (Linnaeus) Sweet Acanthoscelides pallidipennis Acanthoscelides obtectus; Callosobruchus chinensis, C. maculatus, C. phaseoli; Za- Erythrina sandwicensis Deg. brotes subfasciatus Stator limbatus, S. pruininus Laburnum alpinum Brecht. and Presl. Eysenhardtia texana Scheele Bruchidius villosus Acanthoscelides comstocki, A. mixtus Laburnum anagyroides Medik. Galactia striata (Jacq.) Urban Bruchidius villosus Acanthoscelides flavescens Lathyrus aphaca Linnaeus Galactia wrightii Gray Callosobruchus maculatus Acanthoscelides rufovittatus Lathyrus clymenum Linnaeus Gleditsia aquatica Marsh. Callosobruchus maculatus Amblycerus robiniae Lathyrus sativus Linnaeus Gleditsia triacanthos Linnaeus Acanthoscelides obtectus; Bruchus Amblycerus robiniae; Mimosestes nubigens brachialis, B. pisorum; Callosobruchus Glycine max (Linnaeus) chinensis, C. maculatus Acanthoscelides obtectus; Callosobruchus Lens culinaris Medik. chinensis, C. maculatus; Stator limbatus, S. Acanthoscelides obtectus; Callosobruchus pruininus (exp.); Zabrotes subfasciatus chinensis, C. maculatus Glycyrrhiza sp. Lespedeza capitata Michx. Acanthoscelides pallidipennis Acanthoscelides longistilus Glycyrrhiza lepidota Pursh. Lespedeza frutescens (Linnaeus) Elliott Acanthoscelides alboscutellatus, A. aureo- Acanthoscelides longistilus lus complex, A. fraterculus

253 Lespedeza hirta (Linnaeus) Hornem. Merobruchus insolitus, M. knulli; Stator Acanthoscelides longistilus; Meibomeus chihuahua, S. limbatus musculus Lysiloma sabicu Benth. Lespedeza intermedia (S. Wats.) Britt. Merobruchus lysilomae (extralimital) Acanthoscelides longistilus Lysiloma watsoni Rose Lespedeza texana Britt. Merobruchus knulli (extralimital) Acanthoscelides longistilus Mimosa biuncifera Benth. Lespedeza virginica (Linnaeus) Britt. Acanthoscelides chiricahuae, A. speciosus; Acanthoscelides longistilus Stator chihuahua, S. pruininus Leucaena leucocephala (Lam.) de Wit Mimosa borealis Gray Acanthoscelides macrophthalmus; Mimoses- Acanthoscelides chiricahuae, A. schrankiae; tes amicus; Stator limbatus, S. pruininus Merobruchus julianus (questionable) Leucaena pulverulenta (Schlect.) Benth. Mimosa dysocarpa Benth. Acanthoscelides macrophthalmus; Stator Acanthoscelides chiricahuae, A. limbatus compressicornis; Stator pruininus Leucaena retusa Benth. Mimosa grahamii Gray Acanthoscelides macrophthalmus Acanthoscelides chiricahuae, A. compressicornis; Stator pruininus Lotus sp. Mimosa laxiflora Benth. Acanthoscelides pallidipennis; Mimosestes mimosae, M. nubigens; Zabrotes cynthiae Acanthoscelides chiricahuae; Stator (floral record) pruininus Lotus crassifolius (Benth.) Greene Mimosa malacophylla Gray Acanthoscelides fraterculus Acanthoscelides speciosus Lotus mearnsii Britt. Mimosa pigra Linnaeus Acanthoscelides fraterculus Acanthoscelides quadridentatus Lotus purshianus Clements and Clements Mimosa pigra berlandieri (Gray) B.L. Turner Acanthoscelides aureolus complex Acanthoscelides quadridentatus Lotus rigidus (Benth.) Greene Mimosa strigillosa Torr. and Gray Acanthoscelides fraterculus Acanthoscelides distinguendus, A. quadridentatus, A. tridenticulatus Lotus scoparius (Nutt.) Ottleya Mimosa wherryana (N.L. Britt.) Standl. Acanthoscelides aureolus complex, A. fraterculus Acanthoscelides chiricahuae, A. speciosus; Stator chihuahua, S. pruininus Lupinus albus Linnaeus Neptunia plena (Linnaeus) Benth. Acanthoscelides obtectus (doubtful) Stator limbatus, S. pruininus Lysiloma acapulcensis Benth. Nissolia schottii (Torr.) Gray Merobruchus knulli (extralimital) Acanthoscelides mundulus Lysiloma divaricatum (Jacq.) J.F. Macbr. Nissolia wislizenii Gray Merobruchus vacillator Acanthoscelides mundulus Lysiloma latisiliqua (Linnaeus) Benth. Olneya tesota Gray Merobruchus lysilomae Stator pruininus Lysiloma pulverulenta (Schl.) Benth. Onobrychis viciifolia Scop. Stator limbatus Bruchidius cisti Lysiloma microphylla thornberi (Britt. and Rose) Isely Oxytropis lambertii Pursh. Acanthoscelides aureolus complex

254 Oxytropis sericea Nutt. C. pulcher; Caryedon serratus; Zabrotes Acanthoscelides aureolus complex subfasciatus, Z. sylvestris n o t e : Placement of plant species in Parkinso- Pisum elatius Steven ex M. Bieb. nia follows Isely (1975), but these species Bruchus pisorum may be found in botanical literature under Pisum sativum arvense Linnaeus Cercidium. Acanthoscelides obtectus; Bruchus pisorum; Parkinsonia sp. Callosobruchus chinensis, C. maculatus, C. Sennius cruentatus, S. discolor (question- phaseoli; Zabrotes subfasciatus (experimen- able) tal) Parkinsonia aculeata Linnaeus Pithecellobium brevifolium Benth. Mimosestes amicus, M. ulkei; Stator Stator limbatus limbatus, S. pruininus Pithecellobium dulce (Roxb.) Benth. Parkinsonia florida (Benth.) S. Wats. Stator limbatus Mimosestes amicus, M. ulkei; Stator Pithecellobium ebano (Berl.) Barn. and Grimes limbatus Merobruchus julianus, M. major; Stator beali, Parkinsonia microphylla Torr. S. limbatus, S. pruininus Mimosestes amicus, M. protractus; Stator Pithecellobium pallens (Bentham) Standl. limbatus Merobruchus insolitus; Stator limbatus, S. Parkinsonia texana macra (I. Johnston) Isely pruininus Mimosestes amicus; Stator limbatus, S. Pithecellobium saman (Jacq.) Benth. pruininus Stator limbatus, S. pruininus Parkinsonia texana texana (Gray) S. Wats. Pithecellobium unguis-cati (Linnaeus) Benth. Mimosestes amicus, M. protractus, M. ulkei; Stator limbatus Stator limbatus, S. pruininus Prosopis sp. Parryella filifolia Torr. and Gray Sennius discolor (questionable) Acanthoscelides pallidipennis Prosopis alba Griseb. (introduced from Argen- Petteria ramentacea (Seiden) Presl. tina into Arizona) Bruchidius villosus Algarobius prosopis Phaseolus acutifolius Gray Prosopis chilensis (Molina) Stuntz Callosobruchus maculatus; Zabrotes Algarobius prosopis; Mimosestes nubigens subfasciatus Prosopis glandulosa glandulosa Torr. Phaseolus acutifolius latifolius G.F. Freeman Algarobius bottimeri; Mimosestes amicus, Acanthoscelides obtectus M. insularis, M. nubigens, M. protractus; Phaseolus angularis angularis (Willd.) Wight Neltumius arizonensis, N. gibbithorax; Stator Zabrotes subfasciatus limbatus, S. pruininus Phaseolus coccineus Linnaeus Prosopis glandulosa torreyana (L. Benson) M. Acanthoscelides obtectus; Zabrotes Johnston subfasciatus Algarobius prosopis; Mimosestes amicus, Phaseolus lunatus Linnaeus M. nubigens, M. protractus; Neltumius arizonensis, N. gibbithorax; Stator limbatus Acanthoscelides obtectus; Callosobrchus chinensis, C. maculatus, C. phaseoli; Prosopis juliflora (Sw.) DC. Zabrotes subfasciatus Stator limbatus, S. pruininus (exp.); Phaseolus ritensis M.E. Jones Mimosestes insularis (Hawaii only), M. nubigens Acanthoscelides obtectus; Zabrotes amplissimus Prosopis laevigata (Hum. and Bonpl. ex Willd.) M. Johnston Phaseolus vulgaris Linnaeus Algarobius prosopis; Mimosestes protractus Acanthoscelides obtectus; Callosobru- chus chinensis, C. maculatus, C. phaseoli,

255 Prosopis pallida (Humb. and Bonpl. ex Willd.) Acanthoscelides fumatus H.B.K. (Hawaii only) Rhynchosia tomentosa (Linnaeus) Hook and Algarobius bottimeri; Caryedon serra- Arn. tus; Mimosestes amicus, M. insularis, M. Acanthoscelides distinguendus nubigens Robinia pseudoacacia Linnaeus Prosopis pubescens Benth. Amblycerus robiniae (misidentification?); Algarobius prosopis; Mimosestes amicus; Stator pruininus (questionable) Neltumius gibbithorax Schrankia microphylla (Dryand.) F. Macbride Prosopis reptans reptans Benth. Acanthoscelides schrankiae Algarobius bottimeri Schrankia nuttallii (DC.) Standl. Prosopis reptans cinerascens (Gray) Burk. Acanthoscelides schrankiae Algarobius bottimeri, A. prosopis Schrankia roemeriana (Scheele) Blank. Prosopis velutina Wooton Acanthoscelides schrankiae Algarobius prosopis; Mimosestes amicus, Schrankia uncinata Willd. M. protractus; Neltumius arizonensis; Stator pruininus Acanthoscelides schrankiae Psophocarpus tetragonolobus (Linnaeus) DC. Senna alata (Linnaeus) Irwin and Barn. Callosobruchus chinensis Amblycerus nigromarginatus, A. obscurus (both extralimital) Psorothamnus fremontii (Torr. ex Gray) Barn. Senna armata S. Wats. Acanthoscelides daleae Acanthoscelides obrienorum Psorothamnus schotti (Torr.) Barn. Senna bauhinioides Gray Acanthoscelides daleae Sennius morosus; Zabrotes spectabilis Psorothamnus spinosa (Gray) Barn. Senna bicapsularis bicapsularis Linnaeus Acanthoscelides daleae Amblycerus nigromarginatus, A. obscurus Pueraria montana lobata (Willd.) Maes. and (both extralimital); Sennius fallax, S. lebasi, Almeida S. leucostauros Borowiecius ademptus Senna biflora Linnaeus Rhynchosia sp. Sennius fallax Zabrotes obliteratus Senna corymbosa (Lamk.) Irwin and Barn. Rhynchosia americana (Mill.) Metz Amblycerus nigromarginatus Acanthoscelides distinguendus Senna covesii (Gray) Irwin and Barn. Rhynchosia difformis (Elliott) DC. Sennius morosus; Zabrotes spectabilis Acanthoscelides distinguendus Senna durangensis durangensis (Rose) Irwin Rhynchosia erecta (Walter) DC. and Barn. Acanthoscelides distinguendus Sennius morosus; Zabrotes spectabilis Rhynchosia galactoides Endl. ex Walp. Senna hirsuta hirsuta Linnaeus Acanthoscelides distinguendus Sennius medialis Rhynchosia intermedia (Torrey and Gray) Small Senna hirsuta leptocarpa (Benth.) Irwin and Acanthoscelides distinguendus Barn. Rhynchosia latifolia Nutt. ex Torr. and Gray Amblycerus obscurus (extralimital); Sennius Acanthoscelides distinguendus medialis, S. morosus; Zabrotes chavesi Rhynchosia longeracemosa (Mart. and Gal.) Senna lindheimeriana (Scheele) Irwin and Barn. Acanthoscelides flavescens Sennius discolor; Zabrotes spectabilis Rhynchosia minima (Linnaeus) DC. Senna marilandica (Linnaeus) Irwin and Barn. Acanthoscelides flavescens Sennius abbreviatus Rhynchosia texana Torr. and Gray

256 Senna obtusifolia (Linnaeus) Irwin and Barn. Stylosanthes biflora (Linnaeus) B.S.P. Amblycerus obscurus (extralimital); Stylantheus macrocerus Caryedon serratus; Sennius fallax, S. Tamarindus indica Linnaeus morosus Caryedon serratus Senna occidentalis (Linnaeus) Irwin and Barn. Tephrosia ambigua (M.A. Curt.) Chapm. Amblycerus nigromarginatus; Sennius fallax, Acanthoscelides obsoletus S. morosus (extralimital) Tephrosia cinerea (Linnaeus) Pers. Senna pendula advena (Willd.) Acanthoscelides rufovittatus Amblycerus obscurus (extralimital) Tephrosia purpurea (Linnaeus) Pers. Senna pundula ovalifolia (Willd.) Acanthoscelides rufovittatus Amblycerus obscurus (extralimital) Tephrosia spicata (Walt.) Torr. and Gray Senna polyantha (Moc. and Sesse) Irwin and Barn. Acanthoscelides obsoletus Acanthoscelides obrienorum Tephrosia thurberi (Rydb.) C.E. Wood Senna roemeriana (Scheele) Irwin and Barn. Acanthoscelides rufovittatus Sennius discolor, S. morosus; Zabrotes Tephrosia virginiana (Linnaeus) Pers. spectabilis Acanthoscelides distinguendus (misiden- Senna siamea (Lam.) Irwin and Barn. tification),A. obsoletus Stator pruininus (exp.) Trifolium obtusiflorum Hook. Senna spectabilis (DC.) Acanthoscelides pauperculus Zabrotes chavesi Trifolium pratense Linnaeus Senna uniflora (Spreng.) Irwin and Barn. Acanthoscelides pusillimus (suspect) Amblycerus nigromarginatus, A. obscurus Vicia sp. (both extralimital) Acanthoscelides flavescens; Bruchus Senna wislizenii wislizenii (Gray) Irwin and pisorum; Zabrotes bexarensis Barn. Vicia benghalensis Linnaeus Acanthoscelides obrienorum; Sennius dis- Bruchus brachialis color Vicia caroliniana Walt. Sesbania sp. Bruchus brachialis Callosobruchus phaseoli Vicia cracca Linnaeus Sesbania aegyptiaca Pers. Bruchus brachialis Acanthoscelides obtectus; Mimosestes Vicia faba Linnaeus amicus Acanthoscelides obtectus; Bruchus pisorum Sesbania emerus (Aublet) Urban (misidentification?),B. rufimanus; Cal- Acanthoscelides griseolus; Stator pruininus losobruchus chinensis, C. maculatus, C. Sesbania exaltata (Roxb.) Rydb. phaseoli; Zabrotes subfasciatus Acanthoscelides griseolus; Stator pruininus Vicia grandiflora Scop. Sesbania macrocarpa Muhlenberg Bruchus brachialis Stator pruininus Vicia leavenworthii Torr. and Gray Sesbania sesban (Linnaeus) Merr. Zabrotes bexarensis Acanthoscelides obtectus (experimental); Vicia lutea Linnaeus Mimosestes amicus; Stator pruininus Callosobruchus maculatus Sophora chrysophylla (Salisb.) Seem. Vicia pannonica Creantz Stator pruininus Bruchus brachialis Spartium junceum Linnaeus Vicia sativa nigra Linnaeus Bruchidius cisti, B. villosus Bruchus brachialis

257 Vicia sativa sativa Linnaeus Malvaceae Bruchus brachialis, B. rufimanus; Abutilon abutiloides (Jacq.) Garcke ex Britt. and Callosobruchus maculatus P. Wilson Vicia sepium Linnaeus Abutiloneus idoneus; Acanthoscelides sub- Bruchus brachialis aequalis Vicia villosa dasycarpa Ten. Abutilon berlandieri Gray Bruchus brachialis Abutiloneus idoneus; Acanthoscelides aequalis, A. subaequalis Vicia villosa villosa Roth Abutilon hypoleucum Gray Bruchus brachialis Acanthoscelides flavescens Vigna aconitifolia (Jacq.) Marechal Abutilon incanum (Link) Sweet Acanthoscelides obtectus; Callosobruchus chinensis Acanthoscelides subaequalis Vigna angularis (Willd.) Ohwi and Ohashi Abutilon theophrasti Medikus Callosobruchus chinensis, C. phaeseoli, C. Althaeus folkertsi, A. hibisci pulcher; Zabrotes subfasciatus Abutilon trisulcatum Gray Vigna mungo (Linnaeus) Hepper Acanthoscelides subaequalis Callosobruchus chinensis, C. maculatus; Allowissadula holosericea (Scheele) Bates Zabrotes subfasciatus Acanthoscelides aequalis; Abutiloneus Vigna radiata radiata (Linnaeus) Wilcz. idoneus Acanthoscelides obtectus; Bruchus pisorum; Allowissadula lozanii (Rose) Bates Callosobruchus chinensis, C. maculatus, C. Acanthoscelides aequalis phaseoli Herissantia crispa (Linnaeus) Brizicky Vigna subterranea (Linnaeus) Verdc. Acanthoscelides herissantitus Acanthoscelides obtectus; Zabrotes subfas- Hibiscus sp. ciatus Megacerus discoidus (floral record) Vigna umbellata (Thunb.) Ohwi and Ohashi Hibiscus aculeatus Walt. Acanthoscelides obtectus Althaeus hibisci Vigna unguiculata sesquipedalis (Linnaeus) Verdc. Hibiscus militaris Cav. Acanthoscelides obtectus; Zabrotes subfas- Althaeus folkertsi ciatus Hibiscus moscheutos lasiocarpus Cav. Vigna unguiculata unguiculata (Linnaeus) Althaeus folkertsi, A. hibisci Walker Hibiscus moscheutos moscheutos Linnaeus Acanthoscelides obtectus; Callosobru- Althaeus hibisci chus chinensis, C. maculatus; Zabrotes Hibiscus moscheutos palustris (Linnaeus) subfasciatus Althaeus hibisci Lythraceae Kosteletzkya virginica (Linnaeus) K. Presl. ex Lythrum alatum Pursh. Gray Acanthoscelides tenuis Althaeus hibisci, A. steineri Lythrum lineare Linnaeus Malvastrum bicuspidatum (S. Wats.) Rose Acanthoscelides tenuis Acanthoscelides herissantitus

Magnoliaceae Nelumbonaceae Magnolia sp. Nelumbo nucifera Gaertne Gibbobruchus mimus (floral record) Callosobruchus chinensis Magnolia grandiflora Linnaeus Oleaceae Gibbobruchus mimus (floral record) Fraxinus sp. Gibbobruchus mimus 258 Onagraceae Condalia globosa pubescens I. Johnst. Ludwigia alternifolia Linnaeus Neltumius texanus Acanthoscelides alboscutellatus Condalia hookeri hookeri M.C. Johnst. Ludwigia palustris (Linnaeus) Ell. Neltumius texanus Acanthoscelides alboscutellatus Condalia lycioides (Gray) Weberb. Acanthoscelides prosopoides Palmae Condalia obovata Hooker Coccothrinax argentata (Jacq.) L.H. Bailey Neltumius texanus Caryobruchus gleditsiae Condalia spathulata Gray Coccothrinax martii Becc. Neltumius texanus Caryobruchus gleditsiae Condalia warnockii kearneyana M. Johnst. Livingstona chinensis (Jacq.) R. Brown ex Mart. Neltumius texanus Caryobruchus gleditsiae Ziziphus obtusifolia (Hook. ex Torr. and Gray) Phoenix sylvestris Roxb. Gray Caryobruchus gleditsiae Acanthoscelides prosopoides Sabal etoni Swingle ex Nost Caryobruchus gleditsiae Rosaceae Sabal glaucescens Lodd. ex H. Moore Aronia sp. Caryobruchus gleditsiae Acanthoscelides calvus (floral record) Sabal mauritiaeformis (H. Karsten) Griseb and Rubiaceae Wendl. Guettarda sp. Caryobruchus gleditsiae Amblycerus schwarzi (extralimital) Sabal mexicana Mart. Caryobruchus gleditsiae Rutaceae Sabal minor (Jacq.) Pers. Zanthoxylum clava-herculis Linnaeus Caryobruchus gleditsiae Neltumius texanus (likely floral record); Zabrotes victoriensis (likely floral record) Sabal palmetto (Walter) Lodd. ex Schultes Caryobruchus gleditsiae Umbelliferae Sabal parviflora Becc. Daucus carota Linnaeus (floral records) Caryobruchus gleditsiae Acanthoscelides alboscutellatus, A. obtectus; Sabal uresana Trel. Althaeus hibisci; Bruchus brachialis; Mega- cerus discoidus Caryobruchus gleditsiae Taenidia integerrima (Linnaeus) Drude Sabal yapa C. Wright ex Becc. Sennius abbreviatus (floral record) Caryobruchus gleditsiae Serenoa repens (W. Bartram) Small Verbenaceae Caryobruchus gleditsiae Lippia wrightii Gray Washingtonia filifera (Linden ex Andre) H. Mimosestes amicus (floral record) Wendl. Tectona grandis Linnaeus Caryobruchus gleditsiae Amblycerus schwarzi (extralimital)

Rhamnaceae Vitaceae Condalia correllii M. Johnston Vitis arizonica Engelm. Neltumius texanus Amblycerus vitis Condalia globosa globosa I. Johnst. Neltumius texanus Zygophyllaceae Larrea tridentata (Sesse and Mocino) Cov. Neltumius arizonensis (floral record)

259 Appendix V Bracon bruchivorus Muesebeck (Braconidae) Acanthoscelides sp. Natural Enemies of Bruchidae of Bracon kirkpatricki (Wilkinson) (Braconidae) the United States and Canada Caryedon serratus Bracon mellitor Say (Braconidae) The following is a list of predators and Acanthoscelides fraterculus parasitoids of the Bruchidae of the United Bruchobius laticeps (Ashmead) (Pteromalidae) States and Canada. The majority of natural enemies are Hymenoptera. For the cosmo- Acanthoscelides obtectus; Bruchus pisorum; Callosobruchus maculatus politan bruchids, however, predators and parasitoids from other areas of the world Bruchocida orientalis Crawford (EX) are included and marked as “extralimital” (Eupelmidae) (EX). It must be remembered that many Bruchus pisorum, B. rufimanus; Calloso- of these parasitoids were reared from seed bruchus chinensis; Zabrotes subfasciatus pods and that insects of other taxa may Bruchocida vuilleti Crawford (EX) (Eupelmidae) have also invaded the pods and were the Callosobruchus maculatus source of the parasitoids. Catolaccus sp. (Pteromalidae) Principal references consulted were Bonet Acanthoscelides floridae, A. longistilus; et al. 1987; Bridwell 1918; Burks 1954, Mimosestes nubigens 1971; Cushman 1911; De Luca 1962, Catolaccus hunteri Crawford (Pteromalidae) 1965, 1970, 1977, 1980; Hetz and John- Acanthoscelides bisignatus, A. son 1988; Hinckley 1961; Krombein et compressicornis al. 1979; Peck 1963; Pierce 1908; Stef- Cephalonomia gallicola (Ashmead) (Bethylidae) fan 1981; Stein 1983b; and Zacher 1930. Callosobruchus maculatus Names have been checked for accuracy by Cephalonomia hyalinipennis Ashmead members of Systematic Entomology Labo- (Bethylidae) ratory Hymenoptera Unit. Mimosestes nubigens Anisopteromalus calandrae (Howard) Cerceris truncata Cameron (Sphecidae) (Pteromalidae) Algarobius prosopis; Mimosestes amicus, M. Bruchus brachialis, B. pisorum; Callosobru- protractus; Neltumius arizonensis chus chinensis, C. maculatus, C. phaseoli; Stator pruininus; Zabrotes subfasciatus Chaetostricta mukerjii Mani (Trichogrammatidae) Anisopteromalus pratti (Crawford) (Pteromalidae) Callosobruchus maculatus Callosobruchus maculatus Charitopodinus swezeyi (Fullaway and Krauss) (Encyrtidae) Aprostocetus aethiops (Zetterstedt) (EX) (Eulophidae) Bruchus rufimanus; Stator pruininus; Zabrotes subfasciatus Bruchus pisorum Charitopodinus terryi Bridwell (Encyrtidae) Aprostocetus claviger Thomson (EX) (Eulophidae) Algarobius prosopis; Bruchus rufimanus; Stator pruininus; Zabrotes subfasciatus Bruchus pisorum Choetospila elegans Westwood (Pteromalidae) Argiope sp. (Araneae) Bruchus rufimanus; Callosobruchus chinen- Mimosestes nubigens sis, C. maculatus; Zabrotes subfasciatus Bracon sp. (Braconidae) Chremylus elaphus Haliday (Braconidae) Acanthoscelides flavescens; Amblycerus Bruchus rufimanus robiniae; Caryedon serratus; Mimosestes nubigens Chryseida spinola bennetti Burks (Eurytomidae)

260 Acanthoscelides obtectus; Merobruchus Eupelmus inyoensis Girault (Eupelmidae) insolitus Acanthoscelides floridae, A. submuticus Dibrachys cavus (Walker) (Pteromalidae) Eupelmus swezeyi Ashmead (Eupelmidae) Bruchus brachialis Caryedon serratus Dinarmus sp. (Pteromalidae) Eurytoma sp. (Eurytomidae) Megacerus discoidus Acanthoscelides flavescens, A. fraterculus, Dinarmus acutus (Thomson) (Pteromalidae) A. submuticus; Amblycerus robiniae; Bru- Acanthoscelides alboscutellatus; Bruchidius chus pisorum; Lithraeus atronotatus; Mero- cisti, B. villosus; Bruchus brachialis, B. bruchus julianus; Mimosestes nubigens rufimanus Eurytoma obtusa Bugbee (Eurytomidae) Dinarmus colemani Crawford (EX) Bruchus brachialis (Pteromalidae) Eurytoma tylodermatis Ashmead (Eurytomidae) Callosobruchus chinensis Acanthoscelides submuticus; Amblycerus Dinarmus laticeps (Ashmead) (Pteromalidae) robiniae; Bruchus brachialis; Mimosestes Acanthoscelides obtectus; Bruchus pisorum, nubigens B. rufimanus; Callosobruchus chinensis, C. Eurytoma wachtli Mayr (EX) (Eurytomidae) maculatus; Zabrotes subfasciatus Bruchus rufimanus Dinarmus magnus Rohwer (EX) (Pteromalidae) Gastrancistrus undulatus Ratzeburg (EX) Bruchus pisorum, B. rufimanus (Pteromalidae) Dinarmus vagabundus (Timberlake) Bruchus pisorum (Pteromalidae) Glyptocolastes texanus Ashmead (Braconidae) Callosobruchus chinensis, C. maculatus; Algarobius prosopis Zabrotes subfasciatus Habrolepoidea tarsalis Girault (Encyrtidae) Entedon sp. (Eulophidae) Bruchus brachialis Caryedon serratus; Zabrotes subfasciatus Heterospilus bruchi Viereck (Braconidae) Eupelmus sp. (Eupelmidae) Acanthoscelides quadridentatus; Amblyc- Acanthoscelides aureolus complex, A. erus robiniae; Gibbobruchus mimus chiricahuae, A. mixtus, A. obrienorum, A. Heterospilus prosopidis Viereck (Braconidae) quadridentatus; Mimosestes acaciestes Acanthoscelides bisignatus, A. compres- Eupelmus amicus Girault (Eupelmidae) sicornis, A. daleae, A. flavescens, A. qua- Acanthoscelides alboscutellatus, A. floridae, dridentatus, A. submuticus; Algarobius Bruchus brachialis, B. pisorum; Mimosestes bottimeri, A. prosopis; Amblycerus robiniae; acaciestes, M. amicus Callosobruchus chinensis, C. maculatus, C. Eupelmus brevicauda (Crawford) (Eupelmidae) phaseoli; Caryedon serratus; Gibbobruchus Acanthoscelides submuticus mimus; Mimosestes acaciestes, M. amicus, M. nubigens; Stator pruininus; Zabrotes sub- Eupelmus bruchivorus (Crawford) (Eupelmidae) fasciatus Acanthoscelides submuticus; Mimosestes Heterospilus spermophagi Ashmead nubigens (Ichneumonidae) Eupelmus cushmani (Crawford) (Eupelmidae) Amblycerus robiniae Acanthoscelides flavescens, A. obtectus, Horismenus sp. (Eulophidae) A. prosopoides; Merobruchus julianus; Mimosestes nubigens Gibbobruchus mimus; Lithraeus atronotatus; Sennius abbreviatus Eupelmus cyaniceps Ashmead (Eupelmidae) Acanthoscelides bisignatus, A. compressi- Horismenus bruchophagus Burks (Eulophidae) cornis, A. flavescens, A. obtectus, A. submu- Mimosestes nubigens ticus; Amblycerus robiniae; Bruchus bra- Horismenus depressus Gahan (Eulophidae) chialis; Gibbobruchus mimus; Merobruchus Stator pruininus julianus; Mimosestes amicus, M. nubigens

261 Horismenus missouriensis Ashmead pruininus (Eulophidae) Pteromalus leguminis Gahan (Pteromalidae) Acanthoscelides aureolus complex, A. chir- Bruchus pisorum icahuae, A. floridae, A. pallidipennis, A. Pteromalus micans Nees (Pteromalidae) subaequalis, A. submuticus; Algarobius prosopis; Amblycerus robiniae; Merobruchus Bruchus pisorum insolitus; Mimosestes acaciestes; Neltumius Pteromalus piercei (Crawford) (Pteromalidae) texanus; Sennius medialis, S. morosus; Acanthoscelides bisignatus, A. Stator pruininus compressicornis Horismenus productus (Ashmead) (Eulophidae) Pteromalus schwenkei Roomi, Khan, and Khan Acanthoscelides mundulus, A. pallidipennis, (EX) (Pteromalidae) A. submuticus; Algarobius prosopis; Mimos- Callosobruchus chinensis estes amicus Pteromalus sequester Walker (EX) (Pteromali- Laelius utilis Cockerell (Bethylidae) dae) Bruchus brachialis Bruchidius cisti, B. villosus; Bruchus pisorum Lariophagus distinguendus (Foerster) (both European) (Pteromalidae) Pyemotes boylei Krczal (Acarina) Bruchus brachialis, B. rufimanus; Callosobru- Mimosestes nubigens chus chinensis, C. maculatus Pyemotes tritici Lagrèze-Fossat and Montanè Lariophagus texanus Crawford (Pteromalidae) (Acarina) Algarobius bottimeri, A. prosopis; Calloso- Bruchidius cisti; Bruchus rufimanus; bruchus maculatus; Mimosestes nubigens; Callosobruchus chinensis Stator chihuahua, S. pruininus Pyemotes ventricosus Newport (Acarina) Macroneura vesicularis (Retzius) (Eupelmidae) Acanthoscelides obtectus; Bruchus pisorum, Bruchus brachialis B. rufimanus; Callosobruchus chinensis, C. Microdontomerus anthonomi Crawford maculatus; Zabrotes subfasciatus (Torymidae) Sclerodermus immigrans Bridwell (Bethylidae) Acanthoscelides aureolus complex, A. mixtus, A. Caryedon serratus submuticus; Bruchus brachialis, B. pisorum; Senegalella acythopoensis Auriv. (Torymidae) Stator limbatus Callosobruchus maculatus Monomorium sp. (Formicidae) Staphylinus nigrellus Horn (Coleoptera) Mimosestes nubigens Bruchus pisorum Oedaule magnus Rohwer (Pteromalidae) Stenocorse bruchivora (Crawford) (Braconidae) Callosobruchus maculatus Acanthoscelides obrienorum, A. quadridenta- Oedaule stringifrons Waterson (Pteromalidae) tus, A. submuticus; Algarobius bottimeri, A. Caryedon serratus prosopis; Callosobruchus chinensis; Carye- Parasierola distinguenda Kieffer (Bethylidae) don serratus; Gibbobruchus mimus; Mero- Acanthoscelides quadridentatus; Mimos- bruchus julianus; Mimosestes acaciestes, M. estes nubigens amicus, M. nubigens; Stator limbatus Parasierola emigrata Rohwer (Bethylidae) Systasis encyrtoides Walker (EX) (Pteromalidae) Caryedon serratus Bruchus pisorum Phanerotoma sp. (Braconidae) Tetrastichus sp. (Eulophidae) Acanthoscelides quadridentatus Acanthoscelides longistilus; Zabrotes subfasciatus Pseudocatolaccus bruchocida Risbec (EX) (Pteromalidae) Tetrastichus bruchivorus Gahan (Eulophidae) Zabrotes subfasciatus Bruchus brachialis Pteromalus sp. (Pteromalidae) Tetrastichus bruchophagi Gahan (Eulophidae) Acanthoscelides fraterculus; Bruchus bra- Bruchus brachialis chialis; Callosobruchus chinensis; Stator

262 Tetrastichus claviger Thomson (Eulophidae) Mimosestes acaciestes; Neltumius arizon- Bruchus pisorum ensis, N. texanus; Sennius medialis; Stator limbatus; Stylantheus macrocerus Tetrastichus nerio Walker (Eulophidae) Urosigalphus neobruchi Gibson (Braconidae) Bruchus pisorum Acanthoscelides mundulus, A. prosopoides, Torymus atheatus Grissell (Torymidae) A. quadridentatus; Algarobius prosopis; Acanthoscelides obtectus Amblycerus robiniae; Merobruchus insoli- Torymus persicariae Mayr (EX) (Torymidae) tus; Mimosestes acaciestes, M. amicus, M. Bruchidius villosus (questionable record) nubigens; Stator limbatus Triaspis sp. (Braconidae) Uscana caryedoni Viggiani (EX) (Trichogrammatidae) Bruchidius villosus Caryedon serratus Triaspis forbesii (Dalla Torre) (Braconidae) Uscana lariophaga Stephen (EX) Bruchus rufimanus (Trichogrammatidae) Triaspis gibberosus (Szépligeti) (EX) Callosobruchus maculatus (Braconidae) Uscana marilandica Girault Bruchus pisorum, B. rufimanus (Trichogrammatidae) Triaspis luteipes Thomson (Braconidae) Callosobruchus maculatus Bruchus rufimanus Uscana mukerjii (Mani) (EX) Triaspis pallipes Nees (Braconidae) (Trichogrammatidae) Bruchus rufimanus Callosobruchus chinensis, C. maculatus; Triaspis similis (Szépligeti) (EX) (Braconidae) Zabrotes subfasciatus Bruchus rufimanus Uscana semifumipennis Girault Triaspis stictostiba Martin (Braconidae) (Trichogrammatidae) Bruchus rufimanus Acanthoscelides alboscutellatus, A. obtectus; Algarobius prosopis; Althaeus hibisci; Triaspis thoracica (Curtis) (Braconidae) Bruchidius cisti; Bruchus pisorum; Calloso- Bruchus brachialis, B. pisorum, B. rufi- bruchus chinensis, C. maculatus, C. phaseo- manus; Zabrotes subfasciatus li; Caryedon serratus; Megacerus discoidus; Trichogramma sp. (Trichogrammatidae) Mimosestes amicus, M. nubigens; Stator lim- Bruchidius villosus batus, S. pruininus; Zabrotes subfasciatus Trichogrammatidae Uscana senex (Grese) (Trichogrammatidae) Stator pruininus Bruchidius cisti; Bruchus pisorum; Calloso- bruchus maculatus Trichomalopsis leguminis Gahan (Pteromalidae) Zatropis sp. (Pteromalidae) Bruchus pisorum Lithraeus atronotatus Urosigalphus sp. (Braconidae) Zatropis incertus (Ashmead) (Pteromalidae) Amblycerus vitis Abutiloneus idoneus; Acanthoscelides Urosigalphus arizonensis Crawford (Braconidae) chiricahuae, A. longistilus, A. subaequa- Neltumius arizonensis lis, A. submuticus; Bruchus brachialis; Urosigalphus bruchi Crawford (Braconidae) Mimosestes acaciestes Acanthoscelides quadridentatus; Algaro- Zatropis orontas (Walker) (Pteromalidae) bius prosopis; Amblycerus robiniae; Callo- Acanthoscelides submuticus sobruchus chinensis; Caryedon serratus; Merobruchus insolitus, M. julianus; Mimos- Zelus renardii Kolenati (Hemiptera) estes amicus, M. nubigens Bruchus rufimanus; Mimosestes nubigens Urosigalphus bruchivorus Crawford (Braconidae) Acanthoscelides chiricahuae, A. mundulus; Algarobius prosopis; Merobruchus julianus;

263 Glossary of Morphological Cardo: Segment of the maxilla that articu- Terms lates with the head. Carina: Keel; elevated ridge. Some definitions modified from Nichols Caudad: Toward the posterior end of the and Schuh (1989). body. Acicular: Needle-shaped. Caudal: Pertaining to the posterior end of Acuminate: Tapering to a sharp point. the body. Angulate: Meeting at an angle, as two mar- Cephalad: Toward the head end of the gins body. Anterad: Toward the head. Cervical boss: A low tubercle bearing two setiferous punctures on the anterolateral Anterior: In front; toward the head. angle of the pronotum. Anterolateral: Located anteriorly and to the Cervical sulcus: A vertical sulcus parallel- side. ing the anterolateral margin of the prono- Anteroventral: Toward the front on ventral tum in many bruchids. side of body. Closure valve: Valvular structure at the end Antescutellar: Located immediately in front of the internal sac of male genitalia that of the scutellum. controls passage of seminal fluid. Apex: The end of a part farthest from the Clypeolabral suture: A transverse, im- base; tip. pressed line between the clypeus and the Arcuate: Arched; curved like a bow. labrum. Armature: Various chitinous spines, den- Clypeus: Sclerite situated between the ticles, or spicules in internal sac of male frons and the labrum on the front of the genitalia. head. Articulation: Point where two moveable Compressed: Flattened laterally. parts abut; joint. Condyle: A process that forms a point of Asperity: Surface roughening into small articulation of a movable part. elevations. Contiguous: Touching along a common Attenuate: Gradually tapering toward apex. border. Basal lobe: The broad, antescutellar lobe of Convergent: Approaching the same point. the posterior margin of the pronotal disk. Coronal denticles: The crownlike cluster of Basal denticle: A toothlike structure at the denticles at the apex of the tibia in many base of an elytral stria. bruchids. Base: The end of a body part or segment Crescentic: In the shape of a crescent. that is attached to another part; for in- Cristate: Having a prominent carina or stance, the margin of the pronotum adja- ridge on the dorsal surface. cent to the base of the elytra. Cuneate: Elongate triangular shape. Bidentate: Bearing two teeth or denticles. Cuspidate: Pointed, acuminate, coming to Bifid: Divided into two parts; forked. a sharp point. Bifurcate: Separated into two branches. Cuticle: The epidermis covering the entire Boss: Elevated, glabrous raised area, usu- body of an insect. ally on frons. Dentate: Toothed or with toothlike projec- Calcar: A movable spur, usually at apex of tions. tibia (plural, calcaria).

264 Denticle: A small, often acute, tooth. Fovea: An integumental depression with Denticulate: Set with small teeth or den- nearly flat bottom and vertical sides. ticles. Foveola: A small fovea. Depressed: Flattened, as if pressed down. Frons: Area of the head between the eyes Dilated: Widened; expanded. limited posteriorly by the vertex and anteriorly by the clypeus. Dimorphic: Different in form or color; in Bruchidae usually applied to sexual differ- Frontoclypeal suture: Transverse impressed ences. line between the frons and the clypeus. Disk: The central dorsal surface of a body Fusiform: Spindle-shaped; broad in middle part; for example: pronotum, pygidium. and tapered at each end. Distal: Toward the free end of a body part. Galea: The outer lobe of the maxilla, usually having two segments. Dorsad: Toward the upper part or side of a body or body part. Gena: Cheek; side of head below the eye. Dorsolateral: On the dorsal side of a lateral Gibbosity: A hump or protuberance from a face of a body or body part. body surface. Dorsomeson: The middle line of the dorsal Glabrous: Smooth; without vestiture or part of a body or body part. surface structures. Dorsum: The upper surface of a body or Granulose: Grainy; covered with minute, body part. grainlike structures. Elytron: The hard or leathery forewing of a Gula: Median sclerite on ventral side of beetle modified to cover the hind, or flight, head. wings and meeting on the midline in re- Gular suture: Impressed line between the pose (plural, elytra). gena and the gula. Elytral: Pertaining to the elytra. Habitus: General form and appearance. Emarginate: Having an obtuse, rounded, or Host: Organism upon which another or- quadrate notch cut from a margin. ganism feeds. Emargination: A notched or cut-out place Humerus: The shoulder or anterolateral on a margin. angle of a beetle elytron. Epimeron: The caudal sclerite of a thoracic Hypognathous: With the head hanging ver- pleuron. tically. Episternum: The anteriad sclerite of a tho- Hypomeron: The concave lateral face of the racic pleuron. pronotum below bounded dorsally by the Evanescent: Vague; obsolete; nearly disap- lateral carina and ventrally by the proster- pearing. num. Femur: The third segment of an insect leg; Imbricate: Appearing like fish scales or the thigh. shingles. Flabellate: Fanlike; having long, flat or te- Incrassate: Thickened; in bruchids applied rete processes that fold together. to the enlarged, laterally compressed metafemur. Footstalk: The basal sclerite of the maxilla. Impressed: Having shallow depressed ar- Fossa: A pit or deep sulcus. eas. Fossula: A suture on the metacoxal face Internal sac: The invaginated intromittent extending laterad from the fossa of the sac attached at the distal end of the me- trochanter.

265 dian lobe, which is everted during copula- Metepisternal sulcus: Angulate groove on tion. the face of the metepisternum. Integument: The outer skin of the body; Metepisternum: The episternum of the cuticle. metathorax. Interstice: Linear strip of integument be- Microtrichia: Fine, cuticular setae. tween striae, such as the elytral interstic- Mola: Roughened grinding surface of me- es. dian face of the mandible. Interval: Space between surface structures Monophagous: Feeding on or in only one or punctures. host. Labium: The ventral lip of the mouthparts. Mucro: Sharp, pointed process; in bruchids, Labrum: The dorsal lip of the mouthparts. the fixed terminal spine of the metatibia Lageniform: Bottle-shaped with narrow extending from the ventral margin. neck. Notum: Dorsal part of a segment. Lateral: Toward the side; away from the Oblique: Set at or cut off at an angle. midline. Obsolete: Faintly marked or nearly effaced Ligula: Middle lobe of the maxillae. surface feature. Liguliform: Tongue-shaped or straplike. Obtuse: Blunt or rounded apical margin of a Lobed: Having a rounded projection or pro- structure. tuberance. Occipital foramen: Opening in the caudal Maculate: Marked with spots of different part of the head leading into the prothorax color than the background. and through which pass the digestive and nervous systems. Mandibles: Dorsal pair of jaws in mouthparts, principally for biting. Occiput: The posterior part of the epicra- nium between the cranium and the anterior Maxillae: Ventral pair of jaws in the mouth- border of the pronotum. parts; used to taste and to masticate food. Ocular index: Greatest width across eyes Median: In or at the middle. divided by least width between eyes. Mesad: Toward the median line of the Ocular sinus: Emargination of the eye. body. Ogival: Shaped like a double-curved gothic Mesocoxa: The basal segment of the middle arch (from ogee). leg. Opisthognathous: Dependant position of Mesopleuron: Lateral surface of the meso- head but directed caudad and resting on thorax. the procoxae. Mesosternum: Ventral median sclerite of Ovipositor: Tubular or valvular structure at the mesothorax. the end of the female’s abdomen through Metafemur: The third segment of the hind which and by which eggs are placed. leg. Palp, palpus: Jointed appendage of labium Metapleuron: Lateral surface of the meta- or maxilla. thorax. Parasutural sulcus: Sulci paralleling the Metasternum: Ventral median sclerite of metathoracic pleurosternal suture. That on the metathorax. the metepisternum extends from the metepisternal sulcus; that on the metaster- Metatibia: The fourth segment of the hind num is an extension of the postmesocoxal leg. sulcus.

266 Pecten: Subapical denticle or row of den- Recurved: Bowed or bent backward, down- ticles on the ventromesal margin of the ward, or outward. metafemur. Reflexed: With apex of a body part turned, Pectinate: Comblike, especially when ap- bent, or reflected backward. plied to antenna. Reniform: Kidney-shaped. Pedicel: Second segment of the antenna. Sclerite: Any part of the body wall bounded Piceous: Pitchy black with reddish high- by sutures. lights. Scutellum: Triangular or quadrate sclerite Pleuron: The lateral region of the thorax or set between bases of the elytra and the abdomen. posterior margin of the pronotum. Pleurosternal: Pertaining to the area of the Sensillum ampullaceum: Conical sensory thorax surrounding the coxal bases. peg sunk in a pit in the body wall. (In lar- Posterior: Toward the hind end of the body. vae.) Postmesocoxal sulcus: Groove or impressed Sensillum basiconicum: Thin-walled, peg- line on metasternum, usually following shaped sensillum with minute pores; a contour of mesocoxal cavity, sometimes basiconic peg. (In larvae.) angulate or lobed. Sensillum chaeticum: Spinelike or bristle- Postocular lobe: Lobe attached to posterior like sensory organ. (In larvae.) margin of the eye separating it from the Sensillum placodeum: Olfactory sensillum side of the head. in the form of a flat plate. (In larvae.) Prementum: Ventral region of head to which Sensillum trichodeum: Hairlike projection of labium is attached. the cuticle articulated with the body wall Procoxa: First segment of the prothoracic by a membranous socket. (In larvae.) leg. Sensory peg: A thin-walled, peg-shaped Pronotum: Dorsal sclerite of the prothorax. sensillum with minute pores; a basiconic peg. (In larvae.) Prostheca: A setose sclerite in the mesal margin of the mandible in bruchids. Serrate: Sawlike; having a series of acute teeth or angulations along a margin. Protuberance: Any elevation above a surface. Seta: Hair-like projection of the cuticule set in a basal cuticular ring. Proximal: The part of an appendage nearest the body; toward the base of a part. Setiferous, setose: Set with setae. Pubescent: Clothed with short, closely set Sinuate: Wavy or undulating, as a margin. setae. Sinus: A curved break in an otherwise Punctate: Set with impressed points or straight line; a deep indentation in a punctures. margin. Punctulate: Set with extremely fine punc- Spicule: A minute, pointed spine. tures as if pierced with a needle. Spinule: A small spine. Puncture: See “Punctate.” Spur: A spinelike, movable appendage; in Pygidium: The tergum of the last visible bruchids, located at the apex of the metati- segment of the abdomen, morphologically bia in certain groups such as the Amblyc- the seventh segment. erinae. Quadrate: Four-sided, usually with right Sternite: Any subdivision of a sternum. angles. Sternum: The ventral sclerite in a segment.

267 Stipes: The footstalk of the maxilla that Truncate: Cut off squarely at the apex. bears the movable appendages. Tubercle: Small, pimplelike protuberance of Stria: Longitudinal impressed line, usually the body wall. of punctures, on disk of elytra; striae are Umbilicate: Having a central depression separated by interstices. resembling a navel. Striate: Marked with parallel, impressed Umbo: A knob or boss (plural, umbones). lines (for example, elytral striae). Valvular: Valve-like or having the functions Stridulate: To produce sound by rubbing of a valve. one part of the body against another. Venter: The underside of the body. Subapical: Located just anteriad of the apex of a body part. Ventral: Pertaining to the underside of the body. Subbasal gibbosity: An elevation of the integument near the base of the elytron usu- Ventromesal: Ventral side of the body to- ally bearing on its summit one or two den- ward the midline. ticles marking bases of the elytral striae. Vertex: The top of the head between the oc- Subequal: Approximately equal in size or ciput, the eyes, and the frons. length. Vestiture: Hair, scales, or setae covering the Submentum: Basal sclerite of the insect integument of the body. labium. Subparallel: Approximately parallel. Sulcus: Shallow or deep furrow or groove. Suture: Line of juncture of two sclerites. Tarsus: Foot; the jointed appendage attached to the end of the tibia. Tentorium: Internal skeleton of the head. Terete: Circular in cross-section. Tergite: A subdivision of a tergal sclerite. Tergum: A dorsal sclerite of a body segment. Tibia: Fourth segment of a leg. Transverse: Wider than long (when referring to a body part). Transverse sulcus: Vague depression on the front of the head at the dorsal limits of the eyes marking the boundary between vertex and frons. Trochanter: Segment of the leg between the coxa and the femur. Trochantin: A basal sclerite articulating between the coxa and the anterior part of the coxal cavity in the fore and mid legs in some bruchids.

268 References of the immature stages of Callosobruchus maculatus F. (Coleoptera: Bruchidae) in Through September 30, 1997 chickpeas. Revista Colombiana de Entomo- logia 9:27–30. Abeille de Perrin, E. 1888. Tableau synop- tique des Bruchides et Urodonides fran- Aldridge, R.J.W., and R.D. Pope. 1986. cais d’apres m. Fl. Baudi de Selve. Revue The British species of Bruchidius Schilsky d’Entomologie 7:77–90. (Coleoptera: Bruchidae). Entomologists’ Gazette 37:181–193. Agadzhanyan, A.K. 1984. Arginase isoen- zymes and their kinetic properties in larvae Alvarez Marin, D., and J.M. Kingsolver. and pupae of bean weevil Acanthoscelides 1997. A preliminary list of the Bruchidae obtectus. Biologicheskii Zhurnal Armenii (Coleoptera) of Cuba. Entomological News 37:40–45. 108:215–221. Agassiz, J.L.R. 1846. Nomenclatoris zoo- Amaro, J.F., H. Ferrinho, and J.P. Cancela logicus, continens nomina systematic a da Fonseca. 1958. Contribuiçao para estu- generum animalium tam viventium quam do da dinamica das popula es de Caryedon fossilium, secundum ordinem alphabet- gonagra (F.). Garcia de Orta 6:637–647. icum disposita. Fasciculus 11. Jent and Anderson, W.H. 1943. The larva of Holos- Gassmann, Soloduri, Switzerland. tilpna nitens (LeC.) and its relationships Ahmad, I., and H. Murad. 1980a. Male (Coleoptera, Anthribidae). Proceedings of reproductive organs of pulse beetle Cal- the Entomological Society of Washington losobruchus chinensis Linn. (Coleoptera: 45:171–174. Bruchidae). Bulletin of Entomology Annand, P.N., and J.S. Pinckney. 1936. 16(1975):21–30. The vetch bruchid (Bruchus brachialis Ahmad, I., and H. Murad. 1980b. Female Fahraeus). U.S. Department of Agriculture, reproductive organs of pulse beetle Cal- Bureau of Entomology and Plant Quaran- losobruchus chinensis Linn. (Coleoptera: tine, No. E–393. Bruchidae). Bulletin of Entomology Anton, K.-W. 1994. The Bruchidae (Co- 16(1975):31–36. leoptera) of Oman, with desctriptions of a Ahmed, M.Y.Y., Y.S. Salem, and E.A. El- new genus and two new speices. Fauna of badry. 1976. The reproductive system of Saudi Arabia 14:105–112. the southern cowpea weevil Callosobruchus Applebaum, S.W., B.J. Southgate, and H. maculatus F. (Coleoptera: Bruchidae). An- Podoler. 1968. The comparative morphol- nales Zoologie-Ecologie Animale 8:13–16. ogy, specific status and host compatibility Ahmed, S.E., A.H. Kamel, and A.E.A. Wa- of two geographical strains of Calloso- hab. 1978. A study of the duration of dif- bruchus chinensis L. (Coleoptera: Bruchi- ferent stages of Callosobruchus chinensis dae). Journal of Stored Products Research L. and Callosobruchus maculatus F. under 4:135–146. constant conditions. Agricultural Research Arnett, R.H., Jr. 1962. Bruchidae. The Review 56:151–154. bean weevils, part 105. In R.H. Arnett, Jr., Aitken, A.D. 1975. Insect travellers. Vol. 1. The Beetles of the United States, pp. 951– Coleoptera. Pest Infestation Control Labo- 958. Catholic University of America Press, ratory, Technical Bulletin No. 31. HMSO Washington, DC. Press, Edinburgh, Scotland. Arora, G.L. 1971. The male genitalia of Aldana Alfonso, H.M. 1983. Effect of tem- Bruchidae and its significance in generic perature on the development and mortality classification. Proceedings of the 12th International Congress of Entomology,

269 August 2–9, 1968, Moscow (1968) l:226– Baker, C.F. 1895. Biological notes on some 227. Nauka, Leningrad, U.S.S.R. Colorado Coleoptera. Entomological News 6:27–29. Arora, G.L. 1977. Taxonomy of the Bruchi- dae (Coleoptera) of Northwest India. Part Baker, W.L. 1972. Eastern forest insects. 1. Adults. Oriental Insects, Supplement 7. U.S. Department of Agriculture, Miscella- Association for the Study of Oriental In- neous Publication No. 1175. sects, Department of Zoology, University of Baskin, J.M., and C.C. Baskin. 1977. Delhi, India. Predation of Cassia marilandica seeds by Arora, G.L. 1978. Taxonomy of Bruchidae Sennius abbreviatus (Coleoptera: Bruchi- (Coleoptera) of Northwest India. Part 2. dae). Bulletin of the Torrey Botanical Club Larvae. Oriental Insects, Supplement 8. 104:61–64. Association for the Study of Oriental In- Bato, S.M., and F.F. Sanchez. 1972. The sects, Department of Zoology, University of biology and chemical control of Callosobru- Delhi, India. chus chinensis (Linn.) (Coleoptera: Bruchi- Arora, G.L., and H.R. Pajni. 1959. The ef- dae). Philippine Entomologist 2:167–182. fect of temperature and food on the devel- Baudi, F. 1886a. Mylabridum seu Bru- opmental period of Callosobruchus analis chidae (Linn. Schoen. All.) europae et (F.) (Bruchidae). Research Bulletin (Sci- finitimarum regionum faunae recensi- ence) of the Panjab University 10:411–412. tio. Deutsch Entomologische Zeitschrift Arora, G.L., H.R. Pajni, and T. Singh. 1967. 30:385–416. The ambiguity of the abnormal male of Baudi, F. 1886b. Rassegna delle specie Callosobruchus maculatus (F.). Research delle famiglia dei Milabridi (Bruchidi degli Bulletin (Science) of the Panjab University autori) viventi in Europa e regioni finitime. 18:501–505. Naturalista Siciliano 6:1–138. Arora, G.L., and T. Singh. 1971. The biol- Baudi, F. 1887. Mylabridum seu Bru- ogy of Callosobruchus chinensis (L.) (Bru- chidum (Lin. Schon. All.) europae et chidae: Coleoptera). Research Bulletin (Sci- finitimarum regionum Faunae recensi- ence) of the Panjab University 21(1970):55– tio. Deutsche Entomologische Zeitschrift 66. 31:33–80. Ashmead, W.H. 1894. Descriptions of new Bawa, S.R., N.R. Kalla, S. Bansal, and K.C. parasitic Hymenoptera. Transactions of the Kanwar. 1971. Length of spermatozoon nu- American Entomological Society 21:318– clei in normal and “sterile” Calloso-bruchus 344. maculatus (Coleoptera: Bruchidae). Annals Avidov, Z., S.W. Applebaum, and M.J. of the Entomological Society of America Berlinger. 1965. Physiological aspects of 64:952–953. host specificity in the Bruchidae. II. Ovipo- Bawa, S.R., and K.C. Kanwar. 1975. Fine sitional preference and behavior of Calloso- structure of Callosobruchus maculatus bruchus chinensis L. Entomologia Experi- spermatozoon. Journal of Submicroscopic mentalis et Applicata 8:96–106. Cytology 7:71–79. Back, E.A. 1940. Weevils in beans and Bawa, S.R., K.C. Kanwar, and R.K. Gupta. peas. U.S. Department of Agriculture, 1972. Interspecific hybridization and the Farmers’ Bulletin No. 1275. abnormal strain of Callosobruchus macula- Back, E.A., and A.B. Duckett. 1918. Bean tus (Coleoptera: Bruchidae). Annals of the and pea weevils. U.S. Department of Agri- Entomological Society of America 65:1241– culture, Farmers’ Bulletin No. 983. 1242.

270 Bawa, S.R., K.C. Kanwar, and R.K. Mar- Bhattacharya, A.K., P.K. Pathak, and P.K. waha. 1974. Studies on sperm survival in Shah. 1977. The oviposition and develop- fertile and sterile strains of Calloso-bruchus ment of Callosobruchus chinensis (Coleop- maculatus (Coleoptera: Bruchidae). Annals tera: Bruchidae) on several host species. of the Entomological Society of America Bulletin of Grain Technology 15:38–41. 67:519–520. Bianchi, F.A. 1940. Notes on the role of the Bawa, S.R., K.C. Kanwar, and R.K. Mar- self-introduced insects in the economic en- waha. 1975. Paper chromatographic analy- tomology of Hawaii. Proceedings of the Ha- sis of free amino-acids in sterile and fertile waiian Entomological Society 10:377–388. strains of Callosobruchus maculatus. An- Bibby, F.F. 1961. Notes on miscellaneous nals of the Entomological Society of Ameri- insects of Arizona. Journal of Economic ca 67:997–999. Entomology 54:324–333. Bedel, L. 1901. Faune des Coléoptères du Biémont, J.C. 1979a. Influence of host Bassin de la Seine. Annales de la Societé plant and mating on oogenesis in Acan- Entomologique de France 5:341–366. thoscelides obtectus (Coleoptera: Bruchi- Begum, A., M.S. Rahman, and M.A. Bash- dae). Annales de la Societé Entomologique er. 1978. Biology of Callosobruchus chinen- de France 15:93–100. sis (Coleoptera: Bruchidae). Dacca Univer- Biémont, J.C. 1979b. Vitellogenesis in sity Studies, Part B–27, pp. 193–198. Acanthoscelides obtectus (Coleoptera: Bru- Begum, A., M.S. Rahman, and D.R. Seal. chidae) 1. Oocyte development and vitel- 1982. Comparative morphology of the logenin in a European strain. International larval instars of Callosobruchus chinensis Journal of Invertebrate Reproduction (L.) and Callosobruchus analis (F.) (Col.: 1:221–232. Bruchidae). Bangladesh Journal of Zoology Biémont, J.C., and A. Bonet. 1981. The 10:66–79. bean weevil populations from the Acan- Bell, E.A. 1978. Seed toxins. In J.B. Har- thoscelides obtectus Say group living on bourne (ed.), Biochemical Aspects of Plant wild or subspontaneous Phaseolus vul- and Animal Coevolution: Proceedings of the garis L. and Phaseolus coccineus L. and Phytochemical Society Symposium, Read- on Phaseolus vulgaris L. cultivated in the ing, April, 1977, pp. 143–161. Academic Tepoztlan region state of Morelos—Mexico. Press, London and New York. Series Entomologica 19:23–41. Bellows, T.S., Jr. 1982. Simulation models Biémont, J-C., I. Butare, and J. Huignard. for laboratory populations of Calloso-bru- 1987. Grouping and inhibition of oogenesis chus chinensis and Callosobruchus macu- in two strains of Acanthoscelides obtec- latus. Journal of Animal Ecology 51:597– tus Say (Coleoptera: Bruchidae) of differ- 624. ent geographical origin: Specificity of this Bellows, T.S., Jr., and J.R. Hassell. 1984. inhibitory effect. International Journal of Models for interspecific competition in lab- Invertebrate Reproductive Development oratory populations of Callosobruchus spp. 12:185–198. Journal of Animal Ecology 53:831–848. Biémont, J.C., G. Chauvin, and C. Hamon. Berthold, A.A. 1827. Latreille’s naturliche 1981. Ultrastructure and resistance to familien des thierreichs, aus dem Franzö- water loss in eggs of Acanthoscelides obtec- sischen. Mit ammerkungen und zusätzen. tus Say (Coleoptera: Bruchidae). Journal of Weimar, Germany. Insect Physiology 27:667–679.

271 Biémont, J-C., C. Hamon, and G. Chau- California Department of Agriculture Re- vin. 1989. Quelques glandes tegumen- port 65–1. Sacramento, CA. taires abdominales chez la bruche du Blanchard, C.E. 1851. Fauna Chilena. haricot Acanthoscelides obtectus Say (Co- Insectos: Coleopteros. In C. Gay, Historia leoptera: Bruchidae). Bulletin de la Sociétié fisica y politica de Chile. Zoologia, vol. 5, Zoologique de France 114:141. pp. 285–564. Biémont, J.C., and M. Jarry, 1983. Effets Blatchley, W.S. 1910. An illustrated de- inhibiteurs de la présence d’un congé- scriptive catalogue of the Coleoptera or nère sur la maturation des ovocytes chez beetles (exclusive of the Rhynchophora) Acanthoscelides obtectus Say (Coléoptère: known to occur in Indiana. Indiana De- Bruchidae). Canadian Journal of Zoologie partment of Geology and Natural Resourc- 61:2329–2337. es, Bulletin No. 1, Burford, IN. Birch, N., B.J. Southgate, and L.E. Fellows. Blatchley, W.S. 1919. Some new or scarce 1985. Wild and semi-cultivated legumes as Coleoptera from western and southern potential sources of resistance to bruchid Florida.—III. The Canadian Entomologist beetles for crop breeders: A study of Vigna/ 51:65–69. Phaseolus. In G.E. Wickens (ed.), Proceed- ings of the Kew International Conference Blatchley, W.S. 1930a. Notes on the dis- for Arid Lands, Jodrell Laboratory, Royal tribution of Coleoptera in Florida with new Botanic Gardens, Kew, England, 23–27 additions to the fauna of that state. The July, 1984, pp. 303–332. Allen and Unwin, Canadian Entomologist 62:28–35. London. Blatchley, W.S. 1930b. Blatchleyana. Na- Bissell, T.L. 1938. The host plants and ture Publishing Co., Indianapolis. parasites of the cowpea curculio and other Boe, A., B. McDaniel, and K. Robbins. legume infesting weevils. Journal of Eco- 1988. Patterns of American licorice seed nomic Entomology 31:534–536. predation by Acanthoscelides aureolus Bissell, T.L. 1940. Curculionidae, Bru- (Horn) (Coleoptera: Bruchidae) in South chidae, Lepidoptera, and their parasites, Dakota. Journal of Range Management infesting the seeds of cowpea and various 41:342–345. wild plants. Journal of Economic Entomol- Boeving, A.G. 1927. On the classification ogy 33:844–847. of the Mylabridae larvae. Proceedings of Blackwelder, R.E. 1939. Fourth Supple- the Entomological Society of Washington ment 1933 to 1938 (Inclusive) to the Leng 29:133–143. Catalogue of Coleoptera of America, North Boheman, C.H. 1829. Novae coleopterorum of Mexico. John Sherman, Jr., Mount Ver- species. Nouveau Memoires Societe Impe- non, N.Y. rial National Moscou 1:101–133. Blackwelder, R.E. 1946. Checklist of the Boheman, C.H. 1833. [Bruchidae]. In C.J. coleopterous insects of Mexico, Central Schoenherr, ed., Genera et species curcu- America, the West Indies, and South lionidum, cum synonymia hujus familiae: America. Part 4. Bulletin of the United Species novae aut hactenus minus cog- States National Museum 185:551–763. nitae, descriptionibus a Dom. Leonardo Blackwelder, R.E., and R.M. Blackwelder. Gyllenhal, C.H. Boheman, et Entomologis 1948. Fifth Supplement to the Leng Cata- Aliis. Vol. 1, part 1. Roret, Paris. logue of Coleoptera of America, North of Boheman, C.H. 1839. [Bruchidae]. In Mexico. John D. Sherman, Jr., Mount C.J. Schoenherr, ed, Genera et species Vernon, NY. curculionidum, cum synonymia hujus fa- Blanc, F.L. 1965. Zabrotes subfasciatus miliae: Species novae aut hactenus Boheman, new pest species in California.

272 minus cognitae, descriptionibus a Dom. Bottimer, L.J. 1936. Bruchus brachialis in Leonardo Gyllenhal, C.H. Boheman, et En- Georgia. Journal of Economic Entomology tomologis Aliis. Vol. 5, part 1. Roret, Paris. 29:807. Boheman, C.H. 1859. Coleoptera. Species Bottimer, L.J. 1937. Some notes on Bru- novas descripsit. In Kongliga Svenska Fre- chus brachialis Fahraeus. Journal of Eco- gatten Eugenies resa omkring Jorden un- nomic Entomology 30:379. der befal af C.A. Virgin, Aren 1851–1853... Bottimer, L.J. 1944. [Minutes of 540th Zoologi, I, Insecta, Coleopteres. Norstedt meeting, Callosobruchus chinensis in Vicia and Soner, Stockholm, Sweden. faba]. Journal of the Washington Entomo- Bondar, G. 1936. Notas biologicos sobre logical Society 46:23. Bruchideos observados no Brasil. Archivos Bottimer, L.J. 1956. The identity of Bru- do Instituto de Biologia Vegetal 3:1–44. chus arenarius Wolcott. The Coleopterists’ Bonet, A. 1981. Biologia del complejo Bulletin 10:67–68. Acanthoscelides obtectus Say (Col.: Bruchi- Bottimer, L.J. 1961. New United States dae) en poblaciones silvestres y cultivados records in Bruchidae with notes on host de Phaseolus. In Folia Entomologica Mexi- plants and rearing procedures. Annals cana—XIV Congreso Nacional de Entomo- of the Entomological Society of America logia, Abril 6–8, 1981, Mexico, D.F., pp. 54:291–298. 45–46. Bottimer, L.J. 1968a. On the two species of Bonet, A., B. Leroi, J.C. Biémont, et al. Bruchidius established in North America. 1987. Has the Acanthoscelides obtectus The Canadian Entomologist 100:139–145. group evolved in the original zone of its Bottimer, L.J. 1968b. On the location of host plant? In V. Labeyrie, G. Fabres, and types of five species of Bruchidae with D. Lachaise, eds., Insects-Plants, p. 378. notes on early American literature of Acan- W. Junk, Dordrecht, The Netherlands. thoscelides obtectus. The Canadian Ento- Borowiec, L. 1980. Acanthoscelides tar- mologist 100:284–289. nawskii, new species from Bulgaria (Cole- Bottimer, L.J. 1968c. Notes on Bruchidae optera: Bruchidae). Polskie Pismo Entomo- of America north of Mexico with a list of logiczne 50:167–170. world genera. The Canadian Entomologist Borowiec, L. 1987. The genera of seed-bee- 100:1009–1049. tles (Coleoptera, Bruchidae). Polskie Pismo Bottimer, L.J. 1969a. Two new Acantho- Entomologiczne 57:3–207. BBscelides from southern United States Borowiec, L. 1988. Case 2618. Bruchus with notes on related species. The Cana- Linnaeus, 1767, Ptinus Linnaeus, 1767 dian Entomologist 101:975–983. and Mylabris Fabricius, 1775 (Insecta, Co- Bottimer, L.J. 1969b. Bruchidae associ- leoptera): Proposed conservation. Bulletin ated with Mimosa with description of a of Zoological Nomenclature 45:194–196. new species. The Canadian Entomologist Bottimer, L.J. 1931. A vetch bruchid estab- 101:1186–1198. lished in the Middle Atlantic States. U.S. Bottimer, L.J. 1973. Two new American Insect Pest Survey Bulletin 11:347. bruchids in the sordidus group of Sta- Bottimer, L.J. 1935. A new Acanthoscelides tor (Coleoptera: Bruchidae) with notes on from eastern United States. Entomological other species. The Canadian Entomologist News 46:127–129. 105:545–551. Boucher, L., and J. Huignard. 1987. Transfer of male secretions from the sper-

273 matophore to the female insect in Carye- Breitenbacher, J.K. 1925. Variation and don serratus (Ol.): Analysis of the possible heredity in Bruchus quadrimaculatus. The trophic role of these secretions. Journal of Canadian Entomologist 58:131–133. Insect Physiology 33:949–957. Brett, C.H. 1946. Insecticidal properties of Boucher, L., and D. Pierre. 1988. Etude du the indigobush (Amorpha fruticosa). Jour- rhythme d’accouplement chez Caryedon nal of Agricultural Research 73:81–86. serratus (Coleoptera: Bruchidae) en condi- Brewer, I.N., and E. Horber. 1984. Evaluat- tions d’elevage et en conditions naturelles. ing resistance to Callosobruchus chinensis Annales de la Societe Entomologique de Linn. in different seed legumes. In Pro- France 24:151–159. ceedings of the 3rd International Working Boughdad, A., Y. Gillon, and C. Gagnepain. Conference on Stored-Product Entomology, 1987. Trophic value of the amino acids in Kansas State University, Manhattan, KS, the seeds of leguminous plants and larval October 23–28, 1983, pp. 435–443. development of Callosobruchus macula- Bridwell, J.C. 1918. Notes on the Bruchi- tus. Biochemical Systematics and Ecology dae and their parasites in the Hawaiian 15:427–432. Islands. Proceedings of the Hawaiian Ento- Bradley, J.C. 1946. The family name An- mological Society 3:465–509. thribidae, the identity of Amblycerus Thun- Bridwell, J.C. 1919. Some additional notes berg, and the taxonomic position of Eus- on the Bruchidae and their parasites in the phyrus LeConte. Bulletin of the Brooklyn Hawaiian Islands. Proceedings of the Ha- Entomological Society 41:96–99. waiian Entomological Society 4:15–20. Bradley, J.C. 1947. Contributions to our Bridwell, J.C. 1920a. Insects injurious to knowledge of the Mylabridae, seu Bruchi- the algaroba feed industry. The Hawaiian dae with special reference to the fauna of Planter’s Record 22:337–343. northeastern America. Psyche 53:33–42. Bridwell, J.C. 1920b. Notes on the Bru- Brauer, A. 1925. Studies on the embryol- chidae and their parasites in the Hawaiian ogy of Bruchus quadrimaculatus, Fabr. Islands, 3rd paper. The Hawaiian Planter’s Annals of the Entomological Society of Record 4:403–409. America 18:283–305. Bridwell, J.C. 1920c. (Note). Proceedings of Brauer, A. 1928. Spermatogenesis of Bru- the Hawaiian Entomological Society 4:332. chus quadrimaculatus (Coleoptera: Bruchi- dae). Journal of Morphology and Physiol- Bridwell, J.C. 1921a. (Note). Proceedings of ogy 46:217–239. the Hawaiian Entomological Society 4:458 Brauer, A. 1942. Development of bruchid Bridwell, J.C. 1921b. (Note). Proceedings of (Coleoptera) eggs after exposure to low the Hawaiian Entomological Society 4:465. temperature. University of Kentucky Re- Bridwell, J.C. 1923a. The host plants and search Club Bulletin 8:1–5. habits of Acanthoscelides griseolus (Fall). Brauer, A. 1944. Influence of population Proceedings of the Entomological Society of number on egg production in the four- Washington 25:79–80. spotted pea beetle, Bruchus quadrima-cu- Bridwell, J.C. 1923b. Acanthoscelides pul- latus Fabr. Transactions of the Kentucky lus (Fall). Journal of the Washington Acad- Academy of Science 11:56–62. emy of Sciences 13:260. Brauer, A. 1946. Cephalogenesis in rela- Bridwell, J.C. 1923c. [Stator limbatus] in tion to the integration center of the beetle note. Journal of the Washington Academy Callosobruchus maculatus Fabr. Journal of of Sciences 13:261–262. Morphology 78:155–177. Bridwell, J.C. 1925. Bruchidius ater (Marsham), an unrecorded immigrant from

274 Europe. Journal of the Washington Acad- Mexico. Journal of the Washington Acad- emy of Sciences 5:80. emy of Sciences 36:52–57. Bridwell, J.C. 1929a. The cowpea bruchid Bridwell, J.C. 1952. A new genus of Bru- under another name—A plea for one kind chidae affecting Hibiscus in Argentina of entomological specialist. Proceedings of (Bruchinae: Acanthoscelidini). Journal the Entomological Society of Washington of the Washington Academy of Sciences 31:39–44. 42:49–50. Bridwell, J.C. 1929b. Description of a Bridwell, J.C., and L.J. Bottimer. 1933. bruchid immigrant into Hawaii breeding in The hairy-vetch bruchid, Bruchus brachia- the seeds of Convolvulaceae. Proceedings lis Fahraeus, in the United States. Journal of the Entomological Society of Washington of Agricultural Research 46:739–751. 31:112–114. Brimley, C.S. 1938. The Insects of North Bridwell, J.C. 1929c. A preliminary ge- Carolina. North Carolina Department of neric arrangement of the palm bruchids Agriculture, Raleigh, NC. and allies with descriptions of new species. Brindley, T.A., J.C. Chamberlin, F.G. Hin- Proceedings of the Entomological Society of man, and K.W. Gray. 1946. The pea weevil Washington 31:141–160. and methods for its control. U.S. Depart- Bridwell, J.C. 1930. [Amblycerus Thun- ment of Agriculture, Farmer’s Bulletin No. berg]. Footnote to W.D. Pierce, Studies of 1971. the North American weevils belonging to Brindley, T.A., J.C. Chamberlin, and R. the superfamily Platystomidae. Proceedings Schopp. 1952. The pea weevil and methods of the United States National Museum, vol. for its control. U.S. Department of Agricul- 77, no. 2840, art. 17, pp. 29. ture, Farmers’ Bulletin No. 1971. Bridwell, J.C. 1932. The subfamilies of the Britton, W.E. 1897. [Note: Beetles in seeds Bruchidae. Proceedings of the Entomologi- of honey locust]. Entomological News cal Society of Washington 34:100–106. 8:173. Bridwell, J.C. 1935. [Notes of Bruchidae]. Brown, W.J. 1952. Some species of Phy- Proceedings of the Entomological Society of tophaga. The Canadian Entomologist Washington 37:185–187. 84:335–342. Bridwell, J.C. 1938a. Specularius erythri- Brulle, M.A. 1832. Introduction ou consid- nae, a new bruchid affecting seeds of Ery- erations generales sur les animaux arti- thrina (Coleoptera). Journal of the Wash- cules de la Mores et des cyclades. Experi- ington Academy of Sciences 28:69–76. mentales Sciences de Moree 3:124. Bridwell, J.C. 1938b. [Synonymies of Brulle, M.A. 1873. [Cerambycidae (Lami- Acanthoscelides obtectus (Say) and Callo- ini), Bruchidae.] In M. Gemminger and sobruchus maculatus (F.)]. In A.O. Larson E. Harold, Catalogus Coleopterorum hu- and C.K. Fisher, The Bean weevil and the cusque descriptorum synonymicus et southern cowpea Weevil in California, pp. systematicus Cerambycidae (Laminini), 4–5. U.S. Department of Agriculture, Tech- Bruchidae, vol. 10, pp. 2989–3232. Mona- nical Bulletin No. 593. chii, Paris. Bridwell, J.C. 1942. Two new American Burkart, A. 1952. Las leguminosas Argen- bean bruchids (Coleoptera). Revista Chil- tinas, silvestres y cultivados. Acme Agency, ena de Historia Natural Pure y Applicada Buenos Aires, Argentina. 44:249–258. Burkholder, W.E. 1982. Reproductive Bridwell, J.C. 1946. The genera of beetles biology and communication among grain of the family Bruchidae in America north of

275 storage and warehouse beetles. Journal of beetle, Caryedon gonagra (Fab.). Garcia de the Georgia Entomological Society 17(sup- Orta 12:633–643. plement 2):1–10. Cancela da Fonseca, J.P. 1965. Oviposi- Burks, B.D. 1954. Parasitic wasps of the tion and length of adult life in Caryedon Catolaccus group in the Americas. U.S. De- gonagra (Coleoptera: Bruchidae). Bulletin partment of Agriculture, Technical Bulletin of Entomological Research 55:697–707. No. 1093. Cancela da Fonseca, J.P. 1975. Notes sur Burks, B.D. 1971. The Nearctic species of le taux intrinsèque d’accroissement naturel Horismenus Walker (Hymenoptera: Eulo- de la bruche de l’arachide Caryedon serra- phidae). Proceedings of the Entomological tus (Fab.) (Coleoptera, Bruchidae). Terre et Society of Washington 73:68–83. la Vie 29:71–76. Bushnell, R.J. 1936. The development Carr, F.S. 1920. An Annotated List of the and metamorphosis of the mid-intestinal Coleoptera of northern Alberta. Alberta epithelium of Acanthoscelides obtectus Natural History Society, Red Deer, Alberta, (Say) (Coleoptera). Journal of Morphology Canada. 60:221–241. Carvalho, R.P.L. de, and C.J. Rossetto. Bushnell, R.J., and D.C. Boughton. 1940. 1968. Biologia de Zabrotes subfasciatus Longevity and egg production in the com- (Boheman). Revista Brasileira de Entomo- mon bean weevil, Acanthoscelides obtectus logia 13:106–117. (Say). Annals of the Entomological Society Casey, T.L. 1884. Contributions to the of America 33:361–370. Descriptive and Systematic Coleopterology Butare, I., and J-C. Biémont. 1987. of North America. Part 2, pp. 61–198. Col- Les stimuli sensoriels intervenant dans lins, Philadelphia. l’inhibition de l’ovogenèse due a la cohabi- Cassier, P., and J. Huignard. 1979. Etude tation de deux congénères chez Acantho- ultrastructure des glands annexes de scelides obtectus (Say) (Col. Bruchidae). l’appareil génital male chez Acantho- Annales de la Société Entomologique de scelides obtectus Say (Coleoptera: Bruchi- France 23:135–144. dae). International Journal of Insect Mor- Caffrey, D.J. 1943. [Acanthoscelides horni phology and Embryology 8:183–201. (Pic)]. Proceedings of the Entomological Castiglioni, L. 1790. Viaggio negli Stati Society of Washington 45:27. Uniti dell’ America settentrionale fatto negli Caillol, H. 1919. Description d’un Acan- anni 1785, 1786, e 1787. Con algune os- thoscelides nouveau de Timbouctou. Bulle- servazioni sui vegetabli sui utili de qual’ tin de la Société Entomologique de France paese. 2 vols. Milan, Italy. 1919:53–54. Caswell, G.H. 1960. Observations on an Campbell, R.E. 1920. The broad-bean wee- abnormal form of Callosobruchus macula- vil. U.S. Department of Agriculture, Bulle- tus (F.). Bulletin of Entomological Research tin No. 807. 50:671–680. Cancela da Fonseca, J.P. 1956. Contri- Center, T.D., and C.D. Johnson. 1973. buição para o estudo da ecologia de Pachy- Comparative life histories of Sennius (Co- merus acaciae Gyll. (Coleoptera, Bruchi- leoptera: Bruchidae). Environmental Ento- dae). Estudos, Ensaios e Docu-mentos 19, mology 2:669–672. Ministerio do Ultramar, Lisboa, Portugal. Center, T.D., and C.D. Johnson. 1974. Cancela da Fonseca, J.P. 1964. Studies Coevolution of some Bruchidae and their on the larval competition of the bruchid hosts. Ecology 55:1096–1103.

276 Center, T.D., and C.D. Johnson. 1976. Chujo, M. 1937b. Some additions and Host plants and parasites of some Arizona revisions of Bruchidae from the Japanese seed-feeding insects. Annals of the Ento- Empire. Transactions of the Natural His- mological Society of America 69:195–201. tory Society of Formosa 27:189–200. Center, T.D., and R.L. Kipker. 1991. First Clement, S.L., and D.H. Miller. 1982. record in Florida of Acanthoscelides qua- Insect seed predation on Astragalus bisul- dridentatus (Coleoptera: Bruchidae), a catus (Hook.) Gray (Leguminosae). Pan-Pa- potential biological control agent of Mimosa cific Entomologist 58:38–41. pigra. Florida Entomologist 74:159–162. Cock, M.J.W., and H.C. Evans. 1984. Pos- Chen, S.H. 1940. Attempt at a new classifi- sibilities for biological control of Cassia cation of the leaf beetles. Sinensia 11:451– tora and C. obtusifolia. Tropical Pest Man- 481. agement 30:339–350. Chevrolat, L.L.A. 1877. (Les diagnoses de Cockerell, T.D.A. 1902. Record of the nouvelles espèces de bruchides). Bulletin habits of New Mexican Coleoptera. Psyche de la Societé Entomologique de France 9:378–380. 1877: lxxxix–xc, xcvii–xcix, cvi, cxiv–cxv, Conway, J.A. 1983. Notes on the biology cxxv, cxxxiv–cxxxv. and ecology of the groundnut seed beetle Chittenden, F.H. 1902. Some insects inju- Caryedon serratus (Ol.) (Coleoptera: Bru- rious to vegetable crops. U.S. Department chidae) under field conditions in Senegam- of Agriculture, Division of Entomology, bia. Tropical Pest Infestation Laboratory Bulletin No. 33. [Slough, Berkshire, Great Britain], Tropical Chittenden, F.H. 1912a. The broad-bean Stored Product Information Circular No. weevil. U.S. Department of Agriculture, Bu- 45, pp. 11–13. reau of Entomology, Bulletin No. 96:59–82. Costa, C., S.A. Vanin, and S.A. Casari- Chittenden, F.H. 1912b. The cowpea wee- Chen. 1988. Larvas de Coleoptera do Bra- vil. U.S. Department of Agriculture, Bureau sil. Museu de Zoologia Universidade de Sâo of Entomology, Bulletin No. 96:83–94. Paulo, Sâo Paulo, Brasil. Chokouhian, A. 1973. Biology and genera- Craighead, F.C. 1950. Insect enemies of tion number of Callosobruchus maculatus eastern forests. U.S. Department of Ag- F. Entomologie et Phytopathologie Appli- riculture, Miscellaneous Publication No. quees 34:1–4. 657. Choudhuri, D.K., and A. Paul. 1983. Diges- Credland, P.F. 1986. Effect of host avail- tion and utilization of food by the larva and ability on reproductive performance in adult of Callosobruchus chinensis, a pest of Callosobruchus maculatus (F.) (Coleoptera: pulses. Indian Biologist 15:45–47. Bruchidae). Journal of Stored Products Research 22:49–54. Choudhuri, D.K., and A. Paul. 1984. Ef- fects of temperature and relative humidity Credland, P.F., and K.M. Dick. 1987. Food on the fertility and fecundity of Calloso-bru- consumption by larvae of three strains chus chinensis (L.), a serious pest of stored of Callosobruchus maculatus (Coleoptera: pulses. Indian Biologist 16:4–6. Bruchidae). Journal of Stored Products Research 23:31–40. Chujo, M. 1937a. Family Bruchidae, class Insecta, Coleopteroidea-Coleoptera. Fauna Credland, P.F., K.M. Dick, and A.W. Nipponica 10, Fascicle 8(9). Sanseido, To- Wright. 1986. Relationships between lar- kyo. In Japanese. val density, adult size, and egg production in the cowpea seed beetle, Callosobruchus maculatus. Ecological Entomology 11:41– 50.

277 Credland, P.F., and A.W. Wright. 1990. Daniel, S.H., and R.H. Smith. 1994. Func- Oviposition deterrents of Callosobruchus tional anatomy of the egg pore in Calloso- maculatus (Coleoptera: Bruchidae). Physi- bruchus maculatus: a trade-off between gas ological Entomology 15:285–298. exchange and protective functions? Physi- Crotch, G.R. 1867. On the Coleoptera of ological Entomology 19:30–38. the Azores. Proceedings of the Zoological Das, A.K., J. Huignard, M. Barbier, and M. Society of London 1867:359–391. Quesneau-Thierry. 1980. Isolation of the Crotch, G.R. 1870. The genera of Coleop- two paragonial substances deposited into tera studied chronologically (1802–1821). the spermatophores of Acanthoscelides ob- Transactions of the Royal Entomological tectus (Coleoptera, Bruchidae). Experientia Society of London 1870:213–241. 36:918–920. Crowson, R.A. 1938. The metendosternite Davey, P.M. 1958. The groundnut bruchid, in Coleoptera: A comparative study. Trans- Caryedon gonagra (F.). Bulletin of Entomo- actions of the Royal Entomological Society logical Research 49:385–404. of London 87:397–415. Daviault, L. 1928. Sur le développement Crowson, R.A. 1944. Further studies on post-embryonnaire de la bruche du hari- the metendosternite in Coleoptera. Trans- cot: Acanthoscelides obtectus Say. Annales actions of the Royal Entomological Society de la Société Entomologique de France of London 94:273–310. 97:105–132. Crowson, R.A. 1946. A revision of the Davies, J.C. 1972. A note of the occurrence genera of the chrysomelid group Sagrinae of Zabrotes subfasciatus Boh., Coleoptera (Coleoptera). Transactions of the Royal En- (Bruchidae) on legumes in Uganda. East tomological Society of London 97:75–115. African Agricultural and Forestry Journal 37:294–299. Crowson, R.A. 1955. The Natural Classifi- cation of the Families of Coleoptera. Lloyd Davis, C.J. 1967. [Notes and exhibitions]. and Co., London. Proceedings of the Hawaiian Entomological Society 19:344. Crowson, R.A. 1960. The phylogeny of Coleoptera. Annual Review of Entomology Davis, C.J., and N.L.H. Krauss. 1962. Re- 5:111–134. cent introductions for biological control in Hawaii—VIII. Proceedings of the Hawaiian Crowson, R.A. 1984. The use of male ter- Entomological Society 18:245–249. minalia in the higher classification of Cole- optera. Entomologische General 10:53–58. Decelle, J.E. 1951. Contribution à l’étude des Bruchidae du Congo Belge. Revue Curtis, J. 1839. British Entomology; being de Zoologie et de Botanique Africaines illustrations and descriptions of the genera 45:172–192. of insects found in Great Britain and Ire- land; containing coloured figures from na- Decelle, J. 1966. Bruchus serratus Ol., ture of the most rare and beautiful species, 1790, espece-type du genre Caryedon and in many instances of the plants upon Schönherr, 1823. Revue de Zoologie et de which they are found. 16 vol. London. Botanique Africaines 74:169–173. Cushman, R.A. 1911. Notes on the host Decelle, J. 1968. Nouveaux genres et es- plants and parasites of some North Ameri- pèces de Caryedontini (Col.: Bruchidae: can Bruchidae. Journal of Economic Ento- Pachymerinae) d’Afrique et de Madagascar. mology 4:489–510. Bulletin et Annales de la Société Royale Belge d’Entomologie 104:413–426. Decelle, J. 1971. Trois nouvelles especes de Kytorhinus (Col. Bruchidae) du Tibet et du Bhutan. Bulletin et Annales de la

278 Société Royale Entomologique de Belgique De Luca, Y. 1966. Alimentation imaginale 107:105–115. des Bruchidés (Col.). Parasitica 22:26–54. Decelle, J. 1975. Les Bruchides (Coleop- De Luca, Y. 1967a. Hotes larvaires des tera) des Iles Canaries. Bulletin et Annales Bruchides (Col.) sauf Fabacées. Bulletin de la Sociéte Royale d’Entomol-ogique de de la Société Naturelles de l’Ouest de la Belgique 11:109–142. France 64:3–26. Decelle, J. 1981. Bruchidae related to De Luca, Y. 1967b. Notes éthologiques sur grain legumes in the Afro-Tropical area. la ponte et la larve néonate de Bruchidius Series Entomologica 19:193–197. ater (Marsh.) (Col. Bruchidae). Bulletin de Decelle, J. 1990. Algarobius prosopis (Co- la Société Entomologique de France 72:16– leoptera: Bruchidae) dans la péninsule 20. arabique. Bulletin et Annales de la Société De Luca, Y. 1970. Catalogue des meta- Royal Entomologique de Belgique 126:20– zoaires parasites et prédateurs de Bruchi- 22. dae (Coléoptères). Annales de la Société De Geer, C. 1775. Mémoires pour servir à d’Horticulture et d’Histoire Naturelle de l’histoire des insectes. Vol. 5. Hesselberg, l’Herault 110:3–23. Stockholm, Sweden. De Luca, Y. 1972. Catalogue raisonne des Decelle, J., and N. Lodos. 1989. Contri- insectes Antilles Francaises. Annales de bution to the study of legume weevils of Zoologie-Ecologie Animale 4:103–107. Turkey (Coleoptera: Bruchidae). Bulle- De Luca, Y. 1976. Destruction des formes tin et Annales de la Société Royale Belge imaginales d’Acanthoscelides obtectus Say d’Entomologie 125:163–212. (Col. Bruchides) par Neoaplectana carpo- Dejean, P.F.M.A. 1821. Catalogue de la capsae Weiser (Nematoda-Rhabditidae). collection de Coléoptères de M. le Baron Revue de Zoologie Agricole et de Pathologie Dejean. Crevot, Paris. Vegetale 75:127–131. Dejean, P.F.M.A. 1833. Catalogue des De Luca, Y. 1977. Catalogue des meta- coléoptères de la collection de M. le Com- zoaires parasites et prédateurs des bru- te Dejean. Livr. 1:1–96, Livr. 2:97–176. chides (Col.) (Troiseme Note). Bulletin Mequignon-Marvis, Paris de Société Etudes Sciences Naturelle de Nimes 55:5–22. Dejean, P.F.M.A. 1837. Catalogue des coléoptères de la collection de M. le Compte De Luca, Y. 1980. Catalogue des meta- Dejean. Troisième Édition, revue, corrigée zoaires parasites et predateurs des bru- et augmentée. Paris. chides (Col.) (4e note). Bulletin de la So- ciété Etudes Sciences Naturelle de Nimes De Luca, Y. 1959. Les armures génitales 86:37–55. de Bruchus lentis Frölich (Col. Bruchidae). Annales de L’École Nationale d’Agriculture Des Gozis, M. 1881. Quelques rectifica- d’Alger 1:3–7. tions synonymiques touchant différents genera et espèces du coléoptères français, De Luca, Y. 1962. Contribution aux Bru- 1re partie. Bulletin de la Société Entomol- chides (Coleopteres) d’Algérie. Leurs ogique de France 1881:CXII–CXIII. Hôtes—Leurs Parasites—Leurs Stations. Mémoires de la Société d’Histoire Naturelle Des Gozis, M. 1885. Notes et remarques de Afrique du Nord (N.S.) 7:1–15. pour le futur catalogue de la faune Gallo- Rhenane. Revue d’Entomologie 4:116–132. De Luca, Y. 1965. Catalogue des meta- zoaires parasites et prédateurs de Bruchi- Desroches, P. 1983. Developpement lar- dae (Coleoptera). Journal of Stored Prod- vaire d’Acanthoscelides obtectus Say ucts Research 1:51–98. (Col. Bruchidae) en sur peuplement dans les graines de Phaseolus vulgaris et im-

279 portance des facteurs spatiotemporels. Dury, C. 1879. List of the Coleoptera Annales de la Societe Entomologique de observed in the vicinity of Cincinnati. France (N.S.) 19:405–412. Journal of the Cincinnati Society of Natu- Dick, K.M., and P.F. Credland. 1984. Egg ral History 2:162–178. production and development of three Eady, P. 1994. Intraspecific variation in strains of Callosobruchus maculatus (Co- sperm precedence in the bruchid beetle leoptera: Bruchidae). Journal of Stored Callosobruchus maculatus. Ecological Ento- Products Research 20:221–228. mology 19:11–16. Dillon, E.S., and L.S. Dillon. 1961. A Erb, H., and E. Frías. 1983. Efecto de Manual of Common Beetles of Eastern brúquidos (Col.: Bruchidae) sobre la ger- North America. Row, Peterson and Co., minación de semillas de “vinal,” Prosopis Evanston, IL. ruscifolia Gris. (Leguminosae). Cirpon Dobie, P. 1981. The use of resistant variet- 1:167–175. ies of cowpeas (Vigna unguiculata) to re- Erichson, W.F. 1848. Insecten. In R. duce losses due to post-harvest attack by Schomburgk, Versuch einer Fauna und Callosobruchus maculatus. Series Entomo- Flora von Britisch-Guiana, pp. 553–617. logica 19:185–192. Dritter Theil, Leipzig, Germany. Dobie, P., J.V.S. Greve, K. Hothi, and A.M. Errard, C. 1981a. Effet du groupement sur Kilminster. 1979. Inability of storage Bru- l’attractivité des males d’Acanthoscelides chidae to infest winged beans (Psopho-car- obtectus, dans certaines conditions experi- pus tetragonolobus). Entomologia Experi- mentales. Biology of Behaviour 6:229–237. mentalis et Applicata 26:168–174. Errard, C. 1981b. Influence de l’intensité Dohrn, C.A. 1879. [Bruchus abbreviatus da groupement sur l’attractivité des males Melsh.]. Stettiner Entomologische Zeitung d’Acanthoscelides obtectus Say (Coléop- 40:184–189. tère—Bruchidae) vis-a-vis des femelles. Donahaye, E. 1974. Specific identifica- Compte Rendus Hebdomadaire des se- tion within the genus Bruchus (order Co- ances de l’Academie des Sciences Paris leoptera; family Bruchidae) by means of 293:539–544. the genital sclerites. In Progress Report for Essig, E.O. 1926. Insects of Western North the Year 1973/74, pp. 25–32. Ministry of America. Macmillan, New York. Agriculture, Institute for Technology and Essig, E.O. 1929a. Insects of Western Storage of Agriculture Products, Bet Da- North America. Macmillan, New York. gan, Israel. Essig, E.O. 1929b. Origin of the bean Donahaye, E., S. Navarro, and M. Calde- weevil, Mylabris obtectus (Say). Journal of ron. 1966. Observations on the life cycle of Economic Entomology 22:858–861. Caryedon gonagra (F.) on its natural hosts in Israel, Acacia spirocarpa and A. tortilis. Essig, E.O. 1958. Insects and Mites of Tropical Science 8:85–89. Western North America. Macmillan, New York. Downie, N.M. 1950. Notes on the distribution of Bruchus brachialis Fahraeus and Fabricius, J.C. 1775. Systema entomol- Malachius aeneus (L.). The Coleopterists’ ogia, sistens insectorum classes, species, Bulletin 4:20–21. adiectus synonyms, locis, descriptionibus, observationibus. Leipzig, Germany. Doyen, J.T. 1966. The skeletal anatomy of Tenebrio molitor (Coleoptera: Tenebrion- Fabricius, J.C. 1781. Species insectorum, idae). Miscellaneous Publications of the exhibentes eorum differentias specifi- Entomological Society of America 5:101– cas, synonyma auctorum, loca natalia, 150.

280 metamorphosis, adiectis observationibus, Fall, H.C., and T.D.A. Cockerell. 1907. The descriptionibus. Vol. 1. Kiel, Germany. Coleoptera of New Mexico. Transactions Fabricius, J.C. 1787. Mantissa insectorum of the American Entomological Society sistens eorum species nuper detectus adi- 33:145–272. ectis characteribus genericis, differentiis, Faustini, D.L., and D.G.H. Halstead. 1982. specificis, emendationibus, observationi- Setiferous structures of male Coleoptera. bus. Vol. 1. Copenhagen, Denmark. Journal of Morphology 173:43–72. Fabricius, J.C. 1792. Entomologis system- Fernandez, G.C.J., and N.S. Talekar. 1990. atica, emendata et aucta, adijectis synony- Genetics and breeding for bruchid resis- mis, locis, observationibus, descriptioni- tance in Asiatic Vigna species. In K. Fujii, bus. Vol. 1. Copenhagen, Denmark. A.M.R. Gatehouse, C.D. Johnson, et al., Fabricius, J.C. 1798. Supplementum ento- eds., Bruchids and Legumes: Econom- mologiae systematicae. Copenhagen, Den- ics, Ecology, and Coevolution. Proceeding mark. of the 2nd International Symposium on Bruchids and Legumes, Okayama, Japan, Fabricius, J.C. 1801. Systemae eleutherat- September 6–9, 1989, pp. 209–217. Series orum, adjectis synonymis, locis, observa- Entomologica 46. Kluwer, Dordrecht, The tionibus, descriptionibus. Vol. 2. Kiel, Netherlands. Germany. Ferris, G.F. 1942. Some observations Fahraeus, O.I. 1839. [Bruchidae]. In C.J. on the head of insects. Microentomology Schoenherr, ed., Genera et species curcu- 7(Part 2):25–62. lionidum, cum synonymia hujus gamiliae: Filipek, P. 1962. Studies on the bionom- Species novae aut hactenus minus cog- ics and ecology of Acanthoscelides obtectus nitae, descriptionibus a Dom. Leonardo in laboratory conditions. Prace Naukowe Gyllenhal, C.H. Boheman, et entomologis Instytut Ochrony Roslin 4:177–200. aliis. Vol. 5, part 1. Roret, Paris. Fischer von Waldheim, G. 1809. Sur deux Fairmaire, L. 1898. Meteriaux pour la genres nouveaux de coléoptères. Memoires fauna Coleopteres de la region Malagache. Societe Imperial Naturelle Moscou 2:293– Annales de la Societe Entomologique de 304. Belgique 42:222–260. Fitch, A. 1861. The bean weevil, Bruchus Fall, H.C. 1901. List of the Coleoptera of fabae. Transactions of the Rhode Island southern California with notes on habits Society to Encourage Domestic Industry and distribution and descriptions of new 1861:62. species. Occasional Papers of the Califor- nia Academy of Sciences 8:1–282. Fonseca, S.O. 1981. Acerca de la distrib- ucion de Caryobruchus gleditsiae L. (Co- Fall, H.C. 1910. Miscellaneous notes and leoptera: Bruchidae). Folia Entomologica descriptions of North American Coleoptera. Mexicana 50:71–75. Transactions of the American Entomologi- cal Society 36:160–189. Forbes, W.T.M. 1922. The wing venation of the Coleoptera. Annals of the Entomologi- Fall, H.C. 1912. A new Tetropium, two new cal Society of America 15:328–357. Bruchidae, with brief notes on other Cole- Forister, G.W. 1970. Bionomics and ecol- optera. Entomological News 23:320–323. ogy of 11 species of Bruchidae (Coleoptera). Fall, H.C. 1926. A list of the Coleoptera M.S. thesis, Department of Biological Sci- taken in Alaska and adjacent parts of the ences, Northern Arizona University, Flag- Yukon Territory in the summer of 1924. staff. Pan-Pacific Entomologist 2:191–208. Forister, G.W., and C.D. Johnson. 1970. Bionomics of Merobruchus julianus (Co-

281 leoptera: Bruchidae). The Coleopterists’ Frost, C.A. 1931. Mylabris atomus Fall. Bulletin 24:84–87. Bulletin of the Brooklyn Entomological Forister, G.W., and C.D. Johnson. 1971. Society 26:3. Behavior and ecology of Acanthoscelides Fujii, K. 1965. Studies on interspecies prosopoides (Coleoptera: Bruchidae). Pan- competition between the azuki bean wee- Pacific Entomologist 47:224–234. vil and the southern cowpea weevil. I. The Forno, I.W., B. Napompeth, and S. reversal in competition result. Researches Buranapanichpan. 1989. Is biological on Population Ecology 7:43–51. control of Mimosa pigra L. possible? (Ab- Fujii, K., and Khin Mar Wai. 1990. Sex- stract). In Proceedings of the First Asia-Pa- ratio determination in three wasp species cific Conference of Entomology, November ectoparasitic on bean weevil larvae. In K. 8–13, 1989, Chiang Mai, Thailand, p. 265. Fujii, A.M.R. Gatehouse, C.D. Johnson, et Bangkok, Thailand. al., eds., Bruchids and Legumes: Econom- Forster, J.R. 1771. Novas species insec- ics, Ecology and Coevolution. Proceeding torum. Centuria. London. of the 2nd International Symposium on Bruchids and Legumes, Okayama, Japan, Fosberg, F.R. 1966. Miscellaneous notes September 6–9, 1989, pp. 331–340. Series on Hawaiian plants—4. Occasional Papers Entomologica 46. Kluwer, Dordrecht, The of the Bernice P. Bishop Museum 23:129– Netherlands. 138. Fujii, K., A.M.R. Gatehouse, C.D. Johnson, Fox, C.W., and T.A. Mousseau. 1995. et al., eds. 1990. Bruchids and legumes, Determinants of clutch size and seed Economics, Ecology and Coevolution. preference in a seed beetle, Stator beali Proceeding of the 2nd International Sym- (Coleoptera: Bruchidae). Environmental posium on Bruchids and Legumes, Okaya- Entomology 24:1557–1561. ma, Japan, September 6–9, 1989. Series Fox, C.W., and M. Tatar. 1994. Oviposition Entomologica 46. Kluwer, Dordrecht, The substrate affects adult mortality, indepen- Netherlands. dent of reproduction, in the seed beetle Fullaway, D.T. 1913. Report of the ento- Callosobruchus maculatus. Ecological Ento- mologist. In Annual Report of the Hawaiian mology 19:108–110. Agricultural Experiment Station for 1912, Fox, W.B. 1943. Some insects infesting the pp. 24–26. Government Printing Office, “selenium indicator” vetches in Saskatch- Washington, DC. ewan. The Canadian Entomologist 75:206– Fullaway, D.T., and N.L.H. Krauss. 1945. 207. Common Insects of Hawaii. Tongg Publish- Frankenhuyzen, A. van, and F.W. Perquin. ing Co., Honolulu. 1971. Over de levenawijze van Bruchidius Funasaki, G.Y. 1973. New state records ater, een peulmineerder op de Brem. De (Hawaii). U.S. Department of Agricul- Levende Natuur 74:66–68. ture, Cooperative Economic Insect Report Frankenhuyzen, A. van, and F.W. Perquin. 23(12):149. 1972. Over de levenswijze van Bruchidius ater (Marsh, 1802) (sensu Southgate), een Furth, D.G., and K. Suzuki. 1990. The peulmineerder op de brem. Entomologische metatibial extensor and flexor tendons Berichte (Amsterdam) 32:125–129. in Coleoptera. Systematic Entomology 15:443–448. Frick, K.E. 1962. Seed beetle that attacks the seeds of Cytisus scoparius in the east- Furusawa, K. 1987. The lotus seed, ern United States. New Jersey Biocontrol Nelumbo nucifera Gaertn., as a new host Laboratory, Moorestown. Unpublished plant of the adzuki bean weevil Calloso- report. bruchus chinensis L. (Coleoptera: Bruch-

282 idae). Applied Entomology and Zoology insect resistance in Vigna unguiculata. 22:388–389. Journal of Science in Food and Agriculture Gagnepain, C., and J.Y. Rasplus. 1989. 30:948–958. Caryedon serratus and its parasitoids in Gatehouse, A.M.R., S.J. Schackley, K.A. the savanna around Lamto, Ivory Coast. Fenton, et al. 1989. Mechanism of seed Entomophaga 34:559–567. lectin tolerance by a major insect storage pest of Phaseolus vulgaris, Acanthoscelides Garaud, P. 1984. Mise en évidence d’un obtectus. Journal of Science in Food and polymorphisme chromosomique de trans- Agriculture 47:269–280. location dans une population naturelle d’Acanthoscelides obtectus (Coléoptère, Gemminger, M., and E. Harold. 1873. Bruchidae) du Burundi. Genetica 63:85– Catalogus Coleopterorum hucusque 91. descriptorum synonymicus et systematicus, vol. 10, pp. 3218–3232. Monachii, Garaud, P., and P. Lecher. 1982. Etude Paris. biométrique du caryotype de la bruche du haricot Acanthoscelides obtectus, (Coléop- Genaro, J., and J.M. Kingsolver. 1997. tères, Bruchidae). Canadian Journal of Amblycerus schwarzi (Coleoptera: Bruchi- Genetics and Cytology 24:687–692. dae) attacking the seeds of the tropical-al- mond terminalia (Combretaceae) in Cuba. Garg, S.K., S. Mahajan, and S.P. Sharma. Entomological News 108:229–230. 1990. Variations in the life span, enzymes and lipid peroxide levels in aging Caryedon Geoffroy, E.L. 1762. Histoire abrégé des serratus (Coleoptera: Bruchidae). Gerontol- insectes qui se trouvent aux environs de ogy 36:126–131. Paris, dans laquelle ces animaux sont rangés suivant un ordre méthodique, 2 vol. Gatehouse, A.M.R., and J.H. Anstee. 1983. Durand, Paris. The presence and partial characterization of carbohydrase enzymes in the gut Geoffroy, E.L. 1785. [New species]. In A.F. of Callosobruchus maculatus. Experientia Fourcroy, ed., Entomologica parisensis, 39:1013–1015. sive catalogus insectorum, quae in agro parisiensi reperiuntur. Vol. 1. [publisher Gatehouse, A.M.R., L. Barbieri, F. Stirpe, unknown], Paris. and R.R.D. Croy. 1990. Effects of ribosome inactivating proteins on insect develop- George, J., and K.K. Verma. 1997. Vari- ment—differences between Lepidoptera ability in Callosobruchus chinensis (L.) and Coleoptera. Entomologia Experi-men- and evolution of polymorphism in Calloso- talis et Applicata 54:43–51. bruchus (Coleoptera, Bruchidae). Russian Entomological Journal 6:41–48. Gatehouse, A.M.R., and D. Boulter. 1983. Assessment of the antimetabolic effects Germain, J.F., J.P. Monge, and J. Huig- of trypsin inhibitors from cowpea (Vigna nard. 1987. Development of two bruchid unquiculata) and other legumes on develop- populations (Bruchidius atrolineatus (Pic) ment of the bruchid beetle Callosobruchus and Callosobruchus maculatus (Fab.)) in- maculatus. Journal of Science in Food and festing stored cowpea (Vigna unguiculata Agriculture 34:345–350. Walp.) pods in Niger. Journal of Stored Products Research 23:157–162. Gatehouse, A.M.R., K.A. Fenton, and J.H. Anstee. 1985. Carbohydrase and esterase Germar, E.F. 1824. Insectorum species activity in the gut of larval Callosobruchus novae aut minus cognitae, descriptionibus maculatus. Experientia 41:1202–1205. illustratae. Vol. 1. Coleoptera. Hendelu et Filii, Halae, Germany. Gatehouse, A.M.R., J.A. Gatehouse, P. Do- bie, et al. 1979. Biochemical basis of Gibson, A. 1906. Popular and practical entomology. No. 18. The bean weevil

283 (Bruchus obtectus, Say). The Canadian stimuli of legume host seeds in compara- Entomologist 38:365–367. tive ovipositional behaviour of Callosobru- Gibson, L.P. 1972. Revision of the genus chus maculatus (Fab.) and C. chinensis Urosigalphus of the United States and (Linn.). In K. Fujii, A.M.R. Gatehouse, C.D. Canada (Hymenoptera: Braconidae). Mis- Johnson, et al., eds., Bruchids and Le- cellaneous Publications of the Entomologi- gumes: Economics, Ecology, and Coevolu- cal Society of America 8:83–157. tion. Proceeding of the 2nd International Symposium on Bruchids and Legumes, Giga, D.P., and R.H. Smith. 1983. Compar- Okayama, Japan, September 6–9, 1989, ative life history studies of four Callosobru- pp. 45–51. Series Entomologica 46. Klu- chus species infesting cowpeas with special wer, Dordrecht, The Netherlands. reference to Callosobruchus rhodesianus (Pic) (Coleoptera: Bruchidae). Journal of Gokhale, V.G., and B.K. Srivastava. 1975. Stored Products Research 19:189–198. Ovipositional behaviour of Callosobruchus maculatus (Fabricius) (Coleoptera: Bruchi- Giga, D.P., and R.H. Smith. 1985. Oviposi- dae). I. Distribution of eggs and relative tion markers in Callosobruchus maculatus ovipositional preference on several legumi- and Callosobruchus rhodesianus (Coleop- nous seeds. Indian Journal of Entomology tera: Bruchidae): Asymmetry of responses. 37:122–128. Agriculture, Ecosystems and Environment 12:229–239. Green, T.W., and I.G. Palmbald. 1975. Effect of seed predators on Astragalus Gill, J.K., C. Kanwar, and S.R. Bawa. 1971. cibarius and Astragalus utahensis (Legumi- Abnormal “sterile” strain in Callosobruchus nosae). Ecology 56:1435–1440. maculatus (Coleoptera: Bruchidae). Annals of the Entomological Society of America Gupta, D.P., and N. Bhaduri. 1984. Stud- 64:1186–1188. ies on the oviposition of Callosobruchus maculatus. Current Science 53:392–393. Gistel, J. 1848. Naturgeschicte des Thi- erreichs fur hohere Schulen. Hoffmann, Gyllenhal, L. 1833. [Bruchidae]. In C.J. Stuttgart, Germany. Schoenherr, ed., Genera et species curculionidum, cum synonymia hujus familiae: Gistel, J. 1856. Die Mysterien der euro- Species novae aut hactenus minus cogni- paischen insectenwelt. Kempten, Germany. tae, descriptionibus a Dom. Leonardo Gyl- Glick, P.A. 1939. The distribution of in- lenhal, C.H. Boheman, et entomologis aliis. sects, spiders, and mites in the air. U.S. Vol. 1, part 1. Roret, Paris. Department of Agriculture, Technical Bul- Gyllenhal, L. 1839. [Bruchidae]. In C.J. letin No. 673. Schoenherr, ed., Genera et Species Curcu- Glick, P.A. 1957. Collecting insects by air- lionidum, cum Synonymia Hujus Familiae: plane in southern Texas. U.S. Department Species Novae aut Hactenus Minus Cog- of Agriculture, Technical Bulletin No. 1158. nitae, Descriptionibus a Dom. Leonardo Gmelin, J.F. 1790. Linne’s systema na- Gyllenhal, C.H. Boheman, et Entomologis turae, 13th ed., vol. 1, part 4, pp. 1517– Aliis. Vol. 5, part 1. Roret, Paris. 2224. Lipsiae, Liepzig, Germany. Haines, C.P. 1989. Observations on Callo- Goeze, J.A.E. 1777. Entomologische Be- sobruchus analis (F.) in Indonesia, includ- iträge zu des Ritter Linné. Zwolfte Ausgabe ing a key to storage Callosobruchus spp. des Natursystems I. Leipzig, Germany. (Col., Bruchidae). Journal of Stored Prod- ucts Research 25:9–16. Gokhale, V.G., H. Honda, and I. Yama- moto. 1990. Role of physical and chemical Halstead, D.G.H. 1973. Preliminary biological studies on the pheromone produced

284 by male Acanthoscelides obtectus (Say) Hatton, W.H. 1895. Systematic value of (Coleoptera: Bruchidae). Journal of Stored the larva of Spermophagus. The Canadian Products Research 9:109–117. Entomologist 27:290. Hamilton, J. 1892a. Notes on Bruchus al- Hawkeswood, T.J. 1996. Comment on book boscutellatus, Miarus hispidulus, Coelioides review. Australian Journal of Entomology acephalus, and a new Thiobius. Entomo- 35:208, 222. logical News 3:253–255. Heinze, K. 1959. Phytopathogene viren und Hamilton, J. 1892b. Notes on Coleoptera— ihre ueberträger. Dunker and Humblot, No. 10. Canadian Entomologist 24:157– Berlin. 163. Herford, G.H. 1935. A key to the members Hamon, C., J-C. Biémont, and G. Chauvin. of the family Bruchidae (Col.) of economic 1982. Ultrastructure et fonction secrétice importance in Europe. Transactions of the des cellules de la paroi des oviducts laté- Society for British Entomology 2:1–32. raux chez Acanthoscelides obtectus Say Hetz, M., and C.D. Johnson. 1988. Hyme- (Coleoptera: Bruchidae). International nopterous parasites of some bruchid bee- Journal of Insect Morphology and Embry- tles of North and Central America. Journal ology 11:327–339. of Stored Products Research 34:131–143. Hamon, C., J.-C. Biémont, and G. Hewitt, G.B., and W.H. Burleson. 1976. A Chauvin. 1983. Ultrastructure et fonction preliminary survey of the arthropod fauna secrétice des oviductes latéraux le Coleop- of sainfoin in central Montana. Bulletin of tere chez le Coleoptere Bruchidae: Acan- the Montana Agricultural Experiment Sta- thoscelides obtectus. Bulletin de la Société tion No. 693. Entomologique de France 88:249–250. Heyden, L., E. Reitter, and J. Weise. 1883. Harley, K., J. Gillett, J. Winder, et al. 1995. Catalogus coleopterorum Europe et Cau- Natural enemies of Mimosa pigra and M. casi. Berlin, Germany. berlandieri (Mimosaceae) and prospects for control of M. pigra. Environmental Ento- Highland, H.A. 1986. Penetration of pack- mology 24:1664–1678. aging films by the cowpea weevil (Coleop- tera: Bruchidae). Journal of Entomological Harley, K.L.S., W.M. Lonsdale, C.M. Science 21:33–37. Beurle, and R. Segura. 1988. Ecology of Mimosa pigra. Biennial report 1985–87, pp. Hinckley, A.D. 1960. The klu beetle, Mimo- 7.10–7.11. CSIRO, Division of Entomology, sestes sallaei (Sharp), in Hawaii. Proceed- Canberra, Australia. ings of the Hawaiian Entomological Society 17:260–269. Harold, E.V. 1878. Beitrage zur käferfauna von Japan (Viertesstuck): Japanische Käfer Hinckley, A.D. 1961. Comparative ecol- der Berliner Konigi Museum. Deutsche ogy of two beetles established in Hawaii: Entomologische Zeitschrift 22:65–68. an anthribid, Araecerus levipennis, and a bruchid, Mimosestes sallaei. Ecology Harper, R.W. 1966. Bureau of Entomology. 42:526–532. New pest finds. Bulletin of the California Department of Agriculture, vol. 55, no. 2, Hinton, H.E. 1981. Biology of Insect Eggs. pp. 92–98. Pergamon Press, Oxford, NY. 3 vols. Hassan, M.I., N.F. Shaumar, and W.A. Hlavac, T.F. 1972. The prothorax of Cole- Atwa. 1987. Taxonomic study of certain optera: Origin, major features of variation. bruchid lavae [sic] in Egypt (Coleoptera: Psyche 79:123–149. Bruchidae). Bulletin de la Société Royale Hodek, I., A. Bonet, and M. Hodkova. Entomologique d’Egypte 65:13–26. 1981. Some ecological factors affecting

285 diapause in adults of Acanthoscelides Horn, G.H. 1885a. Synonymical notes (no. obtectus from Mexican mountains. Series 3). Entomologica Americana 1:108–113. Entomologica 19:43–55. Horn, G.H. 1885b. Contributions to the Hoffman, A. 1945. Faune de France 44, coleopterology of the United States. Trans- Coleopteres Bruchides et Anthribides. actions of the American Entomological Lechavalier, Paris. Society 12:128–162. Hoffman, A. 1965. Observations sur les Ky- Horn, G.H. 1886. Notes on the “Biologia torhinus et description d’une espece inedite Centrali-Americana.” Transactions of the de la Mongolie centrale (Coleoptera: Bruch- American Entomological Society 13:7–11. idae). Annales de la Societé Entomologique Horn, G.H. 1894. The Coleoptera of Baja de France (N.S.) 1:63–70. California. Proceedings of the California Hoffmann, J.H., F.A.C. Impson, and V.C. Academy of Sciences, 2nd series, 4:302– Moran. 1993. Competitive interactions 449. between two bruchid species (Algarobius Howe, R.W. 1973. Loss of viability of seed spp.) introduced into South Africa for bio- in storage attributable to infestations of logical control of mesquite weeds. Biologi- insects and mites. Seed Science and Tech- cal Control 3:215–220. nology 1:563–586. Hope, J.A., D.F. Horler, and D.G. Row- Howe, R.W., and J.E. Currie. 1964. Some lands. 1967. A possible pheromone of the laboratory observations on the rates of bruchid Acanthoscelides obtectus (Say). development, mortality and oviposition of Journal of Stored Products Research several species of Bruchidae breeding in 3:387. stored pulses. Bulletin of Entomological Hopkins, A.D. 1911. I. Contributions to- Research 55:437–477. ward a monograph of the bark weevils of Hubbard, H.G., and E.A. Schwarz. 1878. the genus Pissodes. U.S. Department of The Coleoptera of Michigan. Proceedings Agriculture, Technical Series No. 20, part of the American Philosophical Society 1, pp. 1–68. 17:593–666. Horber, E. 1978. Resistance to pests of Huignard, J. 1968. Organisation et fonc- grain legumes in the U.S.A. In S.R. Singh, tionnement de l’appareil genital de la H.F. van Emden, and T.A. Taylor, eds., bruche du haricot (Acanthoscelides obtec- Pests of Grain Legumes: Ecology and Con- tus, Coléoptère, Bruchidae). Bulletin Bi- trol, pp. 281–295. Academic Press, Lon- ologique 102:233–248. don. Huignard, J. 1970. Analyse expérimentale Horler, D.F. 1970. (–) methylin-tetradeca- de certains stimuli externes influencant trans-2,4,5-trienoate, an allenic ester pro- l’ovogenèse chez Acanthoscelides obtectus duced by the male dried bean beetle, Acan- Say (Coléoptère, Bruchidae). Colloques thoscelides obtectus (Say). Journal of the Internationaux du Centre National de la Chemical Society C (Organic), pp. 859–862. Recherche Scientifique 189:357–380. Horn, G.H. 1873. Revision of the Bruchi- Huignard, J. 1971. Variations de l’activitié dae of the United States. Transactions of reproductrice des males d’Acanthoscelides the American Entomological Society 4:311– obtectus. Journal of Insect Physiology 342. 17:1245–1255. Horn, G.H. 1875. Synonymical notes and Huignard, J. 1975. Anatomie et histologie description of new species of North Ameri- des glands annexes males au cours de la can Coleoptera. Transactions of the Ameri- vie imaginale chez Acanthoscelides obtec- can Entomological Society 5:126–156. tus. International Journal of Insect Mor- phology and Embryology 4:77–88.

286 Huignard, J. 1976. Interactions between Hummel, A.D. 1827. Essais entomologique the host-plant and mating upon the repro- No. 6. Minister of the Interior, St. Péters- ductive activity of Acanthoscelides obtec- bourg, Russia. tus females (Coleoptera: Bruchidae). In Iablokoff-Khnzorian, S.M. 1966. Con- T. Jermy and Á. Szentesi, eds., The Host sid-érations sur l’édéage des Chrysome- Plant in Relation to Insect Behaviour and lidae et son importance phylogénique. Reproduction, pp. 101–108. Plenum Pub- L’Entomologiste 22:115–137. lishing Co., New York. Iablokoff-Khnzorian, S.M. 1967. Con- Huignard, J. 1983. Transfer and fate of sid-érations sur l’édéage des Chrysome- male secretions deposited in the spermato- lidae et son importance phylogénique. phore of females of Acanthoscelides obtec- L’Entomologiste 23:65–67. tus Say (Coleoptera: Bruchidae). Journal of Insect Physiology 29:55–63. Iablokoff-Khnzorian, S.M., and A.P. Kara- petian. 1973. The ovipositor of Bruchidae Huignard, J. 1984. Transfert, importance and its taxonomic interest. Zoologicheskii physiologique et spécificité des sécrétions Zhurnal 52:1186–1192. males chez les femelles d’Acanthoscelides obtectus (Col., Bruchidae). Bulletin de la International Commission on Zoological Société Entomologique de France 89:953– Nomenclature. 1995. Opinion 1809. Bulle- 962. tin of Zoological Nomenclature 52:208–210. Huignard, J., and J.C. Biémont. 1981. Irwin, H.E., and R.C. Barneby. 1982. The Reproductive polymorphism of populations American Cassiinae. A synoptical revision of Acanthoscelides obtectus from different of Leguminosae tribe Cassieae, Cassiinae Colombian ecosystems. Series Entomologi- in the New World. Memoirs of the New York ca 19:149–164. Botanical Garden 35:1–918. Huignard, J., P. Dupont, and B. Tran. Isely, D. 1973. Leguminosae of the United 1990. Coevolutionary relations between States. I. Subfamily Mimosoideae. Memoirs bruchids and their host plants. The influ- of the New York Botanical Garden 25(1):1– ence on the physiology of the insects. In K. 152. Fujii, A.M.R. Gatehouse, C.D. Johnson, et Isely, D. 1975. Leguminosae of the United al., eds., Bruchids and Legumes: Econom- States. II. Subfamily Caesalpinioideae. ics, Ecology, and Coevolution. Proceeding Memoirs of the New York Botanical Garden of the 2nd International Symposium on 25(2):1–228. Bruchids and Legumes, Okayama, Japan, Isely, D. 1990. Vascular Flora of the South- September 6–9, 1989, pp. 171–179. Series eastern United States, vol. 3, part 2: Le- Entomologica 46. Kluwer, Dordrecht, The guminosae (Fabaceae). University of North Netherlands. Carolina Press, Chapel Hill. Huignard, J., and B. Leroi. 1981. Influence Jacquet, J. 1888. Tableaux analytiques of adult food on the reproduction of vir- Rhynchophores. Échange 4:4–28. gin females of an Acanthoscelides obtectus strain originating from Colombian altopla- Janzen, D.H. 1967. Interaction of the no. Experientia 37:831–833. bull’s-horn acacia (Acacia cornigera L.) with an ant inhabitant (Pseudomyrmex Huignard, J., A. Quesneau-Thierry, and M. ferruginea F. Smith) in eastern Mexico. Barbier. 1977. Isolation, biological action University of Kansas Science Bulletin and evolution of the paragonial substances 47:315–558. contained in the spermatophore of Acan- thoscelides obtectus (Coleoptera). Journal of Insect Physiology 23:351–357.

287 Janzen, D.H. 1969. Seed-eaters versus Attacking Legumes (Pulses), Series Ento- seed size, number, toxicity and dispersal. mologica 19. Proceedings of the Interna- Evolution 23:1–27. tional Symposium, Tours (France), April Janzen, D.H. 1972. Escape of Cassia gran- 16–19, 1980, pp. 131–141. W. Junk, The dis L. beans from predators in time and Hague, The Netherlands. space. Ecology 52:964–979. Jarry, M. 1987. Diet of the adults of Acan- Janzen, D.H. 1975. Interactions of seeds thoscelides obtectus and its effect on the and their insect predators/parasitoids in a spatial pattern of the attacks in the fields tropical deciduous forest. In P.W. Price, ed., of Phaseolus vulgaris. In V. Labeyrie, G. Fa- Evolutionary Strategies of Parasitic Insects bres, and D. Lachaise, eds., Insects-Plants, and Mites, pp. 154–186. Plenum Press, pp. 71–75. W. Junk, Dordrecht, The Neth- New York. erlands. Janzen, D.H. 1977a. The interaction of Jarry, M., and A. Bonet. 1981. Premières seed predators and seed chemistry. Collo- observations sur la contamination par ques Internationaux du Centre National du Zabrotes subfasciatus Boh. (Coleoptera: la Recherche Scientifique 265:415–428. Bruchidae) de gousses de Phaseolus vulgaris L. et P. lunatus L. au Mexique. Acta Janzen, D.H. 1977b. How southern cow- Oecologica/Oecologia Applicata 2:311–315. pea weevil larvae (Bruchidae: Callosobru- chus maculatus) die on non-host seeds. Jarry, M., A. Chacon, and G. Parfait. 1985. Ecology 58:921–927. Influence des fluctations de la temperature et de l’hygrometrie sur “l’activitie” Janzen, D.H. 1978. The ecology and evolu- quotidienne d’Acanthoscelides obtectus sur tionary biology of seed chemistry as relates des plantes de Phaseolus vulgaris en plein to seed predation. In J.B. Harborne, ed., champ. Bulletin d’Sociétié de Zoologique Biochemical Aspects of Plant and Animal France 110:395–402. Coevolution, pp. 163–206. Academic Press, London. Jarry, M., D. Debouzie, and A. Chacon. 1987. Influence de quelques variables liées Janzen, D.H. 1980. Specificity of seed-at- à la plante Phaseolus vulgaris sur la ponte tacking beetles in a Costa Rican deciduous d’Acanthoscelides obtectus dans la nature. forest. Journal of Ecology 68:929–952. Entomologia Expérimentalis et Applicata Janzen, D.H. 1981. Patterns of herbivory 43:105–113. in a tropical deciduous forest. Biotropica Jekel, H. 1855. Insecta Saundersiana: Or 13:271–282. characters of undescribed insects in the Janzen, D.H. 1982. Cenizero tree (Legu- collection of William Wilson Saunders, minosae: Pithecellobium saman) delayed Esq., F.R.S., F.L.S. Part 1. Van Voorst, fruit development in Costa Rican decidu- London. ous forest. American Journal of Botany Johansson, B., and C. Linnaeus. 1763 69:1269–1276. (published 1789). Centuria insectorum. In Janzen, D.H., H.B. Juster, and I.E. Liener. Amoenitates Academicae 6:384–415. J.J. 1976. Insecticidal action of the phytohe- Palm, ed., vols. 1–10 (1785–1790). J.C.D. magglutinin in black beans on a bruchid Schrieber, Erlanger, Germany. beetle. Science 192:795–796. Johnson, C.D. 1963. A taxonomic revision Jarry, M. 1981. Evolution of spatial pat- of the genus Stator (Coleoptera: Bruchi- tern of attacks by Acanthoscelides obtectus dae). Annals of the Entomological Society Say (Coleoptera: Bruchidae) of Phaseolus of America 56:860–865. vulgaris L. pods in south west France. In Johnson, C.D. 1966. Caryedon gonagra V. Labeyrie, ed., The Ecology of Bruchids (Fabricius) established in Mexico (Co-

288 leoptera: Bruchidae). Pan-Pacific Entomol- Johnson, C.D. 1976a. Redescription and ogist 42:162. phylogenetic affinities ofKytorhinus prolix- Johnson, C.D. 1967. Notes on the system- us (Fall) (Coleoptera: Bruchidae: Kytorhini- atics, host plants, and bionomics of the nae). Pan-Pacific Entomologist 52:50–55. bruchid genera Merobruchus and Stator Johnson, C.D. 1976b. Systematics of the (Coleoptera: Bruchidae). Pan-Pacific Ento- genus Stylantheus Bridwell (Coleoptera: mologist 43:264–271. Bruchidae). Journal of the Kansas Ento- Johnson, C.D. 1968. Bruchidae type-speci- mological Society 49:254–261. mens deposited in United States museums Johnson, C.D. 1977a. Two new species of with lectotype designations (Coleoptera). Acanthoscelides (Coleoptera: Bruchidae) Annals of the Entomological Society of from North America, and new bruchid host America 61:1266–1272. records from Desmanthus and Hoffman- Johnson, C.D. 1969a. The status of Bru- seggia (Leguminosae). Pan-Pacific Ento- chus distinguendus Horn (Coleoptera: mologist 53:60–73. Bruchidae). Pan-Pacific Entomologist Johnson, C.D. 1977b. Three new species of 44:279–285. Sennius from Mexico and Central America, Johnson, C.D. 1969b. A redescription of with new host records for other Sennius Acanthoscelides aequalis (Sharp) (Coleop- (Coleoptera: Bruchidae). The Coleopterists’ tera: Bruchidae). Pan-Pacific Entomologist Bulletin 31:117–131. 44:336–339. Johnson, C.D. 1977c. Ecology and behav- Johnson, C.D. 1969c. Horn’s Bruchidae ior of Acanthoscelides mundulus in seeds type material in the Ulke Collection (Cole- of Nissolia schotti (Coleoptera: Bruchidae: optera). Pan-Pacific Entomologist 45:54– Leguminosae). Pan-Pacific Entomologist 56. 53:161–167. Johnson, C.D. 1969d. The lectotype of Bru- Johnson, C.D. 1977d. Notes on the host chus julianus Horn. Annals of the Entomo- plants and distribution of Acanthoscelides logical Society of America 62:676–677. pauperculus (LeConte) (Coleoptera: Bruchi- dae). Pan-Pacific Entomologist 53:303–304. Johnson, C.D. 1969e. The location of the holotypes of Bruchus cubiculus Casey and Johnson, C.D. 1978. Ecology of Neltumius Mylabris wheelocki Blatchley. Pan-Pacific texanus (Coleoptera: Bruchidae) in seeds Entomologist 45:237. of Condalia (Rhamnaceae). Journal of the Kansas Entomological Society 51:432–440. Johnson, C.D. 1970. Biosystematics of the Arizona, California, and Oregon species Johnson, C.D. 1979a. New host records in of the seed beetle genus Acanthoscelides the Bruchidae (Coleoptera). The Coleopter- Schilsky (Coleoptera: Bruchidae). Universi- ists’ Bulletin 33:121–124. ty of California Publications in Entomology Johnson, C.D. 1979b. New host records for 59:1–116. Acanthoscelides (Coleoptera: Bruchidae). Johnson, C.D. 1973. A new Acantho- Pan-Pacific Entomologist 55:61–71. scelides from Indigofera (Coleoptera: Bru- Johnson, C.D. 1980. The use of host chidae). The Coleopterists’ Bulletin 27:169– preferences as taxonomic characters of 174. bruchid beetles (Coleoptera: Bruchidae) Johnson, C.D. 1974. Ecology of two Acan- feeding in the seeds of Cassia (Legumino- thoscelides from Indigofera, with a descrip- sae). Journal of the Kansas Entomological tion of a new species (Coleoptera: Bruchi- Society 53:27–34. dae). Journal of the Kansas Entomological Society 47:268–278.

289 Johnson, C.D. 1981a. Interactions be- Entomological Society of America 77:56– tween bruchid (Coleoptera) feeding guilds 64. and behavioral patterns of pods of the Johnson. C.D. 1984b. New host records Leguminosae. Environmental Entomology and notes on the biology of Stator (Co- 10:249–253. leoptera: Bruchidae). The Coleopterists’ Johnson, C.D. 1981b. Relations of Acan- Bulletin 38:85–90. thoscelides with their plant hosts. Series Johnson, C.D. 1985. Potential use- Entomologica 19:73–81. ful tropical legumes and their relation- Johnson, C.D. 1981c. Host preferences of ships with bruchid beetles. In K.C. Misra, Stator (Coleoptera: Bruchidae) in non-host ed., Ecology and Resource Management seeds. Environmental Entomology 10:857– in Tropics, vol. 1, pp. 206–210. Bhargava 863. Book Depot, Varanasi, India. Johnson, C.D. 1981d. The bionomics of Johnson, C.D. 1986. Caryedon serratus Stator pygidialis (Coleoptera: Bruchidae). (Olivier) (Bruchidae) established in north- The Coleopterists’ Bulletin 35:241–242. ern South America with additional host Johnson, C.D. 1981e. Seed beetle host and locality records from Mexico. The specificity and the systematics of the Le- Coleopterists’ Bulletin 40:264. guminosae. In R.M. Polhill and P.H. Raven, Johnson, C.D. 1987. Relationships be- eds., Advances in Legume Systematics, tween Mimosestes (Coleoptera) and part 2, pp. 995–1027 + 61 pp. microfilm. Acacia (Leguminosae): Is there coevolution Proceedings of the International Legume between these genera? In V. Labeyrie, G. Conference, Royal Botanical Gardens, Kew, Fabres, and D. Lachaise, eds., Insects- England, July 24–29, 1978. Plants, pp. 347–352. W. Junk, Dordrecht, Johnson, C.D. 1983a. Ecosystematics of The Netherlands. Acanthoscelides (Coleoptera: Bruchidae) Johnson, C.D. 1988a. The possible be- of southern Mexico and Central America. ginning of adaptation to a new host by Miscellaneous Publications of the Entomo- bruchid beetles in Venezuela. Biotropica logical Society of America 56:1–370. 20:80–81. Johnson, C.D. 1983b. Mimosestes playa- Johnson, C.D. 1988b. Adaptive radiation zul, new species, with new host records for of Acanthoscelides in seeds: Examples other Mimosestes (Coleoptera: Bruchidae). of legume-bruchid interactions. In C.H. Annals of the Entomological Society of Stirton and J.L. Zarucchi, eds., Advances America 76:816–820. in Legume Biology, pp. 747–779. Mono- Johnson, C.D. 1983c. Handbook on Seed graphs in Systematic Biology. Missouri Insects of Prosopis species. Ecology, Con- Botanical Gardens, St. Louis. trol, and Identification of Seed-Infesting Johnson, C.D. 1990a. Six new species Insects of New World Prosopis (Legumin- of Acanthoscelides from North and Cen- osae). Food and Agriculture Organization of tral America (Coleoptera: Bruchidae). The the United Nations, Rome. Coleopterists’ Bulletin 44:3–18. Johnson, C.D. 1983d. New host records for Johnson, C.D. 1990b. Coevolution of Abutiloneus idoneus Bridwell (Coleoptera: Bruchidae and their hosts: Evidence, Bruchidae). The Coleopterists’ Bulletin conjecture, and conclusions. In K. Fujii, 37:378. A.M.R. Gatehouse, C.D. Johnson, et al., Johnson, C.D. 1984a. Sennius yucatan, eds., Bruchids and Legumes: Economics, new species, a redescription of S. infractus, Ecology, and Coevolution, . Proceeding and new host records for other Sennius of the 2nd International Symposium on (Coleoptera: Bruchidae). Annals of the Bruchids and Legumes, Okayama, Japan, September 6–9, 1989pp. 181–188. Series

290 Entomologica 46. Kluwer, Dordrecht, The (Coleoptera). The Coleopterists’ Bulletin Netherlands. 31:154. Johnson, C.D. 1990c. Systematics of the Johnson, C.D., and J.M. Kingsolver. 1982. seed beetle genus Acanthoscelides (Bruchi- Checklist of the Bruchidae (Coleoptera) of dae) of northern South America. Transac- Canada, United States, Mexico, Central tions of the American Entomological Soci- America, and the West Indies. The Coleop- ety 116:297–618. terists’ Bulletin 35:409–422. Johnson, C.D. 1990d. Confirmation ofHe - Johnson, C.D., J.M. Kingsolver, and A.L. dysarum boreale Nuttall (Leguminosae) as Teran. 1989. Sistematica del genero Stator a host plant for Acanthoscelides fraterculus en Sudamerica. Opera Lilloana 37:1–105. (Horn) (Coleoptera: Bruchidae). Pan-Pacific Johnson, C.D., and R.A. Kistler. 1987. Entomologist 66:175–176. Nutritional ecology of bruchid beetles. In Johnson, C.D. 1990e. New and updated F. Slansky, Jr., and J.G. Rodriguez, eds., host names (Leguminosae: Desmanthus) Nutritional Ecology of Insects, Mites and for some Bruchidae (Coleoptera). Pan-Pa- Spiders, pp. 259–282. John Wiley and cific Entomologist 66:324–325. Sons, New York. Johnson, C.D. 1995a. New host and dis- Johnson, C.D., and T.N. Seeno. 1993. tribution records for Neltumius texanus Mimosestes nubigens (Motschulsky) estab- (Schaeffer) (Coleoptera: Bruchidae). The lished in California (Coleoptera: Bruch- Coleopterists’ Bulletin 49:42. idae). Pan-Pacific Entomologist 69:190. Johnson, C.D. 1995b. New host records Johnson, C.D., and D.H. Siemens. 1995a. from Latin America and new synonymy for Bruchid guilds, host preferences, and new Stator limbatus (Horn) and S. cearanus (Pic) host records from Latin America and Texas (Coleoptera: Bruchidae). The Coleopterists’ for the genus Stator (Coleoptera: Bruchi- Bulletin 49:319–326. dae). The Coleopterists’ Bulletin 49:133– Johnson, C.D., and J.M. Kingsolver. 1971. 142. Descriptions, life histories, and ecology Johnson, C.D., and D.H. Siemens. 1995b. of two new species of Bruchidae infesting Oviposition behavior, guilds, distribution guacima in Mexico. Journal of the Kansas and new host records for the genus Mimo- Entomological Society 44:141–152. sestes Bridwell (Coleoptera: Bruch-idae) Johnson, C.D., and J.M. Kingsolver. 1973. from Venezuela, Colombia and Mexico. The A revision of the genus Sennius of North Coleopterists’ Bulletin 50:155–160. and Central America (Coleoptera: Bru-chi- Johnson, C.D., and D.H. Siemens. 1997a. dae). U.S. Department of Agriculture, Tech- Distribution, oviposition guilds, behavior nical Bulletin No. 1462. and new host records from Latin America Johnson, C.D., and J.M. Kingsolver. 1975. for Algarobius Bridwell, Scutobruchus King- Ecology and redescription of the Arizona solver and Pseudopachymerina spinipes grape bruchid Amblycerus vitis (Coleop- (Erichson) (Coleoptera: Bruch-idae). The tera). The Coleopterists’ Bulletin 29:321– Coleopterists’ Bulletin 51:37–42. 331. Johnson, C.D., and D.H. Siemens. 1997b. Johnson, C.D., and J.M. Kingsolver. 1976. Oviposition behavior, guilds, host relation- Systematics of Stator of North and Central ships and new host and distribution re- America (Coleoptera: Bruchidae). U.S. De- cords for the genus Merobruchus Bridwell partment of Agriculture, Technical Bulletin (Coleoptera: Bruchidae). The Coleopterists’ No. 1537. Bulletin 51:13–21. Johnson, C.D., and J.M. Kingsolver. 1977. Johnson, C.D., and C.N. Slobodchikoff. New taxonomic combinations in Bruchidae 1979. Coevolution of Cassia (Leguminosae)

291 and its seed beetle predators (Bruchidae). Acanthoscelides quadridentatus and A. Environmental Entomology 8:1059–1064. puniceus (Col.:Bruchidae) for biological Johnson, C.D., S. Zona, and J.A. Nilsson. control of Mimosa pigra (with prelimi- 1995. Bruchid beetles and palm seeds: re- nary data on their biology). Entomophaga corded relationships. Principes 39:25–35. 35:85–96. Johnston, M.C. 1962. The North American Keeler, K.H. 1980. The extrafloral nectaries mesquites Prosopis sect. Algarobia (Legumi- of Ipomoea leptophylla (Convolvulaceae). nosae). Brittonia 14:72–90. American Journal of Botany 67:216–222. Jolivet, P. 1957. Recherches sur l’aile des Kempers, K.J.W.B. 1923. Abbildungen von Chrysomeloides (Coleoptera). Memoires Flügelgeäder der Coleopteren. Entomolo- Institut Royal des Sciences Naturelles de gische Mitteilungen 12:71–163. Belgique, Serie 2, Fascicle 51:1–152. Kingsolver, J.M. 1964a. The genus Neltu- Kannan, K.K. 1923. The function of the mius (Coleoptera: Bruchidae). The Coleop- prothoracic plate in bruchid larvae. Mysore terists’ Bulletin 18:105–111. Department of Agriculture, Entomology Kingsolver, J.M. 1964b. A preliminary Service Bulletin, No. 7. key to the species of the genus Bruchus Kasap, H. 1978a. The central nervous (Bruchidae) commonly intercepted in systems of Bruchidae and Chrysomelidae USDA plant quarantine interceptions. (Coleoptera). Hacettepe Bulletin of Natural Plant Quarantine Memo. Sciences and Engineering 7–8:21–29. Kingsolver, J.M. 1965a. On the genus Kasap, H. 1978b. A comparative ana- Abutiloneus Bridwell. The Coleopterists’ tomical study of the alimentary canal of Bulletin 19:125–128. Chrysomeloidea (Coleoptera: Polyphaga). Kingsolver, J.M. 1965b. A new fossil Communications de la Facult des Sciences bruchid genus and its relationships to de l’Universi d’Ankara, Serie C3, Zoologie modern genera (Coleoptera: Bruchidae: 22:53–78. Pachymerinae). The Coleopterists’ Bulletin Kasap, H., and R.A. Crowson. 1979. The 19:25–30. male reproductive organs of Bruchidae and Kingsolver, J.M. 1967. A name change in Chrysomelidae. Turkiye Bitki Korumu Der- Bruchidae (Coleoptera). U.S. Department gisi 3:199–216. of Agriculture, Agricultural Research Ser- Kasap, H., and R.A. Crowson. 1980. The vice. Cooperative Economic Insect Report female reproductive organs of Bruchidae 17(39):900. and Chrysomelidae. Turkiye Bitki Korumu Kingsolver, J.M. 1968. A review of the ob- Dergisi 4:85–102. tectus group in Acanthoscelides Schilsky, Kasap, H., and R.A. Crowson. 1985. The with designations of lectotypes (Coleoptera: studies on the ovipositors and eighth ab- Bruchidae: Bruchinae). Proceedings of dominal segments of some species of Bru- the Entomological Society of Washington chidae and Chrysomelidae (Coleoptera). 70:4–9. Turkiye Bitki Korumu Dergisi 9:131–145. Kingsolver, J.M. 1969a. A key to the spe- Kasap, H., and R.A. Crowson. 1988. Mus- cies of Callosobruchus (Bruchidae) inter- culature of the pre-genital abdominal seg- cepted in USDA plant quarantine inspec- ments of Bruchidae and Chrysomelidae tions. Plant Quarantine Memo 69. (Coleoptera), and its systematic and phylo- Kingsolver, J.M. 1969b. A new species of genetic significance. Turkiye Entomologica neotropical seed weevil affecting pigeon Dergisi 12:141–150. peas, with notes on two closely related spe- Kassulke, R.C., K.L.S. Harley, and G.V. cies (Coleoptera: Bruchidae: Bruch Maynard. 1990. Host specificity of

292 inae). Proceedings of the Entomological Kingsolver, J.M. 1980a. Eighteen new spe- Society of Washington 71:50–55. cies of Bruchidae, principally from Costa Kingsolver, J.M. 1970a. A study of male Rica, with host records and distributional genitalia in Bruchidae (Coleoptera). Pro- notes (Insecta: Coleoptera). Proceedings of ceedings of the Entomological Society of the Entomological Society of Washington Washington 72:370–386. 93:229–283. Kingsolver, J.M. 1970b. A synopsis of the Kingsolver, J.M. 1980b. The quadriden-ta- subfamily Amblycerinae Bridwell in the tus group of Acanthoscelides: descriptions West Indies, with descriptions of new spe- of three new species, notes, synonymies, cies. (Coleoptera: Bruchidae). Transactions and a new name (Coleoptera: Bruchidae). of the American Entomological Society Brenesia 17:281–294. 96:469–497. Kingsolver, J.M. 1986. A taxonomic study Kingsolver, J.M. 1970c. Insects not known of the genus Algarobius (Coleoptera: Bru- to occur in the continental United States. chidae). Entomography 4:109–136. Groundnut bruchid (Caryedon serratus Kingsolver, J.M. 1988. Biosystematics (Olivier)). U.S. Department of Agriculture, of the genus Merobruchus of continental Cooperative Economic Insect Report No. North America and the West Indies (Co- 20, pp. 303–304. leoptera: Bruchidae). U.S. Department of Kingsolver, J.M. 1972a. Synopsis of the ge- Agriculture, Technical Bulletin No. 1744. nus Stator Bridwell in the West Indies, with Kingsolver, J.M. 1990a. New World Bru- descriptions of new species (Coleoptera: chidae: Past, present, future. In K. Fujii, Bruchidae). Proceedings of the Entomologi- A.M.R. Gatehouse, C.D. Johnson, et al., cal Society of Washington 74:219–229. eds., Bruchids and Legumes: Econom- Kingsolver, J.M. 1972b. Description of a ics, Ecology, and Coevolution. Proceeding new species of Algarobius Bridwell (Coleop- of the 2nd International Symposium on tera: Bruchidae). The Coleopterists’ Bulle- Bruchids and Legumes, Okayama, Japan, tin 26:116–120. September 6–9, 1989, pp. 121–129. Series Entomologica 46. Kluwer, Dordrecht, The Kingsolver, J.M. 1975a. New synonymies Netherlands. and combinations in North American Bru- chidae (Coleoptera). Proceedings of the En- Kingsolver, J.M. 1990b. Biosystematics tomological Society of Washington 77:60. of the genus Zabrotes of America north of Mexico (Coleoptera: Bruchidae). Transac- Kingsolver, J.M. 1975b. Amblycerus acap- tions of the American Entomological Soci- ulcensis, a new species of seed beetle from ety 116:135–174. Mexico (Coleoptera: Bruchidae). Journal of the Washington Academy of Sciences Kingsolver, J.M. 1990c. Checklist of Chil- 65:33–35. ean Bruchidae with new synonymies and new combinations (Coleoptera). Revista Kingsolver, J.M. 1979a. New synonymies Chilena de Entomologia 18:49–52. and new combinations in North Ameri- can Bruchidae. The Coleopterists’ Bulletin Kingsolver, J.M. 1992a. Caryedon serratus 33:341–342. (Olivier) new to continental United States (Coleoptera: Bruchidae). Insecta Mundi Kingsolver, J.M. 1979b. A new host record 6:22. for Callosobruchus chinensis (L.) (Coleop- tera: Bruchidae). The Coleopterists’ Bulle- Kingsolver, J.M. 1992b. New records of tin 33:438. Bruchidae (Coleoptera) from the Domini- can Republic. Insecta Mundi 6:78 Kingsolver, J.M. 1995a. New locality records for Bruchidae of Florida and the

293 West Indies (Coleoptera). Insecta Mundi Kingsolver, J.M., and D.R. Whitehead. 9:170. 1976. The North and Central Ameri- Kingsolver, J.M. 1995b. On the family can species of Meibomeus (Coleoptera: Bruchidae. Chrysomela No. 30, p. 2. Bruchidae: Bruchinae). U.S. Department of Agriculture, Technical Bulletin No. 1523. Kingsolver, J.M. 1996. Amblycerus schwarzi Kingsolver (Coleoptera: Bruch- Kirk, V.M. 1969. A list of beetles of South idae) recorded new for North America. Carolina. Part 1- northern Coastal Plain. Insecta Mundi 10:93. South Carolina Agricultural Experiment Station, Technical Bulletin No. 1933. Kingsolver, J.M., and L. Borowiec. 1988. The genus Spermophagus in the New World Kirk, V.M. 1970. A list of the beetles of (Coleoptera, Bruchidae). Elytron 2:81–84. South Carolina. Part 2—Mountain, Pied- mont and southern Coastal Plain. South Kingsolver, J.M., T.J. Gibb, and G.S. Carolina Agricultural Experiment Station, Pfaffenberger. 1989. Synopsis of the bru- Technical Bulletin No. 1038. chid genus Althaeus Bridwell (Coleoptera) with descriptions of two new species. Kirk, V.M., and E.U. Balsbaugh, Jr. 1975. Transactions of the American Entomologi- A list of the beetles of South Dakota. South cal Society 115:57–82. Dakota University Agricultural Experiment Station, Technical Bulletin No. 42. Kingsolver, J.M., and C.D. Johnson. 1978. Systematics of the genus Mimosestes (Co- Kistler, R.A. 1982. Effects of temperature leoptera: Bruchidae). U.S. Department of on six species of seed beetles (Coleoptera: Agriculture, Technical Bulletin No. 1590. Bruchidae): An ecological perspective. Annals of the Entomological Society of Kingsolver, J.M., C.D. Johnson, S.W. America 75:266–271. Swier, and A. Teran. 1977. Prosopis fruits as a resource for invertebrates. In B.B. Kistler, R.A. 1985. Host-age structure Simpson, ed., Mesquite—Its Biology in Two and parasitism in a laboratory system of Desert Ecosystems, pp. 109–122. US/IBP two hymenopterous parasitoids and lar- Synthesis Series. Dowden, Hutchinson & vae of Zabrotes subfasciatus (Coleoptera, Ross, Stroudsburg, PA. Bruchidae). Environmental Entomology 14:507–511. Kingsolver, J.M., J. Romero N., and C.D. Johnson. 1993. Files and scrapers: Cir- Kitamura, K., M. Ishimoto, and S. Ishii. cumstantial evidence for stridulation in 1990. Bruchid resistance factors in three species of Amblycerus, one new Phaseolus and Vigna legumes. In K. Fujii, (Coleoptera: Bruchidae). Pan-Pacific Ento- A.M.R. Gatehouse, C.D. Johnson, et al., mologist 69:122–132. eds., Bruchids and Legumes: Econom- ics, Ecology, and Coevolution. Proceeding Kingsolver, J.M., and P. Silva. 1991. Up- of the 2nd International Symposium on date of scientific names of Bruchidae Bruchids and Legumes, Okayama, Japan, (Coleoptera) listed by Bondar in “Notas Bi- September 6–9, 1989, pp. 229–239. Series ológicos” (1931 and 1932). Anais da Socie- Entomologica 46. Kluwer, Dordrecht, The dade Entomológica do Brasil 20:411–415. Netherlands. Kingsolver, J.M., and D.R. Whitehead. Kitch, L.W., and L.L. Murdock. 1986. Par- 1974. Classification and comparative biol- tial characterization of a major gut thiol ogy of the seed beetle genus Caryedes proteinase from larvae of Callosobruchus Hummel (Coleoptera: Bruchidae). Transac- maculatus. Archives of Insect Biochemistry tions of the American Entomological Soci- and Physiology 3:561–576. ety 100:341–430. Knull, J.N. 1934. Notes on Coleoptera, No. 4. Entomological News 45:207–212.

294 Krauss, N.L.H. 1944. [Note]. Proceedings of the Pacific Northwest with suggestions for the Hawaiian Entomological Society 12:11. its control on seed peas. U.S. Department Krauss, N.L.H. 1945. Notes on some Ha- of Agriculture, Technical Bulletin No. 599. waiian insects. Proceedings of the Hawai- Larson, A.O., and C.K. Fisher. 1924. Lon- ian Entomological Society 11:309–317. gevity and fecundity of Bruchus quadri- Krauss, N.L.H. 1962. Biological control maculatus Fab. as influenced by different investigations on insect, snail and weed foods. Journal of Agricultural Research pests in tropical America, 1961. Proceed- 29:297–305. ings of the Hawaiian Entomological Society Larson, A.O., and C.K. Fisher. 1938. The 18:131–133. bean weevil and the southern cowpea Krauss, N.L.H. 1963. Biological control weevil in California. U.S. Department of investigations on Christmas berry (Schi- Agriculture, Technical Bulletin No. 593. nus terebinthifolius) and emex (Emex sp.). Larson, A.O., and P. Simmons. 1923. Notes Proceedings of the Hawaiian Entomological on the biology of the four-spotted bean Society 18:282–287. weevil, Bruchus quadrimaculatus Fab. Krombein, K.V., P.D. Hurd, Jr., D.R. Smith, Journal of Agricultural Research 26:609– and B.D. Burks. 1979. Catalog of Hy- 616. menoptera in America, nNrth of Mexico. Larson, A.O., and P. Simmons. 1924a. Smithsonian Institution Press, Washing- Insecticidal effect of cold storage on bean ton, DC. 3 vols. weevils. Journal of Agricultural Research Kumar, D., and K.K. Verma. 1980. Aedea- 27:99–105. gal musculature in Phytophaga (Coleop- Larson, A.O., and P. Simmons. 1924b. tera). Journal of Natural History 14:237– Longevity and fecundity of Bruchus 270. quadrimaculatus Fab. as influenced by Küster, H.C. 1850. Die Käfer Europa’s. different foods. Journal of Agricultural Re- Heft 19, Nos. 1–100. Baure und Raspe, search 29:297–305. Nürnberg, Germany. Latreille, P.A. 1802. Histoire naturelle, Labeyrie, V. 1968. Evolution de l’activitié générale et particulière, des crustacés et reproductrice des males d’Acanthoscelides des insectes, vol. III, xii. F. Dufart, Paris. obtectus. Annales d’Epiphyties 19:197. Latreille, P.A. 1810. Considérations Labeyrie, V., ed. 1970. L’influence des générales sur l’ordre naturel del animaux stimuli externes sur la gamétogenè des in- composant les classes des crustacés, des sectes. Colloques internationaux du Centre arachnides, et des insectes; avec un tab- National de la Recherche Scienti-fique. No. leau methodique de leurs genres, disposés 189. Paris. en families. C.F. Schoell, Paris. Labeyrie, V. 1981. Rencontre des sexes Latreille, P.A. 1825. Familles naturelles du d’Acanthoscelides obtectus Say (Insectes règne animal. Baillière, Paris. Coléoptères, Bruchidae) dans des univ- Lawrence, J.F., and A.F. Newton, Jr. 1982. ers expérimentaux monotones. Biology of Evolution and classification of beetles. An- Behaviour 6:59–72. nual Review of Ecology and Systematics Lago, P.K., and M.O. Mann. 1987. Survey 13:261–290. of Coleoptera associated with flowers of LeConte, J.E. 1824. Descriptions of some wild carrot (Daucus carota L.) in northern new species of North American insects. Mississippi. The Coleopterists’ Bulletin Annals of the Lyceum of Natural History 41:1–8. 1:169–173. Larson, A.O., T.A. Brindley, and F.G. Hin- LeConte, J.L. 1857. Report upon the in- man. 1938. Biology of the pea weevil in sects collected on the survey. (Reports of

295 explorations and surveys for a railroad Leroi, B. 1981. Feeding, longevity and route from the Mississippi River to the Pa- reproduction of adults of Acanthoscelides cific Ocean.) War Department, Washington, obtectus Say in laboratory conditions. DC. [Preprint] Series Entomologica 19:101–111. LeConte, J.L. 1858. Descriptions of new Leroi, B., C. Chararas, and J.M. Chipou- species of Coleoptera, chiefly collected by let. 1984. Etude des activités osidasiques the United States and Mexican Boundary du tube digestif des adultes et des larves Commission, under Major W.H. Emory, de la bruche du haricot, Acanthoscelides U.S.A. Proceeding of the Academy of Sci- obtectus. Entomologia Experimentalis et ences, Philadelphia 1858:59–89. Applicata 35:269–273. LeConte, J.L., ed. 1883. The Complete Leroi, B., and M. Jarry. 1981. Relations wWitings of Thomas Say on the Entomol- d’Acanthoscelides obtectus avec différen- ogy of North America. Bailliere Bros., New tes espèces de Phaseolus: Influence sur la York. fécondité et possibilitiés de développment Leech, H.B. 1954. [Stator limbatus (Horn)]. larvaire. Entomologia Experimentalis et Pan-Pacific Entomologist 30:85–86. Applicata 30:73–82. Leech, H.B. 1959. [Note]. Pan-Pacific Ento- Linnaeus, C. 1758. Systema naturae per mologist 35:60. regna tria naturae. 10th ed., vol. 1. Hol- miae, Stockholm, Sweden. Leng, C.W. 1920. Catalogue of Coleop- tera of America, nNoth of Mexico. John D. Linnaeus, C. 1767. Systema naturae per Sherman, Jr., Mount Vernon, NY. regna tria naturae. 12th ed., vol. 1, pars 2, pp. 533–1327. Holmiae, Stockholm, Swe- Leopold, R.A. 1941. The role of male acces- den. sory glands in insect reproduction. Annual Review of Entomology 21:199–221. Linnaeus, C. 1788. Caroli a linne systema naturae per regna tria naturae, secundum Lepesme, P. 1942. Sur l’éclosion et le com- classes, ordines, genera, species, cum portement de la larve néonate chez Acan- characteribus, differentiis, synonymis, thoscelides obtectus Say (Col., Bruchidae). locis. Edito decima tertia, aucta, reformata. Bulletin de la Société Entomologique de 13th ed., vol. 1, pp. 1517–2224. Beer, Lip- France 47:7–9. siae, Germany. Lepesme, P. 1944. Les Coleopteres des Loi, G., and L. Fornasari. 1985. Influenza dendrees alimentaires et des produits della temperatura sullo sviluppo dell’uovo industriels entreposes. Encyclopedie Ento- de Acanthoscelides obtectus (Say) (Co- mologie, Series A 22:198–219. leoptera, Bruchidae). Frustula Entom- Lepesme, P. 1947. Les insectes des palm- ologica (N.S.) 6:407–412. iers. P. Chevalier, Paris. Lucas, M.H. 1858. [Spermophagus Leroi, B. 1978. Alimentation des adultes semifasciatus]. Bulletin Entomologique d’Acanthoscelides obtectus Say (Coléoptère, in Annals de la Société Entomologique de Bruchidae): Influence sur la longévite et la France 1858:28. production ovarienne des individus vi- Luckow, M., and C.D. Johnson. 1987. erges. Annales Zoologie-Ecologie Animales New host records of Bruchidae (Coleop- 10:559–567. tera) from Desmanthus (Leguminosae) from Leroi, B. 1980. Regulation de la production Texas and Mexico. Pan-Pacific Entomolo- ovarienne chez Acanthoscelides obtectus gist 63:48–49. (Coleoptera: Bruchidae): influence de l’age Luk’yanovich, F.K., and M.E. Ter-Minas- des femelles lors de la fecondation et la sian. 1957. Bruchidae. Zoologica Institute presence des grains. Entomologia Experi- Akademie Nauk SSSR, New Series No. 67. mentalis et Applicata 28:132–144. Fauna SSSR, Tome 24, No. 1.

296 McFarlane, J.A., and A.J.S. Wearing. Mathwig, J.E. 1971. Relationships between 1967. A means of differentiating between bruchid beetles (Amblycerus robiniae) and Acanthoscelides obtectus (Say) and honey locust trees (Gleditsia triacanthos). Zabrotes subfasciatus (Boh.) (Coleoptera: Ph.D. thesis, Department of Entomology, Bruchidae) in white haricot at the pupal University of Kansas, Lawrence. stage. Journal of Stored Products Research Mathwig, J.E. 1972. Adult bruchid activity 3:261–262. as indicated by tanglefoot traps. Journal of MacLeod, G.F. 1933. Effects of ultraviolet the Kansas Entomological Society 45:200– radiations on the bean weevil, Bruchus 208. obtectus Say. Annals of the Entomological Maulik, S. 1941. Biology and morphol- Society of America 26:603–615. ogy of the Sagrinae (Chrysomelidae, Co- Mann, J., and R.A. Crowson. 1983a. On leoptera). Annals and Magazine of Natural the occurrence of mid-gut cecae, and History 7:235–256. organs of symbiont transmission, in leaf- Mead, F., ed. 1989. [Seed weevils–new beetles (Coleoptera: Chrysomelidae). The Florida records]. Tri-Ology Technical Re- Coleopterists’ Bulletin 37:1–15. port 28:2. Mann, J.S., and R.A. Crowson. 1983b. Meik, J., and P. Dobie. 1986. The ability of Phylogenetic significance of the ventral Zabrotes subfasciatus to attack cowpeas. nerve cord in the Chrysomelidae. System- Entomologia Experimentalis et Applicata atic Entomology 8:103–108. 42:151–158. Mansour, K. 1934. On the phylogenetic Melsheimer, F.V. 1806. A Catalogue of classification of the Coleoptera. Bulletin de Insects of Pennsylvania. W.D. Lepper, la Société Royale Entomologique d’Egypte Hanover, PA. 4:190–203. Melsheimer, F.E. 1853. Catalogue of the Manter, J.A. 1917. Notes on the bean Described Coleoptera of the United States. weevil. Journal of Economic Entomology Smithsonian Institution, Washington, DC. 10:190–193. Menten, L.A., and J.O.M. Menten. 1984. Marcu, O. 1939. Die Stellung der Bruch- Determination of the period of infesta- idae im System auf Grund vergleichner tion of the bean (Phaseolus vulgaris L.) by Untersuchungen des Flügeladers. Folia Acanthoscelides obtectus (Say, 1813) un- Zoologica et Hydrobiologica (Strand ed.) der field conditions. Turrialba (Turrialba) 9:370–374. 34:333–336. Marsham, T. 1802. Entomologia Brittanica, Menten, L.A.S., F.M. Wiendl, and J.O.M. sistens insects Brittaniae indigena, secun- Menten. 1981. Determination of the flight dum methodum Linnaeanum disposita. range of Acanthoscelides obtectus (Say, Tom. 1, Coleoptera. J. White, London. 1831) (Coleoptera, Bruchidae) using 131I Mathur, P.N., and R.K. Dhadial. 1961. iodine isotope as radioactive tracer. Ener- Studies on the external morphol- gia Nuclear e Agricultura 3:34–43. ogy of Bruchus affinis Frol. (Coleoptera, Menusan, H., Jr. 1934a. Effects of con- Phytophaga, Bruchidae). Part 1, Head stant light, temperature, and humidity on capsule and mouthparts. Entomologisk the rate and total amount of oviposition Tidskrift 82:222–230. of the bean weevil, Bruchus obtectus Say. Mathur, P.N., and R.K. Dhadial. 1963. Journal of Economic Economic Entomol- Morphology of the head capsule and ogy 28:448–453. mouthparts of Caryedon gonagra Fabricius Menusan, H., Jr. 1934b. Effects of tem- (Coleoptera: Bruchidae). Proceedings of perature and humidity on the life pro- the Entomological Society of Washington cesses of the bean weevil, Bruchus obtectus 65:265–273.

297 Say. Annals of the Entomological Society of Messina, F.J., and J.A.A. Renwick. 1985. America 27:515–526. Mechanism of egg recognition by the cow- Menusan, H., Jr. 1936. The influence of pea weevil Callosobruchus maculatus. constant temperatures and humidities on Entomologia Experimentalis et Applicata the rate of growth and relative size of the 37:241–245. bean weevil, Bruchus obtectus Say. Annals Metcalf, C.L., and W.P. Flint. 1939. De- of the Entomological Society of America structive and Useful Insects, Their Habits 29:279–288. and Csontrol. 2nd ed. McGraw-Hill, New Menusan, H., Jr., and G.F. MacLeod. 1938. York. Toxicity of high temperatures to bean wee- Mickey, G.H. 1935. Spermioteleosis of vil eggs. Journal of Economic Entomology Bruchus quadrimaculatus Fabr. Journal of 30:954–958. Morphology 57:147–167. Messina, F.J. 1984. Influence of cowpea Mierzejewska, E. 1982. Mixed infection of pod maturity on the oviposition choices Acanthoscelides obtectus Say with entomo- and larval survival of a bruchid beetle pathogenic fungi. Polish Ecological Studies Callosobruchus maculatus. Entomologia 8:443–447. Experimentalia et Applicata 35:241–248. Miller, S.E., and W.S. Davis. 1986. Insects Messina, F.J. 1985. Ability of ovipositing associated with the flowers of two species seed beetles to discriminate between seeds of Malacothrix (Asteraceae) on San Miguel with differing egg loads. Ecological Ento- Island, California. Psyche 92:547–555. mology 10:225–230. Minney, B.H.P., A.M.R. Gatehouse, P. Messina, F.J. 1987. Genetic contribution Dobie, et al. 1990. Biochemical bases of to the dispersal polymorphism of the cow- seed resistance to Zabrotes subfasciatus pea weevil (Coleoptera: Bruchidae). Annals (bean weevil) in Phaseolus vulgaris (com- of the Entomological Society of America mon bean); a mechanism for arcelin toxic- 80:12–16. ity. Journal of Insect Physiology 36:757– Messina, F.J. 1990. Alternative life-histo- 767. ries in Callosobruchus maculatus. Envi- Mitchell, R. 1975. The evolution of ovipo- ronmental and genetic bases. In K. Fujii, sition tactics in the bean weevil, Calloso- A.M.R. Gatehouse, C.D. Johnson, et al., bruchus maculatus (F.). Ecology 56:696– eds., Bruchids and Legumes: Econom- 702. ics, Ecology, and Coevolution. Proceeding Mitchell, R. 1977. Bruchid beetles and of the 2nd International Symposium on seed packaging by palo verde. Ecology Bruchids and Legumes, Okayama, Japan, 58:644–651. September 6–9, 1989, pp. 303–315. Series Entomologica 46. Kluwer, Dordrecht, The Mitchell, R. 1990. Behavioral ecology of Netherlands. Callosobruchus maculatus. In K. Fujii, A.M.R. Gatehouse, C.D. Johnson, et al., Messina, F.J., J.L. Barmore, and J.A.A. eds., Bruchids and Legumes: Econom- Renwick. 1987a. Oviposition deterrent ics, Ecology, and Coevolution. Proceeding from eggs of Callosobruchus maculatus– of the 2nd International Symposium on spacing mechanism or artifact. Journal of Bruchids and Legumes, Okayama, Japan, Chemical Ecology 13:219–226. September 6–9, 1989, pp. 317–330. Series Messina, F.J., J.L. Barmore, and J.A.A. Entomologica 46. Kluwer, Dordrecht, The Renwick. 1987b. Host selection by ovipos- Netherlands. iting cowpea weevils: Patterning of input Miyamoto, D.M., and J.M. Van Der Meer. from separate sense organs. Entomologia 1982. Early egg contractions and patterned Experimentalis et Applicata 43:169–173. parasynchronous cleavage in a living

298 insect (Callosobruchus maculatus) egg. Fab., a serious pest of stored pulses. In- Wilhelm Roux’s Archives for Developmental dian Journal of Entomology 26:345–351. Biology 191:95–102. Morallo-Rejesus, B. 1990. Callosobru- Mizell, R.F., and T.E. Nebeker. 1982. chus chinensis (Linn.) in grain legumes: Preference and oviposition rates of adult Losses and control in the Philippines. In Thanismus dubius (F.) on three prey spe- T. Yoshida, ed., Loss From and Control of cies. Environmental Entomology 11:139– Bruchids in Developing Countries, Country 143. Report Session. Supplement to Proceeding Mohyuddin, A.I. 1969. The biology and of the Second International Symposium on host spectrum of some stenophagus in- Bruchids and Legumes, Okayama, Japan, sects found on Convolvulus and Calystegia September 6–9, 1990, pp. 54–61. spp. at Belleville, Ontario. Commonwealth Moreno G.I., and F.F. Bibby. 1943. Institute of Biological Control, Technical Ynsectos del algodon y otros Malvaceae en Bulletin No. 12, pp. 131–146. las regiones de Matamoras, Tams., y del Moldenke, A.R. 1971. Host-plant rela- Valle Baja del Rio Bravo. Fitofilo 2:20–110. tions of phytophagous beetles in Mexico Morimoto, K. 1990. A synopsis of the (Coleoptera: Bruchidae, Chrysomelidae, bruchid fauna of Japan. In K. Fujii, A.M.R. Curculionidae). Pan-Pacific Entomologist Gatehouse, C.D. Johnson, et al., eds., 47:105–116. Bruchids and Legumes: Economics, Ecol- Moller, H., R.H. Smith, and R.M. Sibley. ogy, and Coevolution. Proceeding of the 1990. Evolutionary demography of a bru- 2nd International Symposium on Bruchids chid beetle. 3. Correlated responses to and Legumes, Okayama, Japan, Sep- selection and phenotypic plasticity. Func- tember 6–9, 1989, pp. 131–140. Series tional Ecology 4:489–493. Entomologica 46. Kluwer, Dordrecht, The Netherlands. Monga, D. 1972. Spermatheca and associated accessory gland in Callosobruchus Motschulsky, V. 1854. Voyages. Lettre de maculatus (F.). Zoologica Poloniae 22:71– M. de Motschulsky à M. Ménétriés. Études 79. Entomologiques 2:1–15. Monga, D., and M.L. Sareen. 1980. Mor- Motschulsky, V. 1863. Essai d’un cata- phological and histochemical studies on logue des insectes de l’Ile Ceylan. Bulle- the ovary of some of some bruchids (Bru- tin Société Imperial Naturelle de Moscou chidae: Coleoptera). Zoologica Poloniae 36:421–532. 27:483–498. Motschulsky, V. 1874. Enumération des Monge, J-P. 1985. L’importance des con- nouvelles espèces de coléoptères rapportés tacts de la nymph ou de l’imago d’Acantho- de ses voyages. Bulletin Société Imperial scelides obtectus Say avec sa plante-hote Naturelle de Moscou 46:203–252. (Phaseolus vulgaris L.) dans la regulation Muesebeck, C.F.W., Krombein, K.V., and de l’activitié reproductrice des femelles: H.K. Townes. 1951. Hymenoptera of Amer- Quelques donnees preliminaries. Colloqui- ica north of Mexico. Synoptic catalog. U.S. um Recherce Académie des Sciences Paris Department of Agriculture, Agriculture 301:17–20. Monograph 2. Monros, F. 1955. Remarques sur les af- Mukerji, D. 1938. Anatomy of the lar- finities des familles de Cerambycoidea val stages of the bruchid beetle Bruchus (Coleoptera). Institut Royal des Sciences quadrimaculatus Fabr., and the method of Naturelles de Belgique 31:1–7. emergence of the larva from the egg-shell. Mookherjee, P.B., and M.L. Chawla. 1965. Zeitschrift für Angewandte Entomologie Effect of temperature and humidity on the 25:442. development of Callosobruchus maculatus

299 Mukerji, D. 1949. Structure and function predation and differential germination. of bursa copulatrix and the associated Journal of the Kansas Entomological organs in Bruchidae (=Lariidae) (Coleop- Society 56:169–174. tera, Phytophagidae) and their taxonomic Nelson, D.M., and C.D. Johnson. 1983b. significance. Current Sciences, Bangalore Selenium in Astragalus (Leguminosae) and 18:255–257. its effect on host preferences of bruchid Mukerji, D., and M.A.H. Bhuya. 1937. beetles. Journal of the Kansas Entomologi- Reproductive system of the bruchid beetles cal Society 56:267–272. Bruchus quadrimaculatus Fabr., and Bru- Nichols, S.W., and R.T. Schuh. 1989. The chus (Callosobruchus) chinensis L. (Bru- Torre-Bueno Glossary of Entomology. New chidae-Coleoptera). Journal of Morphology York Entomological Society, New York. 61:175–221. Nilsson, J.A., and C.D. Johnson. 1990. A Mukerji, S., and S.N. Chatterjee. 1951. new species of palm bruchid from Cuba Morphology of the genital structure of and a redescription of Caryobruchus some of the Bruchidae (Lariidae) of India gleditsiae (L.) (Coleoptera: Bruchidae: and Ceylon and their taxonomic impor- Pachymerinae). The Coleopterists’ Bulletin tance. Indian Journal of Entomology 13:1– 44:50–59. 28. Nilsson, J.A., and C.D. Johnson. 1993a. A Mukerji, S., M.G.R. Menon, and S.N. Chat- taxonomic revision of the palm bruchids terjee. 1957. The taxonomic position of (Pachymerini) and a description of the Caryedon fuscus (Goeze), C. gonager (Fa- world genera of Pachymerinae (Coleoptera: bricius), and C. languidus (Gyllenhal) (Co- Bruchidae: Pachymerinae). Memoirs of the leoptera: Bruchidae) based on a study of American Entomological Society 41. the genitalia. Proceedings of the Royal En- tomological Society of London 26:103–106. Nilsson, J.A., and C.D. Johnson. 1993b. Laboratory hybridization of Stator beali and Müller, O.F. 1764. Fauna Insectorum S. limbatus, with new host records for S. Fridrichsdalina sive Methodica Descriptio limbatus and Mimosestes amicus (Coleop- Insectorum. Agri Fredrichdalensis. Copen- tera: Bruchidae). The Southwestern Natu- hagen, Denmark, and Leipzig, Germany. ralist 38:385–387. Mulsant, E., and C. Rey. 1858. Etude Cole- Nwanze, K.F., and E. Horber. 1975. How optera du genre Bruchus. Opuscula Ento- seed size affects the occurrence of “active” mologica 8:1–44. and “miniature” forms of Callosobruchus Nakamura, H. 1968. A comparative study maculatus in laboratory populations. Envi- on the ovipositional behavior of two species ronmental Entomology 4:729–732. of Callosobruchus (Coleoptera: Bruchidae). Nwanze, K.F., and E. Horber. 1976. Seed Japanese Journal of Ecology 18:192–197. coat of cowpeas affect oviposition and lar- Neelgund, Y.F., and S.M. Kumari. 1983. val development of Callosobruchus macula- Gut bacterial flora of cowpea weevils. Cur- tus. Environmental Entomology 5:213–218. rent Science 52:140–141. Nwanze, K.F., J.K. Maskarinec, and T.L. Neilson, C.L., and R.H. Handford. 1954. Hopkins. 1976. Lipid composition of the Insects of the season 1953 in the interior normal and flight forms of adult cowpea of British Columbia. Canadian Insect Pest weevils, Callosobruchus maculatus. Journal Review 32:1–11. of Insect Physiology 22:897–899. Nelson, D.M., and C.D. Johnson. 1983a. Ofuya, T.I. 1987. Callosobruchus maculatus Stabilizing selection on seed size in As- (Fabricius) (Coleoptera: Bruchidae) ovipo- tragalus (Leguminosae) due to differential sition behaviour on cowpea seeds. Insect Science and Its Application 8:77–79.

300 Ofuya, T.I. 1989. Effect of larval infesta- maculatus Fab. (Coleoptera, Bruchidae) in tion on the choice of seed for oviposition by cowpea seeds. Journal of Stored Products Callosobruchus maculatus (F.) (Coleoptera: Research 18:1–8. Bruchidae). Insect Science and Its Applica- Ott, J.R. 1991. The biology of Acantho- tion 10:437–440. scelides alboscutellatus (Coleoptera: Ofuya, T.I., and S.O. Agele. 1989. An ex- Bruchidae) on its host plant, Ludwigia amination of the patrol phase of oviposition alternifolia (L.) (Onagraceae). Proceedings behaviour of Callosobruchus maculatus of the Entomological Society of Washington (Fabricius) (Coleoptera: Bruchidae). Jour- 93:641–651. nal of Stored Products Research 25:101– Ott, J.R., and M. Lampo. 1991. Body size 104. selection in Acanthoscelides alboscut- Ogijewicz, B. 1948. The nervous-sensory ellatus. I. Entrapment within the fruit system and the sensual organs of the feet of Ludwigia alternifolia. Oecologia 87(4): of Coleoptera. Studia Societatis Scien- 522–527. tarum Torunensis, Section E (Zoology) Paddock, F.B., and H.J. Reinhard. 1919. 1:121–165. The cowpea weevil. Texas Agricultural Ex- Okamoto, K. 1971. The synchronization periment Station, Bulletin No. 256. of the life cycles between Callosobruchus Pajni, H.R. 1959. Anatomy of Calloso-bru- chinensis and its parasite Anisopteromalus chus analis Fabr. (Bruchidae: Coleoptera). calandrae. Japanese Journal of Ecology Research Bulletin (Science) of the Panjab 20:233–237. University 10:21–24. Olivier, A.G. 1790. Encyclopédie méthod- Pajni, H.R. 1965. Metamorphosis of the ique. Histoire Naturelle. Insectes. Vol. 5, salivary glands in Callosobruchus macula- part 1. Pankouke, Paris. tus (F.) (Coleoptera: Bruchidae). Research Olivier, A.G. 1795. Entomologie, ou his- Bulletin (Science) of the Panjab University toire naturelle des insectes. Avec leurs 16:265–266. caractères génériques et spécifiques, leur Pajni, H.R. 1968a. The development of description, leur synonymie, et leur figure male genital ducts and the associated enluminée ... Coléoptères 4, 79. Lanneau, structures in Callosobruchus maculatus Paris. [Genera separately numbered.] (Bruchidae: Coleoptera). Research Bulletin Olsen, C.E. 1918. [Exhibition of speci- (Science) of the Panjab University 19:81– mens]. Journal of the New York Entomo- 88. logical Society 26:234. Pajni, H.R. 1968b. Development of the Osborn, H.T. 1949. Insect pest survey. female genital ducts and the associated California Department of Agriculture, 30th structures in Callosobruchus maculatus Annual Report. Vol. 38, pp. 159–166. Sac- (F.). Research Bulletin (Science) of the Pan- ramento. jab University 19:341–348. Osborn, T.C., D.C. Alexander, S.S.M. Sun, Pajni, H.R. 1968c. The larval and imaginal et al. 1988. Insecticidal activity and lectin oenocytes of Callosobruchus maculatus. homology of arcelin seed protein. Science Beiträge zur Entomologie 18:233–238. 240:207–210. Pajni, H.R. 1969. Structure and metamor- Oshima, K., H. Honda, and I. Yamamoto. phosis of the tracheal system of Calloso- 1973. Isolation of an oviposition marker bruchus maculatus (F.) (Coleoptera: Bru- from azuki bean weevil, Callosobruchus chidae). Research Bulletin (Science) of the chinensis (L.). Agricultural and Biological Panjab University 20:113–118. Chemistry 37:2679–2680. Pajni, H.R. 1981. Trophic relations and Osuji, F.N.C. 1980. Radiographic studies of ecological status of the adults of Bruchus the development of Callosobruchus

301 pisorum L. and allied field species of Bru- Perez, G., and A. Bonet. 1984. Parasitoid chidae (Coleoptera). Series Entomol-ogica Hymenoptera de Acanthoscelides obtec- 19:125–129. tus (Coleoptera: Bruchidae) en Tepoztlan, Pajni, H.R., and A. Jabbal. 1986. Some ob- Morelos. Folia Entomológica Mexicana servations on the biology of Zabrotes sub- 59:71–78. fasciatus (Boh.) (Bruchidae: Coleop-tera). Perttunen, V. 1972. Humidity and light Research Bulletin (Science) of the Panjab reactions of Sitophilus granarius L., S. University 37:11–16. oryzae L. (Col., Curculionidae), Rhizopertha Pajni, H.R., and S. Sood. 1975. Effect dominica F. (Bostrychidae), and Acan-thos- of peapollen feeding on maturation and celides obtectus Say (Bruchidae). Annales copulation in the beetle, Bruchus pisorum Entomologici Fennici 38:161–176. L. Indian Journal of Experimental Biology Perttunen, V., and T. Häyrinen. 1969. The 13:202–203. effect of temperature and light intensity on Parella, M., and L.T. Kok. 1975. Bindweeds flight initiation and take-off rate in Acan- and their potential for biological control. thoscelides obtectus Say (Col., Bruchidae). Virginia Journal of Science 26:44. Annales Entomologici Fennici 35:190–204. Parnell, J.R. 1964. The external morphol- Perttunen, V., and T. Häyrinen. 1970a. ogy of the larvae and notes on the pupae of Effect of age on take-off activity in Acan- Bruchidius ater (Marsh.) (Col., Bruchidae) tho-scelides obtectus Say (Col., Bruchidae). and Apion fuscirostre F. (Col., Curculi-oni- Annales Entomologici Fennici 36:35–39. dae). Entomologists’ Monthly Magazine Perttunen, V., and T. Häyrinen. 1970b. 100:83–87. Individual variation in the take-off activ- Patton, W.H. 1895. Systematic value of ity of Acanthoscelides obtectus Say (Col., the larva of Spermophagus. The Canadian Bruchidae). Annales Entomologici Fennici Entomologist 27:290. 36:107–110. Pawar, B.L., and K.K. Verma. 1977. Re- Peterson, A. 1951. Larvae of Insects. Part tournement of the aedeagus in Bruchidae 2. Coleoptera, Diptera, Neuroptera, Si- (Coleoptera, Phytophaga). Entomon 2:171– phonaptera, Mecoptera, Trichoptera. Ed- 174. wards Brothers, Columbus, OH. Paykull, G. 1800. Fauna Suecica. Insecta. Pfaffenberger, G.S. 1977. Comparative Vol. 3. Edman, Upsala, Sweden. descriptions of the final larval instar of Bruchus brachialis, B. rufimanus, and B. pi- Payne, T.H. 1913. The acacia weevil in sorum (Coleoptera: Bruchidae). The Coleop- southern California. Bulletin of the South- terists’ Bulletin 31:133–142. ern California Academy of Sciences 12:40– 41. Pfaffenberger, G.S. 1979. Comparative description and bionomics of the first and -fi Peck, O. 1963. A catalogue of the Nearctic nal larval stages of Amblycerus acapul-cen- Chalcidoidea (Insecta; Hymenoptera). The sis Kingsolver and A. robiniae (Fabricius) Canadian Entomologist, Supplement 30. (Coleoptera: Bruchidae). The Coleopterists’ Perez, G. 1982. Himenopteros parasitoides Bulletin 33:229–238. asociados a Acanthoscelides obtectus Pfaffenberger, G.S. 1980. Description, (Say) (Coleoptera: Bruchidae) en Tepozt- bionomics and phylogenetic discussion of lan, Morelos. Thesis for Biologist Degree. the first instar larva ofMegacerus cubicus Escuela Nacional de Estudios Profesion- (Motsch.) (Coleoptera: Bruchidae). Jour- ales, Departamento de Biologia, Iztacala, nal of the Kansas Entomological Society Mexico. 53:350–356.

302 Pfaffenberger, G.S. 1981. A comparative (Coleoptera). U.S. Department of Agricul- description and phenetic analysis of the ture, Technical Bulletin No. 1525. first instar larva of sevenStator species Pfaffenberger, G.S., S. Muruaga de (Coleoptera: Bruchidae). The Coleopterists’ L’Argentier, and A. Teran. 1984. Morpho- Bulletin 35:255–268. logical descriptions and biological discus- Pfaffenberger, G.S. 1984. Morphological sions of the first and final larval instars descriptions and biological and phylo- of four species of Megacerus (Coleoptera: genetic discussions of the first and final Bruchidae). The Coleopterists’ Bulletin instars of four species of Megacerus larvae 38:1–26. (Coleoptera: Bruchidae). The Coleopterists’ Philippi, R.A., and F.H.E. Philippi. 1864. Bulletin 38:1–26. Beschreibung einiger neues Chilenischen Pfaffenberger, G.S. 1985a. Checklist of Käfer. Stettiner Entomologische Zeitung selected world species of described first 25:313–406. and/or final larval instars (Coleoptera: Pic, M. 1902. Coléoptères presumes nou- Bruchidae). The Coleopterists’ Bulletin veaux de la Rhodesia. Revista de Entomo- 39:1–6. logia 21:4–7. Pfaffenberger, G.S. 1985b. Description, Pic, M. 1912. Renseignements généraux differentiation, and biology of the four sur les Bruchidae. Échange 336:91–93. larval instars of Acanthoscelides obtectus (Say) (Coleoptera: Bruchidae). The Coleop- Pic, M. 1913a. Bruchidae. Coleopterorum terists’ Bulletin 39:239–256. Catalogus part 55, pp. 1–74. W. Junk, Berlin. Pfaffenberger, G.S. 1985c. New host record for Acanthoscelides chiricahuae (Fall) Pic, M. 1913b. Coléoptères exotiques in (Coleoptera: Bruchidae). The Coleopterists’ partie nouveaux. Échange 29:106–110. Bulletin 39:256. Pic, M. 1913c. Espèces et variétiés nou- Pfaffenberger, G.S. 1986. Morphology and velles appartenant a diverses families. biology of larval Gibbobruchus mimus (Say) Mélanges Exotico-Entomologiques 6:8–16. (Coleoptera: Bruchidae). The Coleopterists’ Pic, M. 1922. Nouveautés diverses. Bulletin 40:49–61. Mélanges Exotico-Entomologiques 35:1–32. Pfaffenberger, G.S. 1990. A scanning Pic, M. 1924. Nouveautés diverses. electron microscopic view of the final larval Mélanges Exotico Entomologiques 42:1–32. instar of Zabrotes subfasciatus (Coleoptera: Pic, M. 1928. Nouveaux Bruchidae. Bruchidae: Amblycerinae). The Coleopter- Annals and Magazine of Natural History ists’ Bulletin 44:37–49. 11: 297–299. Pfaffenberger, G.S. 1991. Bruchidae Pic, M. 1929. Nouveautés diverses. (Chrysomeloidea). In F.W. Stehr, ed., Im- Mélanges Exotico-Entomologiques 54:1–36. mature Insects, vol. 2 , pp. 561–568. Kendall/Hunt, Dubuque, IA. Pic, M. 1930a. Nouveaux coléoptères exotiques. Bulletin de la Societe Linneenne de Pfaffenberger, G.S., and D.H. Janzen. Lyon 9:36–37. 1984. Life history and morphology of first and last larval instars of Costa Ri- Pic, M. 1930b. Notes, diverses, nouveautés. can Caryedes brasiliensis Thunberg (Co- Échange 46:13–14. leoptera: Bruchidae). The Coleopterists’ Pic, M. 1930c. Nouveautés diverse. Bulletin 38:267–281. Mélanges Exotico-Entomologiques 55:1–36. Pfaffenberger, G.S., and C.D. Johnson. Pic, M. 1932. Nouveautés diverses. 1976. Biosystematics of the first-stage Mélanges Exotico-Entomologiques 59:10– larvae of some North American Bruchidae 36.

303 Pic, M. 1933. Nouveautés diverses. species. Entomologia Experimentalis et Mélanges Exotico-Entomologiques 61:3–36. Applicata 23:152–162. Pic, M. 1935. Neue Bruchidae. Entomolo- Pimbert, M. 1985c. A model of host plant gischer Anzeiger 15:65–66. change of Zabrotes subfasciatus Boh. (Coleoptera: Bruchidae) in a traditional Pic, M. 1938. Nouveautés diverses, muta- bean cropping system in Costa Rica. Bio- tions. Mélanges Exotico-Entomologiques logical Agriculture and Horticulture 3:39– 70:22–25. 54. Pic, M. 1942. Opuscula martialis. Échange Pimbert, M.P., and D. Pierre. 1983. 7:10. Ecophysical aspects of reproduction. I. Pic, M. 1943. Note rectificative (Col. Bruch- The influence of pod maturity and seeds idae). Revue Française d’Entomologie of Phaseolus vulgaris and the influence of 9:148. insemination on the reproductive activity Pic, M. 1954. Bruchidae (Coleoptera). Be- of Zabrotes subfasciatus. Ecological Ento- iträge Fauna Perus 4:184–186. mology 8:87–94. Pichard, B., B. Leroi, and A. Bonet. 1991. Pinckney, J.S. 1937. The vetch bruchid, Comparación des cycles d’Acanthoscelides Bruchus brachialis Fahraeus. Journal of obtectus et d’A. obvelatus (Coleopteres, Economic Entomology 30:621–632. Bruchidae) a Tepoztlan (Mexique). Acta Podoler, H., and S.W. Applebaum. 1971. Oecologica 12:185–201. Basic nutritional requirements of larvae of Pierce, W.D. 1908. A list of parasites the bruchid beetle, Callosobruchus chinen- known to attack American Rhynchophora. sis L. Journal of Stored Products Research Journal of Economic Entomology 1:380– 7:187–193. 396. Pope, R.D. 1956. The family name Pierce, W.D. 1912. Systematic notes and Bruchidae. Entomologists’ Monthly Maga- descriptions of some weevils of economic or zine 92:45–46. biological importance. Proceedings of the Pouzat, J. 1970. Role des organes sen- United States National Museum 42:155– so-riels cephaliques dans l’ovogenese 170. et l’emission chez la bruche du haricot Pierre, D. 1980. Influence of seeds or ripe Acanthoscelides obtectus Say. Colloques pods of Phaseolus vulgaris on the repro- Internationaux du Centre National du la ductive activity of Zabrotes subfasciatus Recherche Scientifique 189:381–400. (Coleoptera: Bruchidae). Compte Rendus Pouzat, J. 1975. Analyse experimentale Hebdomadaire des Seances de l’Academie du role de l’ovitube dans le comportement des Sciences Paris 290:1007–1010. de ponte de la bruche du haricot (Acan- Pierre, D., and M. Pimbert. 1981. Some tho-scelides obtectus Say/--/Coleoptera, data on the reproductive activity of Bruchidae). Behaviour 54:258–277. Zabrotes subfasciatus in the laboratory. Pouzat, J. 1976. Analyse experimentale Series Entomologica 19:113–123. du role de l’ovitube dans le comportement Pimbert, M. 1985a. Comparaison du de ponte de la bruche du haricot (Acan- comportement de ponte de Zabrotes thoscelides obtectus Say). Behaviour 54: subfasciatus Boh. (Col. Bruchidae) en 258–277. presence de gousses ou de graines de Pouzat, J. 1977. Effect of stimulations Phaseolus vulgaris L. Biology of Behaviour coming from the host-plant, the bean 10:309–319. (Phaseolus vulgaris L.) on the egg-laying Pimbert, M. 1985b. Reproduction and behaviour of the bean weevil (Acantho- oviposition preferences of Zabrotes scelides obtectus Say). Colloques Inter- subfasciatus stocks reared from two host

304 nationaux du Centre National du la Re- of the Entomological Society of Nigeria cherche Scientifique 265:115–131. 1:3–6. Pouzat, J. 1978. Host plant chemosensory Prevett, P.F. 1971. The larvae of some influence on oogenesis in the bean wee- Nigerian Bruchidae (Coleoptera). Transac- vil, Acanthoscelides obtectus. Entomologia tions of the Royal Entomological Society of Experimentalis et Applicata 24:601–608. London 123:247–312. Pouzat, J. 1981. The role of sense organs Puri, G., and S.P. Sharma. 1984. Changes in the relations between bruchids and their in trehalose levels with age in the bruchid, host plants. Series Entomologica 19:61–72. Callosobruchus maculatus (Fabr.). Insect Pouzat, J. 1982. The effect of antennal am- Science and Its Application 5:103–105. putation on oogenesis in the bean weevil Qi, Y., and W.E. Burkholder. 1985. Study (Acanthoscelides obtectus). Ento-mologia on sex pheromone biology of the azuki Experimentalis et Applicata 31:333–338. bean weevil, Callosobruchus chinensis (L.). Pouzat, J. 1983. La repartition des oeufs Contributions from the Shanghai Institute pondus par Acanthoscelides obtectus entre of Entomology 5:1–11. des graines identiques de Phaseolus vulgar- Rahman, M.F., and M.-U. Ameen. 1986. is. Biology of Behaviour 3:215–230. Metamorphic changes of anatomy of the Pouzat, J., H. Bilal, D. Nammour, and larval alimentary canal of Callosobruchus M. Pimbert. 1989. A comparative study analis and Callosobruchus chinensis (Bru- of the host plant’s influence on the sex chidae, Coleoptera). Bangladesh Journal of pheromone dynamics of three bruchid spe- Zoology 14:157–166. cies. Acta Oecologia Oecologica Generalis Rahman, M.F., and M.-U. Ameen. 1990. 10:401–410. Metamorphic changes of the malpighian Prevett, P.F. 1965. The genus Caryedon in tubules of Callosobruchus analis Fab. and northern Nigeria, with descriptions of six C. chinensis (L.) (Bruchidae: Coleoptera). new species (Col. Bruchidae). Annales de Bangladesh Journal of Zoology 18:79–90. la Société Entomologique de France (N.S.) Raina, A.K. 1970. Callosobruchus spp. 1:523–547. infesting stored pulses (grain legumes) in Prevett, P.F. 1966a. The identity of the India and a comparative study of their biol- palm kernel borer in Nigeria, with system- ogy. Indian Journal of Entomology 32:303– atic notes on the genus Pachymerus Thun- 310. berg (Coleoptera: Bruchidae). Bulletin of Ramos, R.Y. 1978. Contribucion al conoc- Entomological Research 57:181–192. imiento de los bruquidos (Col. Bruchidae) Prevett, P.F. 1966b. A new genus and spe- del Mediterraneo occidental: I.—Notas cies of Pachymerinae (Coleoptera: Bruchi- taxonomicas sobre el genero Bruchus L. dae) from South America. Proceedings of Nouvelle Revue d’Entomologie 8:315–320. the Royal Entomological Society of London Randolph, N.M., and B.B. Gillespie. 1958. 35:81–83. Notes on the biology of Bruchus brachialis Prevett, P.F. 1967a. The larva of Caryedon Fahr. Journal of Economic Entomology serratus (Ol.): The groundnut seed beetle 51:401–402. (Coleoptera: Bruchidae). Journal of Stored Rathvon, A. 1870. A new bean weevil. Products Research 3:117–123. American Entomologist 2:118–119. Prevett, P.F. 1967b. Notes on the biology, Reid, C. 1996. More on the family Bruchi- food plants, and distribution of Nigerian dae. Chrysomela No. 31:3. Bruchidae (Coleoptera), with particular reference to the northern region. Bulletin

305 Reitter, E. 1912. Fauna Germanica. Die Bruchidae) in a storage grain plague. Käfer des Deutschen Reiches. IV Band. Ciencias Agricultura 23:16–20. Lutz’ Verlag, Stuttgart, Germany. Rogers, C.E., and J.C. Garrison. 1975. Rey, C. 1893. Remarques en passant: Seed destruction in indigobush Amorpha Famille des Bruchides. Échange 9:3,25– by a seed beetle. Journal of Range Manage- 26,37. ment 28:241–242. Ribeiro, C.S. 1989. Estudo fenético de Romero N., J., and C.D. Johnson. 1997. algunos espécies do genero Amblycerus New synonymy of Zabrotes chavesi King- Thunberg, 1815 (Coleoptera: Bruchidae). solver and Zabrotes vandykei Kingsolver, M.S. thesis. Departmento de Zoologia, with new host and distribution records Universidade Federal do Parana, Curitiba, (Coleoptera: Bruchidae: Amblycerinae). The Brazil. Coleopterists’ Bulletin 51:74. Rico-Arce, M. de L. 1991. New species, Romero, J., and C.D. Johnson. 1999. combinations and synonyms for Zygia, Zabrotes sylvestris, a new species from the Cojoba, Marmaroxylon and Pithecellobium United States and Mexico related to Z. sub- (Leguminosae, Mimosoideae, Ingae). Kew fasciatus (Boheman) (Bruchidae: Coleop- Bulletin 46(3):493–521. tera: Amblycerinae). Coleopterists Bulletin Riley, C.V. 1871. Third Annual Report of 53:87–98. the Noxious, Beneficial and Other Insects, Romero, J., C.D. Johnson, and J.M. King- of the State of Missouri. Wilcox, Jefferson solver. 1996. Revision of the genus Amblyc- City, MO. erus of the United States and Mexico (Cole- Riley, C.V. 1892. The first or post-embry- optera: Bruchidae; Amblycer-inae). United onic stage of the pea and bean weevils. The States Department of Agriculture Technical Canadian Entomologist 24:185–186. Bulletin No. 1845. Riley, C.V. 1893. Report of the entomolo- Roomi, M.W., Z.I. Khan, and S.A. Khan. gist. In Report of the Secretary of Agricul- 1973. Pteromalus schwenkei (Hymenop- ture, United States Department of Agricul- tera, Pteromalidae) a new species as a pri- ture (1892). Washington, DC. mary parasite of the bean weevil, Bruchus chinensis L., from Pakistan. Zeitschrift fur Riley, C.V., and L.O. Howard. 1892a. On Angewandte Entomologie 72:394–400. the nomenclature and oviposition of the bean weevil. Insect Life 4:27–33. Rosenfeld, A.H. 1911. Insects in spanish moss. Journal of Economic Entomology Riley C.V., and L.O. Howard. 1892b. The 4:398–409. pea and bean weevils. Insect Life 4:297– 302. Rosenhauer, W.G. 1856. Die Thiere Anda- lusiens nach den Beschreibungen von 249 Riley, C.V., and L.O. Howard. 1892c. neuen oder bis jetzt noch unbe-schrieben Food-plants of North American species of Gattunen und Arten. Erlangen, Blaesing, Bruchus. Insect Life 5:165–166. Germany. Robert, P. 1985. A comparative study of Rup, P.J. 1986. Mating and its attendant some aspects of the reproduction of three behavior in Callosobruchus maculatus Caryedon serratus strains in presence of (Coleoptera: Bruchidae). Journal of Stored its potential host plants. Oecologia 65:425– Products Research 22:77–80. 430. Rup, P.J. 1988. Antenna and antennal Roche, R., M. Gonzalez Valenzuela, sensilla dimorphism in Callosobruchus and M.E. Simanca. 1985. Life cycle of maculatus (F.) (Coleoptera: Bruchidae). Callosobruchus maculatus (Coleoptera:

306 Journal of Stored Products Research Satija, R.C., and R.P. Kaur. 1967a. Brain 24:83–86. during post-embryonic life of Calloso-bru- Rup, P.J., and S.P. Sharma. 1978. Behav- chus maculatus. Research Bulletin (Sci- ioural response of males and females of ence) of the Panjab University 18:475–489. Callosobruchus maculatus (F.) to the sex Satija, R.C., and R.P. Kaur. 1967b. Growth pheromones. Indian Journal of Ecology of the body and the brain during post-em- 5:72–76. bryonic life in Callosobruchus macu-latus. Sakai, A., H. Honda, K. Oshima, and I. Research Bulletin (Science) of the Panjab Yamamoto. 1986. Oviposition marking University 18:357–361. pheromone of two bean weevils, Calloso- Satija, R.C., and R.P. Kaur. 1968. Visual bruchus chinensis and Callosobruchus centres during postembryonic life of Callo- maculatus. Journal of Pesticide Science sobruchus maculatus (F.). Research Bul- 11:163–168. letin (Science) of the Panjab University Samuelson, G.A. 1991. [Bruchid in koa 19:101–102. haole pods—Acanthoscelides macro-phthal- Satija, R.C., K. Sumal, and Sumeet. 1975. mus.] Hawaiian Entomological Society Morphogenetic studies on the brain of Cal- Newsletter 1:2. losobruchus analis (Coleoptera). Research Sandhu, R., and S. Neena. 1982. Obser- Bulletin (Science) of the Panjab University vation of the brain and optic peduncle of 26:61–68. the adult Zabrotes subfasciatus. Research Sawaf, S.K. el- 1956. Some factors affect- Bulletin (Science) of the Panjab University ing the longevity, oviposition, and rate of 33:55–66. development in the southern cowpea wee- Sandner, H., and M. Pankanin. 1974. Ef- vil, Callosobruchus maculatus F. Bulletin fect of the presence of food on egg laying de la Société Entomologique de Egypte by Acanthoscelides obtectus (Coleoptera: 40:29–95. Bruchidae). Polskie Pismo Entomologiczne Say, T. 1824. Descriptions of coleopterous 43:811–817. insects collected in the late expedition to Sano, I. 1967. Density effect and environ- the Rocky Mountains, performed by order mental temperature as the factors produc- of Mr. Calhoun, Secretary of War, under ing the active form of Callosobruchus macu- the command of Major Long. Journal of the latus (F.) (Coleoptera: Bruchidae). Journal Academy of Natural Sciences of Philadel- of Stored Products Research 2:187–195. phia 3:298–331. Sano-Fujii, I. 1984. Effect of bean wa- Say, T. 1831. Descriptions of North Ameri- ter content on the production of the ac- can Curculionides and an arrangement of tive form of Callosobruchus maculatus (F.) some of our known species agreeably to (Coleoptera: Bruchidae). Journal of Stored the method of Schoenherr. [publisher un- Products Research 20:153–161. known], New Harmony, IN. Sano-Fujii, I. 1986. The genetic basis of Schaeffer, C.F.A. 1904. New genera and the production of the active form of Cal- species of Coleoptera. Journal of the New losobruchus maculatus (F.) (Coleoptera: York Entomological Society 12:197–236. Bruchidae). Journal of Stored Products Schaeffer, C.F.A. 1907. New Bruchidae Research 22:115–123. with notes on known species and list of Saplina, G.S. 1980. Callosobruchus species known to occur at Brownsville, maculatus. Zaschita Rastenii 7:41. Texas, and in the Huachuca Mountains, Arizona. Science Bulletin, Museum of the

307 Brooklyn Institute of Arts and Sciences Latin America. In D. White, ed., Proceed- 1:291–306. ings of the 15th International Congress of Schaeffer, C.F.A. 1909. New Coleoptera Entomology, August 19–27, 1976, Wash- chiefly from Arizona. Science Bulletin, Mu- ington, DC, pp. 691–698. Entomological seum of the Brooklyn Institute of Arts and Society of America, College Park, MD. Sciences 1:375–386. Schwarz, E.A. 1878. The Coleoptera of Schilsky, J. 1905. Bruchidae. In H.C. Florida. Proceedings of the American Philo- Küster and G. Kraatz, Die Käfer Europa’s, sophical Society 17:353–372. pp. 41a–f, 41A–MM; Nos. 1–100. Bauer Scopoli, J.A. 1763. Entomologia Carniolica und Raspe, Nürnberg, Germany. exhibens insecta Carnioliae indegena et Schlising, R.A. 1980. Seed destruction of distributa in ordines, genera, species, California morning glories (Convolvulaceae: varietates, methodo Linneana. Trattner, Calystegia) by bruchid beetles. Madrona Vienna, Austria. 27:1–16. Scudder, S.H. 1876. Fossil Coleoptera from Schmitt, M. 1989. On the phylogenetic the Rocky Mountain Tertiaries. Bulletin of position of the bruchidae within the Chrys- the Geological and Geographical Survey of omeloidea (Coleoptera). Entomo-graphy the Territories 2:77–87. 6:531–537. Scullen, H.A., and J.L. Wold. 1969. Biol- Schmitt, M., U. Mischke, and E. Wach- ogy of wasps of the tribe Cercerini, with a mann. 1982. Phylogenetic and functional list of the Coleoptera used as prey. Annals implications of the rhabdom patterns in of the Entomological Society of America the eyes of Chrysomeloidea (Coleoptera). 62:209–214. Zoologica Scripta 11:31–44. Seeliger, R. 1943. Genetische Unter-su- Schoenherr, C.J. 1823. Curculionides. Isis chungen an dem Flügelmuster des Boh- Oken, Zweiter Band. Heft 10. Coleoptera nenkäfers Zabrotes subfasciatus Boh. 1132–1146. Zeitschrift für Induktive Abstammung- sund Vererbungslehre 81:196–251. Schoenherr, C.J., ed. 1833. Genera et species curculionidum, cum synonymia hu- Seurat, L.G. 1900. Sur la morphologie de jus familiae: species novae aut hactenus l’appareil respiratoire de la larve et de la minus cognitae, descriptionibus a Dom. nymph du Bruchus ornatus Boh. Compte Leonardo Gyllenhal, C.H. Boheman, et Rendus Hebdomadaire des Seances de entomologis aliis. Vol. 1, part 1[?]. Roret, l’Academie des Sciences, Paris 131:620– Paris. 623. Schoenherr, C.J., ed. 1839. Genera et spe- Shade, R.E., R.C. Pratt, and M.A. Pome- cies curculionidum, cum synonymia hu- roy. 1987. Development and mortality of jus familiae: species novae aut hactenus the bean weevil, Acanthoscelides obtectus minus cognitae, descriptionibus a Dom. (Coleoptera: Bruchidae), on mature seeds Leonardo Gyllenhal, C.H. Boheman, et of tepary beans, Phaseolus acutifolius, and entomologis aliis. Vol. 5, part 1[?]. Roret, common beans, Phaseolus vulgaris. Envi- Paris. ronmental Entomology 16:1067–1070. Schoof, H.F. 1941. The effects of various Sharma, G., and S.P. Sharma. 1981. Age- relative humidities on the life process of dependent changes in esterases of Cal- the southern cowpea weevil, Calloso-bru- losobruchus maculatus Fab. (Coleoptera: chus maculatus Fab. Ecology 22:297–306. Bruchidae). Experimental Aging Research 7:107–115. Schoonhoven, A.V. 1977. Pests of stored beans and their economic importance in Sharma, S., and P. Sharma. 1979. Age- related protein changes in bruchids

308 Zabrotes subfasciatus and Callosobruchus Shukla, G.S., and S.K. Pandey. 1977. maculatus. Indian Journal of Experimental Sexual dimorphism in Callosobruchus chi- Biology 17:1197–1200. nensis (L.) (Coleoptera: Bruchidae). Ento- Sharma, S.P., I. Jit, and G. Sharma. 1983. mological News 88:265–266. Age-related changes in Callosobruchus Sidhu, D.S., T. Singh, N. Kumar, and S.P. maculatus (Coleoptera) and Zaprionus Kaur. 1984. Lipid composition of the fertile paravittiger (Diptera). Acta Entomologica and sterile strains of Callosobruchus macu- Bohemoslavaca 80:336–340. latus (F.) (Coleoptera: Bruchidae). Journal Sharma, S.P., and N. Rai. 1984. Amino- of Entomological Research 8:98–100. acid variations during aging of Calloso- Singh, K.N., V.P. Varma, and P.K. Srivas- bruchus maculatus (Coleoptera). Zoologica tava. 1980. Change in the development Orient 1:26–29. behaviour of pulse beetles Callosobruchus Sharp, D. 1885. Bruchidae. Biologia Cen- maculatus (Fabricius) and Callosobruchus trali-Americana. Coleoptera 5:437–504. chinensis (Linnaeus). Indian Journal of Entomology 42:304–306. Sharp, D. 1886. On the Bruchidae of Ja- pan. Annals and Magazine of Natural His- Singh, T. 1973. A comparative study of the tory (Ser. 5) 17:34–38. female reproductive organs in Bruchidae (Coleoptera) with a consideration of their Sharp, D., and F.A.G. Muir. 1912. The bearing on classification. Bulletin of Ento- comparative anatomy of the male genital mology 14:66–75. tube in Coleoptera. Transactions of the Entomological Society of London 1912(part Singh, T. 1978a. Comparative morphologi- III):477–642. cal studies on the male reproductive system of Bruchidius (Bruchidae: Coleoptera). Shetler, S.G., and L.E. Skog. 1978. A pro- Entomon 3:69–75. visional checklist of species for flora North America (revised). Monographs in System- Singh, T. 1978b. The male accessory atic Botany. Missouri Botanical Gardens, glands in Bruchidae (Coleoptera) and their St. Louis. taxonomic significance. Entomologica Scandinavica 9:198–203. Shinoda, K., and T. Yoshida. 1984. Rela- tionship between adult feeding and emi- Singh, T. 1979. A key to the north-west gration from beans of azuki bean weevil, Indian Bruchidae (Col.). Entomologist’s Callosobruchus chinensis Linne (Coleop- Monthly Magazine 113:219–231. tera: Bruchidae). Applied Entomology and Singh, T. 1981a. A taxonomic study of the Zoology 18:202–211. wing of Bruchidae (Coleoptera). Oriental Shinoda, K., and T. Yoshida. 1987a. Effect Insects 15:221–225. of fungal feeding on longevity and fecundity Singh, T. 1981b. Field observations on the of the azuki bean weevil, Calloso-bruchus life-history of Callosobruchus chinensis (L.) chinensis (L.) (Coleoptera: Bruch-idae), in (Bruchidae: Coleoptera), a pest of stored the azuki bean field. Applied Entomology grain. Journal of Environmental Research and Zoology 22:465–473. 11:29–31. Shinoda, K., and T. Yoshida. 1987b. Field Singh, T. 1981c. Morphology and muscula- biology of the azuki bean weevil, Calloso- ture of the head capsule and mouth parts bruchus chinensis (L.). Japanese Journal of of Callosobruchus chinensis (L.) (Col.–Bru- Applied Entomology and Zoology 29:14–20. chidae). The Indian Zoologist 5:97–102. Shomar, N.F.H. 1963. A monographic revi- Singh, T. 1982. Comparative studies on the sion of the Bruchidae of Egypt (U.A.R.). male reproductive system of seven species Bulletin of the Entomological Society of Egypt 47:141–196.

309 of Caryedon (Bruchidae: Coleoptera). Ento- Snodgrass, R.E. 1909. The thorax of in- mologists’ Monthly Magazine 117:185–190. sects and the articulation of the wings. Singh, T. 1983. Comparative morphology of Proceedings of the United States National the head capsule of adult Bruchidae (Co- Museum 36:511–595. leoptera) and its taxonomic significance. Snodgrass, R.E. 1935. Principles of Insect Research Bulletin (Science) of the Panjab Morphology. McGraw-Hill, New York. University 33:93–99. Southgate, B.J. 1958. Systematic notes Singh, T., I. Kaur, and M.S. Saini. 1979. on species of Callosobruchus of economic Biology of Zabrotes subfasciatus (Boh.) importance. Bulletin of Entomological (Bruchidae: Coleoptera), pest of sieva Research 49:591–599. beans. Entomon 4:201–203. Southgate, B.J. 1963. The true identity of Singh, T., H.S. Rose, and S. Kaur. 1984. the broom bruchid and synoptic notes on The effect of amputation of antennae on other species of Bruchidae. Annals of the the oviposition of Callosobruchus macula- Entomological Society of America 56:795– tus (Fab.) (Coleoptera: Bruchidae). Bulletin 798. of Pure and Applied Science 3:50–57. Southgate, B.J. 1964. Distribution and Singh, T., and J.S. Yadav. 1979. Charac- hosts of certain Bruchidae in Africa. Tropi- terization of Caryedon Schoenherr (Bruchi- cal Stored Products Information Circular dae: Pachymerinae) with comments on the No. 7, pp. 277–279. status of Pachymerinae. Bulletin of Ento- Southgate, B.J. 1975. Bruchidae. In A. mology 20:169–172. Aitken, ed., Insect travellers, vol. 1, pp. Skaife, S.H. 1925. On variation and hered- 16–19. Ministry of Agriculture, Fisheries ity in the Bruchidae. Transactions of the and Food, Pest Infestation Control Labora- Royal Society of South Africa 12:221–242. tory, Technical Bulletin No. 31, London. Skaife, S.H. 1926. The bionomics of the Southgate, B.J. 1978. The importance of Bruchidae. South African Journal of Sci- the Bruchidae as pests of grain legumes, ence 23:575–588. their distribution and control. In S.R. Slingerland, M.U. 1892. The bean weevil. Singh, H.F. van Emden, and T.A. Taylor, Insect Life 5:86–87. eds., Pests of Grain Legumes: Ecology and Control, pp. 219–229. Academic Press, Slobodchikoff, C.N., and C.D. Johnson. London. 1973. A phenetic and a phylogenetic approach to the classification of a genus of Southgate, B.J. 1979. Biology of the seed beetles (Coleoptera: Bruchidae). Sys- Bruchidae. Annual Review of Entomology tematic Zoology 22:280–294. 24:449–473. Smith, S.G., and J.H. Brower. 1974. Chro- Southgate, B.J. 1981. Univoltine and mosome numbers of stored-product Cole- multivoltine cycles. Series Entomologica optera. Journal of the Kansas Entomologi- 19:17–22. cal Society 47:317–328. Southgate, B.J. 1983. Handbook on Seed Smith, L.L., and D.N. Ueckert. 1974. In- Insects of Acacia Species. Food and Agri- fluence of insects on mesquite seed pro- culture Organization of the United Nations, duction. Journal of Range Management Rome. 27:61–65. Southgate, B.J. 1984. Observations on the Snodgrass, R.E. 1908. A comparative study larval emergence in Callosobruchus chin- of the thorax in Orthoptera, Euplexoptera ensis (Coleoptera: Bruchidae). Entomologia and Coleoptera. Proceedings of the Ento- Generalis 9:177–180. mological Society of Washington 9:95–108.

310 Southgate, B.J., R.W. Howe, and G.A. development of Acanthoscelides obtectus Brett. 1957. The specific status ofCalloso- Say. Series Entomologica 19:165–174. bruchus maculatus (F.) and Callosobruchus Stampoulos, D., and J. Huignard. 1980. analis (F.). Bulletin of Entomological Re- L’influence de diverses parties de la graine search 48:79–89. de haricot (Phaseolus vulgaris) sur le Southgate, B.J., and R.D. Pope. 1957. The développement des larves d’Acanthoscel- groundnut seedbeetle, a study of its iden- ides obtectus Say (Coléoptèra, Bruchidae). tity and taxonomic position. Annals and Entomologia Experimentalia et Applicata Magazine of Natural History 10:669–672. 28:38–46. Spirina, T.S. 1974. The comparative mor- Staneva, E. 1980. Forms of cowpea weevil phology of the male and female genitalia in (Callosobruchus maculatus) and major dif- the two forms of the four-spotted cowpea ferences between them. Rastitelna Zashtita beetle Callosobruchus maculatus (Cole- 28:30–34. optera: Bruchidae). Entomological Revue Staneva, E. 1982. Studies on the food- (English translation of Entomologiskoe plants of the cowpea weevil Callosobru- Obozrenie) 53:22–27. chus maculatus (Coleoptera: Bruchidae). Srivastava, F.A.Z., and S.K. Bhatia. 1959. Rasteniev’dni Nauki 19:111–119. The effect of host species on the oviposition Staneva, E. 1983. Some biological char- of Callosobruchus chinensis Linn. (Cole- acteristics of the four-spotted bean weevil optera: Bruchidae). The Annals of Zoology Callosobruchus maculatus F. Rasteniev’dni 3:37–42. Nauki 20:110–121. Srivastava, U.S. 1953a. Reproductive Steffan, J.R. 1945. Contribution a l’etude organs of certain stored-grain beetles. II: de Zabrotes subfasciatus Boheman. Mem- Male organs of Sitophylus oryzae, Laria oirs of the Museum d’Histoire Natural affinis and Rhizopertha dominica. Proceed- (Paris) 21:55–94. ings of the Natural Academy of Sciences, India, 23:46–65. Steffan, J.R. 1946. La larve primaire de Bruchidius fasciatus Ol. et ses rap- Srivastava, U.S. 1953b. Reproductive ports avec quelques larves neonates de organs of certain stored-grain beetles. III: Bruchides. Bulletin de la Société Ento- Female organs of Sitophylus oryzae, Laria mologique de France 15:12–16. affinis and Rhizopertha dominica. Proceed- ings of the Natural Academy of Sciences, Steffan, J.R. 1981. The parasites of bru- India, 24:21–36. chids. In V. Labeyrie, ed., The Ecology of Bruchids Attacking Legumes (Pulses), Srivastava, U.S. 1966. Maxillary glands of Proceedings of the International Sympo- some coleopteran larvae. Indian Journal of sium, Tours (France), Apr. 16–19, 1980, Entomology 28:547–550. pp. 223–233. Series Entomologica 19. W. Stamopoulos, D.C. 1987. Influence of the Junk, The Hague, Netherlands. Leguminosae secondary substances on the Stein, J.D. 1983a. Insects infesting Acacia ecology and biology of Bruchidae. Ento- koa (Legumosae [sic]) and Metrosideros mologia Hellenica 5:61–67. polymorpha (Myrtaceae) in Hawaii: An- Stamopoulos, D.C. 1989. Effects of pho- notated list. Proceedings of the Hawaiian toperiod on the biology of Acanthoscelides Entomological Society 24:305–316. obtectus Say. Journal of Applied Entomol- Stein, J.D. 1983b. The biology, host range, ogy 107:150–154. parasites and hyperparasites of koa seed Stamopoulos, D., and P. Desroches. 1981. insects in Hawaii: A review. Proceedings Influence of the integument ofPhaseolus of the Hawaiian Entomological Society vulgaris seeds and of larval density on the 24:317–326.

311 Steinhauer, A.L. 1959. The biology and Swezey, O.H. 1954. Forest entomology in seasonal development of the vetch bruchid, Hawaii. Bernice P. Bishop Museum Special Bruchus brachialis Fahr. in Oregon. Jour- Publication No. 44. Honolulu. nal of Economic Entomology 52:955. Swier, S.R. 1974. Comparative seed pre- Stephens, J.F. 1829. A systematic cata- dation strategies of mesquite bruchids in logue of British insects: Being an attempt Arizona, with particular reference to seed to arrange all the hitherto discovered in- height, direction, and density. M.S. thesis, digenous insects in accordance with their Department of Biological Sciences, North- natural affinities, vol. 4:205–215. Baldwin ern Arizona University, Flagstaff. and Craddock, London. Szentesi, A. 1976. The effect of the ampu- Stickney, F.S. 1923. The head-capsule of tation of head appendages on the oviposi- Coleoptera. Illinois Biological Monographs tion of the bean weevil, Acanthoscelides 8:1–104. obtectus Say (Coleoptera: Bruchidae). Sym- Strong, R.G., G.J. Partida, and D.N. War- posia Biologica Hungarica 16:275–281. ner. 1968. Rearing stored-product insects Szentesi, A. 1981. Pheromone-like sub- for laboratory studies: Bean and cowpea stance affecting host-related behaviour of weevils. Journal of Economic Entomology larvae and adults in the drybean weevil, 61:747–751. Acanthoscelides obtectus. Entomologia Ex- Sturm, J. 1843. Catalog meiner Käfer- perimentalis et Applicata 30:219–226. Sammlung. Verfasser, Nürnberg, Germany. Takenouchi, Y. 1955. Chromosomes of Suffrian, E. 1870. Verzeichness der von Dr. three species of bruchids. Japanese Jour- Gundlach auf der Insel Cuba gesammelten nal of Genetics 30:7–9. Rüsselkäfer. Archiv für Naturgeschichte Takenouchi, Y. 1971a. Chromosomes in 36:150–234. males of 5 strains of Callosobruchus macu- Surtees, G. 1961. Spermathecal structure latus (Coleoptera: Bruchidae). Canadian in some Coleoptera associated with stored Journal of Genetics and Cytology 13:708– products. Proceedings of the Royal Ento- 713. mological Society of London (A) 36:144– Takenouchi, Y. 1971b. A further study on 152. the chromosomes of Callosobruchus chi- Suzuki, K. 1969. Comparative morphol- nensis (Coleoptera: Bruchidae). Kontyu ogy and evolution of the hind wings of the 39:332–337. family Chrysomelidae (Coleoptera). Kontyu Tanaka, K., K. Ohsawa, H. Honda, and I. 37:32–40. Yamamoto. 1981. Copulation release pher- Swezey, O.H. 1912. Some recent weevil de- omone erectin from the azuki bean weevil terminations. Proceedings of the Hawaiian Callosobruchus chinensis. Journal of Pesti- Entomological Society 2:167–168. cide Science 6:75–82. Swezey, O.H. 1921. Kauai insect notes and Tanaka, K., K. Ohsawa, H. Honda, and I. records. Proceedings of the Hawaiian Ento- Yamamoto. 1982. Synthesis of erectin, a mological Society 4:521. copulation release pheromone of the azuki bean weevil, Callosobruchus chinensis L. Swezey, O.H. 1925. [Note]. Proceedings of Journal of Pesticide Science 7:535–537. the Hawaiian Entomological Society 6:3–4. Tandon, G.N. 1960. Morphology of the Swezey, O.H. 1936. Fruit-eating and seed- head capsule and mouth parts of Cal- eating insects of Hawaii. Proceedings of losobruchus analis F. (Coleoptera, Phy- the Hawaiian Entomological Society 9:196– tophaga, Bruchidae). Zoologischer Anzeiger 201. 177:380–390.

312 Tanner, V.M. 1927. A preliminary study of superparasitisme chez l’hymenoptere the genitalia of female Coleoptera. Transac- solitaire Bruchocida vuilleti Cwf. (Hym., tions of the American Entomological Soci- Eupelmidae), parasite des stades larvaires ety 53:5–50. de son hote, Callosobruchus maculatus Tantawi, M.A. el-, K.A. Gouhar, M.M. Man- F. (Col., Bruchidae). Zeitschrift fur Ange- sour, and M.W. Guirguis. 1976. Blocking wandte Entomologie 101:243–256. of embryonic development in the southern Terrell, E.E., S.R. Hill, J.H. Wiersema, and cowpea weevil Callosobruchus maculatus W.E. Rice. 1986. A checklist of names for (Coleoptera: Bruchidae) by some juvenile 3,000 vascular plants of economic impor- hormone analogs. Zeitschrift für Ange- tance. U.S. Department of Agriculture, wandte Entomologie 81:37–42. Agricultural Handbook No. 505. Taper, M.L. 1990. Experimental character Thiara, N., R. Wadhwa, and S.J. Sharma. displacement in Callosobruchus maculatus. 1988. Age-related changes in total and mi- In K. Fujii, A.M.R. Gatehouse, C.D. John- tochondrial proteins in Caryedon serratus son, et al., eds., Bruchids and Legumes: Oliver [sic] (Coleoptera: Bruchidae). Jour- Economics, Ecology, and Coevolution. Pro- nal of Animal Morphology and Physiology ceeding of the 2nd International Sympo- 35:31–38. sium on Bruchids and Legumes, Okayama, Thiery, D. 1982a. Influence de la teneur Japan, September 6–9, 1989, pp. 289–301. en eau et de la durete du tegument des Series Entomologica 46. Kluwer, Dordre- graines de Phaseolus vulgaris sur la fre- cht, The Netherlands. quence de penetration des larves neonates Taylor, T.A. 1974. Observations on the d’Acanthoscelides obtectus. Entomologia effects of initial population densities in Experimentalis et Applicata 32:141–145. culture, and humidity on the production of “active” females of Callosobruchus macula- Thiery, D. 1982b. Consequences d’un tus (F.) (Coleoptera: Bruchidae). Journal of “epuisement” des larves neonates d’Acan- Stored Products Research 10:113–122. thoscelides obtectus sur la frequence de penetration dans des graines stockees de Taylor, T.A., and C. Agbaje. 1974. Flight Phaseolus vulgaris. Entomologia Experi- activity in normal and active forms of Cal- mentalis et Applicata 32:195–197. losobruchus maculatus (F.) in a store in Ni- geria. Journal of Stored Products Research Thiery, D., and M. Jarry. 1985. Hatching 10:9–16. rhythm in the bean weevil Acanthoscelides obtectus and larval penetration of Phaseo- Taylor, T.A., and J.I.S. Aludo. 1974. A lus vulgaris seeds. Insect Science and Its further note on the incidence of “active” Application 6:33–36. females of Callosobruchus maculatus (F.) in mature cowpeas in the field in Nige- Thukral, D. 1976. Structural organization ria. Journal of Stored Products Research and functions of spermathecal accessory 10:123–125. glands in Caryedon gonagra L. (Coleop- tera: Bruchidae). Indian Journal of Zoology Teran, A.L. 1967. Observaciónes sobre las 4:39–42. estructuras genitales de los machos de di- versos géneros de Bruchidae (Coleoptera). Thunberg, C.P. 1791. Dissertatio entomo- Acta Zoologia Lilloana 22:307–336. logica novas insectorum species sistens. Museum naturalium academiae Upsalen- Teran, A.L., and J.M. Kingsolver. 1977. sis. Appendix pars 111–122. Revision del genero Megacerus (Coleoptera: Bruchidae). Opera Lilloana 25:1–287. Thunberg, C.P. 1805. Illustrationes generum aliquot insectorum coleoptra- Terrasse, P., and D. Rojas-Rousse. 1986. torum. Göettingische gelehrte Anzeigen Distribution de la ponte et evitement du 28:281–282.

313 Thunberg, C.P. 1815. De Coleopteris ro- U.S. Department of Agriculture, Bureau of stratis. Nova Acta Royale Societie Scienti- Plant Quarantine. 1943. List of intercepted fica Upsala 7:104–125. plant pests. 1942. Thunberg, C.P. 1816. Fyra nya arter af U.S. Department of Agriculture, Bureau of Bruchus-slägtet. Vetenskapsakademien Plant Quarantine. 1944. List of intercepted Akademia Handlingar 1816:43–47. plant pests. 1943. Tikku, K., B.P. Saxena, and O. Koul. 1978. U.S. Department of Agriculture, Bureau of Oogenesis in Callosobruchus chinensis and Plant Quarantine. 1945. List of intercepted induced sterility by Acorus calamus oil va- plant pests. 1944. pors. Annales de Zoologie-Ecologie Animale U.S. Department of Agriculture. 1971. 10:545–551. Distribution of vetch bruchid. Cooperative Townsend, C.H.T. 1895. Some notes on Economic Insect Report No. 21. Bruchus in New Mexico. The Canadian En- U.S. Department of Agriculture. 1973. tomologist 27:277. [Note on Callosobruchus albocallosus, p. Townsend, C.H.T. 1903. Contributions to 149, and C. pulcher, p. 171]. Cooperative a knowledge of the coleopterous fauna of Economic Insect Report No. 23. the lower Rio Grande Valley in Texas and U.S. Department of Agriculture. 1978–97. Tamaulipas, with biological notes and A catalog of the Coleoptera of America special reference to geographical distribu- North of Mexico. Agriculture Handbook No. tion. Transactions of the Texas Academy of 529. [Issued in fascicles.] Science 5(1902):49–101. U.S. Department of Agriculture, Soil Con- Trelease, S.F., and H.M. Trelease. 1937a. servation Service. 1982. National list of Immunity of certain insects to selenium plant names. Vol. 1, List of plant names. poisoning. Science 85:590. Vol. 2, Synonymy. TP–159. Trelease, S.F., and H.M. Trelease. 1937b. Utida, S. 1954. “Phase” dimorphism ob- Toxicity to insects and mammals of foods served in the laboratory population of the containing selenium. American Journal of cowpea weevil Callosobruchus quadri-macu- Botany 24:448–451. latus. Japanese Journal of Applied Zoology Udayagiri, S., and S.R. Wadhi. 1982. A key 18:161–168. to world bruchid genera. National Bureau Utida, S. 1961. Experimental studies on of Plant Genetic Resources (New Delhi), the interaction between the bean weevils Scientific Monograph 5. and their parasitic wasps. In H. Strouhal Udayagiri, S., and S.R. Wadhi. 1989. Cata- and M. Beier, eds. Proceedings of the XI log of Bruchidae. Memoirs of the American International Congress for Entomology, Entomological Institute 45:1–301. Vienna, August 17–25, 1960, pp. 731–734. Ulke, H. 1902. A list of the beetles of the Reisser’s Söhne, Vienna, Austria. District of Columbia. Proceedings of the Utida, S. 1967. Collective oviposition and United States National Museum 25 (No. larval aggregation in Zabrotes subfasciatus 1275). (Boh.) (Coleoptera: Bruchidae). Journal of U.S. Department of Agriculture, Bureau of Stored Products Research 2:315–322. Plant Quarantine. 1940. List of intercepted Utida, S. 1969. Photoperiod as a factor plant pests. 1938. inducing the flight form in the population U.S. Department of Agriculture, Bureau of of the southern cowpea weevil, Calloso- Plant Quarantine. 1942. List of intercepted bruchus maculatus. Japanese Journal of plant pests. 1941. Applied Entomology and Zoology 13:129– 134.

314 Utida, S. 1972. Density dependent poly- tus (Olivier). Revista Facultad National de morphism in the adult of Callosobruchus Agronomia 27:71–74. maculatus (Coleoptera: Bruchidae). Jour- Verma, K.K., and R. Saxena. 1996. The nal of Stored Products Research 8:111– status of Bruchidae as a family. Chrys- 126. omela 32:3. Utida, S. 1974. Polymorphism in the adult Wade, O. 1919. The four-spotted cowpea of Callosobruchus maculatus—a possible weevil (Bruchus quadrimaculatus Fabri- process of evolution to stored product cius). Oklahoma Agricultural Experiment pest. Proceedings of the 1st International Station Bulletin No. 129. Working Conference on Stored Product Entomology, Savannah, GA, October 7–11, Wagner, W.L., D.R. Herbst, and S.H. Sohm- 1974, pp. 686–699. er. 1990. Manual of the Flowering Plants of Hawaii. University of Hawaii Press, Hono- Utida, S. 1981. Polymorphism and phase lulu. dimorphism in Callosobruchus. Series En- tomologica 19:143–147. Wallace, F.L., and R.C. Fox. 1975. A comparative morphological study of the hind Vats, L.K. 1972. Tracheal system in the wing venation of the order Coleoptera, part larvae of the Bruchidae (Coleoptera: Bru- 1. Proceedings of the Entomological Society chidae). Journal of the New York Entomo- of Washington 77:329–354. logical Society 80:12–17. Wallace, F.L., and R.C. Fox. 1980. A com- Vats, L.K. 1973. Taxonomic values of ana- parative morphological study of the hind tomical characters in bruchid larvae (Bru- wing venation of the order Coleoptera, part chidae: Coleoptera). Research Bulletin (Sci- 2. Proceedings of the Entomological Society ence) of the Panjab University 24:167–169. of Washington 82:609–654. Vats, L.K. 1974a. Distinctive characters of Wang, R., and L.T. Kok. 1983. Synchro- the larvae of three species of Calloso-bru- nization of Megacerus discoidus with the chus Pic (Bruchidae: Coleoptera), together hedge bindweed Calystegia sepium, in with a key for their identification. Indian southwestern Virginia. Virginia Journal of Journal of Entomology 36:17–22. Science 34:109. Vats, L.K. 1974b. Method for distinguish- Wang, R., and L.T. Kok. 1986a. Life history ing larvae of Bruchidae (Coleoptera) attack- of Megacerus discoidus (Coleoptera: Bru- ing the same host plant. Entomologists’ chidae), a seed feeder of hedge bindweed. Monthly Magazine 110:142. Annals of the Entomological Society of Vats, L.K. 1976a. The malpighian tubules America 79:359–363. in the larvae of the family Bruchidae (Cole- Wang, R., and L.T. Kok. 1986b. Host speci- optera). Indian Biologist 5:79–82. ficity ofMegacerus discoidus (Coleoptera: Vats, L.K. 1976b. Alimentary canal in Bruchidae) and its impact on hedge bind- bruchid larvae (Bruchidae: Coleoptera). weed, Calystegia sepium. Environmental Research Bulletin (Science) of the Panjab Entomology 15:834–838. University 27:103–106. Ward, C.R., C.W. O’Brien, L.B. O’Brien, et Vayssière, P., and P. Lepesme. 1941. Sur al. 1977. Checklist of New World insects quelques Bruchides nuisibles. Revue Fran- associated with Prosopis (Mesquite). U.S. caise d’Entomologie 8:198–202. [new spe- Department of Agriculture, Technical Bul- cies described on pp. 201–202] letin No. 1557. Velez-Angel, R. 1972. Tres plagas insectiles Warthen, J.D., Jr. 1989. Neem (Azadir- recientemente detectadas en Antioquia. 1. achta indica A. Juss.): Organisms affected El gorgojo del tamarindo, Caryedon serra- and reference list update. Proceedings of

315 the Entomological Society of Washington Wenzel, H. 1912. [Notes, December 11, 91:367–388. 1911, meeting of the American Entomologi- Wasserman, S.S. 1981. Host-induced ovi- cal Society]. Entomological News 23:140. position preference and oviposition mark- Werner, F.G., and G.D. Butler. 1958. A sur- ers in the cowpea weevil Calloso-bruchus vey of the insects on mesquite in southern maculatus. Annals of the Entomological Arizona (preliminary report). Department of Society of America 74:242–245. Entomology, University of Arizona, Tucson. Wasserman, S.S. 1985. Oviposition be- Unpublished manuscript. haviour and its disruption in the southern Wheeler, J., and J.T. Longino. 1988. Ar- cowpea weevil, Callosobruchus maculatus thropods in live oak galls in Texas. Ento- F. (Coleoptera: Bruchidae). Journal of Eco- mological News 99:25–29. nomic Entomology 78:89–92. Wheeler, W.H., J. Hunt, and E.P. Reagan. Wasserman, S.S. 1987. Behavioral analysis 1950. List of intercepted plant pests, 1948. of male-induced interstrain differences in U.S. Department of Agriculture, Bureau of realized fecundity in Callosobruchus macu- Entomology and Plant Quarantine, Wash- latus. In M.D. Huettel, ed., Evolutionary ington, DC. Genetics of Invertebrate Behavior: Progress White, B.E. 1941. A new species of Bru- and Prospects, pp. 145–152. Plenum Press, chus with notes on Bruchus major Fall and New York. julianus Horn. Pan-Pacific Entomologist Wasserman, S.S., and D.J. Futuyma. 17:189–190. 1981. Evolution of host plant utilization White, R.E. 1983. A Field Guide to the in laboratory populations of the southern Beetles of North America. Houghton Mif- cowpea weevil, Callosobruchus maculatus flin, Boston. Fabricius (Coleoptera: Bruchidae). Evolu- tion 35:605–617. Whitehead, D.R., and J.M. Kingsolver. 1975a. Biosystematics of the North and Waterworth, P.D. 1986. Internal seed in- Central American species of Gibbobruchus festing insects. U.S. Department of Agricul- (Coleoptera: Bruchidae: Bruchinae). Trans- ture, Animal and Plant Health Inspection actions of the American Entomological Service, Publication No. 81–48. Society 101:167–225. Weiss, H.B. 1919. The more important Whitehead, D.R., and J.M. Kingsolver. insect enemies of the rose-mallow in New 1975b. Beetles and wasps associated with Jersey. New Jersey Department of Agricul- Cassia biflora L. (Caesalpiniaceae) fruits in ture, Circular No. 25. Costa Rica, with a new species of Sennius Weiss, H.B., and E.L. Dickerson. 1919. (Coleoptera: Bruchidae). Journal of the Insects of the swamp rose-mallow, Hibis- Washington Academy of Sciences 65:154– cus moscheutos L., in New Jersey. Journal 157. of the New York Entomological Society Wickham, H.F. 1895a. On the larvae of 27:39–68. Lucidota, Synoxylon, and Spermophagus. Wendt, H. 1980. Eine für Sudost-Europa Bulletin of the Laboratories of Natural His- neue Samenkäfer-Art (Coleoptera: Bru- tory, State University of Iowa 3:31–35. chidae). Folia Entomologica Hungarica Wickham, H.F. 1895b. A seed weevil (Sper- 42:223–226. mophagus robiniae) in honey locust seeds Wendt, H. 1984. Zür kenntnis der Bru- (Gleditsiae triacanthos). Bulletin of the chidenfauna Bulgariens. Deutsche Ento- Laboratories of Natural History, State Uni- mologische Zeitschrift 31:153–167. versity of Iowa 3:157.

316 Wickham, H.F. 1912. A report on some Rico. Journal of Agriculture of the Univer- recent collections of fossil Coleoptera from sity of Puerto Rico 20:1–627. the Miocene shales of Florissant. Bulletin Wollaston, T.V. 1854. Insecta Maderensia; of the Iowa State University Laboratories of being an account of the insects of the Ma- Natural History 6:3–38. deiran group. Van Voorst, London. Wickham, H.F. 1913a. Fossil Coleoptera Wollaston, T.V. 1870. On the Coleoptera of from Florissant in the United States Na- St. Helene. Annals and Magazine of Natu- tional Museum. Proceedings of the United ral History 5:18–37. States National Museum 45:283–303. Woodruff, R.E. 1968. The palm seed “wee- Wickham, H.F. 1913b. Fossil Coleoptera vil,” Caryobruchus gleditsiae (L.), in Florida from the Wilson Ranch near Florissant, (Coleoptera: Bruchidae). Florida Depart- Colorado. Bulletin of the Laboratories of ment of Agriculture, Entomology Circular Natural History, State University of Iowa No. 73. 4:3–29. Yadav, J.S. 1973. Cytological studies on Wickham, H.F. 1914. New Miocene Coleop- Caryedon Schonh. (Bruchidae: Coleoptera). tera from Florissant. Bulletin of the Muse- The Nucleus 16:126–129. um of Comparative Zoology 58:480–484. Yadav, T.D. 1977. Mechanism of emer- Wickham, H.F. 1917. New species of fossil gence of pulse beetle. Entomologists’ News- beetles from Florissant, Colorado. Proceed- letter 7:36–37. ings of the United States National Museum 52:463–472. Yadav, T.D., and N.C. Pant. 1975. Effect of feeding by developing stages of Calloso- Wiersema, J.H., J.H. Kirkbride, Jr., and bruchus maculatus (Fab.) and C. chinensis C.R. Gunn. 1990. Legume (Fabaceae) (Linn.) on germination of pulse seeds. Seed nomenclature in the USDA germplasm Research 3:107–110. system. U.S. Department of Agriculture, Technical Bulletin No. 1757. Yadav, T.D., and S. Singh. 1977. Respira- tory behaviour of developing stages of Cal- Wigglesworth, V.B. 1947. The Principles of losobruchus maculatus (Fab.) and C. chi- Insect Physiology. Methuen, London. nensis (Linn.) under airtightness. Bulletin Wightman, J.A., and B.J. Southgate. 1982. of Grain Technology 15:67–70. Egg morphology, host and probable re- Yamamoto, I. 1986. New insect pheromone gions of origin of the bruchids (Coleoptera: erectin-like substances. Phytoparasitica Bruchidae) that infest stored pulses—an 14:361. identification aid. New Zealand Journal of Experimental Agriculture 10:95–99. Yamamoto, I. 1990. Chemical ecology of bruchids. In K. Fujii, A.M.R. Gatehouse, Wilson, K. 1988. Egg-laying decisions by C.D. Johnson, et al., eds., Bruchids and the bean weevil Callosobruchus maculatus. Legumes: Economics, Ecology and Coevo- Ecological Entomology 13:107–188. lution. Proceeding of the 2nd International Wittich, F.W. 1940. Allergic rhinitis and Symposium on Bruchids and Legumes, asthma due to sensitization to the Mexican Okayama, Japan, September 6–9, 1989, bean weevil (Zabrotes subfasciatus Boh.). pp. 53–62. Series Entomologica 46. Klu- Journal of Allergy 12:42–45. wer, Dordrecht, The Netherlands. Wolcott, A.B. 1912. Two new species of Co- Yip, J.B. 1936. Insect damage to seeds of leoptera from Illinois. The Canadian Ento- Cracca virginiana L. Journal of Economic mologist 44:161–163. Entomology 29:622–629. Wolcott, A.B. 1936. Insectae Borinquenses. Yoshida, T. 1989. Life history of Calloso- A revised checklist of the insects of Puerto bruchus chinensis one generation per year. In Proceedings of the First Asia-Pacific

317 Conference of Entomology, Chiang Mai, Thailand, November 8–13, 1989, p. 37. Bangkok, Thailand. Yoshida, T., ed. 1990. Loss from and control of bruchids in developing countries, Country Report Session. In Supplement to Proceeding of the Second International Symposium on Bruchids and Legumes, Okayama, Japan, September 6–9, 1990. Zachariae, G. 1959. Das Verhalten des Speisebohnenkäfers Acanthoscelides obtectus Say (Coleoptera: Bruchidae) im Freien in Norddeutschland. Zeitschrift für Ange- wandte Entomologie 43:345–365. Zacher, F. 1929. Nahrungsauswald und biologie der Samenkäfer. Deutsche Ge- sellschaft für Angewandte Entomologie 1928:55–62. Zacher, F. 1930. Untersuchungen zür morphologie und biologie der Samenkäfer. Ar- beiten aus der Biologischen Reichs-anstalt für Land- und Forstwirtschaft, Berlin- Dahlem, 18:233–384. Zacher, F. 1932. Untersuchungen über die anatomie der Geschlechtsorgane und die systematik der Samenkäfer (Bruchidae). Archivio Zoologico Italiano 16:1005–1009. Zacher, F. 1952. Die Nährpflanzen der Samenkäfer. Zeitschrift fur Angewandte Entomologie 33:460–480. Zampetti, M.F. 1981. Posizione sistematica di alcune specie appartenenti al genre Bruchidius Schilsky (Coleoptera: Bruchi- dae). Bolletino del Museo Civico di Storia Naturelle di Verona 8:383–410. Zia, Y. 1936. Comparative studies of the male genital tube in Coleoptera Phytopha- ga. Sinensia 7:326,336–339.

318 Index ater (Marsham), Bruchidius...... 78 atomus (Fall), Acanthoscelides...... 98 abbreviatus (Say), Sennius...... 200 atrocephalus Pic, Acanthoscelides...... 125 aboriginalis Wickham, Bruchus [fossil atronotatus (Pic), Lithraeus...... 171, 226 species]...... 237 auctus (Fall), Sennius...... 206 Abutiloneus Bridwell...... 88 aureolus Horn, Acanthoscelides, species abutilonis (Motschulsky), complex...... 99 Acanthoscelides...... 97 baboquivari Johnson, Acanthoscelides.. 101 acaciestes Kingsolver and Johnson, Mimosestes...... 186 barbicornis (Fabricius), Callosobruchus... 79 Acanthobruchus Ramos, Bruchus...... 69 basicornis Pic, Zabrotes...... 52 Acanthocelides...... 89 beali Johnson, Stator...... 212 acanthocnemus (Jekel), bexarensis Kingsolver, Zabrotes...... 43 Acanthoscelides...... 131 biguttatus Fabricius, Callosobruchus...... 80 Acanthoscelides Schilsky...... 89 biguttellus Schoenherr, Callosobruchus... 80 Acanthoscelidini Bridwell...... 88 binotatus (Pic), Caryedes...... 164 ademptus (Sharp), Callosobruchus...... 76 bisignatus (Horn), Acanthoscelides...... 102 aequalis (Sharp), Acanthoscelides...... 96 bistriatus (Fabricius), Callosobruchus...... 80 albiscutellaris (Ashmead), biustulus (Fall), Acanthoscelides...... 103 Acanthoscelides...... 97 bivulneratus (Horn), Sennius...... 200 alboguttis (Motschulsky), Meibomeus..... 173 borealis (Schoenherr), Gibbobruchus...... 169 alboscutellaris (Zacher), Acanthoscelides...... 97 Borowieius Anton...... 75 alboscutellatus (Horn), Acanthoscelides... 97 bottimeri Kingsolver, Algarobius.....213, 225 Algarobius Bridwell...... 155 bottimeri Kingsolver, Stator...... 156 alternatus Bridwell, Megacerus...... 65 bowditchi Wickham, Bruchus [fossil species]...... 237 Althaeus Bridwell...... 158 brachialis Fahraeus, Bruchus...... 70 ambiguus (Gyllenhal), Callosobruchus..... 83 breweri (Crotch), Mimosestes...... 131 Amblycerinae Bridwell...... 30 Bruchelae Latreille...... 19 Amblycerini Bridwell...... 30 Bruchinae Bridwell...... 75 Amblycerus Thunberg...... 30 Bruchidius Schilsky...... 76 amicus (Horn), Mimosestes...... 187, 226 Bruchinae Pic...... 58 amplissimus Kingsolver, Zabrotes...... 42 Bruchini Bridwell...... 69 Andromisus Gozis...... 163 Bruchoides Ramos, Bruchus...... 69 anilis Scudder, Bruchus [fossil species]...... 237 Bruchus Linnaeus...... 19, 69 antaeus Wickham, Bruchus [fossil brunneostictus (Fall), Acanthoscelides.... 142 species]...... 237 calderensis (Sharp), Caryedes...... 164 Anthotribus Gistel...... 30 californicus (Boheman), Sennius...... 204 arachidis Fahraeus, Callosobruchus...... 83 Callosobruchus Pic...... 79 arenarius (Wolcott), Zabrotes...... 42 calvus (Horn), Acanthoscelides...... 104 arizonensis (Schaeffer), Neltumius...... 195 canus Germar, Bruchidius...... 77 arthriticus (Fabricius), Caryobruchus...... 26 319 carpophiloides Wickham, Bruchus [fossil depressus (Fall), Sennius...... 201 species]...... 237 desertorum (LeConte), Algarobius...... 157 Caryedes Hummel...... 163 desmanthi Johnson, Acanthoscelides.... 109 Caryedon Schoenherr...... 24 desmoportheus Kingsolver and Caryedontini Bridwell...... 24 Whitehead, Meibomeus...... 172 Caryobruchus Bridwell...... 26 discoidens (Dury), Megacerus...... 63 cearanus Pic, Stator...... 215 discoideus (Pic), Megacerus...... 63 celatus (Sharp), Sennius...... 205 discoidii (Motschulsky), Megacerus...... 63 Cercidiestes Bridwell...... 185 discoidus (Say), Megacerus...... 62 chandleri Kingsolver, Zabrotes...... 44 discolor Fall and Cockerell, Sennius...... 207 chavesi Kingsolver, Zabrotes...... 44 discolor (Horn), Sennius...... 202 chihuahua Johnson and Kingsolver, discopterus (Fall), Sennius...... 203 Stator...... 214 distinguendus (Horn) Acanthoscelides... 109 chinensis (Linnaeus), Callosobruchus divaricatae Whitehead and Kingsolver, ...... 80, 226 Gibbobruchus...... 168 chiricahuae (Fall), Acanthoscelides...... 105 dominicanus (Jekel), Mimosestes...... 189 cingulatus (Suffrian), Zabrotes...... 52 dormescens Jekel, Bruchus [fossil cisti (Fabricius), Bruchidius...... 77 species]...... 237 coconino Johnson and Kingsolver, dorsopictus (Lepesme), Zabrotes...... 52 Stator...... 214 eldenensis Kingsolver, Zabrotes...... 47 cognatus (Sharp), Stator...... 215 erythrocerus (Riley), Meibomeus...... 173 collusus (Fall), Acanthoscelides...... 135 (electus) Bridwell, Lithraeus...... 170 compressicornis (Schaeffer), elegans Bridwell, Lithraeus...... 170 Acanthoscelides...... 106 elegans Sturm, Callosobruchus...... 80 comstocki Johnson, Acanthoscelides...... 107 eugenie Bottimer, Megacerus...... 64 conicicollis Fairmaire, Callosobruchus...... 86 eustrophoides (Schaeffer), conspersus (Motschulsky), Amblycerus...... 32 Acanthoscelides...... 97 exhumatus Wickham, Bruchus [fossil coryphae (Olivier), Megacerus...... 59 species]...... 237 crategi (Fahraeus), Gibbobruchus...... 169 exiguus (Horn), Acanthoscelides...... 152 crenatus (Schaeffer), Megacerus...... 68 fabae (Fitch), Acanthoscelides...... 131 cristicollis (Sharp), Gibbobruchus...... 167 fabae Motschulsky, Bruchus...... 74 cruciger Geoffroy, Bruchus...... 72 fabae (Riley), Acanthoscelides...... 131 cruciger Horn, Zabrotes...... 45 fabi (Rathvon), Acanthoscelides...... 131 cruentatus (Horn), Sennius...... 201 fallax (Boheman), Sennius...... 203 cubiculus (Casey), Megacerus...... 60 Falsobruchus Pic...... 163 cubicus (Motschulsky), Megacerus...... 61 fasciatus Olivier, Bruchidius...... 78 cynthiae Kingsolver, Zabrotes...... 46 figuratus Gyllenhal, Callosobruchus...... 86 Cytorhinus Agassiz...... 28 flavescens Fahraeus, daleae Johnson, Acanthoscelides...... 108 Acanthoscelides...... 111 debilis Gyllenhal, Bruchidius...... 77 flavicornis Sharp, Abutiloneus...... 88 debilis Schilsky, Bruchidius...... 77 flexicaulis Schaeffer, Merobruchus...... 181 densus Horn, Zabrotes...... 47 320 floridae (Horn), Acanthoscelides...... 112 insularis Johnson and Kingsolver, florissatensis Wickham, Oligobruchus Mimosestes...... 188, 226 [fossil species]...... 238 intermedius Motschulsky, Bruchus...... 72 folkertsi Kingsolver, Althaeus...... 159 interruptus (Sharp), Stator...... 215 fraterculus (Horn), Acanthoscelides...... 113 ireriae Romero, Johnson, and Kingsolver, fumatus (Schaeffer), Acanthoscelides..... 114 Amblycerus...... 33 fuscus Bedel, Bruchus...... 24 irresectus (Fahraeus), Acanthoscelides ...... 89, 131 gibbithorax (Schaeffer), Neltumius...... 196 julianus (Horn), Merobruchus...... 177 Gibbobruchus Pic...... 166 karasini Fischer, Kytorhinus...... 28 gibbothorax Kingsolver, Neltumius...... 196 kingsolveri Pic, Acanthoscelides...... 119 gilvipes Motschulsky, Acanthoscelides... 131 kiotoensis Johnson, Acanthoscelides..... 115 gleditsiae (Castiglioni), Amblycerus...... 36 knulli (White), Merobruchus...... 180 gleditsiae (Johansson and Linnaeus), Caryobruchus...... 26 Kytorhininae Bridwell...... 28 gonager (Chujo), Caryedon...... 24 Kytorhinus Fischer...... 28 gonagra (Fabricius), Caryedon...... 24 Kytorrhinus Baudi...... 28 granarius (Marsham), Bruchus...... 74 kytorrhinensis (Pic), Megacerus...... 67 griseolus (Fall), Acanthoscelides...... 115 Kytorrhininae Luk’yanovich and Ter-Minassian...... 28 haywardi (Wickham), Oligobruchus [fossil species]...... 238 Kytorhunus Motchulsky...... 28 helianthemum Bottimer, Laria Scopoli...... 20, 69 Acanthoscelides...... 116 lebasi (Fahraeus), Sennius...... 205 helvinus (Motschulsky), Caryedes...... 164 leguminarius Gyllenhal,, henshawi Wickham, Bruchus [fossil Acanthoscelides...... 131 species]...... 237 leucogaster (Sharp), Zabrotes...... 52 herissantitus Johnson, leucosomus (Sharp), Megacerus...... 62 Acanthoscelides...... 117 leucospilus (Sharp), Megacerus...... 65, 226 hibisci (Olivier), Althaeus...... 160 leucostauros Johnson and Kingsolver, hoffmanseggi (Gyllenhal), Amblycerus...... 36 Sennius...... 206 horni (Pic), Acanthoscelides...... 153 limbatus (Horn), Stator...... 215, 226 humboldtae Kingsolver, Zabrotes...... 48 limpidus (Sharp), Merobruchus...... 182 idoneus Bridwell, Abutiloneus...... 88 lineatipennis (Pic), Megacerus...... 65 immunis (Sharp), Mimosestes...... 189 litteratus Schoenherr, Bruchus...... 82 impiger (Horn), Megacerus...... 64 Lithraeus Bridwell...... 170 incensus (Sharp), Caryedes...... 165 lobatus (Fall), Acanthoscelides...... 120 incretus Motschulsky, longistilus (Horn), Acanthoscelides...... 121 Acanthoscelides...... 131 longiventris (Sharp), Mimosestes...... 192 infirmus (Sharp), Sennius...... 203 longus (Pic), Caryedon...... 24 innotatus (Pic), Mimosestes...... 189 lunarius Rey, Bruchus...... 72 inornatus (Horn), Mimosestes...... 189 lysilomae Kingsolver, Merobruchus...... 180 inquisitus (Fall), Acanthoscelides...... 118 macrocerus (Horn), Stylantheus...... 221 insolitus (Sharp), Merobruchus...... 176 macrophthalmus (Schaeffer), Acanthoscelides...... 122 321 mactatus (Pic), Megacerus...... 67 obesulus (Sharp), Sennius...... 208 maculatus (Fabricius), obliteratus (Horn), Zabrotes...... 48 Callosobruchus...... 82, 226 obrienorum Johnson, Acanthoscelides... 128 maculiventris (Fahraeus), Megacerus...... 66 obscurus (Fitch), Acanthoscelides...... 129 major (Fall), Merobruchus...... 181 obscurus (Sharp), Amblycerus...... 34 managuanus (Pic), Sennius...... 203 obsoletus (Say), Acanthoscelides...... 129 margaretae Johnson, Acanthoscelides... 123 obtectus (Say), Acanthoscelides...... 89, 130 medialis (Sharp), Sennius...... 206 ochraceicolor (Pic), Acanthoscelides...... 111 Megacerini Bridwell...... 58 ochraceus (Schaeffer), Megacerus Fahraeus...... 58 Acanthoscelides...... 89, 111 Megacerus, subgenus of Megacerus...... 58 ochreolineatus (Fall), Merobruchus...... 177 Meibomeus Bridwell...... 172 ochrolineatus (Pic), Merobruchus...... 177 Merobruchus Bridwell...... 24, 175 Oligobruchus Kingsolver [fossil genus]... 238 mexicanus (Sharp), Acanthoscelides...... 149 oregonensis Johnson, Acanthoscelides.. 134 mimosae (Fabricius), Mimosestes...... 189 ornatus (Boheman), Callosobruchus...... 82 mimosae Gemminger and Harold, osborni Wickham, Bruchus [fossil Acanthoscelides...... 131 species]...... 238 Mimosestes Bridwell...... 185 Pachybruchus Pic, subgenus of mimus (Say), Gibbobruchus...... 169 Megacerus...... 58 minusculus Pic, Zabrotes...... 52 Pachymera Berthold...... 163 mixtus (Horn), Acanthoscelides...... 124 Pachymerinae Bridwell...... 24 modestus (Sharp), Acanthoscelides...... 125 Pachymerini Bridwell...... 26 morosus (Sharp), Sennius...... 207 Pachymerus Schoenherr...... 163 mucrofer (Bottimer), Stylantheus...... 221 pallidipennis (Motschulsky), Acanthoscelides...... 135 multisparsus (Pic), Megacerus...... 67 pallidipes (Fahraeus), mundulus (Sharp), Acanthoscelides...... 126 Acanthoscelides...... 131 murinus (Schoenherr), Gibbobruchus..... 169 pauperculus (LeConte), musculus (Boheman), Zabrotes...... 52 Acanthoscelides...... 136 musculus (Say), Meibomeus...... 173 pectinicornis (Linnaeus), Callosobruchus...... 80 Mylabris Fabricius...... 20 pectoralis (Horn), Acanthoscelides...... 137 Mylabris Müller...... 20, 69 pectoralis (Sharp), Zabrotes...... 52 napensis Johnson, Acanthoscelides...... 127 Pedalophus Bottimer, Caryedes...... 163 Neltumius Bridwell...... 194 Pedapholus Gistel, Caryedes...... 163 nebulosus (Olivier), Amblycerus...... 31 pedicularius (Sharp), Acanthoscelides.... 138 nesapius Fahraeus, Bruchus...... 52 perforatus (Horn), Acanthoscelides...... 139 nictitans (Motschulsky), Sennius...... 201 perplexus (Fall), Acanthoscelides...... 135 nigrinus (Horn), Sennius...... 201 pescaprae (Fahraeus), Megacerus...... 58 nigromarginatus (Motschulsky), Amblycerus...... 34 phaseoli (Gyllenhal), Callosobruchus ...... 86, 226 nubigens (Motschulsky), Mimosestes, ...... 191, 226 piger (Sharp), Stator...... 216 nugax Motschulsky, Bruchidius...... 77

322 pisi (Linnaeus), Bruchus...... 72 rufovittatus (Schaeffer), pisorum (Linnaeus), Bruchus...... 72, 225 Acanthoscelides...... 145 placidus (Horn), Merobruchus...... 182 rufus (De Geer), Callosobruchus...... 80 planifrons Horn, Zabrotes...... 50 rufus (Motschulsky), Acanthoscelides.... 136 pluto Wickham, Spermophagus salicis Scopoli, Bruchus...... 72 [fossil species]...... 238 sallaei (Sharp), Mimosestes...... 191 primoticus (Wickham), Oligobruchus scabricollis (Chevrolat), Caryedes...... 164 [fossil species]...... 238 schaefferi (Pic), Acanthoscelides...... 146 probus (Sharp), Sennius...... 204 schaefferianus Bridwell, Megacerus...... 68 prolixus (Fall), Kytorhinus...... 28 schrankiae (Horn), Acanthoscelides...... 147 prosopis (LeConte), Algarobius...... 157 schwarzi Kingsolver, Amblycerus...... 37 prosopoides (Schaeffer), scudderi (Wickham), Oligobruchus Acanthoscelides...... 140 [fossil species]...... 238 protractus (Horn), Mimosestes...... 192 scutellaris (Fabricius), Callosobruchus..... 80 pruininus (Horn), Stator...... 211, 216, 226 scutellaris Sharp, Kytorhinus...... 28 Pseudopachymerus Pic...... 163 semicinctus Horn, Zabrotes...... 52 pugiunculus (Fall), Acanthoscelides...... 151 semicolon (Sharp), Stator...... 219 pugiunculus (Pic), Acanthoscelides...... 151 semifasciatus (Boheman), Zabrotes...... 52 pulcher (Pic), Callosobruchus...... 82, 226 seminulum (Horn), Acanthoscelides...... 148 pulicarius (Motschulsky), Sennius Bridwell...... 198 Acanthoscelides...... 136 Serratibruchus Teran and Kingsolver, pulloides (Fall), Acanthoscelides...... 138 subgenus of Megacerus...... 58 pullus (Fall), Acanthoscelides...... 142 serratifemur (Schaeffer), Megacerus...... 66 pulicarius Motschulsky, Bruchidius...... 77 serratus (Olivier), Caryedon...... 24, 226 pusillimus (Sharp), Acanthoscelides...... 143 sharpi (Pic), Acanthoscelides...... 149 pygidialis (Schaeffer), Stator...... 218 sharpianus Bridwell, Kytorhinus...... 28 Pygobruchus Sharp...... 28 sibutensis Pic, Caryedon...... 24 pythonicus (Pic), Stator...... 218 simplex (Motschulsky), quadratus (Motschulsky), Megacerus...... 66 Acanthoscelides...... 136 quadridentatus (Schaeffer), simulans (Schaeffer), Sennius...... 209 Acanthoscelides...... 144 sinuatus (Fahraeus), Callosobruchus...... 83 quadrimaculatus (Fabricius), sordidus (Horn), Stator...... 219 Callosobruchus...... 82 sparsus Fabricius, Bruchus...... 72 ramicornis (Boheman), Megacerus...... 64 Sparteus Bridwell...... 77 ripiphorus (Fahraeus), Megacerus...... 68 speciosus (Schaeffer), Acanthoscelides... 149 robiniae (Fabricius), Amblycerus...... 35 spectabilis Horn, Zabrotes...... 50 rostratus Motschulsky, Bruchidius...... 77 Spermophagini Borowiec...... 39 rufescens (Motschulsky), Sennius...... 205 Spermophagus Schoenherr...... 20, 30, 238 rufescens (Schaeffer), Acanthoscelides...... 146 Stator Bridwell...... 211 rufimanus Boheman, Bruchus...... 74, 225 steineri Kingsolver, Althaeus...... 161 rufobrunneus Wallaston, stephani Kingsolver, Zabrotes...... 51 Callosobruchus...... 80 Stylantheus Bridwell...... 221

323 stylifer (Sharp), Acanthoscelides...... 150 varicornis (Motschulsky), strigatus (Motschulsky), Mimosestes...... 189 Acanthoscelides...... 131 subaeneus (Schaeffer), Stator...... 219 vicinus (Gyllenhal), Callosobruchus...... 82 subaequalis Johnson, Acanthoscelides.. 151 victoriensis Kingsolver, Zabrotes...... 56 subellipticus (Wollaston), villosus (Fabricius), Bruchidius...... 78 Acanthoscelides...... 131 vitis (Schaeffer), Amblycerus...... 38 subfasciatus (Boheman), Zabrotes... 52, 226 vivificatus (Scudder), Spermophagus submersus (Wickham), Oligobruchus [fossil species]...... 238 [fossil species]...... 238 wheelocki (Blatchley), Megacerus...... 68 submuticus (Sharp), Acanthoscelides..... 152 whitei Johnson and Kingsolver, subnitens Horn, Zabrotes...... 55 Sennius...... 210 subrubrosus (Blackwelder), wilsoni (Wickham), Oligobruchus Meibomeus...... 174 [fossil species]...... 238 subrufus (Motschulsky), Mimosestes...... 189 xanthopus (Suffrian), Sennius...... 204 subserripes (Fall), Acanthoscelides...... 106 Zabrotes Horn...... 39 succinctus Wickham, Bruchus [fossil species]...... 238 surrubresus (Pic), Meibomeus...... 174 sylvestris Romero and Johnson, Zabrotes...... 55 tarnawskii Borowiec, Acanthoscelides... 135 tenuis Bottimer, Acanthoscelides...... 154 terani Kingsolver, Merobruchus...... 183 tetricus (Gyllenhal), Acanthoscelides...... 131 texanus (Schaeffer), Neltumius...... 197 trabuti (Caillol), Callosobruchus...... 83 transversus (Say), Althaeus...... 160 triangularifer (Pic), Megacerus...... 67 tridenticulatus Bottimer, Acanthoscelides...... 155 ulkei (Horn), Mimosestes...... 193 unicolor (Olivier), Bruchidius...... 77 uniformis (LeConte), Algarobius...... 157 usticolor (Sharp), Stator...... 219 vachelliae Bottimer, Stator...... 220 vacillator (Sharp), Merobruchus...... 184 vandykei Kingsolver, Zabrotes...... 44

324