HARVARD UNIVERSITY

LIBRARY

OF THE

Museum of Comparative Zoology

/ ft

Quaest

lones

entomologicae

MUS. COM P.

V

A periodical record of entomological investigations,

published at the Department of Entomology, Uni-

versity of Alberta, Edmonton, Canada

VOLUME VI

1970

11

CONTENTS

Editorial - Profit without Honour 1

Erwin — A reclassification of bombardier beetles and a taxonomic revision of

the North and Middle American species (Carabidae: Brachinida) 4

Book Review 217

Corrigenda 219

Book Review 221

Steiner — Solitary wasps from subarctic North America —

I. Pompilidae from the Northwest Territories and Yukon, Canada 223

Chance — The functional morphology of the mouthparts

of blackfly larvae (Diptera: Simuliidae) 245

Editorial - The Patience of Job 285

Chance — A review of chemical control methods for blackfly

larvae (Diptera: Simuliidae) 287

Thomas — Seasonal occurrence and relative abundance of

Tabanidae (Diptera) in three localities in Alberta 293

Book Review 303

Acknowledgements 304

Corrigenda 304

Editorial - How Many is Too Much? 305

Tawfik and Gooding - Observations on mosquitoes during 1969

control operations at Edmonton, Alberta 307

Kevan and Kevan - Collembola as pollen feeders and flower visitors

with observations from the high Arctic 311

El Moursy - The taxonomy of the Nearctic species of the

genus Byrrhas Linnaeus (Coleoptera: Byrrhidae) 327

Reddy The mode of action of insect repellents I: choice chamber

experiments with the German cockroach Blattella germanica (L.) 339

Reddy - The mode of action of insect repellents II:

electrophysiological studies 353

Announcements 364

Corrigenda 364

INDEX

Abate, 287, 307

Abdelnur, O. M., 245, 282

Abies lasiocarpa, 231

Aedes, 307, 310

communis, 308

excrucians, 308

fitchii , 279, 308

flavescens, 308

hexodontus, 308

increpitus, 308

pionips, 308

punctor, 308

riparius, 308

spencerii, 308

stimulans, 308

vexans, 308

Ageniodeus, 242

Agrell, I., 311,323

Ahrens, A., 209

Alluaud, C., 209

Alnus, 141

Altner, H.,314, 326

Ambrosia trifida, 316

Ammosphex, 239, 242

Anderson, J. R., 245, 280, 282

Andrewes, H. E., 189, 194, 209

Aneshansley, D., 324

(see Eisner, T., 321)

Anisochaeta, 167

Anopheles messae, 279

Anoplius, 242

cylindricus, 238

imbellis, 238

ithaca, 237

marginatus, 239

nigerrimus , 238, 242

relativus, 242

tenebrosus, 238

tenuicornis, 238, 242

ventralis, 242

virginiensis , 242

Anoplochares, 240, 242

Anthyllis fulneraria, 313, 320, 322

Antoine, M., 6, 1 1 , 209

Aploa, 13, 17, 22, 27, 42, 45, 175, 185,

187, 194

nobilis, 16, 18, 21, 23, 26, 30, 185

pictus, 42

Aporinellus taeniatus, 242

Aptinidius, 38

Aptinina 11, 17, 25, 28, 32, 37, 173, 192

Aptinoderus, 12, 24, 41, 174, 185, 187, 192

Aptinomimus, 42,46, 174, 185, 187

Aptinomorphus, 11, 13,20, 34,36, 173, 185,

187, 191

Aptinus, 12, 13, 20, 25, 27, 36, 42, 47, 173,

185, 187, 192, 194

americanus, 55

bombarda, 16, 18, 23, 26

displosor, 38, 173

jan th inipennis 156

Arachnospila , 240, 242

Araneidae, 241

Araneus

cornu tus, 241

patagiatus, 241

Arnason, A. P., 245, 287, 283

aspen, 226, 23 1

Aspidistra, 313

elatior, 313, 321

Aty lotus, 293

duplex, 297

incisuralis, 296

Auclair, J. C., 318, 324

Axelrod, D. I., 189, 194, 196, 205, 209

Badcock, R. M., 245, 280, 282

Ball, G. E., 6, 1 1, 20, 166, 184, 194, 196,

198, 202, 207, 209

Ball, G. B., 217

Barnley, G. R., 287, 290

Barreda, E.. A., 287, 290

Bar-Zeev, M., 339, 352

Basilewsky, P., 6, 11, 14, 28, 31 , 169, 189,

194, 209

Bates, H.W., 191,210

Bates, M., 279, 282

Bees, honey, 318

Beetles,

bombardier, 5-208

water, 12

Bell, R. T., 14, 17, 20, 22, 166, 168, 210

Bellinger, P. E., 314, 324

Bembidiines, 167

Bembidion, 25

benzene, 355, 358

hexachloride, 289

Berck, B., 245, 283, 287, 290

Bervoerts, W., 292 (see Wanson, M.) 287, 289

IV

Be tula, 316

Betulaceae, 316

blackfly larvae, 245

bloodsucking species, 277

cibarium, 262, 266

comparison with non-filtering species,

262

feeding, 266-275, 279

filter feeding, 245, 247-262, 280

glands, 278

head capsules, 245, 275

hypostomium, 262, 266

labio-hypopharyngeal complex, 260-262,

266, 278

labrum, 253, 263-264, 277

mandibles, 253-258, 264, 277-278

maxillae, 258-260, 264-266, 278

morphology, 275-279

mouthpart function, 245-284

mouthpart movements, 267-272

non-bloodsucking species, 277

non-filtering species, 262-266

rearing methods, 246

Blackwelder, R. E., 210

Blatchley, W. S.,6,210

Blattella germanica, 339, 355, 358, 360

Block, B. D., 353, 363

Bodvarsson,H , 311,314,316,324

Boeckh, J., 353,362

Boheman, C. H., 2 10

Boistel, J., 353, 362

Bonelli, F. A., 210

Bourletiella aquatica, 316,317

arvalis, 316, 317

batroches, 316, 317

bilineata, 314, 317

fallonae , 316, 3 17

hortensis, 314, 316, 322

lutea, 312, 317, 320, 322, 323

pallipes, 313, 317, 322

pruinosa, 312, 322

repanda ,311,317, 322

russata, 316, 317

signata, 312, 316, 321,322

spinata, 316, 317

Brachinida, 4-2 1 5

Brachinini, 25, 27, 32, 169, 171

Brachinus aabaaba, 5, 54, 157, 160, 164,

177, 182,. 202, 205

Brachinus (cont.)

adustipennis , 51, 74, 76, 78, 81, 177, 201

aeger, 51,74,76, 177,201

af finis, 134

alexiguus, 5, 50, 55, 177, 179, 201

altemans , 52, 84, 86, 88, 177, 180, 201

americanus, 27, 50, 54, 56, 58, 177, 201

amplipennis, 136

arboreus, 51, 74, 76, 78, 177, 201

atbarae, 136

azureipennis , 53,99, 111, 115, 177,181

ballistarius , 123

bigutticeps, 192

brunneus, 50, 68, 70, 177, 180, 201, 205

cancellatus, 85

capnicus, 5, 50, 56, 58, 60, 177, 179, 199

carinulatus, 85

cephalotes, 121, 127

chalchihuitlicue , 5, 51, 74, 76, 78,177,201

chirriador, 5, 51, 74, 76, 78, 80, 177, 201

cibolensis, 5, 50, 94, 96, 177, 201, 206

cinctipennis, 50, 94, 96, 177, 181, 201

conformis, 54, 118, 177, 181,201

connectus, 42

consanguineus , 54, 111, 1 16, 177, 181,201

convexus, 104

cordicollis, 54, 127, 144, 148, 177, 182,

202, 205

costipennis, 17, 19, 49, 84, 166, 177, 180,

201,205

crepitans, 11, 13, 19

cruciatus, 47

cyanipennis, 41,52, 124, 177, 181,201

cyanochroaticus, 53, 144, 147, 177, 182,

202

cyanopterus, 134

deyrollei, 88

distinguendus , 88

distribution patterns, 196-204

rfryas, 27, 175, 179, 194

elongatulus, 52, 54, 60, 64, 177, 179, 177,

201

explosus, 5,51, 157, 159, 177, 182, 199,

202, 205

favicollis, 52, 133, 137, 140, 177, 181,202

fidelis, 104

fulminatus, 54, 151, 177, 182, 202, 206

fumans, 53, 97, 133, 137, 177, 180, 201,

205, 207

V

Brachinus (cont.)

galactoderus, 5, 51, 124, 126, 131, 177,

181, 201, 206

gebhardis, 51, 124, 126, 131, 177, 181,

201, 206

gilvipes, 71

gracilis , 144

grandis, 51, 68, 70, 72, 177, 201, 205

hirsutus, 52, 92, 96, 177, 180, 201

hyalea, 180

ichabodopsis, 5, 54, 144, 148, 150, 177,

182, 199, 202

immaculicornis , 24, 42

imperialensis , 52, 133, 137, 143, 182,

202, 206

imporcitis, 5, 21, 54, 111, 177, 181, 201,

206

italicus, 42

janthinipennis , 4, 52, 152, 154, 156, 177,

181, 201, 206

kansanus , 52, 83, 86, 177, 180, 201, 205

kavanaughi, 54, 99, 102, 105, 108, 111,

177, 181, 201, 206

lateralis, 51, 73, 76, 78, 177, 180,201,205

lecontei, 104

leptocerus, 144

leucoloma, 73

librator, 88

longipalpis, 17

luzonicus, 192

metf/ws, 54, 124, 126, 129, 177, 181, 201

melanarthrus, 19, 50, 68, 70, 177, 201,

206

mexicanus, 7, 51, 99, 102, 104, 166, 177,

181,201,206

microamericanus , 19, 50, 55, 58, 177,179,

201

minutus, 129

moM/s, 5, 17, 50, 85, 157, 159, 162, 177,

182, 199, 202

neglectus, 53,99, 102, 105, 110, 177, 181,

201, 206

newberryi, 164

niger, 179

nigricans , 179

oaxacensis, 5, 52, 116, 120, 177, 181,201

ovipennis, 52, 1 18, 120, 177, 181, 201

oxygonus, 54, 151, 154, 177, 182, 202,

206

Brachinus (cont.)

pallidus , 12, 51, 55, 62, 92, 106, 125,166

177, 180, 201, 206

patruelis, 53, 1 17, 120, 177, 181, 201

perplexus, 52, 133, 137, 141, 177,182,202

phaeocerus, 16, 18, 21, 23, 53, 99, 111,

115, 166, 177, 181, 201, 206

puberulus, 53, 133, 137, 140, 177,182,202

pumilio, 156

puncticollis, 93

pygmaeus, 14, 46

quadrimaculatus , 39

quadripennis , 53, 99, 102, 130, 177, 181,

201, 207

rejectus, 127

repressus, 164

rhytiderus, 5 1 , 60, 63, 177, 179,201, 206

rugipennis , 54, 84, 86, 91, 177, 180, 201

*z/tez, 19, 50, 67, 177, 180, 201, 205

scotomedes, 17

sejungenius, 99

senegalensis , 34

similis, 123

sonorous , 5, 50, 52, 157, 160, 177, 182,

202, 206

spinipes, 69

stenoderus, 175

stenomus, 151

strennus, 88

stygicornis, 99

sublaevis, 53, 144, 148, 177, 182,202,206

suf flans, 134

tabasconus, 136

tenuicollis, 53, 123, 126, 177, 181, 201

texanus, 52, 54, 60, 61, 64, 179, 180,205

thermarum, 32

tormentarius , 88

tschernikhi, 93, 99

vdox, 144

velutinus, 52, 133, 137, 142, 177, 182,

202, 207

verticalis, 34

viridipennis , 13, 51, 84, 86, 88, 90, 177,

180, 201

viridis, 90

vulcanoides, 53, 151, 154, 177, 182, 202

Br achy nap tius , 42

Brachynicius , 42

Brachynidae, 32

VI

Brachynidius , 44

Brachynillus , 14, 28, 31, 18

extimus, 3 1

varendorffi, 12, 1 4, 3 1

Brachyninae, 32

Brachynini, 32

Brachynitae, 32

Brachynolomus , 17, 27, 42, 185, 192

Brachynomorphus , 41, 74

Brachynus, 42

ueger, 77

azureipennis , 115

brevior, 65

carinulatus, 85

cognatus , 85

consanguineus, 1 1 6

funebris, 41

melanarthrus, 72

pulchelus, 149

rugipennis, 91

69

tibialis, 41

Bracken, G. K., 293, 301

Brassica, 3 1 8

Brauns, A., 3 15, 324

Bretherick, O., 3 1 8, 326

Brittain, W. H., 311,313, 324

Brown, A. W. A., 287 , 290

Brown, I. C., 224, 227, 243

Browne, S. G., 289, 290

Brulle, G. A., 210

Brundin, L., 166, 210

Brunn, B., 214 (see Robbins, C. S.), 1

Biidel, A., 320, 324

Burdick, G. E., 287, 289, 290

Burks, B. D., 243

Burton, G. J., 270, 282

Burtt, E. T., 245, 279, 282

Byrrhidae, nearctic, 327

Byrrhus, 327-338

americanus, 327 , 332, 335

angustulus, 329

brunnescens, 329

callidus, 33 1

canterius, 329

centralis, 331

concolor, 327, 332, 335, 338

consuetus, 332

criddlei, 329

Byrrhus (cont.)

cyclophorus, 321, 335

difficilis, 330

egenus, 33 1

eximus, 327, 331,335,337

explicatus, 331

fasciatus, 327, 330, 334, 337

fulvovestitus , 329

geminatus, 327 , 330, 335

imperitus, 329

key, 328

kirbyi, 327, 331,335, 338

laramiensis, 332

perditus, 331

pettiti, 330

picipes, 331

pilula, 327

rigens, 332

stolidus, 330

torpidus, 330

vafer, 332

wickhami, 332

Campanula, 313, 320

barbata, 313,321

Carabidae, 5, 6, 27, 46, 164, 167

advanced, 168

higher, 167

lower, 167

primitive, 168, 173

Carabus crepe tans, 6, 42

exhalans, 42

fumans, 47, 134

scolopeta, 42

carbaryl, 289

Car ex, 293, 297

Carl, J., 311,320, 324

Carlsson, G., 245, 276, 280, 282

Carmichaelia , 316

Carpenter, G. D. H., 168, 210

Carpenter, S. L., 307, 310

Carrel, J. E., 324 (see Eisner, T.), 321

Caryophyllacene, 315

Casey, T. L., 333

Castelnau (de LaPorte), F. L. N. C., 210

Cerastium alpinum, 320

arcticum, 315, 322

Chance, M. M., 245-284, 287-292

Chapman, R. F., 221

Chaudoir, M. de, 6, 210

Vll

Chaudonneret, J., 247, 253, 267 , 282

Chevrolat, L. A. A., 210

Chironomidae, larvae, 279

Chironomus plumosus, 279

Chlaenius, 14

chlorten, 289

Chordospartium, 316

Christiansen, K., 311, 315, 324

Chrysops, 293

excitans, 296

frigidus, 294, 296

furcatus, 294, 296

mit is, 296

nigripes, 296

nubiapex, 294, 296

Cincindela aequinoctialis, 36

Cnecostolus, 17, 27, 42, 47, 174, 185, 187,

194

Cnephia, 252

dacotensis, 245, 250, 257, 259, 262,273

strenua, 248

cockroach, American, 353, 355, 358, 361

German, 345, 350, 353, 358, 359, 361

(see Blattella germanica )

Cohn, T. J„ 194, 196, 206, 210

Coffea, 314, 322

arabica, 3 1 4

Colinvaux, P. A., 243

Collembola, arctic, 31 1-323

flower visitors, 31 1-323

pollen feeders, 31 1-323

Collins, D. L., 288,290, 291, 292

(see Travis, B. V.), 287

Compositae, 313,316,318

alpine, 312, 320

Coniferae, 3 1 5

Connolly, R., 284 (see Williams, T. R.), 245

Cook, E. F., 247, 255, 258, 264, 282

Coope, G. R., 243

Cope, O. B., 287, 289, 290

Coraboef, E., 353, 362

Corbet, P. S., 288

Cory, E. N., 287, 292

Cosway, C. A., 339,352

Courtois, L., 292 (see Wanson, M.), 287, 289

Crafts, R. R., 291 (see Guttman, D.), 287

Craig, D. A., 222, 247, 260, 282

Crepidogaster, 25, 28, 184, 189

bimaculatus , 29

Crepidogaster (cont.)

caffra, 16, 18, 23, 30

numeratus, 3 1

pusillus, 31

rufescens, 29

Crepidogastrillus, 28, 31, 184

curtulus, 3 1

Crepidogastrini, 6-194

Crepidogastrinus , 11, 14, 28, 31, 184

kochi, 3 1

Crepidogastritae, 28

Crepidolomus* 14, 28, 31, 184, 189

extimus, 14

Crepidonellus, 28, 31, 184

Crepidostoma, 29

Crosskey, M. E., 290 (see Davies, J. B.), 287

Crosskey, R. W., 247, 260, 282, 287, 290

Crozetia crozetensis, 277 , 280

Cruciferae, 318

Cryobius , 166

Csiki, E., 6, 211

Culex, 279

Culiseta alaskaensis, 308

inomata , 279

Cupressaceae, 315

Cymindis, 164

Dalmat, H. T„ 287, 292

Darlington, P. J. Jr., 6, 20, 184, 189, 191,

194,211

Davies, D. M., 245, 280, 282, 284, 288, 290

(see Wood, D. M.), 248

Davies, J. B., 287, 290

Davies, L., 245, 253, 263, 278, 282

DDT, i , 287, 307

effects on crustaceans, 288, 289

fish, 288, 289

insects, 288

mammals, 288

mites, 288

molluscs, 288

worms, 288

Dean, H. J., 290 (see Burdick, G. E.), 287,

289

De Bemardi, F., 315, 324

Debot, L., 245, 282

DeFoliart, G. R„ 245, 276, 284, 292

(see Travis, B. V.), 287

Degeeria nivalis, 3 1 2

Dejean, P. F. M. A., 6, 21 1

Vlll

Dennis, R., 3 14, 321 , 324

Deonier, C. C., 288, 291

Desoria glacialis, 3 1 2

Dethier, V. G., 362, 355, 361, 363

Deuterosminthurus , 313

bicinctus, 312, 313

diatoms, 313

diazinon, 289

Dickie, R. J., 245, 280, 282

diethyl toluamide, 355

dimethyl phthalate, 355

Diptera, 245,287,293,301

Doflein, F., 312, 324

Donaldson, P. E. K., 354, 363

Drepanocyrtus, 314, 317, 322

flavovirens, 314, 317, 322

Drooz, A. T., 288,291

DuChanois, F. R„ 287, 291

Dufour, L., 211

Dumbleton, L. J., 245, 247, 278, 280, 283

Dursban, 289, 307

Dyke, G. R., 243 (see Fuller, W. A.), 225

Eabry, H. S., 287, 291

Ebeling, W. ,36 1,363

Eisner, T„ 22, 21 1, 321, 324

El Moursy, A. A., 327-338, 333

Entomobrya assuta, 3 1 6

clitellaria, 3 1 6

comparata, 311,317, 322

nivalis, 312, 313, 314, 317, 322

spectabilis, 312, 317,322

Entomobryinae, 315

Entomobryoides purpurascens , 3 1 6

Episyron, 241

biguttatus, 241

Oregon , 241

quinquenotatus, 241

Erichson, W. F.,211

Erwin, T. L., 5-215

Espenshade, E. B. Jr., 194, 196, 204, 21 1

ether, 355, 358

Evagetes, 236, 242

bradleyi, 237

crassicornis, 236, 242

hyacinthinus, 237 , 242

parvus , 237, 242

sc udder i , 237

subangulatus, 236

Evans, H. E„ 232, 236, 243

Evans, W. G., 303

Evarthrus, 196, 198, 204

Fabricius, J. C., 21 1

Faegri, K., 319, 324

Fairchild, G. B., 287, 290

Fairmaire, L. , 2 1 1

fenthion, 289

fir, alpine, 23 1

Douglas, 3 1 4

Fisher, von Waldheim, G., 21 1

Folsom, J.W., 31 1,313, 324

Folsomia, 314, 317

fimetaria, 313, 317, 321

Forbes, W. T. M., 20, 211

Ford, E. B., 168, 210

forest, boreal, 223

Forster, J. R., 333

Fortner, G., 245, 25 1 , 266, 276, 280, 283

Fraxinus, 141

Fredeen, F. J. H., 245, 279, 283, 287, 290

(see Arnason, A. P.), 288

Freitag, R., 184, 196, 198, 202, 207, 211

Frempong-Boadu, J., 287, 289, 290

Frisa, C., 290 (see Burdick, G. E.), 287

Frost, S., 283,291

Fuller, W. A., 225, 243

Galerita ruficollis , 167

Galeritini, 167

Garms, R., 289, 292

Garnham, P. C. C., 287, 290

Gehringiini, 167

Gigantodax , 252

Gisin, H., 312, 314, 324

Gjullin,C.M., 287,291

Glaucops, 293

fratellus , 297

Glytotendipes glaucus, 279

Goiny, H., 292 (see McMahon, J. P.), 287

golden-rod, 316

Goodhue, L. D„ 340, 352

Gooding, R. H., 307-310

Goulding, R. L., 288, 291

Graham, A., 296, 205, 21 1

Graham, P.,293,301,307,310

Great Slave Lake, 227-231

Greenslade, P. J. M., 202, 212

Grenier, P., 245, 247, 253, 260, 267, 280, 283

Gryllotalpa africana, 12

Gryllotalpidae, 12

IX

Gunn, D. L., 339, 352

Guttman, D., 287, 289, 291

Gymnopais, 263, 264

Gyorkos, H., 284 (see Wood, D. M.), 248

Habu, A.,6, 11, 192,212

Hacker, H. A., 20, 212

Haematopota, 293

americana, 294, 296

Hagan, K. B., 287, 292

Hagmeier, E. M., 202, 204, 212

Halffter, G., 194, 196, 212

Handschin, E., 31 1, 321, 323, 324

Hanec, W., 293, 301

Harris, T. W., 212

Harada, S., 292 (see Suzuki, T.), 289

Harris, E. J., 290 (see Burdick, G. E.), 290

Haydak, M.H.,318, 324

Heer, 0.,212

Heliconia, 69

Hellumorphoides, 167

Henning, W., 10, 166, 169, 177, 181,212

heptachlor, 289

Hershkovitz, P., 191, 195, 212

Heterosminthurus cornutus , 314

Highton, R. B., 292, (see McMahon, J. P.),

287

Hinton, H. E., 253, 270, 283

Hocking, B., 2, 223, 242, 283, 291, 304,

306, 315, 318, 320, 324

Hodgkin, A. L., 354, 356, 363

Hodgson, B. N., 320, 325

Hodgson, E. S., 353, 362

Hoffmann, C. H„ 288, 291

Hope, F. W., 212

Hopewell, W. W., 290 (see Arnason, A. P.) ,

288

Hora, S. L., 253, 270, 283

Horn, G. H., 333

Howden, H. F., 198, 212

Howland, H. C., 324 (see Eisner, T.), 321

Hubei, D. H., 354, 363

Hubenthal, W., 41, 212

Hugel, M.F.,318, 325

Husman, G. N., 288, 291

Hybomitra, 293

affinis, 294, 296, 299

arpadi, 296, 299

astuta, 296

epistates , 294, 297 , 299

Hybomitra (cont.)

frontalis , 294, 296, 299

illota, 294, 296, 299

itasca , 291, 299

lasiophthalma, 294, 296, 299

liorhina, 296, 299

melanorhina, 296, 299

metabola, 296, 299

nuda, 294, 297, 299

opaca, 296, 299

osburni, 296

rupestris, 296, 299

tetrica, 296, 299

typhus, 296, 299

zonalis, 296, 299

Hydrophilius obtusus, 125

Hydrophilidae, 12

Hydropsyche, 279

Hynes, H. B. N., 283, 288, 291 (see Kershaw,

W. E.), 245, 289, 291 (see Williams, T. R.),

245

Hypogastrura ,315,317

purpurascens , 321

socialis, 315, 317

Illiger, J. C. W., 212

insecticide, 2, 281, 289, 307, 310

Isotobryoides ochracius, 316

Isotoma, 315,317

kosi, 314, 317

kosiana, 314, 315, 317

nigra , 314, 317

notabilis, 316

saltans, 312, 317

viridis , 315, 317, 322

Isotomurus palustris, 313,317

Ito, Y., 292 (see Suzuki, T.), 289

Jacquelin du Val, P. N. C., 212

Jamieson, C. A., 318, 324

Jamnback, H., 287, 290, 292 (see Collins,

D. L.), 289 (see Travis, B.V.), 287

Jeannel, R. G., 6, 11, 14, 17,22, 29,189

Jedlicka, A., 191, 212

Johnson, C. G., 303

Johnson, R. D., 291 (see Kershaw, W.E.),

287, 289

Johnston, M. R. L., 290 (see Davies, J.B.),

287

Jolliffe, A. W., 227, 243

Jones, T. R. E., 245, 278, 280, 283

X

Jorgensen, C. B., 245, 279, 283

Juniperus, 315

pachypholea, 3 1 5

Jurassic, 231

Kaiser, P., 279, 283

Kaissling, K. E., 353, 362

Karlstrom, Thor N. V., 217

Katianna, 314, 317

Keller, C., 313, 325

Kershaw, W. E., 245, 283, 287, 289, 291

(see Williams, T. R.), 245

Kevan, P. G., 311-326

Kevan, D. K. McE., 31 1-326

Khan, A. A., 361,363

Kindler, J. B., 287, 292

King, D. B., 196,212

Kirby, W., 333

Knuth, P.,312, 320, 325

Kodiak, refugium, 217

Kolbe, H. J., 213

Korlan, 289

Krombein, K. V., 237, 239, 243 (see

Muesebeck, C. F. W.), 237

Kos, F., 314, 325

Kracht, A., 279, 284

Krog, D., 320, 325

Kuhnelt, W., 314, 325

Kuiken, K. A., 318, 326

Kuzoe, F. A. S., 287, 292

LaCasse, W. J., 307, 310

Lacher, V., 353, 361, 363

Lacordair, J. T., 213

Lafert^-Senectere, F. T., 213

Lapsus calami , 39, 121, 127, 134

Larix laricina , 226

Larson, D. J., 196, 213

larvicide, 245, 281, 288, 290

blackfly, 288, 290

simuliid, 287

Latreille, P. A., 213

Laycock, A. H., 224, 243

Lea, A. O., 287, 292

Lebia, 164

grandis, 164

Lebied, B., 292 (see Wanson, M.), 287, 289

Lebiini, 164

Lecompte, J., 353, 362

LeConte, J. L., 6, 213, 333

Ledum groenlandicum, 293

Leguminosae, 316

Leng, C. W., 333

Leontodon montanus, 312, 322

taraxaci, 312, 320, 322

Lepidocyrtinus domesticus, 316

Lepidocyrtus , 3 1 4, 3 1 6

albicans, 313

cyaneus, 313,317

lanuginosus, 312, 313, 316, 322

Lepidoptera, 321

Lesquerella, 319

arctica, 311,318

Lewis, D. J., 276, 283

Liebke, M.,6, 191,213

Liliaceae, 313

Lindane, 289

Lindroth, C. H., 20, 213

Linnaeus, C., 333

Linnaniemi (Axelson), W. M., 312, 325

Linne, C. von, 6, 213

Linsley, E. G., 10,213

lipids, 318

Lobopleuri, 167

Lophopompilus , 242

Loricerini, 167

Lubbock, Sir J., 312, 325

Lubliner-Mianowska, K., 318, 325

Lunden, R., 318, 325

Lutshnik, V., 213

Lutz, F. E., 321, 325

Lycosidae, 239

MacGinitie, H. D., 196,213

Maciolek, J. A., 272, 283

Macnamara, C., 311,313, 325

Macronema, 279

Maindron, M., 213

Maitland, P. S., 245, 280, 283

Mani, M. S., 315,325

Mannerheim, C. G. von, 213

Martin, P. S., 196, 204, 207, 213

Mason, W. R. M., 237,241,244

Mastacina, 1 1, 15,28,32, 166, 169, 189, 191

Mastax, 11, 13, 17,20, 25,32, 167, 173, 184

?hargreavesi , 16, 18, 21, 23, 26

Matchett, R. E., 291 (see Kershaw, W. E.),

287

Matsuda, R., 247, 283

Matthews, J., 218

Maynard, E. A., 314, 325

XI

Mayr, E. E., 10, 213

McAlpine, J. F., 293, 301

McDuffie, W. C., 291 (see Hocking, B.), 287

McMahon, J. P., 287, 290, 292

Means, R. G., 287, 289, 291

Medicago, 316, 322

Megalothorax minimum, 316, 317

Mehringer, P. J., 196, 213

Merkel, E. P„ 287, 291

methoxychlor, 287, 289

Metriini, 166

Metrius, 167

Miall, L. C., 279, 283

Mills, M. L., 291 (see Kershaw, W.E.)

mimicry, Mullerian, 174

Moreau, R. E., 187, 189, 194, 196, 213

Morita, H., 353, 363

mosquitoes

control, 307-310

ecology, 307, 310

larvae, 1 , 307, 310

Motschulsky, V. von, 214

Muesebeck, C. F. W., 237, 244

Muirhead-Thompson, R. C., 287, 292

Muller, H., 312, 319, 325

Muscidae, 319

muskeg, 294, 297

Naumann, G., 245, 272, 279, 284

Nematocera, 255

Neobrachinus, 5, 175-183

Noel-Buxton, M. B., 245, 284, 287, 292

Notospartium, 3 1 6

Novosieversia glacialis , 320

Omophranini, 166

onchocerciasis, 287

Onychiurus pseudo funetarius , 315, 317

Opisthiini, 167

Or che sella , 3 1 6

bi fas data, 315, 317

hexafasciata, 316

quinquefasciata, 315

villosa, 315, 317

zebra, 316

orchid, 3 1 2

Orthoptera, 12

Osborn, H., 279, 284

Osten Sacken, R., 279, 284

Ozaenini, 166

Pachy teles, 167

Paclt, J., 314,321, 325

Pantin, C. F. A., 246, 284

Parakatianna , 314, 317

albirubrafrons , 316

diversitata, 316

Parapheropsophus , 34

Parisi, V., 315,324

Parker, G. L., 292 (see Swabey, Y. H.), 289

Paulian, R., 195, 214

Paussidae, 167

Pedicularis, 321

Penny, M. M., 245,280, 283

Peringuey, L. A., 6, 191, 214

Periplaneta, 355, 359

americana , 355, 357

permafrost, 225

Peterson, B. V., 245, 270, 278, 280, 284

(see Wood, D. M.), 248

Petrishcheva, P. A., 289, 292

Pewe, T. L., 244

Phelps, R. J., 245, 276, 279, 284

Pheropsophidius , 5, 1 1, 13, 15, 20, 25, 27,

34,36, 164, 173, 185, 187, 191, 194

aequinoctialis, 49, 157, 160, 165

biplagiatus, 49, 157, 160, 162, 165

rivieri , 16, 18, 23, 26

Pheropsophina , 11, 17, 20, 22, 25, 28, 32,

37, 166, 173, 189, 191

old world, 17

Pheropsophini , 33

Pheropsophus, 12, 20, 22,, 25, 27, 33, 184

187, 191, 195

acute costatus , 34

?bimaculatus , 16, 18, 21, 23, 26, 30

biplagiatus, 36, 165

fasciatocollis , 42

jessoensis, 12

microrrhabdus, 42

Phidippus, 240

Philip, C. B., 293, 301

Picea, 293

glauca, 226, 293

mariana, 226

Pickering, L. R., 266, 283

Pijl, L. van der, 312, 318, 324

pine, jack, 223, 226

lodgepole, 23 1

red, 316

Pinus Banksiana, 226, 227

Xll

Pinus (cont.)

lati folia, 23 1

mughus, 314

Platanus, 87, 141

Platybrachinus , 42, 45

Platypatrobus lacustris, 147

Pogonomyioides segnis, 3 1 9

Polykatianna flammea, 316

polytetrafluorethylene, 340

Pomaderris phylacae folia, 3 1 6

Pompilidae (Northwest Territories and

Yukon), 223, 244

Pompilini, 236, 241

Pompilinus, 238, 242

Pompilus, 239, 241

angularis, 239

apicatus, 240

arctus, 240, 242

eureka, 240

fumipennis, 240, 242

imbecillus, 239

luctuosus, 239-242

michiganensis , 240

occidentalis, 239

scelestus, 240

solomus, 242

Poole, T. B., 314, 325

poplar, balsam, 226, 231

Populus, 141, 293

balsamifera, 226, 293

tremuloides, 226, 293

Porta, A., 214

Post, A., 289, 292

Povolny, D., 192, 195, 214

Prevost, A., 287, 292

Priesner, E., 353, 363

Priocnemis notha, 237

Proisotoma , 315, 317

Prosimulium, 246, 248, 252, 255, 260, 262,

275, 277

fontanantum, 250, 252, 277

frohnei, 250, 252, 255, 277

f us cum, 250, 252, 277

multidentatum, 250, 252, 277

travisi, 246, 250, 252, 255, 259, 261

ursimum , 277

Protopheropsophus, 17, 34, 36, 164, 173,

185, 187, 191

biplagiatus, 17

Pseudaptinus , 42, 46

Pseudo katianna lutea, 316

Pseudomorpha, 167

Pterostichus, 20

Pucat, A. M., 245, 284

Puri, I.M., 245,253, 278, 284

Putzeys, M., 214

Quercus, 87, 165

Quisenberry, B. F., 287, 291

ragweed, 316

Ranunculaceae, alpine, 313

Ranunculus glacialis , 312, 320, 322

Rapoport, E. H., 321, 326

Raybould, J. N., 287, 289, 292

Reddy, M. J., 339-352, 353-363

refugium, Kodiak, 217

Regan, F. R., 287, 292

Reitter, E., 6, 214

Rempel, J. C., 245, 283, 287, 290 (see

Arnason, A. P.), 288

repellents, 339-35 1

mosquito, 339

MGK R-874 (2-hydro xyethyl-n-octyl

sulphide), 340, 342, 353, 355

Rhamnaceae, 316

Rhysodini, 167

Richards, W. R., 287, 291

Riley, C. V., 279, 294

Ripper, W., 321, 326

Ritchie, A. H., 314, 326

Robbins, C. S., 198, 214

Roeder, K. D., 356, 363

Roessler, H. P., 353, 363

Romer, A. S., 195, 214

ronnel, 289

Rosadeae, 320

Ross, H. H., 196,202, 207,214

Rossi, P., 214

Roth, L. M., 339, 352

Rowe, J.S., 225, 227, 231, 244

Roys, C. C., 339, 352, 355, 359, 362, 363

Rubtsov, I. A., 245, 248, 251, 253, 258, 260,

277, 280, 284

Sadanaga, K., 11, 12, 212

Safyanova, V. M., 289, 292

Salicaceae, 320

# Salix, 87, 141,320

Salticidae, 240

Savage, R. J. G., 195,214

Xlll

Savile, D. B. O., 318, 326

Saxifragaceae, 315,318

Saxifraga hir cuius, 315, 322

oppositifolia, 318, 320, 322

Say, T., 214

Schaller, F., 315, 326

Schenk, C. F., 292, (see Swabey, Y. H.), 287

289

Schmidt-Goebel, H. M., 191,214

Schneider, D., 353, 362, 363

Schott, H., 312, 314, 326

Schuchman, S. M., 287, 289, 292

Scirpus-Typha, 82, 150

Scott, H. G., 315, 326

Scrophulariaceae, 321

Scudder, S. H., 214

Sedlag, U., 314, 326

Sekhon, S. S., 361, 363

Shamsuddin, M., 293, 301

Shamurin, V. F., 320, 326 (see Tikhomirov,

B. A.), 320

Sharma, G. D., 315, 326

Sharplin, C. D„ 223, 225, 242, 304,320,324

Shelf ord, V. E., 320, 326

Shepard, H. H., 339, 352

Shtepa, V. S., 326 (see Tikhomirov, B. A.),

320

Silberglied, R. E., 324 (see Eisner, T.), 321

Silfer, E. H., 361, 363

Simpson, G. G., 195, 198, 214

Simuliidae, 245, 287

Simuliidae, 253, 278

Simulium, 260, 275, 277

damnosum, 287

decorum, 245, 250, 255, 257, 260, 262,

273, 279

neavei, 287

oviceps, 277 , 280

pictipes, 248, 252

venustum, 245, 250, 255, 257, 260, 262,

273, 279

verecundum, 246

vittatum, 245, 267, 271, 273, 279

Sinella binoculata, 316

Skea, J., 290 (see Burdick, G. E.), 290

Sleeper, D. A., 288, 291

Smart, J., 245,247, 280, 284

Sminthuridae, 315

Lapland, 314

Sminthurides aquaticus, 3 1 3, 3 1 4, 3 1 7

pumilis, 314, 317

Sminthurinus aureus, 3 1 6

elegans, 316

radiculus, 316

Sminthurus, 313, 317

, dorsalis, 316

facialis, 316

fitchi, 316

hortensis, 313

luteus, 312

obscurus, 316

viridis, 313,316,322

SmithC. D., 282, 246

Smith, L. S., 244

Smith, M.H., 353, 363

Snodgrass, R. E., 27, 215, 247, 284

Solidago, 316

Solier, A. J. J.. 21 5

Sommerman, K. M., 245, 252, 284

Sphagnum, 293, 294, 297

spruce, 223, 293

black, 226, 231

white, 226,231,293,297

Stainer, J. R., 316, 326

Stattler, W., 279, 284

Stebbins, L. L., 243

Steel, R. G. D., 343, 345, 352

Stehr, W. C., 215

Steinbock, O., 313, 326

Steiner, A. L., 223-244

Stenaptinus, 17, 20, 25, 34, 173, 185, 187,

191

?kolbei, 24

krichna, 34

Stephens, J. F., 215

Stevens, C. von, 215

Stojanovitch, C. J., 315, 326

Storm, A., 290 (see Cope, O. B.), 287, 289

Strebel, O., 311,313, 321, 323, 326

Strickland, E. H„ 1, 276, 284, 293, 297, 301

Stults, C. D., 202, 204, 212

Styphlodromus, 14, 25, 28, 37, 40, 173, 185,

187, 192

bicolor, 40

Styphlomerinus , 14, 25, 37, 40, 173, 185,

187, 192

Styphlomerus, 12, 20, 25, 27, 37, 173, 185,

187, 192

XIV

Styphlomerus (cont.)

? cilia t us, 16, 18, 21, 23, 30

fuscifrons, 40

Surber, E. W„ 288, 291

Suzuki, T., 289, 292

Swabey, J. H., 287, 289, 292

Sweeney, C., 290 (see Burdick, G. E.), 287,

289

Syme, P. D., 245, 282

Syrphidae, 319

Tabanidae, 293- 301

Tabanus, 293

frontalis, 301

marginalis , 296

taiga, 223

tamarack, 226

Tanner, V. M., 22, 25,215

Taraxacum, 313, 318, 320, 322

Tarentula, 239

Taufflieb, R., 287, 289, 292

Tawfik, M.S., 307-310

Thomas, A. W., 293-301

Thomisidae, 321

Thomson, C. G., 215

Thorsteinson, A. J., 293, 301

Tikhomirov, B. A., 320, 326

Tiller, R. E., 287, 292

Todd, F. E., 318, 326

toluene, 355, 358

Tomocerus flavescens, 316

minor , 315, 321

vulgaris , 315, 317

Torrie, J. H., 345, 352

Townes, H. K., 244, 293, 301

Trachypachus , 167

traps. Malaise, 293

Manitoba fly, 293, 296

Travis, B. V., 287, 289, 292 (see Collins,

J. C.), 289, (see Guttman, D.), 287

Trechinae, 1 1

Trechini, 12, 15, 17

Trichoptera, 279

Trimen, R., 166, 215

Trollius, 3 1 1

europaeus, 3 1 1

Tsuga , 316

Tukey, J. W.,345,352

Tunzi, M. G., 272, 283

Tuomikoski, R., 166, 215

Twinn, C. R., 291 (see Hocking, B.), 287

Twinnia biclavata, 245, 261, 268, 274, 277

tibblesi, 263

Typha, 293

Typha-Scirpus, 143

Tyronia , 14, 28, 31, 184, 189

caeca, 12, 14

humerata, 22

kivuensis, 22

Usinger, R. L , 10, 213

Vogel, S., 312, 326

Walshe, B. M., 245, 279, 284

Walters, M.C., 315,326

Wanson, M., 287, 289, 292

Warkentin, J., 224, 244

wasps, and climate, 225

ecology, 223-225

fossorial, 223

Great Slave Lake, 225-231

seasonal and daily distribution, 234

solitary, subarctic America, 223-244

spider, 223

zoogeography, 241

Weaver, N., 318, 326

Weber, F., 215

Whitcomb, W„ 319, 326

Whitehead, D. R., 6, 196,215

Wickham, H. F., 7, 12, 215

Wilhemia equina, 277

Williams, T. R., 245, 283, 284, 287, 291

(see Kershaw, W. E.),245

Willis, E. R., 339, 352

willow, 23 1

Wilson, H. F., 319, 326

Wilton, D. P„ 287, 292

Wonders, W. C„ 224, 244

Wood, D. M., 245, 247, 252, 260, 264, 267,

270, 284

Woodford, A. O., 195,215

Wright, R. H., 361, 363

Wu, Y. F., 245, 280, 284

Yamashita, S., 353, 363

Yukon, 226, 231, 243

Zahar, A. R., 245, 284

Zim, H. S., 214

Quaestiones

entomologicae

MUS. COMP. ZOOU

library

FEB 19T/1

harvard

UNIVERSITY

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

VOLUME VI

NUMBER 1

JANUARY 1970

QUAESTIONES ENTOMOLOGICAE

A periodical record of entomological investigations published at the Department of

Entomology, University of Alberta, Edmonton, Alberta.

Volume 6 Number 1 20 February, 1970

CONTENTS

Editorial — Profit without honour 1

Erwin — A reclassification of bombardier beetles and a taxonomic revision of

the North and Middle American species (Carabidae:Brachinida) 4

Book Review 217

Corrigenda 219

Editorial — Profit without Honour

Could the widespread use of DDT be a disaster? This question was the title given by the

late E. H. Strickland to a short paper which he wrote in 1945 (Ent. News 56: 85-88). He

answered this rhetorical question with a qualified yes. His qualifications were two. Firstly,

that DDT proved to approach in effectiveness the claims then made for it; secondly that it

be employed on a widespread scale, over large connected areas. Both requirements have

been met. For once, a commercial product had indeed met the claims made for it, at least

as regards initial effectiveness. And DDT has certainly been used over large connected areas.

Do we have a disaster on our hands? Or have we heeded this 25 year old prophesy in time?

When Strickland wrote, little was known of the persistence of DDT and less of its cumu-

lative build-up in food chains. Yet he correctly foresaw the hazards of threatening the main-

tenance of populations of predators — at any level.

It can be estimated that about 4.5 million tons of DDT have been produced and most of

this has been released into the biosphere. This would represent 0.25 lb. per acre of the 55 Vi

million square miles of the land surface of the earth, or 2.4 lb. per arable acre. And for many

years now DDT has not been alone; other chlorinated hydrocarbons with general toxicity

and varying high persistence have joined it. Although it is now known that much of the DDT

is in the atmosphere, much in the oceans (Frost, 1969, Environment 1 1(6): 14), and much

in the top ends of food chains (Peterle, loc. cit. 34) and although its half-life in the soil may

be rather less than 1 0 years, these figures have serious implications. Good control of mos-

quito larvae could be obtained in many areas with an application of 0.03 lb. of DDT per

acre per annum. It seems inescapable that the insect fauna of the world must by now be so

altered that we shall probably never know in any kind of detail what it was like before

1944. This, to an entomologist, is disaster; it will take decades, if not centuries, for a mea-

sure of normality to return; how long the first breakdown product, DDE, remains as a

hazard in the environment is unknown. None can say how many species, potentially valu-

able, have gone for good. Let us hope we have at least learned the lesson that persistent

broad-spectrum insecticides must be rigidly limited to narrow-spectrum applications.

2

Looking back, one wonders just how this came about. A chemical which can stop a typh-

us epidemic instanter and interrupt malaria transmission over large areas of the world, can-

not be wholly bad. No other insecticide could have done these things so cheaply, nor with

such negligible direct hazard to the sprayers — and the sprayed; 500 million people have

lived for years in houses sprayed thrice yearly with DDT with virtually no records of sick-

ness traceable to this. Factory workers making it, day in day out for 20 years, are no less

healthy than other people (Laws, Curley, and Biros 1967, Arch, environ. Health 15: 766).

How much then, of the blame for the disaster rests with the users? How much with the

manufacturers and their salesmen? Profits have been substantial from DDT (Bean, 1963,

Ag. Chem. 18(1): 50-51, 118-119) and the profit motive has been evident in its abuse. The

view that if one pound is good two pounds are twice as good is easily exploited. There is no

honour in this. The marketing of insecticides should require the provision of details of pro-

cedures to detoxify them, in addition to the data on the hazards of their use. None can

know for sure when this may be needed nor how urgently. If the environment must be done

such damage, to keep the economy going, perhaps we have a sick economy and should cure

this first. We are not yet so close to taking over a new planet that we can afford to ignore

what we are doing to our present one.

There is indeed little honour to be gleaned in this situation; perhaps only for those who

discovered the remarkable properties of this material, plus at best a handful of far-sighted

entomologists who warned us early. Did we fail to listen because we have a generation gap

at both ends?

Brian Hocking

Brachinus janthinipennis (Dejean), male, Cornwall, Connecticut.

A RECLASSIFICATION OF BOMBARDIER BEETLES

AND A TAXONOMIC REVISION OF THE NORTH AND MIDDLE AMERICAN

SPECIES (CARABIDAE: BRACHINIDA)

TERRY LEE ERWIN

Museum of Comparative Zoology

Harvard University

Cambridge, Massachusetts Quaestiones entomologicae

U.S.A. 02138 6:4-215 1970

A taxonomic revision of the species of the North and middle American Brachinus Weber

is presented. Sixty-two species are recognized as valid. Neobrachinus new subgenus, is

erected to contain the New World species formerly included in Brachinus (sensu s trie to).

The following 14 taxa are described as new: Brachinus alexiguus, B. capnicus, B. chalchi-

huitlicue, B. chirriador, B. cibolensis, B. javalinopsis, B. imporcitis, B. oaxacensis, B. galac-

toderus, B. ichabodopsis, B. mobilis, B. explosus, B. aabaaba, B. sonorous. Twenty-five

names are reduced to synonymy for the first time.

A key to the species is given. Each species and species group is described and synonymies

are listed. The distribution of each species is presented by locality records and distribution

maps. Structures used in identification are illustrated.

Two species of Pheropsophidius Hubenthal occur in Middle America and are included in

the key to bombardier beetles of North and Middle America, one of these species is des-

cribed. Structures of both species are illustrated.

A reclassification of the genera of the world bombardier beetles is presented. Seventeen

genera are recognized as valid. These include 13 valid subgenera, one of which is new. Two

tribes are included in the division Brachinida. One of these tribes includes four subtribes,

two of which are new. Three generic names are reduced to synonymy for the first time. All

taxa are described and keys for their identification are presented.

A discussion of the general morphology of members of the genera is presented. Structures

discussed are illustrated.

A hypothetical phytogeny is presented for all genera of the division Brachinida and all

species of Neobrachinus new subgenus. The geographical distribution of these taxa is also

discussed. Figures and tables accompany the discussions.

CONTENTS

Materials and methods p. 6

Comparative morphology p. 11

Taxonomy p. 27

Life histories and immature stages p. 166

Phylogeny p. 166

Zoogeography p. 184

References p. 209

6

Erwin

The bombardier beetles are a complex and taxonomically difficult group belonging to the

family Carabidae. In 1862, LeConte stated: “1 must also add that I consider the species of

this genus very decidedly opinionative, and that I am only impelled to the publication of

this note by the necessity of giving names to a certain number of recognized forms, and of

placing as synonyms some which I formerly considered as distinct, but which increased col-

lections have since shewn to be varieties.” The situation had hardly changed 98 years later

when Ball (1960: 164) wrote: “The taxonomy of the North American species of this group

is very poorly understood and it is almost a waste of time at present to attempt to determine

individuals to species.”

Until now, a taxonomic revision of all North and Middle American species of Brachinus

has not been undertaken, probably because of the great morphological similarity among the

species, a lack of traditional characteristics for separating the species, and the unavailability

of much of the “type” material. The first of these obstacles was surmounted after the dis-

covery of external characteristics not used previously. The last problem was resolved with

the help of G. E. Ball who compared my specimens with the type material of Dejean and

Chaudoir in Paris. I was able to compare the LeConte and Blatchley types myself at the

Museum of Comparative Zoology and Purdue University, respectively.

This paper is an extension of my 1 965 publication and deals with the taxonomy and dis-

tribution of Brachinus and Pheropsophidius in North and Middle America. I also propose

here a reclassification of bombardier beetles of the world, including subgeneric components;

discuss phylogenetic and zoogeographic hypotheses; and present a preliminary study of the

comparative morphology of bombardier beetles.

Bombardier beetles were first recognized formally when Linne described the species

Carabus crepitans in 1758. Since that time, more than 600 trivial names have been proposed

for members of the Brachinida. Dejean (1825, 1831), LeConte (1844, 1848, 1858, 1862),

and Chaudoir (1868, 1876) have described most of the species in North and Middle America.

Chaudoir’s paper (1876) has been the only monographic treatment of the Brachinida as a

whole. Basilewsky’s 1959 revision of the Crepidogastrini must be considered monographic

of that tribe. LeConte’s 1862 paper and my revision of the California species (1965) contain

the only new keys to North American species. (Blatchley’s 1910 monumental “Coleoptera

of Indiana” contains a key to the Brachinus species of Indiana, but it was based on LeConte’s

key.)

Many faunal studies have included partial taxonomic treatment of some of the taxa of the

Brachinida. Notable are publications by Antoine (1962), Basilewsky (1962), Darlington

(1968), Habu (1967), Jeannel (1942, 1949), Liebke (1934), Peringuey (1885, 1888, 1896,

1898), and Reitter (1919). No complete taxonomic classification has been proposed for the

group as a whole. Most workers still seem to rely more heavily on the catalogue classification

established by Csiki (1933) than on any other reference source.

MATERIALS AND METHODS

Materials

This study is the result of the examination of 28,633 specimens of North and Middle

American Brachinus, and 2,172 specimens of other bombardier beetles. Most of these speci-

mens have been loaned to me by museums and private collectors in Canada, Europe, Mexico,

and the United States. The others were collected by my wife and me in the United States.

Were it not for the intensive collecting of carabids in Mexico by G. E. Ball and D. R.

Whitehead, this area would have been omitted from this study due to insufficent material.

Bombardier Beetles

7

Larval material was especially difficult to obtain. In addition to material from California

(Erwin, 1967), I have seen one first instar larva of Brachinus mexicanus Dejean, and one

unidentified last instar larva. This is still all that is known in North America, besides that

which H. F. Wickham collected in the 1890’s.

The following abbreviations indicate the various museums and private collections from

which specimens were borrowed:

AMNH American Museum of Natural History, New York, New York 10024, P. Vaurie,

L. Herman Jr.

ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania 19103, H. R. Roberts.

AUAA Auburn University, Auburn, Alabama 36830, K. L. Hays.

BMNH British Museum (Natural History), London, England, R. D. Pope.

CArm C. Armin, 191 West Palm Avenue, Reedley, California 93654.

CAS California Academy of Sciences, San Francisco, California 94118, H. B. Leech.

CBak C. Baker, Boise State College, 1 907 Campus Drive, Boise, Idaho 83707.

CCha C. Chantal, 425 Saint Olivier, Quebec 4, Quebec.

CEWh C. E. White, 2441 East Northview Avenue, Indianapolis, Indiana 46220.

CMPP Carnegie Museum, Pittsburgh, Pennsylvania 15213, G. E. Wallace.

CNC Canadian National Collection of Insects, Entomology Research Institute, Ottawa,

Ontario, E. C. Becker, W. J. Brown.

CNHM Chicago Natural History Museum, Chicago, Illinois 60605, H. Dybas.

CPBo C. Bolivar y Pieltair., Instituto Politecnico Nacional, Mexico D. F.

CUNY Cornell University, Ithaca, New York, 14850, L. L. Pechuman.

CVMA Coachella Valley Mosquito Abatement District Collection, Thermal, California

92274.

DDLa D. J. Larson, University of Calgary, Calgary, Alberta.

DHKa D. H. Kavanaugh, 1121 Garfield Street, Denver, Colorado 80206.

DRWh D. R. Whitehead, University of Alberta, Edmonton 7, Alberta.

DTRT Division of Tropical Research, Tela Railroad Company, La Lima, Honduras,

Central America, W. G. C. Forsyth.

FDAG Florida Department of Agriculture, Gainesville, Florida 32601, R. E. Woodruff.

GRNo G. R. Noonan, University of California, Riverside, California 92502 (including col-

lection of F. Andrews).

HGou H. Goulet, University of Alberta, Edmonton 7, Alberta.

HMO Hope Museum, Oxford, England, E. Taylor.

ISNH Illinois State Natural History Survey, Urbana, Illinois 61803, L. K. Gloyd.

ISUA Iowa State University, Ames, Iowa 50010, J. Laffoon.

JHen J. Hendrichs, Apartado Posta, 1 1-774, Mexico 1 1 , D. F.

JSch J. Schuh, 4039 Shasta Way, Klamath Falls, Oregon 97601 .

KMTB Koninklijk Museum Voor Midden-Afrika-Musee Royal del’ Afrique Centrale,

Tervuren, Belgium, P. Basilewsky.

KSU Kansas State University, Manhattan, Kansas 66502, H. D. Blocker.

LACM Los Angeles County Museum, Exposition Park, Los Angeles, California 90007,

C. L. Hogue.

LBSC Long Beach State College, Long Beach, California 90804, E. L. Sleeper.

LRus L. Russell, Oregon State College, Corvallis, Oregon, 97331.

LSUB Louisiana State University, Baton Rouge, Louisiana 70803, J. B. Chapin.

MSUM Montana State University, Missoula, Montana, 59801, N. Anderson.

MCZ Museum of Comparative Zoology, Cambridge, Massachusetts 02138, P. J.

Darlington, Jr.

8

Erwin

MHNP Museum National d’Histoire Naturelle, Paris, A. Bons.

MMM Moscow Museum, Moscow, U.S.S.R.

NSDA Nevada State Department of Agriculture, Reno, Nevada, 89504, R. C. Bechtel.

OSUC Oregon State University, Corvallis, Oregon 97331, P. O. Ritcher.

OSUS Oklahoma State University, Stillwater, Oklahoma 74075, W. A. Drew.

OUCO Ohio State University, Columbus, Ohio 43210, C. A. Triplehorn.

PSUU Pennsylvania State University, University Park, Pennsylvania 16802, W. W. Boyle,

S. W. Frost.

PUM Purdue University, Lafayette, Indiana 47907, R. H. Arnett.

RCGr R. C. Graves, Bowling Green State University, Bowling Green, Ohio 43402

(including collection of W. Suter).

RESt R. E. Stecker, San Jose State College, San Jose, California 95114.

RFre R. Freitag, Lakehead University, Thunder Bay, Ontario.

ROM The Royal Ontario Museum, University of Toronto, Toronto 5, Ontario, G. B.

Wiggins.

RTBe R. T. Bell, University of Vermont, Burlington, Vermont 05401.

SDSNE1 San Diego Society of Natural History, San Diego, California 92112, C. F. Harbison.

SDSU South Dakota State University, Brookings, South Dakota 57006, E. U. Balsbaugh,

Jr.

SJSC San Jose State College, San Jose, California 95114, J. G. Edwards.

TAMU Texas A & M University, College Station, Texas 77840, H. R. Burke.

TCBa T. C. Barr, Jr., University of Kentucky, Lexington, Kentucky 40506.

TFH1 T. F. Hlavac, Museum of Comparative Zoology, Harvard University, Cambridge,

Massachusetts 02138.

TLEr T. L. Erwin, Museum of Comparative Zoology, Harvard University, Cambridge,

Massachusetts 02138.

TMBH Termeszettudomanyi Muzeum, Musee Hongrois d’Histoire Naturelle, Budapest,

VIII., Baross-u. 13 (Hongrie), Z. Kaszab.

UAFA University of Arkansas, Fayetteville, Arkansas 72701, R. T. Allen, E. P. Rouse.

UASM University of Alberta, Strickland Museum, Edmonton 7, Alberta, G. E. Ball.

UATA University of Arizona, Tucson, Arizona 85721, F. G. Werner.

UBC University of British Columbia, Vancouver, British Columbia.

UCD University of California, Davis, California 95616, R. O. Schuster.

UCR University of California, Riverside, California 92502, S. Frommer.

UIMI University of Idaho, Moscow, Idaho 83843, W. F. Barr.

ULLK University of Louisville, Louisville, Kentucky 40208, C. V. Coveil.

UMAH University of Michigan, Ann Arbor, Michigan 48104, R. D. Alexander.

UMCP University of Maryland, College Park, Maryland 20742, F. E. Wood.

UMSP University of Minnesota, Saint Paul, Minnesota 55101, P. Clausen.

UNCR University of North Carolina, Raleigh, North Carolina 27607, D. A. Young.

UNLN University of Nebraska College of Agriculture, Lincoln, Nebraska 68508, W. T.

Atyeo.

UNSS University of Saskatchewan, Saskatoon, Saskatchewan, N. Church.

UONO University of Oklahoma, Norman, Oklahoma 73069, C. E. Hopla.

USNM Smithsonian Institution, Washington, D. C. 20560, O. L. Cartwright.

USUL Utah State University ,Logan, Utah 84321, W. J. Hanson.

UWLW University of Wyoming, Laramie, Wyoming 82070, N. L. Marston.

UWMW University of Wisconsin, Madison, Wisconsin 53706, R. D. Shenefelt.

UWSW University of Washington, Seattle, Washington 98105, M. H. Hatch.

Bombardier Beetles

9

VMKi V. M. Kirk, North Grain Insects Research Laboratories, Brookings, South Dakota

57006.

VVBa V. V. Baicher, San Jose State College, San Jose, California 95114.

WHTy W. H. Tyson, 823 Cashew Way, Fremont, California 94536.

WSUP Washington State University, Pullman, Washington 99163, M. T. James.

ZMLS Zoological Institute, University of Lund, Lund, Sweden, C. H. Lindroth.

Localities of specimens were determined from labels pinned with the specimens. These

localities are given for each species alphabetically by country, state or province, and countv.

More exact localities are given alphabetically by towns or cities in parentheses behind the

state or county. Behind the parentheses are listed the museums in which the particular

specimens are located.

Methods

Dissecting techniques

Techniques described elsewhere (Erwin, 1965) were used. However, instead of the dis-

sected male genitalia being stored in vials of 70% alcohol they were glued to cards, which

were subsequently pinned beneath the specimens. For storing the female genitalia and

various sclerites (described under comparative morphology) the same procedure was fol-

lowed. The specimens that were completely disarticulated were stored in vials of 70% ethyl

alcohol.

Measurements

Total length measurements were made of representatives of all species. The purpose of

this measurement is to give a general impression of the size range of a species. The great

variation in size of adults within any single species of these beetles is due to differences in

amount of their larval food (Erwin, 1967). The lengths of adult specimens given below were

all obtained in the following manner. From the material at hand, I visually selected both the

largest and the smallest specimen, and measured the head from the anterior edge of the

labrum to the center of the occipital ridge, the pronotum from the anterior margin to the

posterior margin along the center line, and the left elytron from the apex of the scutellum

to the elytral apex, along the suture. These three measurements were added together, pro-

viding a figure that is not affected by expansion and contraction of the membraneous parts

in different killing agents, or by the swelling of the abdomen in gravid females and engorged

beetles. A micrometer eyepiece in a Leitz stereoscopic microscope at a magnification of 50

diameters was used for these measurements. The scale interval represented 0.025 mm.

Illustrations

All drawings were prepared with the aid of a camera lucida on a stereoscopic microscope.

After the outline was drawn with the camera lucida, the specimen was examined under

higher magnification and finer details and shading were added to the drawings. Accompany-

ing scale lines equal 1 .0 mm.

Procedural methods

The 28,633 specimens of North and Middle American bombardier beetles were first

categorized according to smaller geographic areas, for example, southern United States,

northeastern United States, Mexico, Great Plains, etc. This reduced the amount of material

to be concerned with at any one time. The specimens of each geographical area were divided

into population samples on the basis of the label data. Each specimen from a given area was

compared with every other specimen from that area, and comparable specimens were

10

Erwin

grouped into “sets”. Each set was then intensively studied, and dissections of the male

genitalia were made. Those sets with identical genitalia, and at least one concordant external

characteristic in common, were grouped together tentatively until all the material from the

entire area had been analyzed. Only the male genitalia were arbitrarily weighed, all other

characteristics that were studied were considered of equal value. When specimens were

judged to be different, on the basis of their genitalia and at least one external characteristic,

an attempt was made to associate females from the same population sample. No difficulty

was encountered in determining probable male-female associations. Further details concern-

ing the relationships between individuals from different species populations are discussed

under Phylogeny.

Criteria for Species, Subspecies, and supra-specific Taxa

A species may be defined as “a naturally occurring population (or aggregate of popul-

ations) that is reproductively isolated and genetically distinctive” (J. Gordon Edwards, per.

comm.). When the biology of a natural population is fairly well known and the criteria of

this definition have been met, this definition may be applied with reasonable success in the

recognition of species. In the present study, only genitalic and external morphology (includ-

ing pubescence), color, distribution, and some life history data are used, because these are

the only things known about most of the populations treated. When only museum specimens

are utilized, different criteria are necessary, to supplant those given in the definition above,

yet give approximately the same results in recognizing species. As working criteria for these

cases, I use the following: A species is represented by the sum total of the specimens dis-

playing a multidimensional continuum of characteristics, but ultimately delimited from

other such species by gross morphological discontinuities. In this case the characteristics

are external morphology (including pubescence), genitalic morphology, and color. The

determination of the size and nature of the multidimensional discontinuity or “gap” (Mayr,

Linsley,and Usinger, 1953) has traditionally been left to competent taxonomists with exten-

sive experience in the group in question. In most cases, these taxonomists have failed to

state what their criteria were when recognizing these gaps. My criteria for recognizing this

“gap” are the following: two or more similar forms which are naturally sympatric or allopat-

ric are considered separate species, if they differ in genitalic morphology and at least one ex-

ternal characteristic; allopatric forms are considered conspecific if their genitalic morphology

is identical and they show intergradation of external characteristics in geographically proxi-

mal areas (or if no external morphological differences are apparent). Parapatric populations

or aggregates of populations (that is, populations which meet only in a very restricted area,

such as a single valley or single river system) are considered distinct species if there is no

introgression of characteristics near the zones of contact. If there is introgression there, the

forms are considered either to be conspecific or to be two hybridizing species, depending

upon the nature of the introgression.

Naming of subspecies has been avoided, because I feel that more criteria than morpho-

logical characters of museum specimens is necessary for the recognition of limits of taxa

below the species level. Subspecies are geographically or temporally delimited populations

within a species that differ from other such populations (but which are capable of inter-

breeding with those other subspecies). Without extensive information on population dyn-

amics, life histories, and genetic capabilities, subspecies would be ill-defined, and ambiguous

trivial names might be introduced into the literature if such taxa were included in the present

study.

Supra-specific taxa are groups which are established to include all the taxa of the next

lowest rank that demonstrate monophyly. Hennig (1966) argues that all such supra-specific

taxa which demonstrate sister relationships be given equal absolute rank, but the practicality

Bombardier Beetles

11

of this has been questioned (Ball and Erwin, 1969). Many supra-specific taxa presently in

use are paraphyletic. In this paper I have realigned the taxa of the Brachinida in accord with

Hennig’s principles, but I have ranked taxa below the generic level with practicality in mind.

COMPARATIVE MORPHOLOGY

Introduction

Upon discovering the asymmetric anterior tarsal pads in male members of Brachinus

crepitans (Linne), Antoine (1962) stated “il est incomprehensible qu’on ne Tail pas encore

signale ...cet interessant caract^re saute aux yeux quand on examine les tarses par leur face

inferieure.” (It is incomprehensible that no one has yet seen this interesting character which

is so obvious when one examines the ventral surface of the tarsi). This may be said for many

other characteristics of the external morphology. These beetles have been studied by num-

erous European biologists for 200 years or more; even so the taxonomy of the group (and

many other carabid groups) is still in a state of confusion. There can be only one reason.

Those workers viewed the beetles from only the dorsal aspect, because their specimens were

usually glued to cards in that position. I doubt that many of them ever examined the ventral

side of a bombardier beetle. Numerous excellent characteristics have been overlooked by

those using this method of mounting carabid beetles. Had Jeannel seen the aberrant charact-

eristics of Mastax members, he surely would have placed this genus in a family of its own.

In this section I have attempted a beginning of a comprehensive comparative study of

bombardier beetle morphology. The material I have seen that represents species of Old

World genera is not extensive. I have not seen any members of species comprising three of

the monotypic genera, but I believe that I have seen enough material to reach some valid

conclusions. Based on these conclusions, I have proposed certain hypotheses which must be

proved or disproved in subsequent studies. Some of these studies I hope to do myself, yet I

also hope this presentation will provide a foundation for others who are interested in study-

ing the evolution and biology of bombardier beetles.

By presenting my findings on the comparative morphology of bombardier beetles as

Jeannel (1926) did for the Trechinae, I have established a foundation for phylogenetic con-

siderations, using the principles proposed by Hennig ( 1 966). In this section on “comparative

morphology,” I have tried to integrate the descriptive Taxonomy with my interpretations of

the character states.

General characteristics

Size

The bombardier beetles range in size from the very small members of some Mastax and

Crepidogastrinus species (2.0 and 3.0 mm overall length respectively), to the very large

members of some Aptinomorphus species (30.0 mm).

The members of the genus Mastax are all very small, while beetles of the genus Phero-

psophus are large to very large. The remaining genera usually have medium-sized members,

but some kinds may vary from small to large (sometimes even intraspecifically). The Crep-

idogastrini range in size from 3.0 to 16.0 mm, according to Basilewsky (1959). I have seen

specimens of Pheropsophina ranging from 10.0 to 30.0 mm; specimens of Brachinina rang-

ing from 3.0 to 18.2 mm; specimens of Aptinina ranging from 4.5 to 15.0 mm; and speci-

mens of Mastacina ranging from 2.0 to 3.5 mm.

There is considerable variation in size among the members of single species, particularly

in the genera Pheropsophus and Brachinus. I believe that variation in size is due to the ecto-

parasitoid mode of life described for the larvae of these groups (Habu and Sandanaga, 1965;

12

Erwin

Habu, 1967; Wickham, 1893, 1894; Erwin, 1967). I have shown (Erwin, 1967) that larvae

of Brachinus pallidus in California are not host specific, but may attack pupae of at least

two, and probably three, species of hydrophilid water beetles. Since one host is eaten by one

parasitoid larva, the size of the host (larval food available) determines the size of the resul-

tant adult Brachinus pallidus. Habu and Sadanaga (1965) and Habu (1967) have shown that

the larva of Pheropsophus jessoensis Morawitz is an ectoparasitoid on the egg clutches of

Gryllotalpa africana Palisot de Beauvois (Orthoptera, Gryllotalpidae). Those egg clutches

vary in size (number of eggs), and since the larva attacks only one egg clutch the size of the

clutch determines the size of the adult beetle. After the larva begins to undergo hyper-

metamorphosis, it cannot leave the chamber it invaded because of the decrease in its leg

length and the increase in its body size (Erwin, 1967).

It has been repeatedly observed that the proportions of the individual adults remain

constant, regardless of their total size.

General form

The bombardier beetles are a very homogeneous group. The narrow head and prothorax,

together with the wide, truncate elytra and apically protruding abdomen, characterize all

species of the Brachinida. Jeannel (1926) established names for four general body forms

within the tribe Trechini. Three of these types are found in Brachinida, but they are slightly

different from the forms described by Jeannel:

1. Type aile (long-winged type) — pigmented, large eyes, small pronotum, square

elytra with prominent humeri, and large metasternum (includes wing dimorphic

species);

2. Type aptere (short-winged type) - pigmented, small or large eyes, enlarged pro-

notum, elytra short with narrow humeri, and short metasternum;

3. Type anophthalme (eyeless type) — depigmented, blind, enlarged pronotum,

elytra short with narrow humeri, and short metasternum.

The members of Mastacina fit Jeannel’s type aile. In these beetles, the abdomen is not

extended as in the crepidogastrines, but they do have the moniliform antennae of that group.

The members of Pherosophina, Aptinina, and Brachinina all have a similar habitus with

filiform antennae, but in members of Pheropsophidus, Pheropsophus, and Aptinus, there is a

tendency for the abdomen to extend far beyond the apex of the elytra. These subtribes

have some members which correspond to Jeannel’s type aile and others that are type aptere.

The members of the Crepidogastrini are the most primitive of the division Brachinida,

and are slightly different in habitus from other bombardier beetles. These beetles generally

have very short elytra, exposing at least two full abdominal terga, even when the abdomen is

not engorged or gravid. Most members belong to Jeannel’s type aptere, but those of

Tyronia caeca Basilewsky and Brachynillus varendorffi Reitter belong to type anophthalme .

Also, the antennae of the crepidogastrines are moniliform rather than filiform.

Microsculpture

In general the microsculpture of bombardier beetles ranges from regularly to irregularly

isodiametric. Slight variation occurs from this basic pattern.

Throughout the Brachinina, Mastacina, and Crepidogastrini, the isodiametric meshes may

vary into a granulate condition, in which each individual mesh is a convex bump. In members

of Pheropsophus , the meshes are extremely fine and barely impressed, and there is a tend-

ency for them to be stretched and arranged into transverse rows. In Aptinus, the meshes are

also barely impressed, but they are larger than in Pheropsophus and arranged in transverse

rows particularly on the pronotum. Aptinoderus members have the meshes of the head al-

most effaced. Members of Styphlomerus have granulate isodiametric meshes arranged on the

Bombardier Beetles

13

elytra as in members of Mastax, and producing a “satin” appearance.

Pubescence

All bombardier beetles maintain at least some body pubescence in addition to the erect

“depression setae” borne by the members of most species, on the elytra, and other variously

located setae. The members of Crepidogastrini are totally pubescent, while other bombardier

beetles may be totally pubescent or have extensive glabrous areas.

The Mastacina members have glabrous elytra, with the pronotum either glabrous or not.

The elytra have erect setae in depressions 2, 4, 6, and 8. The epipleura are pubescent. The

apical edges of the elytra are devoid of setae.

The members of Pheropsophina have the elytra almost glabrous, but with erect “depres-

sion setae” in all depressions, plus scattered pubescence near the scutellum and along de-

pressions 1 and 8. However, the epipleura are glabrous. The apical edge of the elytra ranges

from smooth in members of Aptinomorphus to densely pubescent in members of Phero-

psophus (sensu stricto). Some members of all the other Pheropsophina subgenera also have

this apical fringe.

Among the Aptinina, the members of Aptinus have elytral pubescence arranged much

like that of members of Pheropsophus, but it is usually more dense. Members of Aptinus

also have the epipleura pubescent. In Styphlomerus and its allies, the elytra and epipleura

are totally pubescent, with an additional apical fringe present (closely spaced setae along the

apex of the elytra).

Among the members of Brachinina, all combinations of elytral pubescence exist, and the

epipleura are always pubescent. These patterns of elytral pubescence are described in detail

below (p.49). The apical fringe is absent from Brachinus subgenera Metabrachinus and

Aploa, but is dense in members of Brachinus viridipennis Dejean, and has long widely spaced

setae in members of Brachinus crepitans (Linne).

The amount of head and pronotum pubescence varies considerably among the species of

Brachinida. The members of Mastax are densely pubescent to glabrous; those of Phero-

psophus, Pheropsophidius, and Aptinus are sparsely setiferous; those of Brachinus are gla-

brous to densely pubescent; and Styphlomerus and its allies are densely pubescent.

The pubescence of the cephalic appendages is quite variable throughout the group. All

members of the Crepidogastrini have dense pubescence on every appendage from base to

the apex. The members of Aptinina, Brachinina, and Mastacina have dense pubescence on

the outer articles of the appendages. This pubescence gradually increases in density from the

base to the apex. The members of Pheropsophina are the least pubescent of the group, with

only stiff spines or setae basally,and sparse pubescence distally. In these beetles and those of

the preceding three subtribes, the antennal pubescence is fairly dense on article three, and

very dense on articles four to eleven. The mandibular scrobes of the members of Crepido-

gastrini and Brachinina are plurisetose, while those of the other groups are unisetose. Be-

sides the short pubescence, these two groups of beetles also have the single large seta in the

scrobe.

The venter of the prothorax is densely pubescent medially in members of Crepidogas-

trini, and is less densely pubescent toward the proepipleura. In the other groups, the pro-

sternum usually has numerous long and scattered setae, with the proepisterna, proepimera,

and proepipleura variably setiferous. The venter of the mesothorax, metathorax, and ab-

domen are pubescent in all groups.

The legs of all groups, except the members of Pheropsophina, are pubescent. On these

beetles the setae are scattered and spine-like, forming rows on members of some species.

Major setae occur in numerous places on bombardier beetles. Members of all groups have

a single long seta in the mandibular scrobe, and a single pair of supraorbital setae. In Mastax,

14

Erwin

however, a second “pseudosupraorbital” pair of setae is behind the eyes, one on either side

of the vertex. With the exception of some crepidogastrines, all groups have one pair of

lateral pronotal setae. The exceptions are members of Tyronia, Crepidolomus, and Crepido-

gastrillus which have a second pair of lateral setae in front of the hind angles. In Crepidogas-

trinus, a series of spine-like setae occurs along the lateral margins of the pronotum from

base to apex.

The major setae of the elytra are in the bottom of depressions between costae, in interval

8, and along depression 1 near the scutellum. Interval 8 and depression 1 have umbilicate

punctures with setae of variable length, depending upon the species. The umbilicate series in

depression 8 is continuous in all groups, except in members of Crepidolomus extimus (Jean-

nel). Basilewsky (1959) indicates this species has members with the umbilicate series divided

into 8 setae anteriorly and 8 posteriorly.

The middle and hind coxae, hind trochanter, and the abdominal sterna 2-5 bear “ambul-

atory setae.” The number and location of these long and widely spaced setae varies with the

species. The setae of the tarsi are discussed in detail below (p. 22).

Head

Cranium

The general shape of the cranium is the same for all groups. In the few species with re-

duced eyes, and in the blind members of Brachynillus, the head is much narrower.

Eyes

Only one species, Brachynillus varendorffi Reitter, is known to have all members eyeless.

Another species, Tyronia caeca Basilewsky, has all its members with highly reduced eyes

with only a few facets, and according to Jeannel (1926), beetles with this type of eye are

blind. The eyes of members of Brachinulus viettei Basilewsky and Brachinus pygmaeus

Dejean are very small, and hardly protrude beyond the sides of the head. These species must

be considered as somewhere between Jeannel’s type aptere and type anophthalme, since they

are “depigmented” in comparison to the normal bright colors of the other bombardier

beetles. All other brachinines have fully functional eyes, although some Brachinoaptinus

members have only a small number of facets.

Antennae

In general, the antennae of members of Brachinina are filiform, while those of Crepidogas-

trini are moniliform. Some groups of the Brachinina, however, have antennae with shortened

articles, as the members of Mastax, Styphlodromus, Styphlomerinus, and Styphlomerus.

In these groups, articles 5-1 1 are almost square, slightly longer than wide, or moniliform. All

groups have a rather robust scape, a very short pedicel, and articles 3-1 1 are subequal. In

members of Neobrachinus, Aptinomorphus, and some Asian and African Brachinina, the

third article is elongate. The length of that third article may prove to be useful in the identi-

fication of Brachinus subgenera (defined on the basis of the internal sac of the male geni-

talia), when the Old World fauna is better known. Bell (1960) used this antennal character

successfully in the genus Chlaenius to distinguish taxa.

Labrum

The labrum of all groups is essentially rectangular and slightly emarginate anteriorly. Six

or eight setae occur along the anterior margin. There are six setae in members of Brachinina,

Pheropsophina, and some Aptinina, and eight setae in Mastax, Crepidogastrini, and some

Aptinus.

Bombardier Beetles

15

Mandibles

There is great divergence in the characteristics of the mandibles of the Brachinida mem-

bers. One group, the Mastacina, has members with highly aberrant mandibles (fig. 6). These

mandibles are no doubt adapted to some specialized feeding behavior, and are correlated

with other modifications of the mouthparts (see below). The members of Mastax have flat,

falciform mandibles, each with a large bifid retinaculum. The scrobe is absent, but the

scrobal seta remains. The basal margin is densely setiferous, as is the ventral groove. This

type of mandible looks very similar to larval mandibles in other bombardier beetles, and may

represent a neotenous condition in these beetles. Jeannel (1926) has suggested neoteny for

other characteristics in the Trechini.

All other groups have mandibles which are basally trigonal and apically subfalciform. All

possess scrobes which are plurisetose in Brachinina (figs. 1, 8) and Crepidogastrini (fig. 7)

members, and unisetose in Aptinina (figs. 2, 3) and Pheropsophina (figs. 4, 5) members.

Members of all species possess a setiferous ventral groove, and in addition the members of

Pheropsophina possess a brush of setae on the basal margin. There appears to be interspecific

variation in the teeth of the cutting edge, but I have not studied this in detail. Intergeneric

mandibular variation occurs as follows: Aptinina members have small rounded cusps, two

terebral and one retinacular; Styphlomerus members have an elongate and ridge-like retina-

culum; Crepidogastrini members have a single sharp terebral tooth and a retinacular ridge;

Pheropsophina members have two types, the first is found in the primitive Pheropsophidius,

and is a small square swelling on the terebrum, while the second is found in members of all

other subgenera, and consists of a single terebral tooth and a bifid retinacular tooth; Brach-

inina members have the most complex pattern of mandibular teeth found in the bombardier

beetles, consisting of several teeth on both the terebral and retinacular edge.

Maxillae

The maxilla (fig. 9) varies little throughout the bombardier beetles. The cardo is small.

The stipes is about three times the length of the cardo. The galea is palpiform and two-

articled, and the lacinia is falciform and sharply pointed at the apex. The dorsal surface of

the lacinia is densely clothed with long setae. Only in members of Mastax is the acute apex

of the lacinia slightly reduced. The palpus is composed of four articles, the last of which is

subcylindrical, except in some members of the Old World Pheropsophina and Brachinulus

(wedge-shaped), Mastax (globose-attenuate), and some Crepidogastrini (securiform, glo-

bose).

Labium

The characteristics of the labium are quite divergent throughout the bombardier beetles.

The ligula probably represents the fused glossae and paraglossae. This is most evident in

members of Mastax (fig. 10) in which the membranous setiferous lateral lobes (= paraglossae)

are twice the size of the convex median sclerotized lobe (= fused glossae). In members of

Mastax this median lobe generally has three or more long setae. The members of Crepido-

gastrini (fig. 1 1 ) possess a very small ligula which has a convex medial sclerotized area and

small membranous lateral lobes, each with two or more setae. The members of Aptinina

(figs. 16, 17) and Brachinina (figs. 13, 15) are generally the same, except the dorsal surfaces

of the lateral lobes are setiferous. In the members of the Old World Pheropsophina (fig. 1 4),

the venter of the ligula is carinate, but in New World Pheropsophina (fig. 12) this area is

globose as in other bombardier beetles. In some cases, interspecific variation occurs in the

disposition of ligular setae and in the shape of the central convexity. These characteristics

can be used for reliable diagnosis of particular species.

In most members of Crepidogastrini (fig. 1 1 ), the last article of the palpus is securiform or

Erwin

8 i— 1

Figs. 1-8. Right mandible, ventral and dorsal aspect. 1. Aploa nobilis Dejean, Chad, Africa. 2. Styphlomerus ?ciliatus

Liebke, Garamba, Congo, Africa. 3. Aptinus bombarda Illiger, Europe. 4. Pheropsophidius rivieri Demay, Ciudad

Bolivar, Venezuela. 5. Pheropsophus ?bimaculatus (Linn6), Mysore, India. 6. Mastax ?hargreavesi Liebke, Abalaliki,

Nigeria. 7. Crepidogaster caffra Peringuey, Cape Point, South Africa. 8. Brachinus phaeocerus Chaudoir, Lake Roberts,

New Mexico. Fig. 9. Right maxilla, dorsal aspect, Aploa nobilis Dejean, Chad, Africa. Accompanying scale lines equal

1.0 mm.

Bombardier Beetles

17

globose, or almost so. In members of most Old World Pheropsophina (fig. 14) and Brachin-

ulus, is is narrowly wedge-shaped. In members of Mastax (fig. 10), it is globose-attenuate,

otherwise it is subcylindrical.

The mentum is generally the same in all groups. It is toothed in members of Aptinina

(figs. 16, 17) and Mastax (fig. 10), but in the latter it may be a result of the deep central pit

surrounded by setae. The lateral lobes of the mentum are usually acute, but in members of

Mastax they are broadly rounded. The deep pit in the center of the mentum is found in all

members of Mastax, Brachinus costipennis, and Brachinus longipalpis. A shallower sulcus is

found in some members of European Brachinus species and in Brachinus (Neobrachinus)

mobilis. Two deep lateral pits are found in the mentum of members of Brachinus sallei (fig.

20) of Mexico and Brachinus scotomedes of Japan. Jeannel (1926) suggested that similar

structures in some members of the Trechini might be acoustical organs and that the inner-

vation should be traced. No one, however, has yet investigated these pits. The setae of the

mentum provide many useful characteristics for species diagnosis. The submentum and its

setae also provide useful characteristics.

The submentum is slightly variable among members of the Brachinida, particularly the

sides that conceal the bases of the cardines (figs. 10-17). In members of Mastax, Brachinus

costipennis (fig. 18), and Brachinus longipalpis , the submentum is very short. This shorten-

ing is concordant with the suicate mentum.

The gular sutures are divergent in all groups except the crepidogastrines (fig. 1 1), in which

they converge to the occipital groove, then diverge.

In summary, the characteristics of the head provide many useful criteria for phylogenetic

considerations and taxonomic diagnoses at all categorical levels.

Pro thorax

Externally the prothoraces of members of Brachinida are very similar (fig. 22). Internally,

however, great divergence is exhibited in the structure of the coxal cavities. Both the uniper-

forate and biperforate conditions exist; that is, one or two holes enter the body cavity for

the passage of muscles. Biperforate coxal cavities (fig. 25) occur in members of Crepidogas-

trini, Mastacina, and Pheropsophina, while the uniporforate condition (fig. 24) occurs in the

Brachinina and Aptinina. The remnants of a bridge (fig. 23) are quite evident in members of

Brachinus (sensu stricto), Brachynolomus, and Aptinina , but are almost absent in other

Brachinus (Neobrachinus).

The proepimeron closes the procoxal cavity behind in all groups except in members of

Protopheropsophus and Stenaptinus. In the members of the latter subgenus, the proepim-

eron barely reaches the prosternal process, and in some specimens there is a noticeable gap.

This variation might not have phylogenetic significance if it were not for the members of

Protopheropsophus biplagiatus Chaudoir, which have a wide and consistent gap behind the

coxal cavity. When the coxal cavities are closed behind, it is usually due to the proepimeron

overlapping the lateral process of the prosternum. However, in Brachynolomus, Brachino-

aptinus, Aploa, and some Cnecostolus and Neobrachinus each lateral process of the proster-

num forms a “socket” or “notch” into which the proepimeron inserts. Bell (1967) points

this out as a generality for Brachinini, but he evidently examined members of Neobrachinus

only.

The only other prothoracic character that shows intergeneric variation is the presence or

absence of a propleural suture. Although the proepisternum and proepimeron are fused, a

ridge runs the length of the fused sclerites in the area of juncture. This ridge occurs in the

Crepidogastrini, Mastacina, and Old World Pheropsophina.

18

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Figs. 10-17. Labium, mentum, and gula, ventral aspect. 10. Mas tax ?hargreavesi Liebke, Abakaliki, Nigeria. 1 1 . Crepi-

dogaster caffra Peringuey, Cape Point, South Africa. 12. Pheropsophidius rivieri Demay, Ciudad Bolivar, Venezuela.

13. Brachinus phaeocerus Chaudoir, Lake Roberts, New Mexico. 14. Pheropsophus ?bimaculatus (Linne), Mysore,

India. 15. Aploa nobilis Dejean, Chad, Africa. 16. Styphlomerus ?ciliatus Liebke, Garamba, Congo, Africa. 17. Ap-

tinus bombarda llliger, Europe. Accompanying scale lines equal 1.0 mm.

Bombardier Beetles

1

18

20

Figs. 18-21. Mentum and submentum, ventral aspect. 18. Brachinus costipennis Motschulsky, 12.2 miles south of El

Banco, Durango, Mexico. 19. Brachinus melanarthrus Chaudoir, 5.0 miles northwest of Acayucan, Veracruz, Mexico.

20. Brachinus sallei Chaudoir, Cozumel Island, Quintana Roo, Mexico. 21. Brachinus microamericanus Erwin, Dun-

dee, Mississippi. Fig. 22. Diagrammatic illustration of prothorax, left lateral aspect. Figs. 23-25. Procoxal cavities,

ventral aspect. 23. Brachinus crepitans (Linne), Switzerland. 24. Brachinus phaeocerus Chaudoir, Lake Roberts, New

Mexico. 25. Pheropsophus ?bimaculatus (Linne), Mysore, India. Fig. 26. Diagrammatic illustration of middle and hind

coxae and metastemum, ventral aspect. Accompanying scale line equals 1.0 mm (pp = proepipleuron, pe = proepi-

stemum, ps = prostemum).

20

Erwin

Meso thorax

The mesothorax is usually unmodified externally throughout the members of Brachinida,

with a few important exceptions. In members of the Crepidogastrini, the mesepimeron is

lost or is very narrow, and is internal. The mesepisternum is in contact with the metepister-

num. I discuss the importance of this character state under flightlessness. The coxal cavities

are conjunct-separate in members of Mastax, Pheropsophidius, and Pheropsophus, and

conjunct-confluent in the rest of the bombardier beetles. Bell (1967) defines coxal ter-

minology.

The elytra exhibit considerable intergeneric variation. Generally, each elytron is rect-

angular (or hemi-ovate in some wingless species) with a moderately wide epipleuron. The

truncate apex is either perpendicular to the suture or is angulate. In the latter case, the

length along the suture is shorter than the epipleural length. The costae of the elytra are

present in all groups, but differently modified in some. The members of the Old World

Pheropsophina have carinate costae, separated by flat, macrosculptured depressions. The

costae are narrow in Aptinomorphus, and wide in Stenaptinus and Pheropsophus (sensu

stricto). The other bombardiers have rounded, low or moderately-high costae, alternating

with the strial depressions. In members of some Brachinus, Styphlomerus, and Crepidogas-

trini, the costae are absent, but the erect “depression setae” mark the location of the striae

commonly found in other carabid beetles.

Meta thorax

The metathorax is similar in all bombardiers, except in members of Mastax. This aberrant

genus has members with the anterior metasternal process broadly rounded, rather than acute

as in other bombardiers. Further, the members of this genus have the metacoxal cavities

lobate-separate, rather than lobate-confluent as in other bombardiers.

In members of wingless Brachinida species, or those with reduced wing membranes (not

wing dimorphic), there is a reduction in the length of the metasternum and metepisterna. In

all cases where there has been wing reduction or loss of flight (even when not necessarily

from reduction of wing membrane), the length of the metasternum behind the mesocoxa

is subequal to or shorter than the diameter of the mesocoxa (fig. 26). The metepisternum has

not become square as in some Pterostichus (Ball, 1960; Hacker, 1968), except in beetles

with wings entirely absent as in members of Aptinus and Crepidogastrini. With the reduction

in the length of these “flight components”, the lobe of the metepimeron becomes larger,

especially in Aptinus. In species which apparently have been flightless for a long time, such

as the members of Crepidogastrini, the mesepimeron disappears, at least externally.

I believe that the relative ages of wingless species of bombardiers (and possibly other

carabids) may be determined by comparisons of metathoracic structures. Hypothetically,

the longer a species has been flightless, the shorter the metasternum, metepisterna, and

wings, the more sloping the humeri, and the larger the lobe of the metepimeron. Ultimately,

the elytra fuse along the suture, but this has not occurred in bombardier beetles. Of course

the reduction probably begins with wing dimorphism as described by Lindroth (1945, 1963)

and Darlington (1936, 1943), but after all members of the species have become flightless,

then reduction in the mesothoracic and metathoracic flight components begins, and a build-

ing up of thoracic “ambulatory components” takes place.

Most groups of bombardier beetles have some wingless members, but I have not yet seen

nor heard of any wingless members of Mastax nor of Styphlomerus and its allies. Among

those bombardier beetles with wings (figs. 27-32), there is great similarity in the venation

and wing-folding pattern. Both are clearly “caraboid” in nature (see Forbes, 1926), except in

members of Mastax. This aberrant group has all the wing veins reduced to vague sclerotized

fields on the membrane. The oblong cell has almost disappeared, and the wedge cell and

Figs. 27-32. Left wing, dorsal aspect. 27. Styphlomerus ?ciliatus Liebke, Garamba, Congo, Africa. 28. Mas tax ?harg-

reavesi Liebke, Abakaliki, Nigeria. 29. Brachinus phaeocerus Chaudoir, Lake Roberts, New Mexico. 30. Pheropsophus

?bimaculatus (Linne), Mysore, India. 31. Aploa nobilis Dejean, Chad, Africa. 32. Brachinus imporcitis new species,

Pinal Creek, Arizona. Accompanying scale lines equal 1.0 mm.

22

Erwin

first and second anal complexes are absent. Further, the wing is rounded apically, rather

than being acute as in the other bombardier groups.

Legs

The legs of bombardier beetles are quite similar throughout the various groups, but the

pubescence and setae vary interspecifically. Only the anterior tibiae and anterior male tarsi

offer obvious characters for taxonomic diagnoses and phylogenetic considerations. Possibly

after enough study of additional representative material, the middle and hind legs may offer

good characteristics in some groups (but not in Neobrachinus).

The anterior tibia has an antennal comb one third the distance from apex to base. The

position of this comb, and the relative positions of its two associated spurs, have been used

as the bases of some proposed classifications (Jeannel, 1941 ; Bell, 1967). In the bombardier

beetles, the proximal spur (figs. 33-39) is located on the upper edge of the comb, either in-

ternally (Aptinina, Crepidogastrini), intermediately positioned ( Aploa , Brachinoaptinus,

Pheropsophus ), or externally. It is absent from members of Mastax (fig. 40). The distal spur

is located at the tibial apex, behind the tarsal insertion.

The vestiture of the male anterior tarsal articles is intergenerically quite diverse. The mem-

bers of Crepidogastrini (fig. 50) have circular pads of setae, spongy in appearance, beneath

either two or three basal tarsal articles.

An Indian species, Tyronia humerata (Chaudoir), has members with three articles with

pads of vestiture, while the African Tyronia kivuensis Basilewsky has only two articles with

vestiture. Members of Mastax have each of the two basal articles bearing two parallel rows of

spatulate setae (fig. 49). The rows are symmetrically arranged on the underside of the ar-

ticles. This same two-rowed pattern is seen in some Brachinina (fig. 42), and in all Pherop-

sopina (figs. 44, 45), except that in the latter two groups three articles have the vestiture.

In members of the Aptinina (figs. 43, 47) and some Brachinina (fig. 41), the two parallel

rows are diagonally placed on the three basal asymmetrical tarsal articles.

Abdomen

The abdomen of the bombardier beetles exhibits very little variation. The females have

seven visible abdominal sterna, while the males have eight. In the male the eighth sternum is

divided into two lateral halves, while in both sexes the ninth tergum is modified into twin

“crepitation chambers”, separated by a median keel. At the base of the keel, on either side,

lies the outlet from the mixing chamber of the crepitating mechanism. When the mechanism

is not being fired, the eighth tergum overlies the ninth. During crepitation the ninth tergum

is telescoped caudally from under the eighth tergum, exposing the twin chambers. Each

chamber consists of a smooth L-shaped trough bordered internally by the median keel, and

bordered caudally by the up-turned apex of tergum nine. The foot of the L lies transversely

at the base of the upturned apex of tergum nine. This construction allows the beetle to

“aim” the chemical components of its irritant toward the predator. Eisner (1958) has

studied the directional aspects of this defensive spray, and he shows it can be aimed. I have

observed the aiming capabilities of these beetles while collecting them and while observing

them in the laboratory. The extensible abdomen and truncate elytra allow free movement of

the abdominal apex for aiming.

Genitalia

Female ovipositor and bursa copulatrix

Tanner (1927) discussed the family characteristics of carabid female genitalia. The bom-

bardier beetles fit the basic description Tanner gives, with the exception of Mastax and

Bombardier Beetles

23

37 38

39 40

Figs. 33-40. Right front tibia of male, lateral aspect. 33. Brachinus phaeocerus Chaudoir, Lake Roberts, New Mexico.

34 . Pheropsophus ?bimaculatus (Linne), Mysore, India. 35. Pheropsophidius rivieri Demay, Ciudad Bolivar, Venezuela.

36. Aptinus bombarda Illiger, Europe. 37. Aploa nobilis Dejean, Chad, Africa. 38. Crepidogaster caffra Peringuey,

Cape Point, South Africa. 39. Styphlomerus ?ciliatus Liebke, Garamba, Congo, Africa. 40. Mastax ?hargreavesi

Liebke, Abakaliki, Nigeria. Accompanying scale lines equal 1.0 mm.

Erwin

Figs. 41-47, 49, 50. Right front tarsus of male, ventral aspect. 41. Brachinus immaculicornis Dejean, France. 42. Bra-

chinus phaeocerus Chaudoir, Lake Roberts, New Mexico. 43. Aptinus bombarda Illiger, Europe. 44. Pheropsophus

?bimaculatus (Linne), Mysore, India. 45. Pheropsophidius rivieri Demay, Ciudad Bolivar, Venezuela. 46. Aploa nobilis

Dejean, Chad, Africa. 47. Styphlomerus ?ciliatus Liebke, Garamba, Congo, Africa. 49. Mastax ?hargreavesi Liebke,

Abakaliki, Nigeria. 50. Crepidogaster caffra Peringuey, Cape Point, South Africa. Figs. 48, 51, 52. Female ovipositor,

ventral aspect. 48. Styphlomerus ?ciliatus Liebke, Garamba, Congo, Africa. 51. Stenaptinus ?kolbei Hubenthal,

Cameroon, Africa. 52. Crepidogaster caffra Peringuey, Cape Point, South Africa. Accompanying scale lines equal 1.0

Bombardier Beetles

25

Crepidogaster . In members of Mastax (fig. 62) there are two lateral, elongate sclerites in the

position Tanner illustrates for the ninth sterna. In the position Tanner illustrates for the

tenth sternite behind the bursal opening, there are two small transverse rods. Further, mem-

bers of Mastax have a single seta on the coxite that exceeds the length of the stylus. In mem-

bers of Crepidogaster (fig. 52), the coxite and stylus are fused and articulated with the

valvifer. Peculiarly, the base of the valvifer is plurisetose.

The members of Aptinina have the normal caraboid three piece ovipositor. In Aptinus

members (fig. 58), the apical-medial corner of the valvifer is plurisetose, while in members of

Styphlomerus, Styphlomerinus, and Styphlodromus (fig. 48), the apical edge of the valvifer

has several flattened spine-like setae. The styli of the Aptinina members are strongly curved

blades, which are usually acute apically. The members of the remaining genera have various-

ly modified styli, depending upon the species. In the members of Pheropsophina (fig. 5 1 ,

59, 61), the stylus blade is variously modified among the subgenera. Pheropsophus (sensu

stricto) members have very elongate and narrow styli, some with microspinules along the

inner edge (fig. 61 ). In Stenaptinus members, the styli are long and spatulate (fig. 5 1 ). In the

members of Pheropsophidius , they are similar to those in members of Brachinus, but lack

the setae (fig. 59).

The various members of Brachinus and Aptinoderus have the normal carabid three piece

ovipositor. The stylus is flattened and triangular, but varies considerably between the species.

The members of Brachinus have a single small seta on the ventral side near the apex of the

spatulate stylus.

The female members of Aptinus (fig. 55) are the only Brachinida which possess bursal

sclerites. The entire walls of the bursa are thinly sclerotized in a laminate fashion. A single

large rod-like sclerite is dorsal to the entrance of the bursa. Laterad, near the base of this rod

on the wall of the bursa, is a large and concave sclerite. At this point the entire bursa is trans-

verse, with a large forked sclerite in a pouch near the concave sclerite.

Male genitalia

The male genitalia are interspecifically distinct, and they can be used to distinguish be-

tween species in at least Neobrachinus. They are composed of the following parts: median

lobe, with the apex, shaft, ligule, basal bend, and basal portion; two parameres (the left one

large and the right one small) attached to the ventral comer of the basal portion of the

median lobe; and an endophallus, which is a membranous sac containing the gonopore,

which in turn may or may not provide attachment for various apical or subapical sclerites

and microtrichial fields.

The members of Crepidogastrini possess a very primitive type of male genitalia. The

median lobe is tubular, straight or bent, sometimes with the shaft contorted. The parameres

are not balteate, but are asymmetric. The left paramere is lobate, and is alongside the median

lobe. The smaller right paramere is lobate and is held close to the shaft. The ligule of the

shaft is absent. The endophallus is more than half the length of the median lobe, and in

members of some Crepidogaster, the apex has microtrichial fields.

In members of Brachinini, the parameres are always balteate, the left is larger than the

right, and both are wrapped around the basal portion of the median lobe and connected by

a membrane.

The members of Mastax have the most aberrant median lobe and endophallus of all the

Brachinida (fig. 53). The apex and shaft are similar to some Bembidion. The endophallus

possesses a coiled ribbon-like sclerite, as well as one basal and one apical sclerotized plate.

The parameres are smaller than in any other group, and they adhere tightly to the base of

the basal portion of the median lobe.

The remaining groups have typical brachinine genitalia with balteate parameres, a more

Erwin

26

Fig. 53. Male genitalia of Mastax ?hargreavesi Liebke, Abakaliki, Nigeria, lateral aspect. Fig. 54. Ventral aspect of

same. Fig. 55. Bursal sclerites of female Aptinus bombarda Illiger, Europe, dorsal aspect. Fig. 56. Male genitalia of

Aptinus bombarda Illiger, Europe, lateral aspect. Fig. 57. Dorsal aspect of same. Figs. 58-62. Female ovipositor, ven-

tral aspect. 58. Aptinus bombarda Illiger, Europe. 59. Pheropsophidius rivieri Demay, Ciudad Bolivar, Venezuela.

60. Aploa nobilis Dejean, Chad, Africa. 61. Pheropsophus ?bimaculatus (Linn6), Mysore, India. 62. Mastax ?hargrea-

vesi Liebke, Abakaliki, Nigeria. Accompanying scale lines equal 1.0 mm. (go = gonopore, ds = dorsal sclerite. Is =

lateral sclerites, tp = transverse pouch, bs = basal sclerite, sp = spermatheca, ov = oviduct).

Bombardier Beetles

27

or less tubular median lobe, and generally a moderate-sized endophallus. Some of these

groups have microtrichia or sclerites on the internal sac. The members of Pheropsophus

(fig. 69) and Pheropsophidius (figs. 437-442) have fields of microtrichia on a very long

endophallus. Styphlomerus members which I have seen have a shorter endophallus, but

still with microtrichia (but I have seen very few examples). Aptinus members have very

contorted median lobes with a very short endophallus. The endophallus has one basal and

two apical sclerites. Although I have not studied Aptinus males and females in copula , I

believe there is a definite correlation between the female bursal sclerites, and the male med-

ian lobe. A study of this relationship might prove fruitful.

Within the genus Brachinus, there have evolved many types of endophallic sclerites. The

members of Neobrachinus, Brachynolomus, Cnecostolus, and some African lineages, have

developed an apical sclerite surrounding the gonopore. Snodgrass (1935) has termed this

sclerite the “virga”. The members of Brachinoaptinus, Brachinus (sensu stricto), Metabrach-

inus, other African lineages, and at least one Oriental lineage, do not have any sclerites on

the endophallus. The members of one Oriental lineage have a long endophallus with a tube

at the apex which bears microtrichia. The members of Aploa, and those of at least one west

African lineage, have a subapical endophallic sclerite. The variation and homologies of these

virgae and sclerites are extremely important aids for persons attempting to understand the

relationships and hence, the dispersal movements of the members of the genus Brachinus.

Because of the extreme similarity in the external features of these beetles, the genitalic

diversity often must be studied carefully to determine the species differences (see below

under Neobrachinus).

The virgae of all North and Middle American species of Brachinus with known males were

studied in great detail. As a result of this study, certain patterns of evolution became appar-

ent. From a simple sclerotized tip of the endophallus (as in extant Brachinus dryas Andrewes

and members of the americanus group) more complex types evolved as indicated in fig. 461 .

TAXONOMY

Division Brachinida Bonelli

Brachinii Bonelli, 1809: tab. syn.

Brachinida Ball, 1960: 164.

The following combination of characteristics is diagnostic of this division of higher Cara-

bidae: anterior tibia anisochaetous middle coxae conjunct; lobe of metepimeron present;

elytra truncate, costate, striae obscure; abdomen with seven or eight sterna normally ex-

posed; outlets of the crepitating mechanism medial, in front of modified ninth tergum; ninth

tergum modified into twin crepitating chambers in both males and females; male genitalia

with an endophallus; parameres asymmetrical, the right minute, the left larger; cuticle of

mature adult pliable.

This division includes the tribes Crepidogastrini and Brachinini.

Key to tribes and subtribes of the division Brachinida

1 Mesepimera absent or almost so; ventral vestiture of male anterior tarsal articles

spongy and pad-like; terminal palpal articles swollen, usually securiform; gular

sutures convergent behind Crepidogastrini

1’ Mesepimera present, broad, clearly visible; vestiture beneath male anterior tarsal

articles consisting of two parallel rows of modified setae; terminal palpal articles

subcylindrical or wedge-shaped ; gular sutures divergent behind .... Brachinini 2

2(1’) Hind coxae contiguous; metasternal process acute; middle coxae not widely sep-

28 Erwin

arated; mandible with a scrobe; upper spur of anterior tibia present 3

2’ Hind coxae widely separated; metasternal process broadly rounded; middle coxae

widely separated; mandible without a scrobe; upper spur absent

Mastacina, new subtribe

3~(2) Mandibular scrobe unisetose 4

3’ Mandibular scrobe plurisetose Brachinina sensu stricto

4(3) Vestiture of male anterior tarsal articles arranged diagonally on asymmetric ar-

ticles; mentum with a tooth on anterior edge (absent in Styphlodromus ); upper

spur on anterior tibia internal (fig. 36) Aptinina, new subtribe

4’ Vestiture of male anterior tarsal articles not diagonal, articles symmetric; mentum

without a tooth; upper spur external or almost so (fig. 35)^

Pheropsophina, new subtribe

Tribe Crepidogastrini Jeannel

Crepidogastritae Jeannel, 1949: 1080.

Crepidogastrini Basilewsky, 1959: 233.

Type genus. - Crepidogaster Boheman, 1848: 68.

The following combination of characteristics is diagnostic of this tribe: labrum with eight

setae; mandibular scrobe plurisetose; palpi with terminal articles swollen or securiform;

gular sutures convergent behind; anterior coxal cavities biperforate, closed behind; pro-

pleural suture present; upper spur of anterior tibia internal; ventral vestiture of male anterior

tarsal articles spongy and pad-like; middle coxal cavities conjunct-confluent; mesepimera

absent or almost so; apical elytral membrane absent; hind coxal cavities confluent; meta-

sternum and metepisterna greatly reduced; wings absent; male parameres not balteate; val-

vifers of female ovipositor setiferous anteriorly ; coxite and stylus of female ovipositor fused.

This tribe contains the genera Crepidogaster, Tyronia, Crepidogastrinus, Crepidonellus,

Crepidogastrillus, Brachynillus, and Crepidolomus.

Key to the genera of the tribe Crepidogastrini (translated and modified from Basilewsky,

1959)

1 Lateral edge of pronotum with many spine-like setae; lateral edge of elytron also

with spine-like setae Crepidogastrinus Basilewsky

1’ Lateral edge of pronotum with one or two pairs of setae; depression 8 of elytron

with umbilicate setae only 2

2(1’) Elytra short and wide, maximum width greater than length at suture 3

2’ Elytra long and narrow, maximum width shorter than length at suture 4

3(2) Pronotum with one pair of lateral setae, near the middle; terminal article of labial

palpus securiform Crepidonellus Basilewsky

3’ Pronotum with two pairs of lateral setae, one just before hind angles and one near

the middle; last article of labial palpus globose Crepidogastrillus Basilewsky

4(2) Pronotum with one pair of lateral setae, near the middle 5

4’ Pronotum with two pairs of lateral setae, one just before hind angles and one near

the middle 6

5(4) Terminal article of labial palpus strongly swollen or securiform; terminal article of

maxillary palpus very swollen or securiform Crepidogaster Boheman

5’ Terminal article of labial palpus fusiform or subcylindrical; terminal article of the

maxillary palpus globose, but not securiform; body slender and elongate

Brachynillus Reitter

6(4’) Terminal article of labial palpus strongly swollen or securiform; terminal article of

maxillary palpus very swollen or securiform Tyronia Liebke

Bombardier Beetles

29

6’ Terminal article of labial palpus fusiform or subcylindrical; terminal article of the

maxillary palpus globose, but not securiform; body slender and elongate

Crepidolomus Basilewsky

Genus Crepidogaster Boheman

(Figs. 7, 11,38,50,52, 63,64)

Crepidogaster Boheman 1848: 68. Type species. — Crepidogaster bimaculatus Boheman

1848: 68 (subsequent designation, Jeannel, 1949: IQ81).

Crepidostoma Motschulsky 1862: 54. Type species. — Crepidogaster rufescens Motschulsky

1862 : 54 (by monotypy).

Diagnostic combination. — Terminal article of labial palpus strongly swollen or securi-

form; terminal article of maxillary palpus very swollen or securiform; pronotum with one

pair of setae, near the middle; elytra long and narrow, their combined maximum width

shorter than their length along the suture.

Description. — Small to large-sized beetles, 4.0 to 16.0 mm (from Basilewsky, 1959).

Color. Generally ferrugineous with brown or black elytra which are sometimes spotted.

Microsculpture. Isodiametric on head, pronotum, and elytra.

Macrosculpture. Entire dorsal surface covered by small setiferous punctulae.

Head. Labrum entire, with eight evenly spaced setae transversely arrayed on disc. Clypeus

rectangular, with two large setae, one inside each anterior angle. Front punctate with numer-

ous setae scattered throughout; furrows shallow; one supraorbital seta over each eye; bead

over eye ending midway the length of eye. Eyes protruding, but not prominent. Antennae

short, robust, pubescent throughout. Mandibles (fig. 7) each broad, triangular, curved at tip;

cutting edge with one small retinacular tooth; ventral groove with short golden setae; scrobe

plurisetose, but with one larger seta. Labial palpi (fig. 1 1) each with last article securiform;

all articles with numerous small setae. Maxillary palpi each more or less filiform, last article

swollen; all articles with numerous small setae. Ligula (fig. 1 1) membranous and square with

anterior angles produced, and center ventrally produced into a plurisetose sclerotized con-

vexity. Mentum (fig. 11) well developed without tooth. Gula (fig. 11) narrowed behind.

Prothorax. As described for the tribe Crepidogastrini, plus numerous punctulae on disc.

Anterior tibiae (fig. 38) each with upper spur internal, at middle of comb emargination.

Male anterior tarsal articles (fig. 50) symmetrical, vestiture beneath first and second consist-

ing of apically widened setae forming pads. Female anterior tarsal articles without vestiture.

Pterothorax. Mesepimera absent, or almost so. Metathoracic process acute. Metepimeron

lobe rounded apically. Elytral humeri sloping, lateral bead discontinuous before base of

humerus. Depression setae present between well developed carinae or costae. Epipleura

pubescent, wide anteriorly and medially, but narrowed toward apex. Wings absent.

Abdomen. As described for the tribe, plus a pair of ambulatory setae arising laterad, at

apex of sterna 3-6.

Genitalia. Male (figs. 63, 64) with small parameres, not balteate, asymmetrical. Median

lobe various, ligule absent. Endophallus of medium length, with two apical microtrichiate

fields and one basal field; virga absent. Female (fig. 52) with elongate, narrow, spatulate, and

unarmed styli; coxites small, continuous with stylus; valvifers setose anteriorly; bursa un-

armed.

Etymology . — Latin, crepo = rattle; Greek, gaster = stomach; referring to the beetles’

ability to crepitate.

Distribution. — The species of Crepidogasten occur south of the equator in Africa and

Madagascar, with only two populations of one species known north of the equator in east

Africa.

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63 64

Figs. 63-70. Male genitalia. 63. Crepidogaster caffra Peringuey, Cape Point, South Africa, ventral aspect: 64. Lateral

aspect of same. 65. Aploa nobilis Dejean, Chad, Africa, ventral aspect. 66. Lateral aspect of same. 67 . Styphlomerus

?ciliatus Liebke, Garamba, Congo, Africa, ventral aspect. 68. Lateral aspect of same. 69. Pheropsophus ?bimaculatus

(Linne), Mysore, India, ventral aspect. 70. Lateral aspect of same. Accompanying scale lines equal 1.0 mm.

Bombardier Beetles

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Genus Tyronia Liebke

Tyronia Liebke, 1934: 18. Type species. — Crepidogaster numeratus Chaudoir, 1876: 96

(original designation).

Diagnostic combination. — As given in the key.

Description. — Small-sized beetles, 3.5 to 7.0 mm (from Basilewsky, 1959). For details

see Basilewsky (1959).

Distribution. — The species of Tyronia occur in south and middle Africa on the eastern

side of the continent. Two species occur in the Oriental Region, one in India, and one in

Ceylon.

Genus Brachynillus Reitter

Brachynillus Reitter, 1904: 178. Type species. — Brachynillus varendorffi Reitter 1904 :

178 (by monotypy).

Diagnostic combination. — As given in the key.

Description. - Small-sized beetles, 5.0 to 5.5 mm (from Basilewsky, 1959). For details

see Basilewsky (1959).

Distribution. — The type species is found as a troglobiont in the “Grottes de Kulumuzi”

in Tanganyika. A second species placed in the genus by Basilewsky is known from one speci-

men labelled “Cape Town”.

Genus Crepidolomus Basilewsky

Crepidolomus Basilewsky, 1959: 33 1 . Type species. — Brachynillus extimus Jeannel, 1955 :

62 (original designation).

Diagnostic combination. — As given in the key.

Description. — Small-sized beetles, 5.5 mm (from Basilewsky, 1959). For details see

Basilewsky (1959).

Distribution. — One female specimen is known from Andohahelo, on the southern

tip of Madagascar.

Genus Crepidonellus Basilewsky

Crepidonellus Basilewsky, 1959: 334. Type species. - Crepidogaster pusillus Peringuey,

1888: 76 (original designation).

Diagnostic combination. — As given in the key.

Description. - Very small-sized beetles, 3.6 to 4.3 mm (from Basilewsky, 1959). For

details see Basilewsky (1959).

Distribution. - The members of Crepidonellus are found in South West Africa and Cape

Province.

Genus Crepidogastrillus Basilewsky

Crepidogastrillus Basilewsky, 1959: 337. Type species. — Crepidogastrillus curtulus Basil-

ewsky, 1959: 338 (original designation).

Diagnostic combination. — As given in the key.

Description. — Very small-sized beetles, 3.4 to 3.6 mm (from Basilewsky, 1959). For

details see Basilewsky (1959).

Distribution. — The range of this genus extends from Cape Province north into South

West Africa, and east to northwest Rhodesia.

Genus Crepidogastrinus Basilewsky

Crepidogastrinus Basilewsky, 1957: 1 17. Type species. — Crepidogastrinus kochi Basilewsky,

1957: 1 18 (by original designation and monotypy).

Diagnostic combination. — As given in the key.

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Erwin

Description. — Very small-sized beetles, 3.0 mm (from Basilewsky, 1959). For details see

Basilewsky (1957, 1959).

Distribution. — The range of this species is confined to southern Angola and south West

Africa.

Tribe Brachinini Bonelli

Brachinii Bonelli, 1809: tab. syn.

Brachynidae Stephens, 1827: 5.

Brachynini Erichson, 1837: 25 (Brachinini, of authors).

Brachinida Heer, 1838: 4.

Brachinides Lacordaire, 1854: 97.

Brachinites Jacquelin du Val, 1857: 55.

Brachinina Thomson, 1859: 6 (Brachynina, of authors).

Brachyninae Kolbe, 1898 : 60.

Brachynitae Alluaud, 1916: 50.

Brachinidae Jeannel, 1942: 1102.

The following combination of characteristics is diagnostic of this tribe: mandibular scrobe

plurisetose or unisetose; labial palpus with terminal article wedge-shaped, subcylindrical,or

globose-attenuate; gular sutures divergent behind; anterior coxal cavities uniperforate or bi-

perforate; propleural suture present or absent; upper spur of anterior tibia present or absent,

internal or external; ventral vestiture of male anterior tarsal articles composed of two parallel

rows of modified setae; mesepimeron present, wide, but not reaching middle coxa; male

parameres balteate.

This tribe contains the subtribes Mastacina new subtribe, Pheropsophina Jeannel, Aptinina

new subtribe, and Brachinina Bonelli.

Subtribe Mastacina new subtribe

Type genus. — Mastax Fischer von Waldheim, here designated.

This subtribe contains a single genus. The diagnostic characteristics are given below.

Genus Mastax Fischer von Waldheim

(Figs. 6, 10, 28, 40, 49, 53, 54, 62)

Mastax Fischer von Waldheim, 1825-28: 111. Type species. — Brachinus thermarum Stevens,

1806: 166 (subsequent designation by Jedlicka, 1963: 547).

Brachinus (of authors).

Diagnostic combination. - Mandibular scrobes reduced, unisetose; last article of palpi

fusiform with subulate tip; mentum with single tooth and deep pit (fig. 10); mandibles each

with large bifid terebral tooth; anterior coxal cavities biperforate-separate, closed behind;

propleural sutures present; pronotum with two longitudinal ridges on disc; upper spur of

anterior tibia absent; middle coxae conjunct-separate; metasternal process (between middle

coxae) broadly rounded; hind coxae widely separated; wing (fig. 28); parameres of male

genitalia extremely small, adherent to base of median lobe; ligule absent; coxite of female

genitalia unisetose, stylus short.

Description. — Small-sized beetles, 2.5 to 4.0 mm.

Color. Head, prothorax, and legs generally ferrugineous to brown. Abdomen, antennae,

and “knees” infuscated. Elytra brown with white spots.

Microsculpture. Isodiametric on head and pronotum, except where obliterated by macro-

sculpture. Longitudinally-stretched meshes on elytra.

Macrosculpture. Longitudinal wrinkles on head and pronotum. Minute chevron-shaped

wrinkles on costae of elytra, surface “satin-like” in appearance.

Bombardier Beetles

33

Head. Labrum emarginate, with eight evenly spaced setae transversely arranged on disc.

Clypeus rectangular, with six evenly spaced setae transversely arranged on disc. Front longit-

udinally wrinkled with numerous scattered setae; furrows shallow, reflexed strongly at sides

over antennal bases; one supraorbital seta over each eye, one pseudosupraorbital seta behind

each eye; bead over eye carinate and prolonged on side of vertex behind pseudosupraorbital

seta; shallowly depressed at middle. Eyes prominent. Antennae robust; articles 5-10 as wide

as long, moniliform; article 1 1 fusiform and subulate at apex; all articles pubescent. Man-

dibles (fig. 6) arcuate each with a strong bifid terebral tooth; basal margin with dense brush

of fine golden setae, also a row of fine setae in ventral groove; scrobes effaced, but unisetose.

Palpi (fig. 10) each with terminal article fusiform, tip subulate; all articles pubescent. Ligula

bilobed; lobes thin and densely setiferous; median portion slightly swollen with two large

setae. Mentum (fig. 10) short, with single tooth medially on anterior edge; center of men-

tum excavated and surrounded by setae. Gula (fig. 10) widened behind.

Prothorax. As described for the subtribe Masticina, and disc with two longitudinal carinae

on each side of midline. Anterior tibia (fig. 40) without upper spur. Male and female with

first two tarsal articles expanded, the male (fig. 49) with modified vestiture beneath these

two articles. Male with a curved nonarticulated spine (fig. 40) on right side of glabrous an-

tennal comb-trough.

Pterothorax. Metathoracic process (between middle coxae) broadly rounded, middle cox-

ae widely separated. Hind coxae widely separated. Metepimeron lobe small, but rounded

caudally. Elytra quadrate, humeri prominent, lateral bead prolonged inside humerus to

base of fifth stria. Setae present between weakly raised costae in striae 2, 4, 6, 8. Epipleura

wide throughout; pubescent. Wings as in fig. 28.

Abdomen. As described for the division Brachinida, except with one pair of ambulatory

setae at apex of sterna 2-5.

Genitalia. Male (fig. 53, 54) with extremely small parameres adherent to base. Median

lobe various, but usually slightly arcuate and blunt at apex. Endophallus extremely com-

plicated with (in repose) two sclerites apically and one sclerite basally; membrane reinforced

with a thin, spiraled, sclerotized rod. Female (fig. 62) with short, narrow, arcuate and un-

armed styli; valvifer transverse, narrow, and elongate; coxite unisetose; bursa unarmed; sper-

matheca present, sausage-shaped.

Etymology . — Greek, mastokos = jaw, mouth; referring to the huge retinacular tooth on

the mandibles of these beetles.

Distribution. — (Fig. 457). Species of Mastax occur on both sides of the equator, but are

not confined to the tropics. Species are known from south of the equator in South Africa

and southern Sumatra. North of the equator some species occur in western, northern, and

eastern Africa, the Congo, Saudia Arabia, Iraq (on both sides of the Caspian Sea), western

India, Ceylon, southeast Asia, Hainan, and Formosa.

Subtribe Pheropsophina Jeannel

Pheropsophini Jeannel, 1948: 1084.

Type genus. — Pheropsophus Solier, 1833: 46.

The following combination of characteristics is diagnostic for this subtribe; mandibular

scrobe unisetose; terminal palpal article wedge-shaped; anterior coxal cavities open or closed

behind; propleural suture present or absent; upper spur of anterior tibia slightly internal;

male anterior tarsal articles symmetrical or almost so; middle coxal cavities separate-con-

junct; apical elytral membrane absent; at least some depressions of elytra microrugose; hind

coxae contiguous-separate; lobe of metepimeron large; parameres of male genitalia small,

balteate; dorsal surface of median lobe at basal bend notched; valvifer of female ovipositor

glabrous.

34

Erwin

This subtribe includes the genera Pheropsophus Solier and Pheropsophidius Hubenthal.

Key to the genera and subgenera of the subtribe Pheropsophina

1 Elytra costate, with costae low and rounded, evenly confluent with depressions;

propleural suture absent Pheropsophidius 2

V Elytra carinate, carinae sharp or rounded, well defined and abruptly confluent

with depressions; propleural sutures present Pheropsophus 3

2(1) Anterior coxal cavities narrowly opened behind; depressions on disc of elytra

without macrosculpture (except near scutellum); apex of elytra obliquely trun-

cate, sutural length shorter than lateral length

subgenus Protopheropsophus Hubenthal

2’ Anterior coxal cavities closed behind; depressions on disc with macrosculpture;

apex of elytron almost truncate; costae well defined

subgenus Pheropsophidius Hubenthal

3(1’) Apical border of elytron with numerous long setae

subgenus Pheropsophus Solier

3’ Apical border of elytron without long setae, with or without very small ones

(barely visible at 50 X) 4

4(3’) Carinae of elytra broader than the depressions, and rounded dorsally

Stenaptinus Maindron

4’ Carinae of elytra very narrow, sharp dorsally (Madagascar)

Aptinomorphus Jeannel

Genus Pheropsophus Solier

(Figs. 5, 14, 25, 30, 34, 44, 51, 61 , 69, 70)

Pheropsophus Solier, 1833: 463. Type species. — Brachinus senegalensis Dejean, 1825: 308

(subsequent designation by Jeannel, 1949: 1084).

Stenaptinus Maindron, 1906: 15. Type species. — Stenaptinus krichna Maindron, 1906:

15 (subsequent designation by Jeannel, 1949: 1084).

Parapheropsophus Hubenthal, 1914: 442. Type species. — Brachinus verticalis Dejean,

1825: 302 (subsequent designation by Jeannel, 1949: 1084; Darlington, 1968: 234).

Aptinomorphus Jeannel, 1949: 1091. Type species. — Pheropsophus acutecostatus Fair-

maire, 1892: 168 (original designation).

Brachinus (of authors).

Diagnostic combination. — Elytral costae carinate, carinae sharp or rounded, well defined

and more strongly contrasting with depressions; propleural sutures present.

This genus includes the subgenera Pheropsophus Solier, Stenaptinus Maindron, and

Aptinomorphus Jeannel.

Subgenus Pheropsophus Solier

(Figs. 5,14, 25, 30, 34, 44, 61, 69, 70)

Parapheropsophus Hubenthal, 1914: 442. Darlington, 1968: 234.

Diagnostic combination. — As given for genus Pheropsophus , plus apical border of elytra

with fringe of long closely spaced setae.

Description. — Medium to large-sized beetles, 15.0 to 25.0 mm.

Color. Generally yellow or ferrugineous, with black markings.

Microsculpture. Isodiametric on head, but slightly stretched transversely on pronotum

and elytral carinae.

Macrosculpture. Depressions between elytral carinae with numerous longitudinal micro-

rugosities.

Bombardier Beetles

35

Head. Labrum entire, with six evenly spaced setae transversely arranged near anterior

margin. Clypeus rectangular, with two setae on each side near middle. Front smooth, gla-

brous; furrows very shallow; one supraorbital seta over each eye; bead over eye almost ab-

sent; eyes prominent. Antennae long, robust, first two articles setiferous, 3-1 1 pubescent.

Mandibles (fig. 5) each broad, arcuate; cutting edge with two small terebral teeth and two

retinacular teeth; ventral groove with short golden setae; basal margin with penicillus;

scrobe unisetose. Labial palpi (fig. 14) each with terminal article narrowly wedge-shaped;

terminal and penultimate articles with numerous stiff setae. Maxillary palpi each more or

less filiform, terminal three articles with numerous stiff setae. Ligula (fig. 14) membranous

and square, with anterior angles barely produced; center ventrally produced into a bisetose

sclerotized carina. Mentum (fig. 14) well developed, without a tooth. Gula (fig. 14) wid-

ened behind.

Prothorax. As described under subtribe Pheropsophina. Anterior tibiae each with subter-

minal spur in intermediate position at top of comb emargination. Male anterior tarsal ar-

ticles 1-3 slightly asymmetrical, vestiture beneath consisting of two parallel rows of spatulate

setae. Female anterior tarsal articles symmetrical, vestiture absent. Anterior coxal cavities

barely closed behind.

Pterothorax. Anterior metathoracic process acute. Middle coxal cavities contiguous-

separate. Metepimeron lobe large, rounded apically. Elytral humeri square, lateral bead

entire to base of humerus. Strial setae present between well developed carinae. Epipleura

wide anteriorly and medially, but narrowing toward apex; not pubescent. Wings present (fig.

30).

Abdomen. As described under division Brachinida, except one pair of ambulatory setae

at apices of sterna 3-6.

Genitalia. Male (fig. 69, 70) with small balteate, asymmetrical parameres; median lobe

variable, but symmetrical, and notched near base; ligule double, narrow, and spatulate;

endophallus short with various fields of microtrichia; virga absent. Female (fig. 61) with

elongate, narrow, slightly curved styli, usually armed with small spines; coxites small, ro-

bust; valvifers glabrous; bursa unarmed, except an elongate sclerite at entrance of bursa.

Etymology . — Greek, phero = bearer; psoph os = sound, or noise; hence “noise bearer”

referring to the crepitating behavior of these beetles.

Distribution. — The range of this subgenus extends over much of Africa and Madagascar,

with the exception of the desert region in the north of Africa, and eastward across Asia

Minor, India, and eastern Asia, into the Malay Archipelago, New Guinea, New Britain, New

Ireland, the Solomons, and Australia.

Subgenus Stenaptinus Maindron

(Fig. 51)

Diagnostic combination. — As in subgenus Pheropsoph us, except wings absent, anterior

coxal cavities slightly open behind; female styli shorter, more spatulate, unarmed.

Description. - Medium to large-sized beetles, about 10.0 to 15.0 mm.

Color. Ferrugineous venter and legs, black above, sometimes head and pronotum also

ferrugineous.

Microsculpture. As in subgenus Pheropsophus .

Macrosculpture. As in subgenus Pheropsophus .

Head. As in subgenus Pheropsophus .

Prothorax. As in subgenus Pheropsophus, except coxal cavity slightly open behind.

Pterothorax. As in subgenus Pheropsophus, except metasternum very short, shorter than

longitudinal diameter of middle coxa (fig. 26); metepisterna almost square; elytral humeri

absent; wings absent.

36

Erwin

Abdomen. As in subgenus Pheropsophus.

Genitalia. Male as in subgenus Pheropsophus. Female (fig. 51) styli unarmed, wider, and

shorter than subgenus Pheropsophus, otherwise similar.

Etymology . — Greek, stenos = narrow; Aptinus = name of another brachinine genus; re-

ferring to the very narrow humeri of these beetles.

Distribution. — The range of this subgenus is the Old World Tropics, including tropical

Africa, India, Ceylon, southeastern Asia, Celebes, Taiwan, Philippines, and possibly New

Guinea.

Subgenus Aptinomorphus Jeannel

Diagnostic combination. — As in subgenus Pheropsophus, except for the following charac-

ters: wings absent; elytral carinae very narrow, ridge-like; apex of elytra without setae;

female styli very short and broad.

Description. — Medium to large-sized beetles, 12.0 to 30.0 mm.

Color. Ferrugineous venter and legs; head and pronotum ferrugineous or brown; elytra

black or brown.

Microsculpture. As in subgenus Pheropsophus.

Macrosculpture. As in subgenus Pheropsophus.

Head. As in subgenus Pheropsophus.

Prothorax. As in subgenus Pheropsophus.

Pterothorax. As in subgenus Stenaptinus.

Abdomen. As in subgenus Pheropsophus .

Genitalia. Male as in subgenus Pheropsophus. Female styli very short and broad, unarmed,

otherwise as in subgenus Pheropsophus .

Etymology. — Aptinus = another brachinine genus; Greek, morphe = form or shape;

referring to the narrow, wingless condition of these beetles, reminding one of members of

Aptinus.

Distribution. — The species of this subgenus are confined to Madagascar.

Genus Pheropsophidius Hubenthal

(Figs. 4, 12, 35, 59, 423, 427, 437-442, 444, 445, 446)

Pheropsophidius Hubenthal, 1911: 547. Type species. — Cicindela aequinoctialis Linne,

1763:395 (subsequent designation by Jeannel, 1949: 1084).

Protopheropsophus Hubenthal, 1911: 548. Type species. — Pheropsophus biplagiatus

Chaudoir, 1876: 18 (by monotypy).

Brachinus (of authors).

Diagnostic combination. — Elytra costate, costae low and rounded, evenly flared into

depressions; propleural suture absent.

This genus includes the subgenera Pheropsophidius Hubenthal and Protopheropsophus

Hubenthal.

Subgenus Pheropsophidius Hubenthal

(Figs. 4, 12, 35, 59, 423, 440, 441, 442, 445)

Diagnostic combination. — As in subgenus Pheropsophus, except costae of elytra not car-

inate, lower, rounded, and evenly confluent with depressions; propleural sutures absent;

female styli short and broad.

Description. — Medium to large-sized beetles, about 12.0 to 20.0 mm.

Color. Yellow or ferrugineous with black markings.

Microsculpture. As in subgenus Pheropsophus .

Macrosculpture. As in subgenus Pheropsophus, except discal depressions with fewer micro-

Bombardier Beetles

37

rugosities.

Head. As in subgenus Pheropsophus.

Prothorax. As in subgenus Pheropsophus, except propleural sutures absent.

Pterothorax. As in subgenus Pheropsophus , except elytral costae as described above.

Abdomen. As in subgenus Pheropsophus .

Genitalia. Male (figs. 440, 441, 442) as in subgenus Pheropsophus. Female (fig. 59) styli

short and broad, otherwise as in subgenus Pheropsophus.

Etymology . — Greek, phero = bearer; psophos = sound; idion = little; referring to the

smaller, but Pheropsophus- like members of this genus.

Distribution. — The range of this subgenus extends from southern Mexico to South

America, as far south as 40° S in Argentina.

Subgenus Protopheropsophus Hubenthal

(Figs. 427, 437, 438, 439, 444, 446)

Diagnostic combination. — As in subgenus Pheropsophidius, except elytra scarcely cos-

tate on disc; anterior coxal cavities open behind; wings absent; humeri absent; apex of elytra

obliquely truncate; microrugosities of elytral depressions absent from disc.

Description. — Small to large-sized beetles, 1 1 .0 to 14.0 mm.

Color. Head, prothorax, venter around coxae, and legs ferrugineous. Elytra and epipleura

dull black, each with a large orange spot.

Microsculpture. As in subgenus Pheropsophus.

Macrosculpture. As in subgenus Pheropsophus, except microrugosities absent from disc.

Head. As in subgenus Pheropsophus .

Prothorax. As in subgenus Pheropsophidius, except anterior coxal cavities open behind.

Pterothorax. As in subgenus Stenaptinus, except elytra as described above.

Abdomen. As in subgenus Pheropsophus.

Genitalia. Male (figs. 437, 438, 439) as in subgenus Pheropsophus . Female styli (fig. 444)

short and broad, otherwise as in subgenus Pheropsophus.

Etymology. - Greek, proto = first; Pheropsophus, another genus of brachinines; referring

to Hubenthal’s idea that this is the most primitive group of Pheropsophina.

Distribution . — (Fig. 446). The members of this monotypic subgenus are known only

from the southern slopes of the Sierra Madre del Sur in Oaxaca, Mexico.

Subtribe Aptinina new subtribe

Type genus. — Aptinus Bonelli, here designated.

The following combination of characteristics is diagnostic of this subtribe: mandibular

scrobe unisetose; mentum with tooth on anterior edge; anterior coxal cavities uniperforate;

propleural suture absent; antennal comb spur internal; male anterior tarsi with articles 1-3

asymmetrical (figs. 43, 47); vestiture of male anterior tarsal articles diagonally arranged;

middle coxal cavities confluent; apical elytral membrane absent; metacoxal cavities conflu-

ent; coxites of female ovipositor setiferous apically.

This subtribe includes the genera Aptinus Bonelli, Styphlomerus Chaudoir, Styphlomer-

inus Jeannel, and Styphlodromus Basilewsky.

Key to the genera of the subtribe Aptinina

1 Integument black; dorsal pubescence sparse; wingless; humeri absent

Aptinus Bonelli

F Integument ferrugineous or yellow, elytra with black areas; dorsal pubescence

dense; winged; humeri square and prominent 2

2(1’) Mentum with large tooth on anterior edge 3

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2’ Mentum without large tooth on anterior edge Styphlodromus Basilewsky

3(2) Dorsal surface with large punctures forming a very rugose surface; elytra concolor-

ous Styphlomerinus Jeannel

3’ Dorsal surface finely and densely punctate; elytra bicolored

Styphlomerus Chaudoir

Genus Aptinus Bonelli

(Figs. 3, 17,36, 43,55,56,57, 58)

Aptinus Bonelli, 1810: tab. syn. Type species . — Brachinus bombarda Illiger, 1800: 112

(subsequent designation by Jeannel, 1942: 1116).

Aptinidius Jeannel, 1942: 1116. Type species. — Aptinus displosor Dufour, 1811: 70

(original designation). NEW SYNONYMY. I do not believe this species is sufficiently

different to warrant its placement in a separate monotypic subgenus.

Diagnostic combination. — Integument black (red prothorax in members of Aptinus

displosor Dufour); dorsal pubescence sparse; humeri absent; wings absent; male genitalia

variously contorted, endophallus with sclerite “complex”; female coxites plurisetose, setae

unmodified, bursa with sclerite “complex”.

Description. — Medium-sized beetles, 8.0 to 14.0 mm.

Color. Generally black, some species with rufous appendages and prothorax.

Microsculpture. Isodiametric on head, pronotum, and elytra, slightly transversely stretched

on pronotum.

Macrosculpture. Pronotum usually with deep setiferous punctures.

Head. Labrum entire to slightly lobate, with six or eight evenly spaced setae transversely

arranged near anterior margin. Clypeus rectangular, with numerous setiferous punctures

scattered on disc. Front smooth, but with numerous setae; furrows very shallow, with deep

groove separating them from antennal base cover and eye; one supraorbital seta over each

eye; bead over eye ending at hind margin of eye. Eyes prominent. Antennae long, narrow,

pubescent throughout. Mandibles (fig. 3) each broad, triangular, curved at tip; cutting edge

with one small terebral tooth and three retinacular teeth; ventral groove with short golden

setae; scrobe unisetose. Labial palpi (fig. 17) each with last article narrowly wedge-shaped;

terminal and penultimate article bearing numerous stiff setae. Maxillary palpi more or less

filiform; last three articles with numerous stiff setae. Ligula (fig. 17) membranous with an-

terior angles produced, and center ventrally produced into a plurisetose sclerotized convex-

ity. Mentum (fig. 17) well developed with a single entire or emarginate tooth at anterior

edge. Gula (fig. 17) widened behind.

Prothorax. As described under subtribe Aptinina, except that numerous, large, deep, seti-

ferous, punctures occur on disc. Anterior tibiae (fig. 36) each with subterminal spur internal,

at middle of comb emargination. Male anterior tarsal articles (fig. 43) 1-3 asymmetrical,

with ventral vestiture consisting of two parallel rows of spatulate setae, diagonally arranged

beneath each article. Female anterior tarsal articles symmetrical, vestiture absent.

Pterothorax. Anterior metathoracic process acute, mesocoxae contiguous-confluent. Hind

coxae contiguous. Metepimeron lobe very large, rounded apically. Elytral humeri strongly

sloping, lateral bead entire to base of humerus. Depression setae present between well devel-

oped costae. Epipleura wide medially and anteriorly, but narrowing toward apex; entirely

pubescent. Wings absent.

Abdomen. As described under division Brachinida, except one pair of ambulatory setae

at apices of sternites 2-5.

Genitalia. Male (figs. 56, 57) with small balteate, asymmetrical parameres; median lobe

variable, but highly asymmetrical and contorted; ligule single, large, and spatulate; endo-

phallus short, with two ventral microtuberculate plates and an apical sclerite. Female (fig.

Bombardier Beetles

39

58) with very short, broad, arcuate, and unarmed styli; valvifers robust and plurisetose;

bursa with complicated arrangement of sclerites (fig. 55).

Etymology . — Greek, apten = unable to fly; referring to the winglessness of all known

species of this genus.

Distribution. — The species of Aptinus occur in the southern parts of Europe from Spain

to the Black Sea, mostly in the mountainous regions of the Alps and Pyrenees.

Genus Styphlomerus Chaudoir

(Figs. 2, 16, 27, 39, 47, 48, 67, 68)

Styphlomerus Chaudoir, 1875: in Rapport sur un memoire de M. le Baron de Chaudoir, by

M. Putzeys. Type species. — Brachinus quadrimaculatus Dejean (subsequent designation

by Andrewes, 1939: 138)

Styphromerus Chaudoir, 1876: 88. Lapsus calami.

Diagnostic combination. — Mandibular scrobe unisetose; labial palpus with terminal article

cylindrical; mentum with a large median tooth; mandibles simple, each with one terebral

tooth and one terebral ridge; propleural suture absent; anterior tibiae each with subterminal

spur internal at middle of comb emargination; male anterior tarsal articles asymmetrical;

middle coxae contiguous-confluent; elytra not costate; elytral epipleura wide or narrow;

apical membrane of elytra absent; hind coxae confluent; metasternal process (between mid-

dle coxae) acute; wings present; parameres of male genitalia small, balteate; ligule double;

valvifers of female genitalia plurisetose; styli short, narrow, and arcuate.

Description. — Medium-sized beetles, 4.5 to 12.0 mm.

Color. Generally testaceous to yellow, many species with blackish elytra with testaceous

or yellow spots.

Microsculpture. Isodiametric on head, pronotum, and elytra.

Macrosculpture. Dorsal surface of head, pronotum, and elytra densely microrugose and

punctate.

Head. Labrum entire, with six evenly spaced setae transversely arranged on disc. Clypeus

rectangular with a single seta at each anterior corner and several small setae scattered over

disc. Front microrugose, with numerous setiferous punctures; furrows very shallow; one

supraorbital seta over each eye; bead over eye incomplete. Eyes prominent. Antennae long,

robust, pubescent throughout. Mandible (fig. 2) broad, triangular, curved at tip; terebral

margin with an elongate terebral ridge, and two retinacular teeth; ventral groove with short

golden setae; scrobe unisetiferous. Labial palpi (fig. 16) each with terminal article slightly

wedge-shaped; terminal and penultimate articles plurisetose, penultimate also bearing two

large setae. Maxillary palpi each more or less filiform; last three articles plurisetose. Ligula

(fig. 16) membranous and square with anterior angles produced, center ventrally produced

into plurisetose sclerotized convexity. Mentum (fig. 16) well developed, with a single tooth

at anterior edge. Gula (fig. 16) widened behind.

Prothorax. As described under subtribe Aptinina, except disc with microrugosities and

scattered punctures. Anterior tibiae (fig. 39) with subapical spur internal at middle of comb

emargination. Male anterior tarsal articles (fig. 47) 1-3 asymmetrical, vestiture beneath con-

sisting of two parallel rows of spatulate setae. Female anterior tarsal articles symmetrical,

vestiture absent.

Pterothorax. Anterior metathoracic process (between middle coxae) acute, prolonged,

middle coxae contiguous-confluent. Hind coxae confluent. Metepimeron lobe small, paral-

lel sided. Elytral humeri square, lateral bead entire to base of humerus. Depression setae

present, erect. Epipleura pubescent; wide medially and anteriorly, but either narrowing to-

ward apex or wide throughout. Wings present (fig. 27).

Abdomen. As described under division Brachinida, except one or more pairs of ambul-

40

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atory setae at apices of sterna 2-5.

Genitalia. Male (figs. 67, 68) with small, balteate, asymmetrical parameres; median lobe

various, but usually elongate, narrow, curved; apical sclerotization poorly defined, but

present; ligule double; endophallus short with numerous microtrichia scattered over sur-

face. Females with short, narrow, and arcuate, unarmed styli; coxites trapezoidal, thin;

valvifers plurisetose; bursa without sclerites.

Etymology . - Greek, styphlos = rough; meros = thigh; referring to the punctate femora

of these beetles.

Distribution. — The range of this genus extends on both sides of the equator in southern

and middle Africa.

Genus Styphlomerinus Jeannel

Styphlomerinus Jeannel, 1949: 1118. Type species. — Styphlomerus fuscifrons Fairmaire

1897: 367 (original designation).

Diagnostic combination. — As in Styphlomerus, except dorsum more strongly macropunc-

tate; elytra without spots (in known species); ligule single.

Description. — Small to medium-sized beetles, about 5.0 to 8.5 mm (Jeannel, 1949).

Color. Head, prothorax, venter, and legs ferrugineous; sometimes head and various scler-

ites infuscated. Elytra and epipleura dark grey or black.

Microsculpture. As in genus -Styphlomerus.

Macrosculpture. Dorsal surface coarsely punctate and rugose.

Head. As in genus Styphlomerus.

Prothorax. As in genus Styphlomerus.

Pterothorax. As in genus Styphlomerus, except epipleura not wide throughout then-

length.

Abdomen. As in genus Styphlomerus.

Genitalia. Male as in genus Styphlomerus, except ligule single. Female as in genus

Styphlomerus.

Etymology . — Greek, styphlos = rough, meros = thigh, inus = having the nature of; refer-

ring to the similarity between members of this genus and Styphlomerus.

Distribution. — The range of this genus extends from India to Japan, and south in the

Malayan Archipelago to Java, and on the islands of Madagascar, Hainan, and Taiwan. Accor-

ding to Jeannel (1949), no species inhabit continental Africa.

Genus Styphlodromus Basilewsky

Styphlodromus Basilewsky, 1959a: 240. Type species. — Styphlodromus bicolor Basilewsky,

1959a: 241 (original designation).

Diagnostic combination. - As in Styphlomerus, except mentum without tooth; dorsal

surface macropunctate; ligule doubled or not (nature of doubling different than in Styph-

lomerus)', elytra with stripes or spots.

Description. — Small to medium-sized beetles, 4.5 to 7.0 mm.

Color. Head, prothorax, venter, and legs ferrugineous. Elytra and epipleura dark gray to

black, variously spotted or striped.

Microsculpture. As in genus Styphlomerus.

Macrosculpture. As in genus Styphlomerinus.

Head. As in genus Styphlomerus.

Prothorax. As in genus Styphlomerus.

Pterothorax. As in genus Styphlomerus, except epipleuron not wide throughout its length.

Abdomen. As in genus Styphlomerus.

Genitalia. Male as in genus Styphlomerus, except ligule single or double, and endophallus

Bombardier Beetles

41

folding pattern more complex. Female as in genus Styphlomerus.

Etymology . — Greek, styphlos = rough; dromeus = runner; referring to the rapidity with

which these Styphlomerus-like beetles run.

Distribution. — The range of this genus extends from middle to southern Africa.

Remarks. — Although this genus is presently monotypic in the literature, I have seen

specimens representing four more species. I have not described and named these, because it

is possible they already have names and have been placed wrongly in the genus Styphlom-

erus or even Brachinus.

Subtribe Brachinina Bonelli

Type genus. — Brachinus Weber.

The following combination of characteristics is diagnostic of this sub tribe: mandibular

scrobes plurisetose; mentum without a tooth on anterior edge; anterior coxal cavities uni-

perforate, closed behind; propleural suture absent; anterior tibiae each with subterminal

spur external or only slightly internal at top of comb emargination; male anterior tarsal

articles either symmetrical or asymmetrical; vestiture beneath articles 1-3 longitudinally or

diagonally arranged; middle coxal cavities confluent; apical elytral membrane present; hind

coxal cavities confluent; coxites of female ovipositor glabrous apically.

This subtribe includes the genera Brachinus Weber, Aptinoderus Hubenthal, and Brachin-

ulus Basilewsky.

Key to the genera of the sub tribe Brachinina

1 Apical membrane of elytron fully developed, wider than sutural flange near apex

Brachinus Weber

1 ’ Apical membrane of elytron narrow, narrower than sutural flange near apex ... 2

2(1’) Integument black, apical articles of labial and maxillary palpi cylindrical

Aptinoderus Hubenthal

2’ Integument brown to ferrugineous, apical articles of labial and maxillary palpi

swollen proximally, acuminate and fusiform toward apex

Brachinulus Basilewsky

Genus Aptinoderus Hubenthal

Aptinoderus Hubenthal, 1919: 332. Type species. — Brachynus funebris Peringuey, 1898:

320 (here designated).

Brachynomorphus Hubenthal, 1919: 335. Type species. - Brachynus tibialis Peringuey,

1898: 321 (here designated). NEW SYNONYMY. I do not believe this species is suf-

ficiently different to warrant its placement in a separate genus.

Diagnostic combination . — Integument black; dorsum of head and pronotum with large,

deeply impressed setiferous punctures; antennal article 3 subequal to articles 1 and 2 com-

bined; anterior coxal cavities closed behind, proepimeron inserted into notch on prosternal

process; male anterior tarsal articles asymmetrical, vestiture beneath diagonally arranged;

subterminal spur of anterior tibia external on top of comb emargination; apterous; humeri

strongly sloped; apical membrane of elytra narrower than sutural flange; endophallus with-

out virga or sclerites.

Description. — Medium-sized beetles, about 10.0 to 1 1.0 mm (Hubenthal, 1919).

Color. Black, with legs and outer antennal articles ferrugineous. Elytra black (except in

cyanipennis, according to Hubenthal, 1919).

Microsculpture. Isodiametric, mostly effaced on head and pronotum, slightly beaded on

elytra.

Macrosculpture. Dorsum of head and pronotum coarsely and densely punctate, each

42

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puncture setiferous.

Head. As in subgenus Brachinus, except center of mentum convex.

Prothorax. As in subgenus Brachinus , except anterior coxal cavities closed behind, pro-

epimeron inserted into notch of prosternal process.

Pterothorax. As in subgenus Brachinus , except metasternum short, shorter than longi-

tudinal diameter of middle coxae; metepisternum short; apterous; humeri strongly sloped;

apical fringe of elytra consisting of short, closely spaced setae.

Abdomen. As in subgenus Brachinus.

Genitalia. As in subgenus Brachinus , except both parameres small, more balteate. Female

as in subgenus Brachinus.

Etymology. — Aptinus = another genus of bombardier beetles; Greek, deros = skin; refer-

ring to the black, Aptinus-hke integument of these beetles.

Distribution. — The range of this genus is restricted to extreme southern Africa.

Genus Brachinulus Basilewsky

Brachinulus Basilewsky, 1958: 96. Type species. - Brachinulus viettei Basilewsky, 1958:

97 (original designation and by monotypy). For details see above and Basilewsky 1962a:

126.

Diagnostic combination. — Terminal palpal article swollen proximally, acuminate and

fusiform toward apex; apical elytral membrane narrower than sutural flange, and with num-

erous small, and closely spaced apical setae.

Description. — Small-sized beetles, 5.0 mm. I have not seen specimens of this group, nor

does Basilewsky provide information on the characteristics which I have utilized for other

species.

Etymology. — Brachinus = another genus of bombardier beetles; Latin, ulus = small;

referring to the small, but Brachinus-Mke, form of these beetles.

Distribution . — These beetles are found only on Isla Principe, off the west coast of Africa.

Genus Brachinus Weber

Brachinus Weber, 1801: 22. Type species. — Carabus crepitans Linn6, 1758: 414 (subse-

quent designation by Latreille, 1810: 426).

Brachynus (of authors). (Unjustified emendation by Ahrens, 1812: t. 9)

Aploa Hope, 1833: 91. Type species. - Aploa pictus Hope, 1833: 91 (by monotypy).

Pseudaptinus Porta, 1909: 90 (nec Castelnau). Type species. — Brachinus italicus Dejean,

1831: 409 (subsequent designation by Jeannel, 1942: 1105).

Cnecostolus Reitter, 1919: 133. Type species. — Carabus exhalans Rossi, 1792: 84

(subsequent designation by Jeannel, 1942: 1105).

Brachynolomus Reitter, 1919: 133. Type species. — Brachinus immaculicornis Dejean,

1826: 466 (subsequent designation by Jeannel, 1942: 1105).

Brachynidius Reitter, 1919: 133. Type species. — Carabus scolopeta Fabricius, 1792: 136

(subsequent designation by Jeannel, 1942: 1105).

Brachinoaptinus Lutshnik, 1926: 43. New name for Pseudaptinus Porta (nec Castelnau).

Aptinomimus Alluaud, 1935: 25. Type species. — Pheropsophus microrrhabdus Alluaud,

1899: 381 (original designation).

Br achy nap tinus Csiki, 1933: 1628 (unjustified emendation).

Metabrachimv Jeannel, 1949: 1100. Type species. — Brachinus connectus Dejean, 1831:

417 (original designation).

Platy brachinus Jeannel, 1 949: 1 100. Type species. — Pheropsophus fasciatocollis Fairmaire,

1901 : 129 (original designation and by monotypy).

Note. — Much confusion has resulted from the two different ways in which Brachinus

Bombardier Beetles

43

(= Brachynus ) is spelled. The name is derived from the Greek brachyus, meaning short, and

refers to the truncate elytra. A correct transliteration of the word results in the name Brach-

ynus, however, Weber used Brachinus when he erected the genus (1801: 22). According to

the International Rules of Zoological Nomenclature (Articles 31a, 32c, and 33a), the Law of

Priority dictates that the name used must be as Weber first used it, that is Brachinus.

Diagnostic combination. — As given in the key.

This genus includes the subgenera Brachinus Weber, Aploa Hope, Cnecostolus Reitter,

Brachynolomus Reitter, Brachinoaptinus Lutshnik, Aptinomimus Alluaud, Metabrachinus

Jeannel, and Neobrachinus new subgenus.

Key to the subgenera of the genus Brachinus

Note. — At least four unnamed lineages are not included in this key, see Zoogeography

for details.

1 Apical edge of apical membrane of elytron with long widely spaced setae, the lat-

ter longer than width of membrane, and membrane with closely spaced fringe of

shorter setae Brachinus (sensu stricto)

1’ Apical membrane without long, widely spaced setae; fringe of shorter setae pre-

sent or not 2

2( 1 ’) Elytra with spotted or variegated color pattern 3

2’ Elytra concolorous or with sutural stripe contrasting in color with disc 5

3(2) Antennal article 3 longer than articles 1 and 2 combined; anterior coxal cavities

closed behind with prosternal process overlapped by proepimera

Metabrachinus Jeannel

3’ Antennal article 3 subequal to articles 1 and 2 combined; anterior coxal cavities

closed behind with proepimeron inserted into notch or socket of prosternal pro-

cess 4

4(3’) Upper spur of anterior tibia external Cnecostolus Reitter

4’ Upper spur slightly internal or in intermediate position Aploa Hope

5(2’) Antennal article 3 longer than articles 1 and 2 combined 6

5’ Antennal article 3 subequal to articles 1 and 2 combined 7

6(5) Range limited to New World and Sikkim Neobrachinus new subgenus

6’ Range limited to Madagascar. Aptinomimus Alluaud

7(5’) Upper spur of anterior tibia slightly internal; apical membrane of elytron without

setae; humeri sloped; wingless Brachinoaptinus Lutshnik

7’ Upper spur of anterior tibia external; apical membrane of elytron with fringe of

setae; humeri square, prominent; winged Brachynolomus Reitter

Subgenus Brachinus Weber (Fig. 23)

Diagnostic combination. — Antennal article 3 subequal to articles 1 and 2 combined;

anterior coxal cavities closed behind, proepimeron overlapping prosternal process; male

anterior tarsal articles 1-3 with vestiture beneath diagonally arranged; apical membranes of

elytra with several widely spaced setae, each longer than the apical membrane; left para-

mere large and triangulate; internal sac without virga or other sclerites.

Description. — Small to medium-sized beetles, 4.6 to 9.0 mm.

Color. Ferrugineous, with metallic blue elytra and epipleura. Various sclerites infuscated

or not.

Microsculpture. Isodiametric on head, pronotum, and elytra.

Macrosculpture. Head at vertex, pronotum, and elytral depressions usually rugose and

punctate.

Head. Labrum entire, with six setae transversely arranged on disc. Clypeus rectangular,

with two setae, one each side near middle, and numerous smaller setae scattered over disc.

44

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Front smooth, with a few scattered setae; furrows shallow; bead over eye continuous to be-

hind eye; one supraorbital seta over each eye; eyes usually prominent. Antennae long, fili-

form, pubescent throughout, pubescence denser on articles 3-1 1 than on 1 and 2. Article 3

subequal to articles 1 and 2 combined. Mandibles (as in fig. 8) each broad, triangular, curved

at tip, cutting edge with one terebral tooth, and four retinacular teeth; ventral groove with

short golden setae; scrobes plurisetose. Labial palpi (as in fig. 13) each filiform; terminal

article slightly swollen, with scattered pubescence; penultimate article plurisetose. Maxillary

palpi (as in fig. 9) each filiform, all articles pubescent. Ligula (as in fig. 13) membranous and

square, center ventrally produced into a setose sclerotized convexity. Mentum (as in fig. 13)

well developed, without tooth; center slightly concave or convex. Gula (as in fig. 13) wid-

ened behind.

Prothorax. As described under subtribe Brachinina. Anterior tibia (as in fig. 33) with sub-

terminal spur slightly internal. Male anterior tarsal articles (as in fig. 41) 1-3 asymmetrical,

vestiture beneath consisting of two diagonal, but parallel rows of setae. Setae of vestiture

spatulate, truncate, and rolled at apex. Female anterior tarsal articles symmetrical, vestiture

absent.

Pterothorax. As described under subtribe Brachinina.

Abdomen. As described under division Brachinida.

Genitalia. Male with asymmetrical balteate parameres, the left large and triangulate, the

right minute; median lobe various, but nearly symmetrical; ligule single, usually well de-

fined; endophallus moderately long, without sclerites or virga. Female as in subgenus Neo-

brachinus.

Etymology. — Greek, brachyus = short; referring to the short truncate elytra of these

beetles.

Distribution . — The range of this subgenus extends throughout Europe and North Africa.

Subgenus Brachynolomus Reitter

(Fig. 41)

Brachynidius Reitter, 1919: 133. NEW SYNONYMY. I do not consider the species placed

in this subgenus by various workers sufficiently different to warrant a separate subgeneric

taxon.

Diagnostic combination. - Antennal article 3 subequal to articles 1 and 2 combined;

anterior coxal cavities closed behind, proepimeron overlapping prosternal process; apical

membrane of elytra with numerous closely spaced setae, shorter than apical membrane;

both parameres small, balteate; male anterior tarsal articles asymmetrical; vestiture beneath

diagonally arranged; internal sac of median lobe with virga surrounding the gonopore.

Description. — Small to medium-sized beetles, about 4.5 to 10.0 mm (Jeannel, 1942).

Color. As in subgenus Brachinus, except some species with a ferrugineous sutural stripe

on elytra, and elytra usually vividly metallic.

Microsculpture. As in subgenus Brachinus.

Macrosculpture. As in subgenus Brachinus.

Head. As in subgenus Brachinus.

Prothorax. As in subgenus Brachinus.

Pterothorax. As in subgenus Brachinus.

Abdomen. As in subgenus Brachinus.

Genitalia. Male as in subgenus Brachinus, except both parameres small, more balteate;

endophallus with an apical virga. Female as in subgenus Neobrachinus.

Etymology . — Greek, brachyus = short; lomo = fringe; referring to the fringe of setae on

the apical membrane of the short elytra.

Distribution . — The range of this subgenus extends from northwestern Africa into south-

Bombardier Beetles

45

ern Europe, and east to Japan.

Subgenus Me tabrachinus Jeannel

Platy brachinus] eannel, 1949: 1100. NEW SYNONYMY. I do not believe the small forms

placed in this subgenus are sufficiently different to warrant their separation from the lar-

ger Me tabrachinus.

Diagnostic combination. — Antennal article 3 longer than 1 and 2 combined; anterior

coxal cavities closed behind, proepimeron overlapping prosternal process; male anterior tar-

sal articles asymmetrical, but less so than in subgenus Brachinus', elytra spotted; apical mem-

brane of elytron without setae; apical elytral membrane wide, wider than sutural flange;

internal sac without virga or sclerites.

Description. — Medium to large-sized beetles, 7.0 to 20.0 mm (Jeannel, 1949).

Color. As in subgenus Brachinus, except elytra with spotted color pattern, epipleura

usually yellow.

Microsculpture. As in subgenus Brachinus.

Macrosculpture. As in subgenus Brachinus.

Head. As in subgenus Brachinus, except mentum always convex at middle, and antennal

article 3 as described above.

Prothorax. As in subgenus Brachinus.

Pterothorax. As in subgenus Brachinus, except costae of elytra sharper, more elevated.

Abdomen. As in subgenus Brachinus.

Genitalia. Male as in subgenus Brachinus, except left paramere small and more balteate.

Female as in subgenus Neobrachinus.

Etymology . — Greek, meta, implying change; Brachinus, the common European genus;

referring to Jeannel’s idea that these beetles had diverged from the main Brachinus line of

evolution.

Distribution. — The range of this subgenus extends throughout southern Africa and Mad-

agascar.

Subgenus Aploa Hope

(Figs. 1,9, 15,31,37,46,60, 65,66)

Diagnostic combination. — Antennal article 3 subequal to articles 1 and 2 combined;

anterior coxal cavities closed behind, proepimeron inserted into notch of prosternal pro-

cess; subterminal spur of anterior tibia intermediately positioned on top of comb emargin-

ation; male anterior tarsal articles symmetrical; apical membrane of elytra without setae;

parameres of male genitalia small, balteate; ligule single, small; endophallus with heavily

sclerotized and pigmented subapical knife-shaped sclerite; valvifer of female genitalia glab-

rous; styli short, spatulate.

Description. — Medium-sized beetles, 12.0 to 14.0 mm.

Color. Generally yellow to testaceous, with black elytral markings.

Microsculpture. Isodiametric on head, pronotum, and elytra, almost effaced on head.

Macrosculpture. None.

Prothorax. As in subgenus Brachinus, except anterior tibiae (fig. 37) each with subtermin-

al spur in intermediate position at top of comb emargination. Male anterior tarsal articles

(fig. 46) 1-3 symmetrical, vestiture beneath consisting of two parallel rows of spatulate

setae. Female anterior tarsal articles symmetrical, vestiture absent.

Pterothorax. As in subgenus Brachinus.

Abdomen. As described for division Brachinida.

Genitalia. Male (fig. 65, 66) with small balteate, asymmetrical, parameres; median lobe

variable, but nearly symmetrical; ligule single, small, and narrow; endophallus short, with

46

Erwin

one large sclerite near apex; virga absent. Female with short, narrow, spatulate, slightly

curved, unarmed styli; coxites small, robust; valvifers glabrous; bursa unarmed.

Etymology. - Greek, haploos = single or simple; probably in reference to Hope’s know-

ledge of only one species of this genus.

Distribution . — The range of this subgenus extends on both sides of the equator in the Old

World. South of the equator the species are confined to southern Africa. In the north they

occur around the periphery of the Sahara Desert, and in (parts of) the Middle East, southern

Europe, India, and Ceylon.

Subgenus Brachinoaptinus Lutshnik

Pseudaptinus Porta, 1909: 90 (nec Castelnau). Primary homonym of Pseudaptinus Castel-

nau, 1834: 36, another genus of Carabidae.

Diagnostic combination. — Antennal article 3 subequal to articles 1 and 2 combined;

anterior coxal cavities closed behind, proepimeron inserted into socket of prosternal process;

subterminal spur of anterior tibia slightly internal on top of comb emargination; male anter-

ior tarsal articles symmetrical; apical membrane of elytra without setae; apterous, humeri

strongly sloping; both parameres of male genitalia small, balteate; endophallus without virga

or sclerites.

Description. — Small to medium-sized beetles 4.5 to 1 1.0 mm.

Color. As in subgenus Brachinus. At least one species ( Brachinus pygmaeus Dejean) has

members which are partially depigmented, pale brown throughout.

Microsculpture. As in subgenus Brachinus.

Macrosculpture. As in subgenus Brachinus.

Head. As in subgenus Brachinus , except eyes smaller, not so convex; extremely reduced

in Brachinus pygmaeus Dejean.

Prothorax. As in subgenus Brachinus , except as noted in diagnosis.

Pterothorax. As in subgenus Brachinus , except metasternum short, subequal to or shorter

than longitudinal diameter of middle coxa; humeri sloping, wings absent or greatly reduced.

Abdomen. As in subgenus Brachinus.

Genitalia. Male as in subgenus Brachinus , except both parameres small, strongly balteate;

endophallus without virga or sclerites. Female as in subgenus Brachinus.

Etymology. — Greek, brachyus = short; apten, unable to fly; referring to the Brachinus

species that cannot fly.

Distribution. — The range of this subgenus extends around the Mediterranean Sea.

Subgenus Aptinomimus Alluaud

Diagnostic combination. — Antennal article 3 elongate, longer than 1 and 2 combined;

anterior coxal cavities closed behind, proepimeron inserted into notch in prosternal process:

subterminal spur of anterior tibia external on top of comb emargination; male anterior

tarsal articles with ventral vestiture longitudinally arranged (according to Jeannel, 1949;;

endophallus not seen.

Description. — Medium to large-sized beetles, 9.5 to 22.0 mm (Jeannel, 1949).

Color. Brown, sometimes head and pronotum more ferrugineous; legs ferrugineous; elytra

and epipleura brown.

Microsculpture. As in subgenus Brachinus.

Macrosculpture. As in subgenus Brachinus.

Head. As in Brachinus (sensu s trie to), except terminal palpal article widened toward apex;

antennal article 3 longer than articles 1 and 2 combined.

Prothorax. As in subgenus Brachinus, except anterior coxal cavities closed behind, pro-

epimeron inserted into “notch” in prosternal process.

Bombardier Beetles

47

Pterothorax. As in Brachinus (sensu stricto), except metasternum short, subequal or shor-

ter than longitudinal diameter of middle coxa; metepisternum short; humeri strongly sloping;

wings absent; apical membrane with very short closely spaced setae.

Abdomen. As in subgenus Brachinus.

Genitalia. Male (see Jeannel, 1949: 1 1 12, fig. 544 a-f) as in subgenus Brachinus except

both parameres small. Female as in subgenus Brachinus.

Etymology. - Greek, apten, unable to fly; mimos = imitator, mimic; referring to the

superficially similar appearance of these beetles to members of the genus Aptinus.

Distribution. — This subgenus is restricted to Madagascar.

Subgenus Cnecostolus Reitter

Diagnostic combination. — Antennal article 3 subequal to articles 1 and 2 combined;

anterior coxal cavities closed behind, proepimeron inserted into notch of prosternal pro-

cess; subterminal spur of anterior tibia external on top of comb emargination; male anterior

tarsal articles with ventral vestiture longitudinally arranged; apical membrane of elytron

without setae; both parameres of male genitalia small, balteate; internal sac with virga around

gonopore.

Description. — Small to medium-sized beetles, about 4.0 to 12.0 mm (Jeannel, 1942;

Reitter, 1919).

Color. As in subgenus Brachinus, except elytra with spotted color pattern, at least one

species {Brachinus cruciatus Quensel) with brown head and pronotum.

Microsculpture. As in subgenus Brachinus.

Macrosculpture. As in subgenus Brachinus.

Head. As in subgenus Brachinus.

Prothorax. As in subgenus Brachinus, except anterior coxal cavities closed by notch inser-

tion of proepimeron into prosternal process; male anterior tarsal articles with longitudinally

arranged vestiture beneath.

Pterothorax. As in subgenus Brachinus, except apical membrane of elytra without fringe •

of setae.

Abdomen. As in subgenus Brachinus.

Genitalia. Male as in subgenus Brachinus, except both parameres small, more balteate;

endophallus with apical virga around gonopore. Female as in subgenus Brachinus.

Etymology. - Greek, knekos, yellow; stola, folds; referring to the yellow spots of the

costate elytra of these beetles.

Distribution. — The range of this subgenus extends from southern Europe (France), east

to at least Mongolia, mostly in more northerly latitudes.

Subgenus Neobrachinus new subgenus

Type species. - Carabus fumans Fabricius, 1781: 307, here designated.

Diagnostic combination. — Antennal article 3 longer than articles 1 and 2 combined;

anterior coxal cavities closed behind; proepimeron inserted into socket of prosternal pro-

cess; male anterior tarsal articles with ventral vestiture longitudinally arranged; apical mem-

brane of elytra with numerous, closely spaced, short setae; both parameres small, balteate,

the left larger than the right; endophallus with a virga surrounding the gonopore.

Description. - Small to large-sized beetles, 4.8 to 18.5 mm.

Color. Variable, but usually head and prothorax ferrugineous and elytra blue or brown.

Microsculpture. As in subgenus Brachinus.

Macrosculpture. As in subgenus Brachinus.

Head. As in subgenus Brachinus, except mentum of some species with one or two pits

surrounded by setae; antennal article 3 longer than articles 1 and 2 combined.

48

Erwin

Prothorax. As in subgenus Brachinus, except anterior coxal cavities closed behind, pro-

epimeron inserted into socket in prosternal process.

Pterothorax. As in subgenus Brachinus, except some species apterous as in members of

subgenus Brachinoaptinus .

Abdomen. As in subgenus Brachinus.

Genitalia. As in subgenus Brachinus, except both parameres small, more balteate, and left

larger than right; virga present at tip of endophallus. Female as in subgenus Brachinus.

Etymology . — Greek, neo, new or most recent; brachyus, short; referring to the pro-

bability that these beetles are the most recently derived of an Old World stock that invaded

the New World.

Distribution. — The range of this subgenus extends throughout the New World from

about 52°N to 40° S latitude. One relict species occurs in Sikkim, in the Himalaya Mountains.

Key to bombardier beetles of North and Middle America

Characteristics used in the keys

The more difficult to use characteristics in the key are further discussed here (see also

comparative morphology).

Color differences are very useful in the identification of bombardier beetles, especially

those of North and Middle America. For the most part, I have used the presence or absence

of a color on various sclerites, rather than comparing shades of colors. Shades of color are

used only for the elytra. In a very few species elytral color is so variable as to be useless for

identification. Color combinations are usually constant. Very teneral adults do not have

pigments deposited, but these are rarely collected.

In North and Middle America the head of Brachinus beetles is usually ferrugineous, but

four Neotropical species have brown heads, and one eastern United States species has a black

head. The mouth parts generally match the color of the head, but in four species with ferru-

gineous head color, the palpi are infuscated or black. The antennal articles 2 to 4 are quite

variable, ranging from ferrugineous to black, with intermediate stages of infuscated apically

and totally infuscated. The scape matches the head color. Antennal articles 5-1 1 are usually

dusky in ferrugineous colored species, but specimens of some species have infuscated lateral

stripes on each article, or each article is entirely infuscated.

The prothorax always matches the color of the head. The elytra are either blue, brown,

black, green, or slate-colored. In some species the epipleura of some specimens are test-

aceous, but usually the epipleura match the elytral color. The sutural costa varies from fer-

rugineous to black. In two species in the study area, the ferrugineous color includes two

costae and their adjacent depressions.

The venter is usually ferrugineous at the middle of the mesosterna and metasterna. The

sides of the metasterna, mesepisterna, mesepimera, metepisterna, and the abdominal sterna

and terga may be ferrugineous, infuscated, black, or (in two species) black with metallic

blue luster. In species without the ferrugineous ground color, the various ventral sclerites

are either infuscated or not. In species with black abdomens, the paramedial “dimples” are

usually ferrugineous. The legs may be either ferrugineous, testaceous, brown, or black.

Usually, testaceous colored legs have infuscated femoral apices (“knees”). This knee color

also occurs in some ferrugineous-legged species. In some ferrugineous species, the tibiae and

tarsi may be infuscated or black, and occasionally the femora also may be black.

Characteristics of pubescence and density of setae are very useful in species identification.

The number of setae on the mentum and submentum of North and Middle American bom-

bardier beetles is variable. Usually, the mentum has two long setae (fig. 19), and the sub-

mentum has about 8 to 12 setae transversely arranged (fig. 19). A second type of setal ar-

rangement exists with the center of the mentum having a small patch of setae (fig. 21). One

Bombardier Beetles

49

or two setae do not constitute a patch. A third type of setal arrangement exists when the

setae of the submentum are twice as abundant (about 16-24). A fourth type of setal arrange-

ment exists when there is a patch on the mentum, and also a doubling of the number of sub-

mental setae (fig. 2 1 ).

The setal arrangement on the proepisternum and proepipleura (= hypomera, Habu,1967)

is useful for identifying members of many species. Several combinations of setal patterns

may occur on these sclerites. In members of some species, the setae are so numerous that

they constitute pubescence. In others, only one or two setae occur on the entire lateral side

of the prothorax. Members of some species have a few scattered setae both anteriorly and

posteriorly on each lateral sclerite of the prothorax. In other species, the proepipleuron is

glabrous, while the proepisternum has setae both anteriorly and posteriorly. In members of

a few species, the proepisternum is completely pubescent, while the proepipleuron is gla-

brous.

The pattern of short pubescence on the elytra is used very frequently in the following key

to North and Middle American bombardier beetles. In all species, erect depression setae,

eighth interval umbilicate setiferous punctures, and scutellar umbilicate setiferous punctures

are present, in addition to the short, densely placed setae that are referred to below as “pub-

escence”. Several elytral patterns of pubescence are exhibited. In most species the elytra are

completely pubescent. In others, the pubescence is restricted to the depressions between the

costae, while the costae are smooth and glabrous. In some western species, the pubescence is

restricted to intervals (area between rows of erect depression setae) 6, 7, and 8 at the middle

of the elytra, but also occurs around the scutellar region and extends completely across the

elytra in the apical sixth. An even greater decrease in the amount of pubescence is seen in

other western and tropical species, where it is restricted to the 8th interval, and the scutellar

region. Only the members of Brachinus costipennis Motschulsky have entirely glabrous

elytra.

The macrosculpture of the anterior surface of the anterior tibia is used here occasionally

to differentiate species. Normally the surface is densely strigose with the strigae extending

longitudinally, however, in a few species the surface is smooth except for some small punc-

tures. In others, the punctures are elongate, and coalescent, looking like shallow strigae.

The height of the elytral costae is used to differentiate a few species. It is a difficult char-

acter to describe, but is not hard to see.

The most important characteristics are those of the male genitalia. To identify males of

many species, the apex of the shaft is all that need be observed. To identify males of some

species the virga of the endophallus must be examined. I have illustrated the male genitalia

of all species in which the male is known. The illustrations should be used in conjunction

with the descriptions provided with the subgenus Neobrachinus, and can be relied upon to

provide an absolute identification.

Key to the species of Brachinus and Pheropsophidius of North and Middle America

1 Mandibles with unisetose scrobes; suture present between proepisternum and pro-

epimeron (. Pheropsophidius ) 2

1’ Mandibles with plurisetose scrobes; no suture between proepisternum and pro-

episternum and proepimeron ( Brachinus ) 3

2(1) Elytron with all intervals strongly costate; depressions between costae minutely

but coarsely sculptured; wings fully developed

P. (Pheropsophidius) aequinoctialis (Linne), p. 165

2’ Elytron with intervals 5-8 strongly costate at apex, and interval 8 costate just

behind humerus, otherwise scarcely costate; each elytron with a large orange spot

at center; wingless. P. (Protopheropsophus) biplagiatus Chaudoir, p. 165

50

Erwin

3(1’) Elytron with ferrugineous sutural stripe, contrasting strongly with the otherwise

darkly colored disc; elytral epipleura pale 4

3’ Elytra concolorous, epipleura pale or dark 5

4(3) Legs (at least tibiae) pale, with dark knees; palpi pale; elytra usually greenish . . . .

B. cinctipennis Chevrolat, p. 97

4’ Legs totally black; palpi infuscated to black; elytra usually bright blue

B. cibolensis new species, p. 98

5(3’) Head, prothorax, and elytra concolorous, either black or brown 6

5’ Head and prothorax testaceous or ferrugineous, strongly contrasting with darkly

colored elytra 9

6(5) Body completely black; elytra black with blue luster; wingless

B. capnicus new species, p. 60

6’ Body brownish with testaceous legs and infuscated knees; winged 7

7(6’) Mentum with two pits surrounded by numerous long setae (fig. 20)

B. sallei Chaudoir, p. 69

7’ Mentum flat to shallowly biconcave, with only two setae 8

8(7’) Elytral pubescence confined to depression 8 (Mexico)

B. melanarthrus Chaudoir, p. 72

8’ Elytral pubescence not confined to depression 8 (Greater Antilles)

B. brunneus Castelnau, p. 71

9(5’) Mentum with a large deep median sulcus surrounded by a ring of setae (fig. 18) . .

10

9’ Mentum flat to shallowly biconcave OR with two shallow lateral pits (fig. 19) . . .

11

10(9) Elytra glabrous; venter ferrugineous B. costipennis Motschulsky, p. 85

10’ Elytra pubescent; venter mostly infuscated B. mobilis new species, p. 159

11(9’) Wings rudimentary, each a narrow elongate pad; metasternum short between

middle and hind coxae, no longer than diameter of middle coxa (fig. 26); humeri

strongly sloped (fig. 72) 12

11’ Wing with at least reflexed apex; metasternum longer than diameter of middle

coxa; humeri sloped, square (fig. 98) or protruding 14

1 2( 1 1 ) Abdomen, metepistema, and sides of metasternum infuscated to black; abdominal

sterna with ferrugineous paramedian dimples; mentum without central setal patch

(fig. 19) 13

12’ Abdomen infuscated only at sides, rest of venter ferrugineous; paramedian dimples

not of a contrasting color; mentum with central setal patch (fig. 21)

B. microumericcnus Erwin

13(12) Submentum densely setiferous (16 or more setae, fig. 21); larger, more robust

beetles; antennae ferrugineous B. americanus ((LeConte), p. 55

13’ Submentum sparsely setiferous (10 or less, fig. 19); smaller beetles; antennal

articles 3 and 4 infuscated B. alexiguus new species, p. 57

1 4( 11’) Elytra brown; legs pale testaceous with dark knees 15

14’ Elytra blue, blue-black, greenish-blue, or slate colored; legs concolorous without

darkly colored knees, or ferrugineous with darkly colored knees 22

15(14) Proepipleura and proepisterna pubescent throughout; venter pale; elytra not true

brown (in part) B. sonorous new species, p. 163

15’ Proepipleura glabrous; proepisterna with only a few scattered setae anteriorly

and/or posteriorly, or glabrous 16

16(15’) Elytral epipleuron at least at humerus pale testaceous, strongly contrasting with

elytral color 17

1 6’ Elytral epipleura and disc of elytra concolorous 19

Bombardier Beetles

51

17(16)

17’

18(17’)

18’

19(16’)

19’

20(19’)

20’

21(20’)

21’

22(14’)

22’

23(22)

23’

24(23)

24’

25(23’)

25’

26(25’)

26’

27(26’)

27’

28(22’)

28’

29(28)

29’

Metasternum at sides usually infuscated; elytral disc glabrous; median lobe as in

figs. 156, 157, 158; stylus as in fig, 145; in United States, west of Rocky Mount-

ains only B. lateralis Dejean, p. 73

Metasternum not infuscated at sides; elytral disc usually sparsely pubescent; in

United States, east of Rocky Mountains only 18

Anterior third of proepisternum with a few setae; median lobe as in figs. 153, 1 54,

155; stylus as in fig. 142; range — United States, Mexico, Central America, and

Cuba B. adustipennis Erwin, p. 81

Anterior third of proepisternum glabrous; median lobe as in figs. 147, 148, 149;

stylus as in fig. 144; range — Mexico, South America . . B. aeger Chaudoir, p. 77

Larger beetles, greater than 15.0 mm in length B. grandis Brulle, p. 73

Smaller beetles, less than 12.0 mm in length 20

Elytra prominently costate; median lobe as in figs. 159, 160, 161 ; stylus as in fig.

143 B. chalchihuitlicue new species, p. 79

Elytra almost smooth, costae barely elevated 21

Median lobe as in figs. 150, 151, 152; stylus as in fig. 146; elytral pubescence

usually extensive, covering apical third of elytra . . . B. arboreus Chevrolat, p. 79

Median lobe as in figs. 162, 163, 164; stylus as in fig. 141; elytral pubescence

usually discontinuous near middle of apical third of elytra

B. chirriador new species, p. 80

Elytral pubescence in outer depressions 6, 7, 8, or just in 8; less dense pubescence

usually across apical third of elytra and sometimes in vicinity of scutellum .... 23

Elytral pubescence all along depressions between costae, usually costae pubescent.

28

Elytral pubescence confined to depression 8, behind humerus and at middle,

although some setae scattered across apical sixth of elytron 24

Elytral pubescence in depressions 6, 7, and 8, and across apical sixth of elytron . .

25

Venter mostly ferrugineous, except for infuscated metepisterna and (in some

specimens) sides of abdomen; elytral costae prominent; microsculpture of pro-

notum isodiametric, surface rugose and shining B. gebhardis Erwin, p. 131

Venter infuscated; elytral costae barely elevated; microsculpture of pronotum

granulate, surface rugose and dull, color milky ferrugineous

B. galactoderus new species, p. 132

Tarsi, tibiae, and apices of femora infuscated to black

(in part) B. rhytiderus Chaudoir, p. 63

Tarsi, tibiae, and femora ferrugineous 26

Venter and antennae ferrugineous B. pallidus Erwin, p. 95

Venter and usually antennal articles 3 and 4 infuscated to piceous 27

Elytra shiny black, very convex B. explosus new species, p. 161

Elytra blue, micro sculpture coarse, surface dull

B. mexicanus Dejean, p. 104

Median elevated portion of mentum with a dense patch of setae (fig. 21); submen-

tum densely setiferous (more than 20 setae) 29

Median raised portion of mentum either glabrous or with one or two small setae

(fig. 19); submentum various 33

Metasternum anterior to antecoxal piece, subequal in length to longitudinal dia-

meter of middle coxa (fig. 26); antennal articles 3 and 4 infuscated at least apical-

ly; elytra usually greenish B. viridipennis Dejean, p. P0

Metasternum longer than middle coxa; antennal articles 3 and 4 usually ferrugin-

eous (except some populations of B. alternans)\ elytra bluish 30

52

Erwin

30(29’)

30’

31(30)

31’

32(31)

32’

33(28’)

33’

34(33)

34’

35(34’)

35’

36(35)

36’

37(35’)

37’

38(37)

38’

39(38)

39’

40(37’)

40’

41(40)

41’

42(41)

42’

43(42)

43’

44(42’)

Proepipleura pubescent throughout their length; proepisterna completely pubes-

cent 31

Proepipleura and proepisterna pubescent only anteriorly and posteriorly, glabrous

at middle B. perplexus Dejean, p. 141

Anterior tibia with anterior surface coarsely strigose; elytral costae barely ele-

vated 32

Anterior tibia with anterior surface punctate, rarely with punctures coalescing;

elytral costae very pronounced, easily visible with unaided eye

B. altemans Dejean, p. 88

Humeral angles square (fig. 98) prominent; elytra broad and quadrate with lateral

margins behind humeri straight to at least middle

B. imperialensis Erwin, p. 143

Humeri rounded, not at all prominent; elytra narrow, with lateral margins behind

humeri evenly arcuate entirely to apex B. velutinus Erwin, p. 142

Abdominal sterna entirely ferrugineous OR infuscated only marginally, with a

pale center extending to apex OR mostly ferrugineous except for infuscated later-

al margins and sternum 6 32

Abdominal sterna infuscated to piceous, rarely center of sterna 2 and 3 paler, but

usually not equal in color to the hind coxae 46

Erect depression setae at least twice as long as elytral pubescence

B. cyanipennis Say, p. 127

Erect setae subequal to or shorter than elytral pubescence 35

Abdominal sterna completely ferrugineous 36

Abdominal sterna with infuscated margins 37

Proepipleura glabrous; proepisterna with a few scattered setae; antennae ferrug-

ineous B janthinipennis (Dejean), p. 156

Proepipleura pubescent; proepisterna pubescent; antennal articles 3 and 4 infus-

cated (in part) B. sonorous new species, p. 163

Pronotum without lateral setae at middle 38

Pronotum with lateral setae present 40

Proepipleura densely pubescent throughout their length 39

Proepipleura with at most a few setae anteriorly

B. kansanus LeConte, p. 83

Elytral depression 1 with erect depression setae at least twice as long as elytral

pubescence; pronotum not densely pubescent

B. oaxacensis new species, p. 1 17

Elytral depression 1 with short depression setae; pronotum densely pubescent . . .

B. hirsutus Bates, p. 93

Elytra moderately to strongly costate 41

Elytra barely costate, almost smooth B. ovipennis LeConte, p. 121

Pronotum completely pubescent 42

Pronotum mostly glabrous, at most with a few scattered setae

(in part) B. rhytiderus Chaudoir, p. 63

Proepipleura glabrous; proepisterna with scattered setae both anteriorly and pos-

teriorly, glabrous medially 43

Proepipleura with at least some setae posteriorly, proepisterna (except fumans)

pubescent throughout 44

Elytra bright blue, usually metallic .... (in part) B. elongatulus Chaudoir, p. 65

Elytra slate-grey with greenish luster. (in part) B. texanus Chaudoir, p. 60

Pronotum (fig. 336) densely covered with large pits, surface very rugose; mentum

with numerous accessory setae scattered over surface

B. favicollis Erwin ,p. 1 40

Bombardier Beetles

53

44’ Pronotum (figs. 334-337) at most with fine punctures, not at all rugose; mentum

without accessory setae (occasionally one or two small setae at middle) 45

45(44’) Anterior tibia with anterior surface strigose; elytra with costae moderately ele-

vated B. puberulus Chaudoir, p. 140

45’ Anterior tibia with anterior surface punctate, the punctulae sometimes coalescent,

but not forming strigae; elytra with highly elevated costae

B. fumans Fabricius, p. 134

46(33’) Venter with metallic blue luster, tibiae and tarsi black; antennal articles 2-1 1 pic-

eous to black; palpi black; elytra brilliant metallic blue

B. azureipennis Chaudoir, p. 1 15

46’ Venter without metallic blue luster, infuscated to blackish or not, otherwise com-

bination of characteristics not as above 47

47(46’) Proepipleura glabrous 48

47’ Proepipleura with at least a few setae, either throughout their length or at both

ends (observe both sides of beetle) 66

48(47) Metasternum infuscated at sides 49

48’ Metasternum not infuscated at sides 61

49(48) Tibiae and tarsi infuscated, at least darker than femora 50

49’ Tibiae and tarsi concolorous with femora 51

50(49) Metasternal process (between middle coxae) usually infuscated; antennal article

3 shorter than diameter of eye; median lobe not ridged ventrally (fig. 257); stylus

narrow (fig. 275) B. phaeocerus Chaudoir, p. 1 1 1

50’ Metasternal process ferrugineous; antennal article 3 longer than diameter of eye;

median lobe ridged ventrally (fig. 240); stylus broad, spatulate (fig. 247)

(in part) B. quadripennis Dejean, p. 99

51(49’) Mesepisterna infuscated to black; elytra strongly costate

B. tenuicollis LeConte, p. 123

51’ Mesepisterna not darkly infuscated (if at all); elytra not or only moderately

costate 52

52(51’) Elytra with costae easily visible; elytral pubescence not dense 53

52’ Elytra without costae; elytral pubescence very dense; elytral color dull slate-blue .

B. sublaevis Chaudoir, p. 149

53(52) Metasternum subequal in length to diameter of middle coxa; humeri sloped; later-

al margin of elytron behind humerus arcuate to apex

B. patruelis LeConte, p. 1 17

53’ Metasternum longer than diameter of middle coxa; humeri square or prominent,

margin behind humerus straight at least to middle of elytra 54

54(53’) Anterior tibia with anterior surface punctate, punctulae small, rarely coalescing . .

55

54’ Anterior tibia with anterior surface strigose 56

55(54) Median lobe with ventral ridge (fig. 237); stylus acute (fig. 250)

B. neglectus LeConte, p. 1 10

55’ Median lobe without ventral ridge (fig. 404); stylus rounded apically (fig. 410) . .

B. vulcanoides Erwin, p. 155

56(54’) Proepipleura and proepisterna glabrous (proepisterna rarely with one to three

setae along anterior edge) (in part) B. medius Harris, p. 129

56’ Proepipleura and proepisterna with pubescence both anteriorly and posteriorly,

glabrcus medially 57

57(56’) Antennal article 3 infuscated throughout 58

57’ Antennal article 3 infuscated apically B. cyanochroaticus Erwin, p. 147

58(57) Median lobe with ventral depression (fig. 383) OR ridged ventrally (fig. 240);

54

Erwin

stylus very broad, spatulate (figs. 247, 386) 59

58’ Median lobe without depression or ridge; stylus acute apically 60

59(58) Median lobe with ventral depression (fig. 383); very long, narrow beetles with long

legs and antennae; elytra elongate; pronotum (fig. 370)

B. ichabodopsis new species, p. 150

59’ Median lobe with ventral ridge (fig. 240); short broad beetles; elytra quadrate;

pronotum (fig. 230) B. quadripennis Dejean, p. 99

60(58’) Range — north of latitude 35° N from New England to Indiana; median lobe (figs.

407, 408, 409); stylus (fig. 411) B. fulminatus Erwin, p. 153

60’ Range — south of latitude 35° N from Florida to Missouri; median lobe (figs. 398,

399, 400); stylus (fig. 412) B. oxygonus Chaudoir, p. 151

61(48’) Elytra with major part of pubescence in rows between costae 62

61’ Elytra with pubescence evenly scattered over surface, costae also pubescent . . 63

62(61) Elytra bright blue, usually metallic .... (in part) B. elongatulus Chaudoir, p. 65

62’ Elytra slate-grey with greenish luster (in part) B. texanus Chaudoir, p. 60

63(61’) Proepisterna glabrous, at most with 1-3 setae near anterior edge

(in part) B. medius Harris, p. 129

63’ Proepisterna pubescent, at least anteriorly and posteriorly 64

64(63’) Antennal article 3 extensively infuscated (sometimes also article 2) 65

64’ Antennal article 3 ferrugineous (apex of 4 sometimes lightly infuscated)

(in part) B. cordicollis Dejean, p. 144

65(64) Humeral angle square, elytral margin behind humerus straight, at least to middle;

range — Florida. B. conformis Dejean, p. 1 19

65’ Humeri sloped, margin behind humerus arcuate to apex; range — Arizona

B. imporcitis new species, p. 1 14

66(47’) Metasternum infuscated at sides 67

66’ Metasternum not infuscated at sides 69

67(66) Submentum densely setiferous (more than 20 setae); larger beetles, longer than

12.0 mm and wider than 5.0 mm across elytra at widest part

B. javalinopsis new species, p. 109

67’ Submentum sparsely setiferous (10 or less); smaller beetles, less than 1 1.5 mm in

length and 4.9 mm in width 68

68(67’) Knees infuscated; elytral pubescence evenly distributed over surface; pronotum

narrow (fig. 421) B. aabaaba new species, p. 161

68’ Knees not infuscated; elytral pubescence in rows between barely elevated costae;

pronotum cordiform (fig. 226) B. kavanaughi Erwin, p. 108

69(66’) Outer antennal articles and sutural costae of elytra black, strongly contrasting

with color of elytra B. consanguineus Chaudoir, p. 1 16

69’ Outer antennal articles ferrugineous and sutural costae of elytra bluish, not con-

trasting with color of elytra 70

70(69’) Proepisterna pubescent anteriorly and posteriorly, glabrous medially; ligule of

median lobe broad, spalutate (fig. 182); stylus very narrow, elongate (fig. 192)

B. rugipennis Chaudoir, p. 91

70’ Proepisterna pubescent throughout; ligule of median lobe paralleliform (fig. 378);

stylus wider, shorter (fig. 386 B. cordicollis Dejean, p. 144

The subgenus Neobrachinus new subgenus

in North and Middle America

The americanus group

The members of this group are characterized by the following: pouch-shaped virga, en-

larged prothorax, and reduced wings together with the modifications of the metathorax.

Bombardier Beetles

55

Four species are included, in two subgroups.

The americanus subgroup

The three species included here, B. americanus (LeConte), B. microamericanus Erwin,

and B. alexiguus new species differ from the species included in the following subgroup

only in overall color, but this color difference is of major importance. The specks included

here are similar in color to the majority of North American brachinines, that is, head and

pronotum ferrugineous, elytra blue, with various sclerites infuscated, depending upon the

species.

Brachinus americanus (LeConte)

(Figs. 71,79,80, 81,87,88)

Aptinus americanus LeConte, 1844: 48. Lectotype, here selected a female, MCZ red type

label number 5839, further labelled with a yellow disc and “72”. Type locality. —

Georgia, as originally given by LeConte.

Aptinus americanus Dejean, 1836: 13. NOMEN NUDUM.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Medium-sized beetles, 8.0 to 13.0 mm.

Color. Metepisterna, sometimes metasternum at sides, and abdominal sterna and terga

infuscated, otherwise ferrugineous. Dorsal surface and epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows rugose and punctate. Surface of pronotum with numer-

ous setiferous punctures, punctures moderately impressed.

Head. Frontal furrows moderately impressed. Antennal scape nearly cylindrical, widened

a little apically. Ligula with sclerotized center area ellipsoid-convex with numerous setae

scattered over its surface. Mentum and submentum without accessory setae.

Prothorax. Pronotum (fig. 71) slightly convex, flattened along center line, sides slightly

reflexed. Proepipleura glabrous. Proepisterna with a few setae anteriorly and posteriorly,

glabrous medially. Anterior tibia with anterior surface punctate.

Pterothorax. Elytra elongate, narrow, moderately costate. Humeri narrow, strongly

sloped. Costae smooth, depressions pubescent. Hind wings reduced, elongate pads. Meta-

sternum short, subequal to or shorter than diameter of middle coxa (fig. 26).

Abdomen. As described for genus.

Genitalia. Male (figs. 79, 80, 81). Median lobe with plane of shaft barely rotated from

plane of basal bend. Basal bend moderately long. Shaft slightly swollen medially. Apex of

shaft narrowed, rounded apically. Ligule short, narrow, rounded apically. Virga (figs. 79,

80). Female (fig. 87). Stylus short, narrow, narrowly rounded at apex.

Variation. — Besides the intrapopulational variation in the shape of the pronotum and

total size, these beetles vary in the amount of reduction of the elytral humeri and in the

amount of infuscation of the metasterna at the sides.

Flight. — The reduction of wing membrane makes these beetles incapable of flight.

Etymology . — The latinized form of America, the place where the types were collected.

Collecting notes. — W. Whitcomb (per. comm.) collected these beetles in Arkansas corn

fields. There is no reason to believe that these beetles are restricted to areas near water (as

are other species of Brachinus in North America).

Life history. — Members of this species have been collected from February to October. I

have seen one teneral specimen collected in September from Hot Springs, Arkansas. Over-

wintering is probably as an adult as in B. pallidus Erwin (Erwin, 1967).

Distribution. - (Fig. 88). The range of this species extends from Texas north to Minn-

esota, east to New York, and south to Florida. I have seen 440 specimens from the follow-

56

Erwin

79 80 81 82 83 84

Figs. 71, 73, 74, 78. Pronotum, right half, dorsal aspect. 71. Brachinus americanus (LeConte), Ludlow, Mississippi.

73. Brachinus capnicus new species, Smokemont, North Carolina. 74. Brachinus alexiguus new species, College Sta-

tion, Texas. 78. Brachinus microamericanus Erwin, Dundee, Mississippi. Fig. 72. Right elytron, dorsal aspect of

humeral angle, Brachinus capnicus new species, Smokemont, North Carolina. Figs. 75-77 , 79-84. Male genitalia.

75. Brachinus alexiguus new species, College Station Texas, ventral aspect. 76. Lateral aspect of same. 77. Dorsal as-

pect of same. 79. Brachinus americanus (LeConte), Washington County, Arkansas, ventral aspect. 80. Lateral aspect

of same. 81. Dorsal aspect of same. 82. Brachinus microamericanus Erwin, Dundee, Mississippi, ventral aspect.

83. Lateral aspect of same. 84. Dorsal aspect of same. Accompanying scale lines equal 1.0 mm.

Bombardier Beetles

57

ing localities:

UNITED STATES

ALABAMA: Blount County (Blount Springs) CMPP; Fayette County (Berry) UASM; Lee County (5.0 miles north of

Auburn) AUAA; Tuscaloosa County (Peterson) UASM. ARKANSAS: Benton County (Rogers) KSU; Garland County

(Hot Springs) CAS, SJSC; (Hot Springs-Lake Hamilton) SJSC; Hempstead County (Hope) UMAH; Izard County UAFA;

Lawrence County (Imboden) LACM, MCZ; Logan County (Mount Magazine) CNHM; Phillips County (West Helena)

UMAH; Polas County (Ouachita Mountains) UASM; Sebastian County (Fort Chaffee) RCGr; Scott County UAFA; Wash-

ington County ISNH, UAFA; (Cove Creek Valley) RFre, UAFA; (Goshen) DRWh; (Mount Sequoyah) ISNH; County

unknown (Knob Hill Ranch, Ozark Mountains) CAS. FLORIDA: (No locality given) WSUP. GEORGIA: (No locality

given) MCZ. ILLINOIS: Alexander County (Olive Branch) CNHM; Champaign County (Urbana) ISNH; Cook County

(Chicago area) CNHM, (La Grange) CAS, UMAH,USNM, (Palos Park) CAS, CNHM, UMAH, (Riverside) UMAH, (Summit)

CNHM, ISNH, (Willow Springs) CAS, CNHM, UASM, UMAH, ZMLS; Hardin County (Junction Highway 34-146) RTBe;

La Salle County RTBe; Putnam County ISNH; Richland County (Olney) ISNH; Will County (Beecher) CNHM; Counties

unknown (Bowmanville) CAS, (Falling Spring) LACM. INDIANA: Crawford County CAS; Franklin County (Metamora)

UMAH; Harrison County PUM; Jefferson County (Clifty Falls State Park) UASM; Knox County PUM; Lake County (Pine)

USNM; Lawrence County (Bedford) PUM ; Monroe County ISUA, (Bloomington) UMAH; Morgan County (Morgan-Monroe

county line) CEWh; Posey County (Hovey Lake) PUM; Ripley County (Versailles) CEWh; Vigo County PUM. IOWA:

Boone County (Ledges State Park) ISUA; Dickinson County (Lake Okoboji) ISUA; Johnson County (Iowa City) MCZ;

Linn County (Palisades) USNM; County unknown (Foster) USNM. KANSAS: Douglas County (Lawrence) UMAH; Frank-

lin County UMAH. KENTUCKY: Cumberland County (Franklin Branch) TCBa; Edmonson County (2.0 miles from Mam-

moth Cave) MCZ; Jessamine County UASM; Meade County (Fort Knox) UASM; Oldham County (Sleepy Hollow) ULLK;

Wayne County (Wolf Creek Lake) ULLK; County unknown (Sanborn) MCZ. MICHIGAN: Monroe County (Monroe) PUM.

MINNESOTA: Winona County UMSP. MISSISSIPPI: Leake County (near Ludlow) RCGr; Perry County (Richton) CUNY.

MISSOURI: Boone County (Columbia) UNLN; Camden County (Camdenton) UMAH; (Ozark Lake) CAS; Jefferson Coun-

ty (Kimmswick) UMAH; Polk County (Aldrich) CUNY; Saint Charles County (Weldon Springs) UASM; Saint Francis

County (Flat River) USNM; Saint Louis County JShu, UASM; (Rockwoods Reservoir) UASM; Teney County (Branson)

CAS; Counties unknown (Mincy) ISUA, (Willard) UASM. NORTH CAROLINA: Orange County (Chapel Hill) CUNY;

Wake County (Raleigh) UNCR. OHIO: Franklin County (Columbus) PUM; Mercer County (Mendon) UMAH; Ottawa

County (near Marblehead) UMAH; Washington County (New Matamoras) OUCO; County unknown (Georgeville) OUCO.

OKLAHOMA: Latimer County CAS, OSUS; Mayes County (Grand) OSUS; Payne County (Stillwater) OSUS; Pontotoc

County OSUS; Tulsa County (Catoosa) CAS. PENNSYLVANIA: Allegheny County CUNY, (Pittsburgh) CMPP; Fayette

County (Ohiopyle) CMPP. TENNESSEE: Davidson County (Nashville) OUCO, USNM. TEXAS: Brazos County (College

Station) MCZ, TAMU; Colorado County (Columbus) USNM; Cooke County (Gainesville) USNM; Dallas County (Dallas)

MCZ; Kendall County (Comfort) CMPP; Lee County CMPP; McLennan County (Waco) MCZ; Newton County

(Call) USNM. WISCONSIN: Green County (Albany) CEWh; La Crosse County (La Crosse) RESt.

Brachinus alexiguus new species

(Figs. 74, 75, 76, 77, 89)

Type locality. — College Station, Texas.

Type specimens. — The holotype male and one paratype male are in the entomological

museum at MCZ and CAS, respectively. The holotype was collected at the type locality on

March 22, 1927. The paratype was collected in Latimer County, Oklahoma by R. D. Bird on

April 25, 1931.

Diagnostic combination . — The diagnostic characteristics are given in the key.

Description. — Small-sized beetles, 7.0 to 8.0 mm.

Color. Antennal articles 3 and 4, mesepisterna, metepisterna, metasternum at sides, and

abdominal sterna and terga infuscated, otherwise ferrugineous. Dorsal surface and epipleura

sterna and terga infuscated, otherwise ferrugineous. Dorsal surface and epipleura of elytra

blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and surface of pronotum punctate, punctures shallowly

impressed.

Head. As in americanus.

Prothorax. As in americanus. Pronotum (fig. 74).

Pterothorax. As in americanus.

Abdomen. As described for genus.

Genitalia. Male (figs. 75, 76, 77). As in americanus, except median lobe slightly wider to-

ward apex, and ligule more elongate and narrower. Virga (figs. 75, 76). Female unknown.

58

Erwin

/

/

85

h

H

MAP SCALE

0 1000 MILES

1 I

0 1400 KILOMETERS

LAMBERT AZIMUTHAL EQUAL-AREA PROJECTION

Figs. 85-87. Right stylus of female ovipositor, ventral aspect. 85. Brachinus capnicus new species, Smokemont, North

Carolina. 86. Brachinus microamericanus Erwin, Dundee, Mississippi. 97. Brachinus americanus (LeConte), Washing-

ton County, Arkansas. Accompanying scale lines equal 1.0 mm. Figs. 88-91. Geographical distribution maps. 88.

Brachinus americanus (LeConte). 89. Brachinus alexiguus new species. 90. Brachinus microamericanus Erwin. 91.

Brachinus capnicus new species. Map scale given here, used on all maps under Taxonomy; triangles (A) indicate state

locality only.

Bombardier Beetles

59

Variation. — Too few specimens are known to evaluate the geographic variation.

Flight. — The reduction of wing membrane makes these beetles incapable of flight.

Etymology. — Latin, ala , wing; exiguus, small, short or scanty; referring to the reduced

wings of these beetles.

Life history. — The two specimens were collected in March and April, respectively, but

neither were teneral.

Distribution. — (Fig. 89). I have seen two specimens from the following localities:

UNITED STATES

OKLAHOMA: Latimer County CAS. TEXAS: Brazos County (College Station) MCZ.

Brachinus microamericanus Erwin

(Figs. 78, 82, 83, 84, 86, 90)

Type locality. — Dundee, Mississippi.

Type specimens. — The holotype male and allotype female are in the entomological mu-

seum at UMAH, both were collected by T. H. Hubbell at the type locality on August 13,

1929. Two paratypes are in each of the following collections: CAS, MCZ, TLEr, UASM,

UMAH.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Small-sized beetles, 6.0 to 8.7 mm.

Color. Metepisterna and sides of abdomen infuscated, otherwise ferrugineous. Dorsal sur-

face and epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and surface of pronotum slightly rugose and punctate,

punctures shallowly impressed.

Head. As in americanus, except antennal scape robust, widened apically, ligula with three

setae in two rows on each side of center area, and mentum and submentum with accessory

setae.

Prothorax. As in americanus, except proepipleura with a few setae anteriorly and poster-

iorly, glabrous medially. Pronotum (fig. 78). Anterior tibia with anterior edge strigose.

Pterothorax. As in americanus.

Abdomen. As described for genus.

Genitalia. Male (figs. 82, 83, 84). Median lobe with plane of shaft rotated about 45° from

plane of basal bend. Basal bend short. Apex of shaft narrowed, narrowly rounded apically.

Ligule short, narrow, truncate. Virga (figs. 82, 83). Female (fig. 86). Stylus short, angulate,

widened apically.

Variation. — The series from Dundee is quite constant in all characteristics, except the

accessory setae of the mentum and submentum which vary in number. However, the single

specimen from Woodrow is considerably smaller and the shape of its pronotum is different.

This specimen may not be conspecific with the others, but further material will have to be

obtained before its relationship can be established.

Flight. — As in americanus.

Etymology. - Greek, mikros, small; americanus, nominate species of the group; referring

to the resemblance, but smaller form of this species to the nominate species.

Life history. — The Dundee specimens were collected in August and the Woodrow speci-

men in May, and none were teneral.

Distribution. — (Fig. 90). I have seen 17 specimens from the following localities:

UNITED STATES

MICHIGAN: Huron County (Charity Island) UMAH. MISSISSIPPI: Tunica County (Dundee) UMAH. MISSOURI: County

unknown (Woodrow) USNM.

60

Erwin

The capnicus subgroup

The single species included here differs from the previous subgroup in its overall black

color. This is the only known black Neobrachinus, although some species have black parts,

and others are very dark brown.

Brachinus capnicus new species

(Figs. 72,73,85,91)

Type locality. — Smokemont, Great Smoky Mountains National Park, North Carolina.

Type specimen. — The holotype female is in the entomological museum at CUNY. The

single known specimen was collected by W. B. Jones et al. in 1938.

Diagnostic combination. — This is the only known species with all black members.

Description. — Medium-sized beetle, 10.0 mm.

Color. Black. Elytra black with metallic blue luster.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows rugose and punctate, disc of pronotum punctate, punc-

tures moderately impressed.

Head. As in americanus, except submentum with accessory setae.

Prothorax. As in americanus, except center of pronotum concave along midline. Pron-

otum (fig. 73).

Pterothorax. As in americanus.

Abdomen. As described for genus.

Genitalia. Male unknown. Female (fig. 85). Stylus short, narrow, acute at apex.

Flight. — As in americanus.

Etymology . — Greek, kapnikos, smoky; referring to the darkly colored integument of

these beetles, the place where the type was collected, and the ability of these beetles to crep-

itate, producing a cloud of “smoke”.

Distribution. - (Fig. 91).

UNITED STATES

NORTH CAROLINA: Swain County (Smokemont) CUNY.

The texanus group

The members of this group are characterized as follows: virga of endophallus extensively

pigmented and oriented horizontally on the internal sac; antennal articles robust; stylus of

the female ovipositor very narrow and acute; elytral pubescence occurring in rows between

costae. Three species, B. texanus Chaudoir, B. rhytiderus Chaudoir, and B. elongatulus

Chaudoir, are included here, but certain South American species also belong to this group.

Brachinus texanus Chaudoir

(Figs. 93,95,96, 97 107,108)

Brachinus texanus Chaudoir, 1868: 299. Lectotype, here selected, a male, MHNP, labelled

“Tejas” and “Ex Museo Chaudoir”. Type locality. — Texas, as originally given by Chau-

doir.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Medium-sized beetles, 7.3 to 9.9 mm.

Color. Mesepisterna, metepisterna, usually metasternum at sides, sides of abdominal ster-

na, sternum 6, and abdominal terga infuscated, usually antennal articles 3 and 4 infuscated,

otherwise ferrugineous. Dorsal surface and epipleura of elytra slate-grey with greenish luster.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrow and surface of pronotum rugose and punctate, punctures

moderately impressed.

Bombardier Beetles

61

Figs. 92-94. Pronotum right half, dorsal aspect. 92. Brachinus rhytiderus Chaudoir, Chuminopolis, Yucatan, Mexico.

93. Brachinus texanus Chaudoir, Mineral Wells, Texas. 94. Brachinus elongatulus Chaudoir, South West Research

Station, Arizona. Figs. 95-97 , 99-104. Male genitalia. 95. Brachinus texanus Chaudoir, Raymond, Mississippi, ventral

aspect. 96. Lateral aspect of same. 97. Dorsal aspect of same. 99. Brachinus rhytiderus Chaudoir, 1.8 miles north of

El Naranjo, San Luis Potosi, Mexico, ventral aspect. 100. Lateral aspect of same. 101. Dorsal aspect of same.

102. Brachinus elongatulus Chaudoir, Madera Canyon, Arizona, ventral aspect. 103. Lateral aspect of same. 104. Dor-

sal aspect of same. Fig. 98. Right elytron, dorsal aspect of humeral angle, Brachinus elongatulus Chaudoir, South West

Research Station, Arizona. Accompanying scale lines equal 1.0 mm.

62

Erwin

Head. Frontal furrows moderately impressed. Antennal scape robust, widest at middle.

Ligula with sclerotized center area ellipsoid-convex with two paramedian setae and two api-

cal setae. Mentum and submentum usually with accessory setae.

Prothorax. Pronotum (fig. 93), convex, flattened along center line, sides slightly reflexed.

Proepipleura glabrous. Proepisterna with setae anteriorly and posteriorly, glabrous medially.

Anterior tibia with anterior surface strigose.

Pterothorax. Elytra elongate, narrow, moderately costate. Humeral angles square. Costae

smooth, glabrous, depressions pubescent. Hind wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 95, 96, 97). Median lobe with plane of shaft rotated slightly from

plane of basal bend. Basal bend short. Shaft nearly straight, swollen slightly at middle, apex

acute and narrow. Ligule broad, moderately long, truncate. Virga (figs. 95, 96). Female (fig.

107). Stylus narrow, long and acute at apex.

Variation. — Intrapopulational variation occurs in the following characteristics: shape of

the pronotum; total length; extent of infuscation on the venter; and intensity of infuscation

of the antennal articles 3 and 4. In all specimens the sides of the abdomen and the metepis-

terna are infuscated. Usually, sternum 6 is more darkly infuscated than the other abdominal

sterna, but usually the latter are also quite dark, except at the very middle. In the very dark-

est specimens only the middle of sternum 2 between the hind coxae is ferrugineous. Concor-

dantly, the sides of the metasternum are also infuscated.

Flight. — These beetles have been repeatedly collected at lights in Texas.

Etymology . — The latinized form of Texas, the place where the types were collected.

Life history. — Members of this species have been collected from March to November. I

have seen a teneral adult collected in July, at Austin, Texas. Overwintering probably takes

place in the adult stage, as in B. pallidus.

Distribution. — (Fig. 108). The range of this species extends from New York to Alberta,

Canada, south to southern Texas. The northern records from Massachusetts, Virginia, Mich-

igan, Minnesota, and Alberta seem to be disjunctions from the main range. I have seen 1,549

specimens from the following localities:

CANADA

ALBERTA: Chin Coulee (Oldman River) UASM.

UNITED STATES

ALABAMA: Dallas County (Marion Junction) CNHM; Sumter County (Livingston) AMNH; Tuscaloosa County (Tusca-

loosa) UASM; County unknown (Dilchamps) CAS. ARKANSAS: Chicot County (Lake Village) CUNY; Faulkner County

(Conway) UAFA; Hempstead County UAFA, (Hope) CAS, MCZ, UMAH; Hot Springs County (Malvern) JSch; Pulaski

County UAFA, (8.0 miles north of Camp Robinson) CNHM, (Little Rock) MCZ; Sebastian County (Fort Chaffee) RCGr;

Washington County ISNH, (Cove Creek Valley) UAFA, (Fayetteville) UAFA. FLORIDA: Pinellas County (Dunedin)

TAMU. KANSAS: Douglas County (Lawrence) PUM; Montgomery County (Independence) AUAA; Rooks County KSU.

LOUISIANA: Caddo-Bossier Parishes (Bossier City) CNC; De Soto Parish (Mansfield) USNM; East Baton Rouge Parish

(Baton Rouge) UAFA; Franklin Parish (Chance) UAFA, (Chase) UAFA; Jefferson Davis Parish (Hathaway) UAFA; Mad-

ison Parish (Tallulah) MSUM; Natchitoches Parish (Vowells Mill) USNM; Ouachita Parish (Calhoun) UAFA; Vermilion

Parish (Gueydan) USNM; Vernon Parish (Rosepine) UAFA; Parish unknown (Auston) UATA. MASSACHUSETTS: Ply-

mouth County (Plymouth) CNHM. MICHIGAN: Allegan County (Allegan) CAS. MISSISSIPPI: Copiah County (Crystal

Springs) MCZ; Hinds County (Jackson) TAMU, (Raymond) RCGr; Pike County (McComb) UWSW; County unknown

(McCormick) UWSW. MISSOURI: Barry County (Monett) USNM; Pulaski County (Fort Leonard Wood) CAS; Saint Louis

County (Saint Louis) CAS; Vernon County (Nevada) FDAG, UASM. NEBRASKA: Lancaster County (Lincoln) UNLN.

OKLAHOMA: Canadian County (El Reno) CNHM; Choctaw County (Hugo) AMNH; Cleveland County (Norman) CAS,

UONO; Custer County (Clinton) AMNH; Grandy County (Chickasha) OSUS; Grayson County (Juniper Point, Lake Tex-

oma, 12.0 miles north of Whitesboro) RCGr; Latimer County OSUS; Lawton County (Fort Sill) ISUA; Logan County

(Guthrie) CNHM; McClaine County UONO; Marshall County (Lake Texoma, Willis) RCGr, (Lake Texoma, 2.0 miles east

of Willis) RCGr, UCD, (Madill) RCGr; Muskogee County (Muskogee) USUL; Nowata County (13.0 miles west of Vinita)

RFre; Oklahoma County (Oklahoma City) CAS; Payne County (Stillwater) OSUS; Tulsa County (Tulsa) CAS, DHKa.

TENNESSEE: Davidson County (Nashville) TCBa; Lincoln County (Fayetteville) GRNo; Madison County (Jackson) CNC;

Putnam County (Cookville) TCBa. TEXAS: Baylor County (8.0 miles south Seymour) CNHM; Bell County (Temple)

CNHM; Bexar County (San Antonio) CAS, CEWh, MCZ, OSUS, TAMU, TLEr, UCR; Blanco County (Cypress Mill) USNM,

(2.0 miles south of Round Mountain) UASM; Brazos County (College Station) CAS, TAMU, (Texas Experiment Station)

Bombardier Beetles

63

TAMU; Cameron County (Brownsville) USNM; Cass County (Linden) CNC; Cherokee County (Alto) JSch; Comal County

(New Braunfels) TCBa, UASM, USNM; Cooke County (Gainesville) USNM; Dallas County (Dallas) CAS, CUNY, ISNH,

MCZ, PSUU, UASM, UMAH, UMSP, USNM; Denton County CAS; Dimmit County (Texas Experiment Station) TAMU;

Eastland County UMSP; El Paso County (El Paso) CMPP; Ennis County (Ennis) AMNH; Erath County (Dublin) ISUA;

Grayson County (Sherman) AMNH; Hunt County (Wolfe City) CUNY; Jefferson County (Port Arthur) AMNH, (Sabine

Pass) WSUP; Jones County (Stanford) AMNH; Karnes County (Gillett) CUNY; Kerr County (Kerrville) CNC; Kleberg

County (Kingsville) CUNY, ISUA; Leon County CAS; Liberty County (Liberty) UWSW; McLennan County (China Spring)

CNHM; Montague County (2.5 miles southwest of Forestburg) CNHM; Montgomery County (Willis) OSUC; Oldham Coun-

ty (Matador Ranch) UWSW; Palo Pinto County (Mineral Wells) TCBa; Panola County (Carthage) AMNH; Potter County

(Amarillo) UWSW; Robertson County (Heame) CAS; Scurry County (Snyder) TCBa; Stephens County (Breckenridge)

TCBa; Tarrant County OSUS, UMAH, (Fort Worth) CUNY ; Taylor County (Abilene) UATA; Travis County (Austin) CAS,

FDAG, MCZ, OUCO.UASM, UWMW, WSUP, ZMLS; Val Verde County OUCO; Victoria County (Victoria) UASM, UMAH,

USNM; Washington County (Brenham) USNM;Webb County (Laredo) UWSW; Williamson County (Elm Water Cave) TCBa;

Zavalla County (Nueces) USNM; Counties unknown (Belfrage) MCZ, (Camp Barkely) OSUC, (Carancahua) USNM, (Fuller)

USNM, (Virginia Point) USNM. VIRGINIA: (No locality given) ANSP. WISCONSIN: Iron County (Mercer) MCZ, UWMW.

Brachinus rhytiderus Chaudoir

(Figs. 92,99, 100, 101, 105, 110)

Brachynus rhytiderus Chaudoir, 1876: 76. Lectotype, here selected, a male, MHNP, labelled

“Mexique” and “Ex Museo Chaudoir”, standing first in a series of seven specimens. Type

locality. — Mexico, as originally given by Chaudoir, but herewith restricted to San Luis

Potosi, Mexico.

Diagnostic combination . — The diagnostic characteristics are given in the key.

Description. — Small to medium-sized beetles, 5.2 to 9.1 mm.

Color. Except for the constantly darkened terga, the infuscated areas of these beetles are

ill-defined. Usually the antennal articles 3 and 4 darker than 1 and 2. Metepisterna and sides

of abdomen usually quite dark, but center of abdominal sterna slightly infuscated, darker

than color of pronotum. Tibiae infuscated, degree varied with specimens. Dorsal surface and

epipleura of elytra slate-colored.

Microsculpture. As described for genus.

Macrosculpture. As in texanus.

Head. As in texanus, except mentum and submentum without accessory setae.

Prothorax. As in texanus. Pronotum (fig. 92).

Pterothorax. As in texanus, except pubescence usually very sparse in discal depressions.

Abdomen. As described for genus.

Genitalia. Male (figs. 99, 100, 101). Median lobe as in texanus, slightly more arcuate.

Virga (figs. 99, 100). Female (fig. 105). Stylus elongate, narrow, apically acute, but not as

sharp as in texanus.

Variation. — As in elongatulus. The intensity of infuscation seems to vary clinally with

darker specimens on the Yucatan Peninsula, in the southern part of Mexico, and in Central

America, while the paler individuals are in Texas.

Flight. — The flight of these beetles has been recorded at lights repeatedly in Mexico.

Etymology . — Greek, rhytido, wrinkle, deros, skin; referring to the costate elytra of

these beetles.

Collecting notes. — G. E. Ball and D. R. Whitehead have collected these beetles from un-

der litter in palm forests at the edge of a large grassy swamp in San Luis Potosi; at the margin

of the Rio Tula, in Hidalgo; and near the edge of a small stream (in litter) in Queretaro.

They also collected specimens in roadside litter (vegetation) near Comitan, Chiapas. In

Nicaragua, these beetles have been collected repeatedly in cotton fields.

Life history. — Members of this species have been collected in all months of the year, ex-

cept January and February. I have seen a teneral adult collected in September in the state of

Chiapas, Mexico. Overwintering or aestivation probably takes place in the adult stage.

Distribution. — (Fig. 110). The known range of this species extends from Texas south

along the eastern side of Mexico to the Canal Zone, Panama. I have seen 625 specimens from

64

Erwin

109

Figs. 105-107. Right stylus of female ovipositor, ventral aspect. 105. Brachinus rhytiderus Chaudoir, 1.8 miles north

of El Naranjo, San Luis Potosi, Mexico. 106. Brachinus elongatulus Chaudoir, South West Research Station, Arizona.

107. Brachinus texanus Chaudoir, Raymond, Mississippi. Figs. 108-1 10. Geographical distribution maps. 108. Brach-

inus texanus Chaudoir. 109. Brachinus elongatulus Chaudoir. 110. Brachinus rhytiderus Chaudoir. Accompanying

scale lines equal 1 .0 mm.

Bombardier Beetles

65

the following localities:

CENTRAL AMERICA

CANAL ZONE: (Barro Colorado) MCZ. COSTA RICA: (5.0 kilometers north of Caflas) GRNo; (La Carpentera) USNM;

(San Jose) MCZ, TCBa; (San Pedro) GRNo; (Tres Rios) MCZ; (Turrialba) CAS; (Zent, Limon) MCZ. GUATEMALA: (Agua

Caliente) MCZ; (Antigua) AMNH, MCZ; (Los Amates) MCZ; (Palin) USNM; (Panzos) MCZ; (Tamahu) AMNH. NICARAG-

UA: (La Calera) USNM. HONDURAS: CUNY, (Copan) MCZ; (El Paraiso) UCD; (La Lima) DTRT; (Rancho Chiquito,

Department of Comayagua) FDAG. PANAMA: (Rovira, Chiriqui) SJSC.

MEXICO

DURANGO: MCZ. CHIAPAS: (3.2 miles north of Arriaga) UASM; (15.6 miles west of Comitan) UASM; (32.5 miles east of

Comitan) UASM; (El Rosario, northwest of Comitan) RTBe; (4.9 miles north of Frontera Comalapa) UASM; (7.7 miles

north of Frontera Comalapa) UASM; (16.3 miles southwest of Las Cruces) UASM; (11.6 miles north of Ocozocuautla)

UASM; (Palenque ruins) UASM; (20.0 miles south of Tuxtla Gutierrez) UCD. HIDALGO: (Jacala) CNC. MICHOACAN:

(Sahuayo) WSUP. NUEVO LEON: (Huasteca Canon, near Monterrey) CNC; (14.8 miles west of Linares) UASM; (Mesa de

Chipinque, near Monterrey) AMNH; (Montemorelos) CAS; (Monterrey) CNC; (5.0 miles south of Monterrey)

CNC; (6.0 miles south of Monterrey) CNC, FDAG; (Rio Elizondo, near Monterrey) MCZ; (Santa Rosa Canon, 14.8 miles

west of Linares) UASM. OAXACA: (La Ventosa) UCR; (50.0 miles north of La Ventosa) ISUA; (Oaxaca) BMNH; (Puente

Zanatepec, near Zanatepec) UASM; (Rio Malatengo, 11.1 miles north of Matias Romero) UASM; (Rio Niltepec, 18.4 miles

west of Zanatepec) UASM; (Tapanatepec) UASM; (3.0 miles northwest of Tapanatepec) ISUA; (17.0 miles northwest of

Zanatepec) ISUA. PUEBLA: (Villa Juarez) JHen. QUERETARO: (Landa de Matamoros) UASM. SAN LUIS POTOSI:

(Ciudad de Valles) WSUP; (5.0 miles northeast of Ciudad del Maiz) CNC; (El Naranjo) OSUC; (1.8 miles north of El

Naranjo) UASM; (3.6 miles west of El Naranjo) UASM; (El Salto Falls) FDAG; (2.7 miles west of Santa Catarina) UASM;

(Tamazunchale) AMNH, DRWh, MCZ. TAMAULIPAS: (2.0 miles west of Antiguo Morelos) DRWh; (14.0 miles west of

Antiguo Morelos) UASM; (Ciudad Mante) CNC; (47.0 kilometers south of Ciudad Victoria) MCZ; (Gomez Farias and

vicinity) AMNH, CUNY; (73.1 miles north of Manuel) UASM; (101.1 miles north of Manuel) UASM; (20.0 miles south of

Victoria) TCBa, UCD; (20.6 miles east of Villa de Casas) UASM; (23.1 miles east of Villa de Casas) UASM. VERACRUZ:

(5.0 miles northwest of Acayucan) UCD; (30.0 miles south of Acayucan) UCD; (Cordoba) AMNH, CAS; (10.0 miles east

of Cordoba) GRNo; (Cotaxtla Experiment Station) CAS; (Coyame, Lake Catemaco) DRWh; (Fortin de las Flores) CUNY,

DRWh, UASM; (20.0 miles northwest of Huatusco) FDAG; (Jalapa) AMNH, ANSP, BMNH, CAS, MCZ; (3.0 miles north-

west of Jalapa) GRNo; (Los Tuxtlas Range) TAMU; (Orizaba) MCZ; (2.5 miles west of Sontecomapan) UASM; (Tinajas)

UCD; (Veracruz) UASM. YUCATAN: (Chuminopolis) AMNH; (Merida) AMNH; (Piste) SJSC; (Ruinas de Kabah) UASM.

UNITED STATES

TEXAS: Bee County (BeeviUe) USNM; Cameron County CAS, MCZ, USNM, UWMW, (Brownsville) USNM, WHTy; Comal

County (New Braunfels) UASM, USNM; Kendall County (5-10.0 miles north of Boeme) UASM; Kleberg County (Kings-

ville) CUNY; Travis County (Austin) CAS, WSUP; Victoria County (Victoria) USNM; County unknown (Belfrage) USNM.

Brachinus elongatulus Chaudoir

(Figs. 94, 98, 102, 103, 104, 106, 109)

Brachynus elongatulus Chaudoir, 1876: 75. Lectotype, here selected, a female, MHNP, lab-

elled “40” and “Ex Museo Chaudoir”, standing first in a series of eleven specimens in

front of label “B. elongatulus Chaudoir”. Type locality. — Orizaba, Mexico, as originally

given by Chaudoir.

Brachynus brevior Chaudoir, 1876: 75. Lectotype, here selected, a female, MHNP, labelled

“Mexique” and “Ex Museo Chaudoir”. Type locality. - Oaxaca, Mexico, as originally

given by Chaudoir. Blackwelder, 1944: 71.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Small to medium-sized beetles, 4.8 to 10.6 mm.

Color. Antennal articles 3 and 4, metepisterna, and sides of abdominal sterna infuscated.

Tibiae, and tarsi usually infuscated, otherwise beetles ferrugineous. Dorsal surface and epi-

pleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. As in texanus, except punctures more deeply impressed.

Head. As in texanus.

Prothorax. As in texanus. Pronotum (fig. 94).

Pterothorax. As in texanus, except elytra more elongate and more strongly costate.

Abdomen. As described for genus.

Genitalia. Male (figs. 102, 103, 104). Median lobe with plane of shaft slightly rotated from

plane of basal bend. Basal bend short. Shaft and apex as in texanus. Ligule short, broad,

66

Erwin

truncate. Virga (figs. 102, 103). Female (fig. 106). Stylus narrow, moderately long, rounded

apically.

Variation. — Intrapopulational variation occurs in the shape of the pronotum and in the

total size, and the tibiae and tarsi may or may not be infuscated. Rarely the metepisterna are

ferrugineous.

Flight. - These beetles have been collected repeatedly at lights throughout the range of

the species.

Etymology . — Latin, elongatus, prolonged; referring to the elongate elytra of these

beetles.

Collecting notes. — My wife and I collected these beetles from beneath stones along

streams in many localities. At Herb Martyr Dam near Portal, Arizona, these beetles were

beneath stones piled on top of other stones which were embedded in loamy soil. The stones

were covered with oak leaf litter and were next to flowing water.

Life history. - Members of this species have been collected during all months of the year.

I have seen teneral adults collected in August from Arizona; in July from Michoacan; in

November from Jalisco; and in October from Acapulco. Overwintering probably takes place

in the adult stage.

Distribution. — (Fig. 109). The range of this species extends from Arizona south to

Oaxaca and into Baja California. Two specimens have been recorded from the Pacific Coast,

one from Oregon and one from California. I have seen 3,810 specimens from the following

localities:

MEXICO

AGUASCALIENTES: (Aguascalientes) AMNH, JHen; (4.0 miles southwest of Aguascalientes) AMNH; (11.0 miles west of

Aguascalientes) UASM; (Mai Paso, 7.0 miles east of Calvillo) AMNH; (15.0 miles west of Pabelon) UMAH. BAJA CALI-

FORNIA: (Big Canon, Sierra Laguna) CAS; (La Laguna) GRNo; (Las Animas Canon Ensenada) SDSNH; (Las Animas,

Sierra Laguna) CAS; (12.0 miles northwest of San Bartolo) CAS. CHIHUAHUA: (Buena Vista) AMNH: (Carta Blanca, 16.0

miles west of Matachic) AMNH; (Catarinas) AMNH; (Chihuahua) CNC; (25.0 miles northwest of Chihuahua) CNC; (10.0

miles east of Cuaohtemoc) ISUA; (32.0 miles south of Hidalgo de Parral) CAS; (1.0 mile east of La Sauceda) AMNH; (8.0

miles west of Matachic) AMNH; (Mesa del Huracan, 108° 15’ 30° 4’) CNC; (23.0 miles south of Minaca) UASM; (Ojo

Laguna) AMNH; (Primavera) AMNH; (San Jose Babicora) AMNH; (San Rafael) AMNH; (Santa Barbara) AMNH; (Santa

Clara Canon, 5.0 miles west of Parrita) AMNH; (Santa Clara, Namiquipa District) AMNH; (Sombreretillo) CAS. DISTRITO

FEDERAL: (Creek at Lomas de Chapultepec) MCZ; (Mexico City) JHen, MCZ, WSUP; (Mixcoac) JHen; (Penon Viejo)

MCZ; (Tacubaya) JHen; (Villa Guadalupe) JHen. DURANGO: (Arroyo El Sauz, 33.0 miles north of Durango) MCZ; (Dur-

ango City) AMNH, ANSP, CUNY, MCZ, WSUP; (5.0 miles west of Durango) CNC; (10.0 miles west of Durango) CNC,

ISUA; (15.0 miles west of Durango) CNC; (20.0 miles west of Durango) CNC; (23.0 miles south of Durango) CNC; (25.0

miles west of Durango) CNC; (27.5 miles west of Durango) UASM; (18.0 miles east of El Salto) AMNH; (Las Puentes)

AMNH; (Nombre de Dios) AMNH; (Rio Chico, 15.7 miles west of Durango) UASM; (Rio Florido, near Las Nieves) UASM.

GUANAJUATO: (2.0 miles north of Irapuato) CNC; (Rio Guanajuato, 9.8 miles south Silao) UASM; (San Miguel Allende)

AMNH. GUERRERO: (Acapulco) JHen, MCZ; (Apipilulco) MCZ; (Cacahuamilpa) JHen; (Chilpancingo) CNC; (8.4 miles

west of Chilpancingo) UASM; (Rio Mezcala, 23.7 miles north of Zumpango) UASM; (Rio Papagayo, 41.0 miles north of

Acapulco) UASM; (Xilitla) AMNH; (9.0 miles north of Zumpango) ISUA; (30.8 miles north of Zumpango del Rio) UASM.

HIDALGO: (Guadalupe) MCZ; (Huichapan) LACM; (Rio Tula, near Tasquillo) UASM; (San Miguel) MCZ. JALISCO-NAY-

ARIT: (No locality given) AMNH. JALISCO: (Ajijic) AMNH, JHen, UATA; (23.2 miles south of Autlan) UASM; (Aten-

quique) CAS; (3.0 miles north of Barra de Navidad, Bahia de Coastecomate) UATA; (Barranquillas) UCR; (0.4 miles west

of Cocula) UASM;(Cruzero de Malpaso, 17.7 miles northwest of Los Volcanes) UASM; (Guadalajara) AMNH, CUNY, MCZ,

RTBe; (61.0 kilometers southwest of Guadalajara) UATA; (Huascato) USNM; (10.0 miles west of Jiquilpan) CAS; (15-20.0

miles west of Jiquilpan) CAS; (8.5 miles north of Juchitlan) UASM; (13.0 miles southeast of Lagos de Moreno) UASM;

(17.9 miles west of Magdalena) UASM; (Mountains west of Tecalitlan) UATA; (Ocotlan) CAS; (Puente Caquixtle, 9.7

miles east of Encamacion de Diaz) UASM; (Puente La Garita, near La Garita) UASM; (Puerto Vallarta) CNC; (Rio Grande

de Santiago, 12.5 miles west of Ixtlahuacan del Rio) UASM; (4.0 miles south of Talpa de Allende) UASM; (6.5 miles south

of Talpa de Allende) UASM; (21.0 miles northeast of Tepatitlan) CAS; (Tuxpan) MCZ; (Savula) USNM; Valle de Guada-

loupe) CAS. MEXICO: (Lago Zumpango, near San Juan Zitlaltepec) UASM; (Los Remedios) JHen; (Serros Guadalupe)

JHen; (Temascaltepec, Tejupilco) CAS. MICHOACAN: (Comanja) CAS; (1.3 miles east of Comanja) UASM; (5.0 kilo-

meters west of Ciudad Hidalgo) MCZ; (1 1.0 miles west of Hidalgo) UCD; (Huajumbaro) UASM; (10.0 miles west of Jiquil-

pan) DRWh; (10.6 miles south of La Huerta) UASM; (Morelia) AMNH, UASM; (9.5 miles west of Morelia) UASM; (25.0

kilometers east of Morelia) RTBe; (3.0 miles west of Quiroga) AMNH; (6.0 kilometers east of Quiroga) UASM; (Tuxpan)

CAS, RTBe; (near Tzintzuntzan) UASM; (8.1 miles east of Vallamar) UASM; (10.0 miles east of Zamora) GRNo; (14.0

miles northwest of Zitacuaro) UCD; (50.0 miles west of Zitacuaro) MCZ. MORELOS: (Canon de Lobos, 9.1 miles east of

Cuernavaca) UASM; (Cuautla) JHen; (Cuernavaca) CAS, CNC, CUNY, MCZ, RCGr, TMBH; (3.6 miles east of Cuernavaca)

UASM; (5.4 miles east of Cuernavaca) UASM; (Progreso) CUNY; (Tejalpa) DRWh, FDAG; (Tepoztlan) RCGr; (Xochi-

Bombardier Beetles

67

tepee) JHen; (Yautepec) UCD. NAYARIT: (5.1 miles north of Chapalilla) UASM; (20.3 miles west of Compostela) GRNo;

(Ixtlan del Rio) CAS; (San Bias) UCD; (8.7 miles east of San Bias) GRNo; (13.8 miles east of San Bias) UASM; (Tepic)

AMNH, UATA; (19.0 miles southeast of Tepic) CAS; (24.0 miles southeast of Tepic) CAS. OAXACA: (25.0 miles south of

Mitla) ISUA; (Oaxaca) UMAH; (Rio Atoyac, near Juchatengo) UASM; (9.3 miles north of Sola de Vega) UASM; (72.5 miles

south of Valle Nacional, Rio Grande) UASM. PUEBLA: (9.0 miles north of Amatitlan) CAS; (5.0 miles south of Izucar de

Matamoros) UCD; (near Petlalcingo) UASM; (3.0 miles northwest of Petlalcingo) UCD; (Puente Tepexco, near Tepexco)

UASM; (Tehuacan) CAS, JHen; (near Tehuitzingo) UASM; (near Tepexco) UASM. QUERETARO: (6.0 miles east of

Celaya) UASM; (near Palmillas) UASM; (6.4 miles east of Pinal de Amoles) UASM; (Queretaro) UASM; (3.0 miles west of

Queretaro) AMNH. SAN LUIS POTOSI: (25.0 kilometers east of Santa Domingo) MCZ. SINALOA: (Camino Real de

Piaxtla) AMNH; (3.4 miles west, 5.0 miles south of Culiacan) GRNo; (3.0 miles east of Culiacancito) GRNo; (2.1 miles

northeast of El Fuerte) GRNo; (Mazatlan) AMNH, MCZ, UCR; (5.0 miles north of Mazatlan) GRNo; (10.0 miles north of

Mazatlan) GRNo; (Rio Baluarte, near Rosario) UASM; (Quiroroba, Alamosa) CAS; (Rio Panuco 11.2 miles northeast of

Concordia) UASM; (Rio Piaxtla, 1.0 miles east of Route 15) UASM; (Rosario) CAS; (20.0 miles east of Villa Union) UCR;

(21.0 miles east of Villa Union) CNC; (27.0 miles east of Villa Union) CNC; (33.0 miles east of Villa Union) CNC; (Vene-

dillo) CAS;(Zenzontle, Culiacan) AMNH. SONORA: (65.0 miles southeast of Agua Prieta) ISUA; (Alamos) CAS; (4.0 miles

west of Alamos) GRNo; (5.0 miles west of Alamos) UATA; (7.2 miles southeast of Alamos) GRNo; (Caramechi, Rio Mayo)

CAS; (8.0 miles east of Imuris) CAS; (San Bernardo, Rio Mayo) CAS; (Yecora) CNC. TAMAULIPAS: (Ciudad Victoria)

CNHM; (Sotano de La Joya de Salas) TCBa. TLAXCALA: (21.0 miles west of Apizaco) CNC. ZACATECAS: (29.0 miles

northwest of Fresnillo) UASM; (Presa Choquen) JHen; (Rio Juchipila, 0.9 miles north of Jalpa) UASM; (1.3 miles southeast

Sain Alto) UASM.

UNITED STATES

ARIZONA: Apache County (White Mountains) CAS; Cochise County (Benson) CAS, UWSW, (Bisbee) CAS, (Canelo)

UATA, (Canelo, Pyeatt’s Ranch) CUNY, (Cave Creek) AMNH, (Cave Creek Canyon) CNHM, (Cave Creek, South Fork)

GRNo, (Chiricahua Mountains) CAS, OUCO, UASM, USNM, (Chiricahua Mountains, Cave Creek Ranch) TCBa, UASM,

(Chiricahua Mountains, 15.0 miles west of Portal) UCD, (Chiricahua Mountains, Redrock Canyon) CAS, (Chiricahua Moun-

tains, Rustlers Park) CNC, (Chiricahua Mountains, Silver Creek wash, 3.3 miles west by northwest of Portal) CUNY, (Chiri-

cahua Mountains, Texas Canyon) CAS, (Chiricahua National Monument) AMNH, CAS, (Cochise Stronghold) TLEr, UATA,

(Douglas) CAS, CNHM, (10.0 miles west of Douglas) TLEr, (Guadalupe Canyon, 32.0 miles east of Douglas) CUNY, UCR,

(Martyr Dam, South West Research Station) TLEr, (Huachuca Mountains) AMNH, CAS, TAMU, UATA; USNM, (Huachuca

Mountains, Carr Canyon) AMNH, CAS, TLEr, (Huachuca Mountains, Fort Huachuca) LACM, (Huachuca Mountains, Gar-

den Canyon) CAS, USNM, (Huachuca Mountains, Huachuca Canyon) LACM, (Huachuca Mountains, Miller Canyon) CAS,

MCZ, OUCO, UCR, (Huachuca Mountains, Sunnyside Canyon) CAS, LACM, (Palmerlee) ANSP, CAS, UMAH, (Paradise)

UASM, (Chiricahua Mountains, Pinery Creek) CNHM, (Portal) AMNH, RCGr, SJSC, TLEr, WHTy, WSUP, UCR, (Ramsey

Canyon, Huachuca Mountains) CAS, SJSC, UATA, UMHN, USNM, (Chiricahua Mountains, Rucker Lake) UASM, (San Ped-

ro near Palominas) UASM, (San Pedro River east of Sierra Vista) UATA, (South Fork Forest Camp, 4.5 miles west of Por-

tal) CAS, (South West Research Station, Portal) AMNH, CNC, CUNY, FDAG, OSUC, SJSC, TLEr, UATA, UCD, UCR,

UIMI, (Wood Canyon, Bisbee) UATA; Gila County (Globe) KSU, UCD, (base of Pinal Mountains) CAS, UCD, (Roosevelt

Lake) CNC, (Winkelman) UATA; Graham County (Aravaipa) CAS, (Aravaipa Creek, between Klondyke and Aravaipa)

DRWh, (Galiuro Mountains, Powers’ Garden) UASM, (Galiuro Mountains, Reservoir, east slope) UASM, (Graham Moun-

tains, Shennon Camp) GRNo; Navajo County (8-15.0 miles northeast of Whiteriver) AMNH; Pima County (Baboquivari

Mountains) CAS, CUNY, LACM, (Baboquivari Mountains, Browns Canyon) AMNH, CAS, UATA, (Baboquivari Mountains,

Schalffer Canyon) ANSP, (Baboquivari Mountains, Sabino Canyon Elkhom Ranch) CAS, (East slope of Coyote Mountains,

0.5 miles north of Mendoza Canyon) UATA, (10.0 miles east of Continental) UATA, (4.0 miles south of Mountain View)

CAS, (Sahuarita) MCZ, (Saint Xavier Mountains, Tucson) CAS, (Santa Catalina Mountains) CAS, UATA, USNM, (Santa

Catalina Mountains, Mount Lemmon, Bear Canyon) CAS, CUNY, UATA, (Santa Catalina Mountains, Peppersauce Canyon)

CAS, UASM, UATA, (Santa Catalina Mountains, Molino Basin) CUNY, TLEr, (Santa Rita Mountains) CAS, UATA, UMSP,

USNM, (Santa Rita Mountains, Gardner Canyon) OSUC, (Tanque Verde) UATA, (Tucson) CAS, MCZ, UASM, UATA,

UIMI, UMSP, USNM, (8.0 miles north of Vail) UATA ; Pinal County (Oracle) USNM, (14.0 miles east of Oracle) CAS ; Santa

Cruz County OSUC, (Calabasas Canyon, east Tumacacori Mountains) UASM, (Canille) AMNH, UATA, (Luis Springs, San

Pedro River) UASM, (Nogales) CAS, CNHM, LACM, MCZ, UCD, USNM, ZMLS, (3.0 miles north of Nogales) LACM, (10.0

miles west of Nogales) UCD, (15.0 miles northwest of Nogales) UCD, (Pajaritos Mountains, Yanks Spring, 4.0 miles south-

east of Ruby) AMNH, CAS, (Patagonia) CAS, CMPP, CNHM, CUNY, UATA, UCD, (Pena Blanca) CNC, CUNY, UASM,

UATA, (Santa Rita Mountains, Madera Canyon) AMNH, ANSP, CAS, CUNY, GRNo, LACM, SJSC, RCGr, UASM, UATA,

UCD, UWSW, WHTy, (Sonoita) WHTy, (Sonoita River, Patagonia) AMNH, (4.0 miles west of Sombrero Butte, Cherry

Creek) USNM, (Tumacacori Mountains, Sycamore Canyon) CUNY, (Tumacacori Mountains, Sycamore Canyon, near Ruby)

CUNY, UATA, (2.0 miles west of Washington Camp) UATA; Maricopa County (Gillespie’s Dam) CAS; Yavapai County

(Mayer) GRNo, (Prescott) CAS; Yuma County (15.0 miles east of Yuma) CNHM; Counties unknown (Florida) UATA,

(Reef) MCZ, (near San Fernando) USNM, (Santa Cruz River) UATA, (Tortolita Mountains) CAS, (Wake Field Mine) PUM.

CALIFORNIA: CMPP, ISNH, TMBH; Santa Cruz County (Amado) UMSP. NEW MEXICO: Catron County (Mogollon

Mountains, Big Dry Creek) CAS, (Mogollon Mountains, Willow Creek) TLEr; Grant County (Silver City) MCZ; Hidalgo

County (Guadalupe Canyon) GRNo, (Rodeo) UCR, (12.0 miles southeast of Rodeo, Post Office Canyon) SJSC. OREGON:

Lane County (Siltcoos Lake) CAS. TEXAS: Jeff Davis County (Limpia Canyon) TLEr.

The sallei group

The members of this group are characterized by their complete brown color, testaceous

legs with infuscated knees, lateral pits of the mentum, and swollen median lobe. A single

species, B. sallei Chaudoir is included here.

Figs. 1 1 1-1 14. Pronotum, right half, dorsal aspect. 111. Brachinus melanarthrus Chaudoir, Tamazunchale, San Luis

Potosi, Mexico. 1 12. Brachinus sallei Chaudoir, Edzna, Campeche, Mexico. 1 13. Brachinus grandis Brulle, Macultepec,

Tabasco, Mexico. 114. Brachinus brunneus Castelnau, Laguna Guanico, Puerto Rico. Figs. 115-126. Male genitalia.

115. Brachinus melanarthrus Chaudoir, Veracruz, Veracruz, Mexico, ventral aspect. 116. Lateral aspect of same.

1 17. Dorsal aspect of same. 1 18. Brachinus grandis Brulle, Macultepec, Tabasco, Mexico, ventral aspect. 1 19. Lateral

aspect of same. 120. Dorsal aspect of same. 121 . Brachinus sallei Chaudoir, Coyame, Lake Catemaco, Veracruz, Mex-

ico, ventral aspect. 122. Lateral aspect of same. 123. Dorsal aspect of same. 124. Brachinus brunneus Castelnau, La

Guanica, Puerto Rico, ventral aspect. 125. Lateral aspect of same. 126. Dorsal aspect of same. Accompanying scale

line equals 1.0 mm.

Bombardier Beetles

69

Brachinus sallei Chaudoir

(Figs. 20, 112, 121, 122, 123, 128, 133)

Brachynus sallei Chaudoir, 1876: 85. G. E. Ball could not locate Chaudoir’s specimens in

MHNP. Chaudoir wrote (1876: 85) that other specimens were in Chevrolat’s collection.

E. Taylor informs me that these specimens are no longer in HMO. Therefore, I presume

the type is lost.

Type locality. — Mexico, as originally given by Chaudoir, but herewith restricted to Tab-

asco, Mexico.

Notes. — Chaudoir compared B. sallei with Chevrolat’s B. spinipes, in the original descrip-

tion of B. sallei. This name was never published by Chevrolat, and therefore not really in ex-

istence, but I add this note to stop possible confusion in the future.

Diagnostic combination. — See group characteristics and key.

Description. — Large-sized beetles, 12.1 to 15.1 mm.

Color. Dark brown, mouthparts and antennae somewhat paler. Legs testaceous with

brown “knees” (femoral apices plus tibial bases). Dorsal surface of elytra dark brown, epi-

pleura paler.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows rugose, sparsely punctate. Disc of pronotum finely ru-

gose along center line, punctures barely impressed.

Head. Frontal furrows deeply impressed. Antennal scape quite robust, widest at middle.

Ligula with sclerotized center area ellipsoid-convex with two apical setae. Mentum (fig. 20)

with two lateral pits each surrounded by numerous setae. Submentum with numerous acces-

sory setae.

Prothorax. Pronotum (fig. 1 12) convex, sides narrowly reflexed. Anterior and posterior

margins with a few shallowly impressed setiferous punctures. Proepipleura glabrous. Pro-

episterna with a few small setae both anteriorly and posteriorly, glabrous medially. Anterior

tibia with anterior edge finely punctate.

Pterothorax. Elytra elongate, moderately costate. Humeral angle square. Pubescence con-

fined to eighth depression, except near apex. Depressions punctate. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 121, 122, 123). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend short. Shaft swollen at basal third, flared and flattened

at apical third. Ligule short, rounded apically. Virga (figs. 121, 122). Female (fig. 128). Sty-

lus narrow, elongate, arcuate, acute at apex.

Variation. — Too few specimens are known for me to evaluate geographic variation, but

the sixteen specimens I have seen are rather constant.

Flight. — J. G. Edwards collected a specimen at lights on Cozumel Island, Quintana Roo,

Mexico.

Etymology . — Patronym for August Salle, in whose collection Chaudoir first saw speci-

mens of this species.

Collecting notes. — On the north shore of Lake Catemaco in Veracruz, Mexico, these

beetles live in heavily shaded marshes about 10.0 meters from the lake edge. The predom-

inant plant was a Heliconia species.

Life history. — Members of this species have been collected from December to April and

August. One teneral adult was collected in January in the state of Campeche, Mexico.

Distribution. — (Fig. 133). The known range of this species extends from southern San

Luis Potosi to southern Chiapas and the Yucatan peninsula. The species also occurs on Isla

Cozumel, off the coast of Quintana Roo. I have seen 16 specimens from the following local-

ities:

70

Erwin

Figs. 127-130. Right stylus of female ovipositor, ventral aspect. 127. Brachinus grandis Brulle, Venedillo, Sinaloa,

Mexico. 128. Brachinus sallei Chaudoir, Edzna, Campeche, Mexico. 129. Brachinus melanarthrus , Tela, Honduras.

130. Brachinus brunneus Castelnau, Humacao, Puerto Rico. Figs. 131-134. Geographical distribution maps. 131.

Brachinus grandis Brulle. 132. Brachinus melanarthrus Chaudoir. 1 33. Brachinus sallei Chaudoir. 134. Brachinus brun-

neus Castelnau. Accompanying scale line equals 1.0 mm.

Bombardier Beetles

71

MEXICO

CAMPECHE: (Edzna) UASM. CHIAPAS: (San Quintin, Chiquita Miramar) UASM. QUINTANA ROO TERRITORY:

(Cozumel Island) SJSC. SAN LUIS POTOSI: (Valles) CPBo. TABASCO: (Macultepec) JHen; (Tierra Colorado de Itzmate)

JHen. VERACRUZ: (Coyame, Lake Catemaco) UASM; (San Andres Tuxtla) UCR.

The brunneus group

The members of this group are characterized by the form of the virga, brown color of the

body, and testaceous legs and infuscated knees. These beetles lack the lateral mental pits

found in B. sallei. I have seen specimens of South American species that belong to this group.

It is possible that when the South American fauna is better known, this group will have to

be redefined. Two species, B. brunneus Castelnau, and B. melanarthrus Chaudoir are in-

cluded here.

Brachinus brunneus Castelnau

(Figs. 114, 124, 125, 126, 130, 134)

Brachinus brunneus Castelnau, 1834: 59. Lectotype, here selected, a male, MHNP, un-

labelled, but standing third in a series of seven in front of box label “brunneus Castelnau

Antilles Mus. Berl”. Chaudoir (1876: 84) mentions that he received four specimens from

Castelnau’s type series. Type locality. — Cayenne, French Guiana as originally given by

Castelnau.

Brachinus gilvipes Mannerheim, 1837: 41. Lectotype, here selected, a female, MHNP, lab-

elled “St. Thomas, Mannerheim” and standing fifth in a series of seven specimens in front

of box label “brunneus Castelnau Antilles Mus. Berl”. Type locality. - St. Thomas Island,

Antilles as given on Mannerheim’s label, and indicated by Chaudoir (1876: 84). Chaudoir,

1876: 84.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Small to medium-sized beetles, 7.0 to 9.0 mm.

Color. As in sallei.

Microsculpture. As described for genus.

Macrosculpture. As in sallei.

Head. As in sallei, except antennal scape not as robust, mentum without pits or accessory

setae, submentum without accessory setae.

Prothorax. As in sallei. Pronotum (fig. 1 14)

Pterothorax. As in sallei, except elytral pubescence extending along depressions 6,1, and

8, and costae barely Elevated.

Abdomen. As described for genus.

Genitalia. Male (figs. 124, 125, 126). Median lobe with plane of shaft rotated 45° from

plane of basal bend. Basal bend short. Apex of shaft rounded. Ligule short, narrow, rounded

apically. Virga (figs. 124, 125). Female (fig. 130). Stylus narrow, acute at apex.

Variation. — Besides the intrapopulational variation in the shape of the pronotum and the

total size, these beetles exhibit color differences of the third and fourth antennal articles.

The range is from testaceous with infuscated bases to completely infuscated, and occurs

within single population samples.

Flight. — The flight of these beetles has not been recorded.

Etymology . — Medieval Latin, brunneus, brown; referring to the color of these beetles.

Life history. — Members of this species have been collected in October, January, March,

and May. No teneral adults have been seen.

Distribution. — (Fig. 134). The known range of this species extends from French Guiana

to Puerto Rico and Hispaniola (Haiti). I have seen 108 specimens from the following local-

ities:

72

Erwin

GREATER ANTILLES

CUBA: (Baiagua) MCZ; (Colon) MCZ. DOMINICAN REPUBLIC: (Barahona) MCZ; (Monte Cristi) MCZ. HAITI: (Etang

Lachaux southwest peninsula) MCZ. PUERTO RICO: (Ensenada) MCZ; (Humacao) CUNY, MCZ; (Laguna Guanica) MCZ;

(Mereidita Finca Ponce) MCZ; (Mayaguez) CUNY; (Toa-Baja) CUNY, MCZ.

VIRGIN ISLANDS

St. Croix Island, MCZ; St. Thomas Island, MHNP.

Brachinus melanarthrus Chaudoir

(Figs. 19, 111, 115, 116, 117, 129, 132)

Brachynus melanarthrus Chaudoir, 1876: 84. Lectotype, here selected, a male, MHNP,

unlabelled, but standing first in front of box label “melanarthrus, Chaud. Caracas, Sall6”.

Type locality. — Mexico, as originally given by Chaudoir, but herewith restricted to

Veracruz, Mexico.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Small to medium-sized beetles, 6.6 to 11.2 mm.

Color. As in sallei.

Microsculpture. As described for genus.

Macrosculpture. As in sallei.

Head. As in sallei, except mentum (fig. 19) without deep lateral pits and accessory setae

and submentum without accessory setae.

Prothorax. As in sallei. Pronotum (fig. 111).

Pterothorax. As in sallei.

Abdomen. As described for genus.

Genitalia. Male (figs. 115, 116, 117). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend long. Apex of shaft narrowed, elongate. Ligule short,

broadened apically. Virga (figs. 115, 116). Female (fig. 129). Stylus narrow, parallel-sided,

rounded apically.

Variation. — Too few specimens are known of this species to analyze the geographical

variation. However, the size may vary clinally, the largest specimens in Mexico, the smallest

specimens in Honduras. One female from Veracruz is darker in color and the surface of the

pronotum is dull due to very sense rugosities.

Flight. - These beetles have been collected at lights in the state of San Luis Potosi, Mex-

ico.

Etymology . — Greek,, melanos , black; arthron, joint; referring to the infuscated knees

of these beetles.

Life history. — Members of this species have been collected in May, July, and August, but

no teneral adults were seen.

Distribution. — (Fig. 132). The known range of this species extends from southern San

Luis Potosi, Mexico, to northern Honduras. I have seen seven specimens from the following

localities:

CENTRAL AMERICA

HONDURAS: (La Lima) DTRT ; (Tela) DTRT.

MEXICO

SAN LUIS POTOSI: (Tamazunchale) MCZ. VERACRUZ: (5.0 miles northwest of Acayucan) UCD; (Veracruz) CNC.

The grandis group

The members of this group are characterized by very large size and darkly pigmented and

elongate virgae. One species, B. grandis Brulle, is included.

Bombardier Beetles

73

Brachinus grandis Brulle

(Figs. 113, 118, 119, 120, 127, 131)

Brachinus grandis Brulle, 1838: 19. Lectotype, here selected, a female, MHNP, unlabelled,

but standing next to a pin with a point and the label “grandis Brulle”. The point holds

the labrum of the specimen. Type locality. — Bolivia, as originally given by Brulle.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Very large-sized beetles, 16.4 to 18.5 mm.

Color. Mesepisterna, metepisterna, metasternum at sides, abdominal sterna and terga, and

knees infuscated, otherwsie ferrugineous. Dorsal surface and epipleura of elytra brown.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows rugose, sparsely punctate, punctures barely impressed.

Head. Frontal furrows moderately impressed. Antennal scape robust, widest apically.

Ligula with sclerotized center area ellipsoid-convex with two apical setae. Mentum and sub-

mentum without accessory setae.

Prothorax. Pronotum (fig. 113), slightly flattened along center line, sides narrowly re-

flexed. Proepipleura glabrous, proepisterna with a few scattered setae both anteriorly and

posteriorly. Anterior tibia with anterior surface sparsely punctate.

Pterothorax. Elytra elongate, broad, moderately costate. Humeral angle square. Pubes-

cence confined to outer intervals, except in apical third. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 118, 119, 120). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend long. Apex of shaft narrowed to apex. Ligule poorly

defined, short, truncate. Virga (figs. 1 18, 1 19). Female (fig. 127). Stylus very large, broad,

narrowly rounded at apex.

Variation. — Too few specimens are known to evaluate geographic variation.

Flight. — The flight of these beetles has not been recorded.

Etymology . — Latin, grandis , large; referring to the very large size of these beetles com-

pared to others in the genus.

Life history. — Members of this species have been collected in March, May, and June, but

no teneral adults were seen.

Distribution. — (Fig. 131). The known range of this species extends from Sinaloa and San

Luis Potosi in Mexico to Bolivia, South America, but this has been determined by only four

localities. It is probable that this species has discontinuous and local populations. I have

seen five specimens from the following localities:

MEXICO

SINALOA :(Venedillo), CAS. SAN LUIS POTOSI: (El Pujal) CPBo. TABASCO: (Macultepec) JHen.

The lateralis group

The members of this group are characterized by their brown elytra. It is a provisional

group pending the outcome of a revision of the South American species many of which have

brown elytra.

Brachinus lateralis Dejean

(Figs. 137, 145, 156, 157, 158, 169)

Brachinus lateralis Dejean, 1831: 424. Lectotype, here selected, a male, MHNP, labelled

“male, lateralis m. in Amer. bor.” on green paper, “Ex Museo Chaudoir” and “Type” on

red paper. Type locality. — North America, as originally given by Dejean, but herewith

restricted to Imperial County, California.

Brachinus leucoloma Chaudoir, 1868: 301. Lectotype, here selected, a male, MHNP, lab-

elled “Californie, R. Gila, LeConte” and “Ex Museo Chaudoir.” Type locality. - Gila

74

Erwin

Figs. 135-140. Pronotum, right half, dorsal aspect. 135. Brachinus chalchihuitlicue new species, San Bias, Nayarit,

Mexico. 136. Brachinus chirriador new species, Puente La Garita, La Garita, Jalisco, Mexico. 137. Brachinus lateralis

Dejean, 20.9 miles north of Arriaga, Chiapas, Mexico. 138. Brachinus arboreus Chevrolat, Mazatlan, Sinaloa, Mexico.

139. Brachinus adustipennis Erwin, 20.0 miles west of Rosario, Sinaloa, Mexico. 140. Brachinus aeger Chaudoir,

Paso Antonio, Escuintla, Guatemala. Figs. 141-146. Right stylus of female ovipositor, ventral aspect. 141. Brachinus

chirriador new species, Puente La Garita, La Garita, Jalisco, Mexico. 142. Brachinus adustipennis Erwin, 20.0

miies west of Rosario, Sinoloa, Mexico. 143. Brachinus chalchihuitlicue new species, Mazatlan, Sinaloa, Mexico.

144. Brachinus aeger Chaudoir, Paso Antonio, Escuintla, Guatemala. 145. Brachinus lateralis Dejean, Rio Niltepec,

Oaxaca, Mexico. 146. Brachinus arboreus Chevrolat, Mazatlan, Sinaloa, Mexico. Accompanying scale lines equal 1.0

mm.

Bombardier Beetles

75

River, Imperial County, California, as originally given on Chaudoir’s labelled specimen.

Erwin, 1965: 7.

Diagnostic combination. — The brown elytra with pale epipleura and glabrous elytral disc

separate members of this species from all others west of the continental divide in the United

States. In Mexico and further south, only the genitalic characteristics provide reliable diag-

nostic characteristics (see key couplet 17).

Description. — Medium-sized beetles, 6.1 to 9.3 mm.

Color. Mesepisterna, metepisterna, abdomen at least at sides, and “knees” infuscated.

Head and prothorax ferrugineous. Usually mouth parts, antennal articles 1-4, legs, and mid-

dle of venter testaceous. Dorsal surface of elytra brown, epipleura testaceous.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows rugose, sparsely punctate. Surface of pronotum smooth

with a few scattered shallowly impressed setiferous punctures.

Head. Frontal furrows shallowly impressed. Antennal scape robust, widest at middle.

Ligula with sclerotized center area ellipsoid-convex with two apical setae, and a few small

inconspicuous setae. Mentum and submentum without accessory setae.

Prothorax. Pronotum (fig. 137) convex, sides narrowly reflexed. Proepipleura glabrous.

Proepisterna with a few setae anteriorly. Anterior tibia with anterior surface finely punc-

tate.

Pterothorax. Elytra moderately long, barely costate. Humeral angle square. Pubescence

usually confined to outer depressions and costae with some scattered patches occasionally

on disc. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 156, 157, 158). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend moderately long. Apex of shaft flattened, ridged

around edge and with a median ventral keel. Shaft swollen slightly at middle. Ligule short,

truncate. Virga (figs. 156, 157). Female (fig. 145). Stylus long, narrow, apically rounded.

Variation. — Intrapopulational variation occurs in the following characteristics: the pres-

ence or absence of pubescence on the elytral disc, shade of brown color of the elytra, the

height of the costae, the shape of the pronotum, and body size.

Flight. — The flight of these beetles has been recorded at lights throughout the range of

the species.

Etymology . — Latin, lateralis, of the side; referring to the pale epipleura of the elytra on

these beetles.

Collecting notes. — These beetles occur in a number of habitats. In California, they have

been collected on the shores of the Salton Sea (saline) and Lake Elsinore (fresh water). In

Arizona, they have been collected at the edges of lakes ancf in wet meadows. In Mexico,

they have been collected in gravel beneath larger stones at the edges of streams.

Life history. - Members of this species have been collected in all months of the year. I

have seen teneral adults collected in March in Puebla, in May and September in Sonora, and

in August in California. These beetles probably overwinter (or aestivate) as adults.

Distribution. — (Fig. 169). The range of this species extends from northern Arizona south

to Chiapas, Mexico, and populations occur in Baja California. I have seen 458 specimens

from the following localities:

MEXICO

BAJA CALIFORNIA: (20.0 miles north of Comondu) CAS; (Estero, at mouth of Arroyo Rosario) CAS; (San Ignacio)

CAS; (65.0 kilometers south of Tijuana) LACM. CHIAPAS: (20.9 miles north of Arriaga) UASM. DURANGO: (Durango

City) AMNH, MCZ. GUANAJUATO: (Lago Yuriria, near Yuriria) UASM. GUERRERO: (Cacahuamilpa) JHen; (2.0

miles north of El Mogote) UASM; (Rio Mezcala, 23.7 miles north of Zumpango) UASM; (9.0 miles north of Zumpango)

ISUA. JALISCO: (Ajijic) JHen, UATA; (Guadalajara) AMNH. MICHOACAN: (Morelia) AMNH, UASM; (near Tzintzunt-

zan) UASM. MORELOS: (Canon de Lobos, 9.1 miles east of Cuernavaca) UASM; (Cuernavaca) ANSP, BMNH, MCZ, RTBe;

(Progreso) CUNY, WSUP; (Xochictepec) JHen. NAYARIT: (San Bias) CAS; (Tepic) UATA; (19.0 miles southeast of

Figs. 147-164. Male genitalia. 147. Brachinus aeger Chaudoir, Rio Teapa, Tabasco, Mexico, ventral aspect. 148. Lateral

aspect of same. 149. Dorsal aspect of same. 150. Brachinus arboreus Chevrolat, 32.0 miles south of Acaponeta,

Nayarit, Mexico, ventral aspect. 151. Lateral aspect of same. 152. Dorsal aspect of same. 153. Brachinus adustipennis

Erwin, Manatee Springs State Park, Florida, ventral aspect. 154. Lateral aspect of same. 155. Dorsal aspect of

same. 156. Brachinus lateralis Dejean, near Petlalcingo, Puebla, Mexico, ventral aspect. 157. Lateral aspect of same.

158. Dorsal aspect of same. 159. Brachinus chalchihuitlicue new species, San Bias, Nayarit, Mexico, ventral aspect.

160. Lateral aspect of same. 161. Dorsal aspect of same. 162. Brachinus chirriador new species, Puente La Garita,

La Garita, Jalisco, Mexico, ventral aspect. 163. Lateral aspect of same. 164. Dorsal aspect of same. Accompanying

scale lines equal 1.0 mm.

Bombardier Beetles

77

Tepic) CAS. OAXACA: (25.0 miles south of Mitla) ISUA; (Puente Zanatepec, near Zanatepec) UASM; (Rio Atoyac,

near Juchatengo) UASM; (Rio Niltepec, 18.4 miles west of Zanatepec) UASM; (72.5 miles south of Valle Nacional) UASM.

PUEBLA: (Puente Tepexco, near Tepexco) UASM; (near Petlalcingo) UASM; (near Tepexco) UASM. SINALOA: (13.0

miles north of Guamuchil) GRNo; (Los Mochis) CAS, GRNo; (Mazatlan) AMNH, CNC. SONORA: (10.0 miles west of

Alamos) AMNH; (Ciudad Obregon) CNC; (16.0 miles northeast of Ciudad Obregon) CNC; (35.0 miles northeast of Ciudad

Obregon) CNC; (14.0 miles southeast of Empalme) CAS; (Hermosillo) AMNH; (La Atascosa) UASM; (Navajoa) GRNo,

JHen; (Pesqueria) CAS; (Rio Yagui 12.0 miles west of Ciudad Obregon) CNC; (Sonoyta) AMNH. VERACRUZ: (Jalapa)

BMNH.

UNITED STATES

ARIZONA: Cochise County (Benson) CAS, (San Bernardino Ranch) KSU, LACM; Graham County (Thatcher) UCD;

Maricopa County (Phoenix) MCZ, UATA, USNM; Mohave County (Littlefield) UCD; Pima County (Arivaca Creek at

Arivaca) CAS, (Baboquivari Mountains, Browns Canyon) AMNH, (Santa Catalina Mountains) CAS, UATA, (Tucson)

AMNH, ANSP, CAS, USNM; Pinal County (Picacho) CAS; Santa Cruz County (6.0 miles north of Nogales) UASM, (15.0

miles northwest of Nogales) UCD, (Pajarita Mountains) CAS, (Patagonia) CUNY, UATA, (Pena Blanca) UASM; Yuma

County (Cibola) CAS, (Ehrenburg) UATA, (Fort Yuma) USNM, (Yuma) CAS, ISNH, MCZ, USNM; County unknown

(Senator) AMNH. CALIFORNIA: Imperial County (Calpatria) CAS, (El Centro) CAS; Los Angeles County PSUU, USNM,

ZMLS, (Cypress) LACM, (East Manhattan) LACM, (Lake Hodges) SDNHM, (Long Beach) CAS, (Los Angeles) CAS,

(Pasadena) CAS, TLEr; Orange County (Anaheim) SDNHM, (Laguna Canyon) UCD; Riverside County (Blythe) LACM,

UIMI, (Corona) UCD, (Elsinore) CAS, (Lake Elsinore) CAS, CMPP, UIMI, UNLN, USNM, (Mecca) CVMA, (Riverside)

CUNY, LACM, UMAH, USNM, (Salton Sea) WBa, (Salton Sea, Mecca) CNC, USNM; San Bernardino County (Needles)

CAS, (Ontario) CAS, (San Bernardino) MCZ, OUCO, (Saratoga Springs, Death Valley) UCD; San Diego County ANSP,

CCha, CUNY, (Oceanside) CAS, (San Felipe Creek 14.0 miles east of Julian) UASM, (San Diego) SDNHM, (San Juan

Capistrano) UIMI, (Sweetwater Valley) SDNHM. NEVADA: Clark County (Logandale) NSDA.

Brachinus aeger Chaudoir

(Figs. 140, 144, 147, 148, 149, 168)

Brachynus aeger Chaudoir, 1876: 82. Lectotype, here selected, a female, MHNP, labelled

“Nouve Grenade,” and standing first in a series of five specimens. Type locality. — Col-

ombia, South America, as originally given by Chaudoir.

Diagnostic combination. — Only the genitalia provide reliable diagnosis, but see key coup-

let 18.

Description. — Small to medium-sized beetles, 5.8 to 8.9 mm.

Color. As in lateralis, except mesepisterna usually pale.

Microsculpture. As described for genus.

Macrosculpture. As in lateralis.

Head. As in lateralis, except ligula without inconspicuous setae.

Prothorax. As in lateralis, except costae weaker.

Pterothorax. As in lateralis, except costae weaker.

Abdomen. As described for genus.

Genitalia. Male (figs. 147, 148, 149). Median lobe with plane of shaft barely rotated from

plane of basal bend. Basal bend long. Shaft bulbous at basal third; narrowed to acute boot-

shaped apex; venter ridged medially. Ligule moderately long, broad, rounded apically. Virga

(figs. 147, 148). Female (fig. 144). Stylus short, parallel-sided, narrowly rounded at apex.

Variation. - As in lateralis, except disc of elytra generally pubescent.

Flight. — The flight of these beetles has not been recorded.

Etymology . — Latin, aeger,, sick, troubled; referring, I think, to the pale colors of these

beetles compared to others in the genus.

Collecting notes. — G. E. Ball and D. R. Whitehead have collected these beetles on gravel

banks of several rivers in Mexico.

Life history. — Members of this species have been collected from April to June and

November to January, but no teneral adults were seen.

Distribution. — (Fig. 168). The range of this species extends from Sonora, Mexico, south

to Colombia, South America. Most records are from the western coast of Mexico, but one

specimen was collected in San Luis Potosi, Mexico. I have seen 15 specimens from the fol-

lowing localities:

CENTRAL AMERICA

GUATEMALA: (Paso Antonio, Escuintla) BMNH.

78

Erwin

Figs. 165-170. Geographical distribution maps. 165. Brachinus chirriador new species. 166. Brachinus adustipennis

Erwin. 167. Brachinus chalchihuitlicue new species. 168. Brachinus aeger Chaudoir. 169. Brachinus lateralis Dejean.

170. Brachinus aboreus Chevrolat.

Bombardier Beetles

79

MEXICO

CHIAPAS: (Tuxtla Gutierrez) BMNH. GUERRERO: (Rio Mezcala, 23.7 miles north of Zumpango) UASM. NAYARIT:

(Rio Acaponeta, 2.4 miles south of Acaponeta) UASM; (19.0 miles southeast of Tepic) CAS. OAXACA: (Oaxaca City)

BMNH. SAN LUIS POTOSI: (El Pujal) CPBo. SINALOA: (Camino, Real de Piaxtla) AMNH. SONORA: (Ciudad Obregori)

CNC; (Cocorit) UCR. TABASCO: (Rio Teapa, near Teapa) UASM. VERACRUZ: (San Andres Tuxtla) BMNH.

Brachinus chalchihuitlicue new species

(Figs. 135, 143, 159, 160, 161, 167)

Type locality. — San Bias, Nayarit, Mexico.

Type specimens. — The holotype male and allotype female are in the entomological col-

lections at CAS; both were collected at the type locality by B. Malkin on September 17-21,

1953. Twenty paratypes collected at various localities and on various dates are in AMNH,

CAS, MCZ, TLEr, UASM.

Diagnostic combination. — Although members of this species have their elytral costae

elevated more than any other in the group, the only reliable diagnostic characters are the

genitalia, but also see key couplet 20.

Description. — Medium-sized beetles, 8.7 to 1 1.0 mm.

Color. Antennal articles 3 and 4, mesepimera, metepisterna, abdominal sterna at least at

sides, abdominal terga, and knees infuscated, otherwise ferrugineous. Legs testaceous. Dorsal

surface of elytra brown, epipleura slightly paler.

Microsculpture. As described for genus.

Macrosculpture. As in lateralis.

Head. As in lateralis, except ligula without minute setae.

Prothorax, ks'm lateralis. Pronotum (fig. 135).

Pterothorax. As in lateralis, except costae more highly elevated.

Abdomen. As described for genus.

Genitalia. Male (figs. 159, 160, 161). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend long. Apex of shaft very narrow, elongate. Ligule

long and broad, rounded apically. Virga (figs. 159, 160). Female (fig. 143). Stylus broad,

tapering to almost acute apex.

Variation. — Besides the intrapopulational variation in shape of the pronotum and in

body size, these beetles vary locally in the height of their costae.

Flight. — The flight of these beetles has not been recorded.

Etymology. - Chalchihuitlicue, the goddess of runoff waters, streams, lakes, and the sea,

in early Teotihuacan and Nahuat-Toltec cultures of central Mexico. This goddess is also the

wife of Tlaloc, the god of rain and thunder. The name refers to the habitat in which most

Brachinus species are found.

Life history. — Members of this species have been collected from July to September. Ten-

eral adults were collected in July in Sinaloa and in August in Guerrero.

Distribution. — (Fig. 167). The known range of this species extends from Sinaloa to

Guerrero on the west coast of Mexico. I have seen 30 specimens from the following local-

ities:

MEXICO

GUERRERO: (Acapulco) MCZ. NAYARIT: (San Bias) CAS; (Tepic) AMNH. SINALOA: (Mazatlan) AMNH, GRNo,

UCR; (Venedio) CAS. STATE UNKNOWN: (Saltillo) MCZ.

Brachinus arboreus Chevrolat

(Figs. 138, 146, 150, 151, 152, 170)

Brachinus arboreus Chevrolat, 1834: 42. Lectotype, here selected, a male, HMO, Type

number COL. 114 1/3, further labelled “Brachinus arboreus Chev. Col. Mex. 1 cent No. 2

Mexico Salle.” Type locality. - Orizaba, Mexico, as originally given by Chevrolat.

80

Erwin

Diagnostic combination. — Only the genitalia provide reliable diagnosis, but see also key

couplet 21 .

Description. — Small to medium-sized beetles, 6.1 to 9.2 mm.

Color. Base of antennal articles 3 and 4, mesepisterna, metepisterna, abdominal sterna at

least at sides, abdominal terga, and knees infuscated, otherwise ferrugineous. Dorsal surface

of elytra brown, epipleura seldom paler.

Microsculpture. As described for genus.

Macro sculpture. As in lateralis.

Head. As in lateralis, except ligula without inconspicuous setae.

Prothorax. As in lateralis. Pronotum (fig. 138).

Pterothorax. As in lateralis.

Abdomen. As described for genus.

Genitalia. Male (figs. 150, 151, 152). Median lobe with plane of shaft barely rotated from

plane of basal bend. Basal bend short. Apex of shaft narrowed, acute, ridged ventrally at

middle and at sides forming two small sulci. Ligule short, broad, truncate. Virga (figs. 150,

151). Female (fig. 146). Stylus narrow, parallel-sided, slightly curved, narrowly rounded

apically.

Variation. — As in lateralis.

Flight. — The flight of these beetles has not been recorded.

Etymology . — Latin, arboreus, of the trees. The reason Chevrolat gave this name is un-

explained in his description.

Collecting notes. — D. R. Whitehead has collected these beetles along a small stream in

Jalisco.

Life history. — Members of this species have been collected in January, March, June, and

November, but no teneral adults were seen.

Distribution . — (Fig. 170). The range of this species extends from Sinaloa south to Hon-

duras. I have seen 57 specimens from the following localities:

CENTRAL AMERICA

HONDURAS: (Copan) MCZ; (La Lima) DTRT.

MEXICO

JALISCO: (9.0 miles east of Guadalajara) AMNH. NAYARIT: (32.0 miles south of Acaponeta) CAS; (19.0 miles southeast

of Tepic) CAS. MORELOS: (5.0 miles east of Cuernavaca) UCD. SINALOA: (Mazatlan) MCZ, TLEr; (Rosario) CAS.

VERACRUZ: (Cordoba) BMNH; (Jalapa) BMNH, MCZ.

Brachinus chirriador new species

(Figs. 136, 141, 162, 163, 164, 165)

Type locality. — Six miles west of Cintalapa, Route 190, Chiapas, Mexico.

Type specimens. - The holotype male and allotype female are in the MCZ. The holotype

was collected by G. E. Ball and D. R. Whitehead at the type locality on September 7, 1965.

The allotype was collected at Jacala, Hidalgo by R. Haag on June 23, 1939. Eight paratypes

collected on various dates at various localities are in CAS, CPBo, TLEr, UASM.

Diagnostic combination. — The restriction of elytral pubescence to depressions 7 and 8 in

the basal two-thirds of the elytra will usually separate these beetles from others with brown

elytra in Mexico and Central America. Only the genitalia provide reliable diagnosis, however.

Description. — Medium-sized beetles, 7.2 to 10.0 mm.

Color. As in lateralis, except epipleura dark.

Microsculpture. As described for genus.

Macrosculpture. As in lateralis.

Head. As in lateralis.

Prothorax. As in lateralis, except anterior tibia with anterior surface finely strigose.

Bombardier Beetles

81

Pronotum (fig. 136).

Pterothorax. As in lateralis , except pubescence restricted to depressions 7 and 8.

Abdomen. As described for genus.

Genitalia. Male (figs. 162, 163, 164). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend short. Apex of shaft narrow, slightly elongate. Ligule

short, broad, truncate. Virga (figs. 162, 163). Female (fig. 141). Stylus very broad, tapering

apically to narrowly rounded apex.

Variation. — Too few specimens are known to evaluate geographic variation.

Flight. - The flight of these beetles has not been recorded.

Etymology . — Mexican-Spanish, chirriador, the one who crepitates.

Collecting notes. — G. E. Ball and D. R. Whitehead collected these beetles at the edge of a

pond at the type locality by treading coarse grass and emergent vegetation.

Life history \ — Members of this species have been collected from April to June, and Aug-

ust to December, but no teneral adults were seen.

Distribution. — (Fig. 165). The known range of this species extends from northern Tam-

aulipas, Mexico, south along the east coast to Honduras and on the west coast of Mexico in

Sinaloa and Nayarit. I have seen 1 1 specimens from the following localities:

CENTRAL AMERICA

HONDURAS: (La Lima) DTRT.

MEXICO

CHIAPAS: (6.0 miles west of Cintalapa) UASM. JALISCO: (Puente La Garita, near La Garita) UASM. NAYARIT: (San

Bias) CAS. SAN LUIS POTOSI: (El Pujal) CPBo. TAMAULIPAS: (La Coma, Aldama) CPBo. VERACRUZ: (San Rafael,

Jicaltepec) MCZ; (Veracruz) UCD.

Brachinus adustipennis Erwin

(Figs. 139, 142, 153, 154, 155, 166)

Type locality. — Myakka River, Myakka River State Park, Sarasota County, Florida.

Type specimens. — The holotype male and allotype female are in MCZ; both were

collected at the type locality by T. L. and L. J. Erwin on May 29, 1968. Thirty paratypes

collected at various localities and on various dates are in AMNH, CAS, DRWh, MCZ, TLEr,

and UASM.

Diagnostic combination. — The brown elytra and pale epipleura, and pubescent elytral

disc separate members of this species from all others east of the continental divide in the

United States. In Mexico and further south only the genitalic characters provide reliable

separation, but see also key couplet 17.

Description. — Medium-sized beetles, 6.0 to 10.2 mm.

Color. Metepisterna, knees and sides of abdominal sterna infuscated. Antennal articles 3

and 4, usually terga slightly infuscated, otherwsie body and limbs testaceous to ferrugineous.

Dorsal surface of elytra brown, epipleura usually paler.

Microsculpture. As described for genus.

Macrosculpture. As in lateralis.

Head. As in lateralis, except minute ligula setae apparently absent.

Prothorax. As in lateralis, except anterior tibia with anterior surface strigose. Pronotum

(fig. 139).

Pterothorax. As in lateralis, except disc of elytra usually pubescent, and costae weaker.

Abdomen. As described for genus.

Genitalia. Male (figs. 153, 154, 155). Median lobe with plane of shaft barely rotated from

plane of basal bend. Basal bend moderately long. Apex of shaft narrowed, rounded, and

ridged ventrally; lateral ridges less prominent than in lateralis, ending before apex; apex

slightly bent dorsally. Ligule short, narrowed toward rounded apex. Virga (figs. 153, 154).

82

Erwin

Female (fig. 142). Stylus narrow, broadened slightly at apical third, rounded apically.

Variation. — As in lateralis, except disc of elytra usually pubescent.

Flight. — The flight of these beetles has been recorded at lights throughout the range of

the species.

Etymology. — Latin, adustus, tanned, brown; pennis, wing; referring to the tan-colored

elytra of these beetles.

Collecting notes. — My wife and I collected these beetles in a number of localities in the

southern United States. Along the Myakka River, at the type locality, they occurred beneath

boards and stones on the river’s grassy banks. At Juniper Springs, Florida, and in the

Okefenokee Swamp, we found them in the roots of grass clumps in Scirpus-Typha marshes.

In Texas they occurred on the muddy banks of the San Jacinto River in grassy clumps.

Life history. — Members of this species have been collected from March to December.

Teneral adults were collected in May in Florida, and in August in Tennessee. Overwintering

(or aestivating) probably occurs in the adult state.

Distribution . — (Fig. 166). The range of this species extends from New York and Michi-

gan, west to the west coast of Mexico, south to Panama. I have seen 552 specimens from the

following localities:

CENTRAL AMERICA

PANAMA: (La Chorrera) CAS.

GREATER ANTILLES

CUBA: (No locality given) TMBH; (Camoa, Havana) MCZ; (Soledad) MCZ.

MEXICO

AGUASCALIENTE: (15.0 miles west of Pabellon) UMAH. NAYARIT: (Acaponeta) AMNH; (32.0 miles south of

Acaponeta) CAS. SINALOA: (Los Mochis) CAS; (Mazatlan) AMNH, CAS, UASM; (20.0 miles west of Rosario) UCR;

(Venedillo) CAS. SONORA: (16.0 miles northeast of Ciudad Obregon) CNC; (Hermosillo) CNHM; (Rio Yagui, 12.0 miles

west of Ciudad Obregon) CNC. TABASCO: (San Juan Bautista) BMNH; (Villa Hermosa) FDAG. TAMAULIPAS: (9.9 miles

west of La Pesca) UASM. VERACRUZ: (Veracruz) UASM. YUCATAN: (Uxmal) TLEr.

UNITED STATES

ALABAMA: Mobile County (Mobile) ANSP, USNM. ARKANSAS: Arkansas County (Almyra) USNM; Desha County (No

locality given) UAFA; Hempstead County (Hope) CAS, MCZ; Poinsett County (No locality given) UAFA; Pulaski County

(Little Rock) AMNH; Washington County (No locality given) UAFA. FLORIDA: Alachua County (Gainesville) FDAG,

UMAH, (Payne’s Prairie) UMAH; Brevard County (Indian River) USNM; Citrus County CAS, USNM, (Inverness) MCZ;

Collier County (Naples) CUNY ; Columbia County (Lake City) DRWh; Dade County (Florida City) UASM, (Miami) CMPP,

(Royal Palm State Park) PUM; Escambia County (Pensacola) FDAG; Hemando County (Brooksville) CAS; Highlands

County (Archbold Biology Station) CEWh, PSUU; Hillsborough County (Tampa) CAS, UMSP; Lake County USNM,

(Fruitland Park) ANSP, UMAH, (Groveland) FDAG, (5.6 miles east of Juniper Springs) TLEr; Lee County (Fort Myers)

PUM; Levy County (Cedar Keys) USNM, (Manatee Springs State Park) RFre, UASM; Manatee County (Oneco) UASM;

Marion County (No locality given) MCZ; Osceola County (Kissimmee) MCZ, PUM, UCD, USNM; Palm Beach County

(Lake Worth) CAS; Pasco County (Dade City) FDAG; Pinellas County (Clearwater) CNHM, (Dunedin) AMNH, CAS,

CUNY, UMAH, UWMW, (Tarpon Springs) AMNH, CNHM; Putnam County (Crescent City) USNM; Sarasota County

(Myakka River State Park) TLEr, UASM, (Sarasota) PUM; Seminole County (Lake Harney) USNM; Volusia County

(Enterprise) CAS, ISNH, OUCO, UMSP, USNM; County unknown (Fort Capron) ISNH. GEORGIA: Baker County

(Newton) CNC; Tift County (Tifton) OUCO; Ware County (8.0 miles northeast of Fargo) TLEr. ILLINOIS: Rich-

land and Lawrence Counties (Wabash) MCZ. INDIANA: Floyd County (New Albany) CEWh; Posey County (No

locality given) PUM, KANSAS: Douglas County (Lawrence) PUM. LOUISIANA: USNM, Acadia Parish (Crowley)

CAS; Allen Parish (Kinder) UASM; Calcasieu Parish (Lake Charles) USNM, (Sam Houston State Park) CUNY, TLEr;

East Baton Rouge Parish UAFA, (Baton Rouge) LSUB, UMAH; Evangeline Parish (Lake Chicot State Park) TLEr; Franklin

Parish (Chase) UAFA; Iberia Parish (Avery Island) ANSP; Jefferson Parish (Harahan) CNHM; Livingston Parish (Denham

Springs) LSUB; Madison Parish (Tallulah) TAMU; Orleans Parish (New Orleans) ANSP, CAS, LACM, MCZ, USNM, UWMW,

WSUP; Ouachita Parish (Calhoun) UAFA; Saint John the Baptist Parish (Garyville) LSUB; Saint Martin Parish (No locality

given) UAFA; Vermilion Parish (Gueydan) USNM; Vernon Parish (Rosepine) UAFA; Parish unknown (Mound) FDAG.

MICHIGAN: Benzie County (No locality given) PUM. MISSISSIPPI: Hinds County (Jackson) UCR; County unknown

(McCormick) UWSW. MISSOURI: Boone County (Columbia) UCD; Saint Louis County (Saint Louis) UCR. NEW MEX-

ICO: Chaves County (Roswell) UWSW. NEW YORK: Westchester County (Peekskill) MCZ. OKLAHOMA: Carter County

(Ardmore) OSUS. TENNESSEE: Davidson County (Nashville) USNM; Madison County (Jackson) CNC. TEXAS: Atascosa

County (Pleasanton) TAMU; Blanco County (Cypress Mills) USNM; Brazos County (College Station) TAMU; Cameron

County (Brownsville) CAS, CNC, CUNY, OUCO, TLEr, USNM; Colorado County (No locality given) UMSP; Dallas County

(Dallas) MCZ; El Paso County (El Paso) CMPP; Fayette County (Engle) CAS; Frio County (5.0 miles north of Dilley)

UASM; Hardin County (9.0 miles west of Beaumont) OSUC; Harris County USNM, (Highway 59 at San Jacinto River,

near Houston) TLEr; Harrison County (near Sabine River) TCBa; Hidalgo County (Mercedes) USNM, (Weslaco) TAMU;

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83

Jefferson County (20.0 miles southwest of Sabine Pass) RCGr; Kerr County (Kerrville) CNC; Kleberg County (Kingsville)

CUNY; Lee County (No locality given) UMAH; Smith County (Lake Palestine, 17.0 miles southwest of Tyler) DRWh;

Travis County (Austin) CAS; Val Verde County (Del Rio) CNC.

The kansanus group

This group is characterized by its large shield-like virga, sharply costate elytra and lack of

lateral pronotal setae. A single species, B. kansanus LeConte, is included.

Brachinus kansanus LeConte

(Figs. 175, 176, 177, 178, 195, 198)

Brachinus kansanus LeConte, 1862: 524. Lectotype, here selected, a male, MCZ red type

label number 5851, further labelled with a green disc. Type locality. — Kansas, as origin-

ally given by LeConte.

Diagnostic combination. — This species is best characterized by its sharply costate elytra,

lack of lateral pronotal setae, very narrowly reflexed sides of the pronotum, and almost

glabrous proepipleura.

Description. — Medium-sized beetles, 8.7 to 11.5 mm.

Color. Ferrugineous, sides of abdomen of some specimens slightly infuscated. Dorsal sur-

face and epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Head behind eyes and frontal furrows rugose and shallowly punctate.

Disc of pronotum with numerous shallowly impressed setiferous punctures.

Head. Frontal furrows moderately impressed. Antennal scape robust, widened apically.

Ligula with sclerotized center area ellipsoid-convex with two apical setae. Mentum and sub-

mentum various, with or without accessory setae.

Prothorax. Pronotum (fig. 175) convex, flattened along center line, sides very narrowly

reflexed. Proepipleura and proepistema with a few scattered setae. Anterior tibia with an-

terior edge strigose.

Pterothorax. Elytra elongate, narrow, strongly costate. Humeral angle square. Costae

smooth and glabrous, depressions between costae pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 176, 177, 178). Median lobe with plane of shaft rotated about 45°

from plane of basal bend. Basal bend short. Median lobe nearly straight, slightly swollen at

middle.* Apex of shaft broadly rounded. Ligule short, very broad, truncate. Virga (figs.

176, 177). Female (fig. 195). Stylus elongate, narrow, parallel-sided, almost blunt apically.

Variation. — Besides the intrapopulational variation in body size and shape of the prono-

tum, these beetles vary in number of accessory setae of the mentum and submentum, the

setal number of the ligula, and in the color of the elytra (blue to blue-green). The range of

variation in all characteristics is seen in single population samples.

Flight. — The flight of these beetles has not been recorded.

Etymology . — The latinized form of Kansas, the place where the type was collected.

Collecting notes. — This species occurs along river and stream courses in very sandy areas

beneath the broadleaf deciduous forest which follows these water courses into the Great

Plains. K. L. Hays has also collected these beetles in the sand dunes near Manhattan, Kansas.

Life history. — Members of this species have been collected from June to October. One

teneral adult was collected in July in Oklahoma. Overwintering is probably in the adult

stage.

Distribution. — (Fig. 198). This is the only species restricted to the Great Plains area. It

occurs along eastward flowing tributaries of the Mississippi and Missouri Rivers. I have seen

95 specimens from the following localities:

186 187 188 189 190 191

Figs. 171-175, 179. Pronotum, right half, dorsal aspect. 1 7 1 . Brachinus rugipennis Chaudoir, Highlands County, Flor-

ida. 172. Brachinus altemans Dejean, Dallas County, Texas. 173. Same, Dallas County, Texas. 174 . Brachinus costi-

pennis Motschulsky, 12.2 miles south of El Banco, Durango, Mexico. 175. Brachinus kansanus LeConte, Scandia,

Kansas. 179. Brachinus viridipennis Dejean, Mobile, Alabama. Figs. 176-178, 180-191. Male genitalia. 17 6. Brachinus

kansanus LeConte, Scandia, Kansas, ventral aspect. 177. Lateral aspect of same. 178. Dorsal aspect of same. 180. Bra-

chinus rugipennis Chaudoir, Archbold Research Station, Florida, ventral aspect. 181. Lateral aspect of same. 182. Dor-

sal aspect of same. 183. Brachinus viridipennis Dejean, Mobile, Alabama, ventral aspect. 184. Lateral aspect of same.

185. Dorsal aspect of same. 186. Brachinus costipennis Motschulsky, 12.2 miles south of El Banco, Durango, Mexico,

ventral aspect. 187 & 188. Lateral & dorsal aspects of same. 189. Brachinus altemans Dejean, Okefenokee Swamp,

Georgia, ventral aspect. 190 & 191. Lateral & dorsal aspects of same. Accompanying scale lines equal 1.0 mm.

Bombardier Beetles

85

UNITED STATES

ARKANSAS: Crawford County (No locality given) USNM. ? DAKOTA: (No locality given) MCZ. KANSAS: Atchison

County (No locality given) CAS; Douglas County (No locality given) CMPP; Republic County (Republican River, Scandia)

CAS; Riley County KSU, (Manhattan) USNM. ILLINOIS: Adams County (Quincy) CNHM; Calhoun County (Kamps-

ville) MCZ. IOWA: Boone County (Ledges State Park) ISUA. MINNESOTA: (No locality given) UMSP. MISSOURI: (no

locality given) ANSP, ISNH, MCZ, UMSP. NEBRASKA: Dodge County (Fremont) UNLN; Otoe County (Nebraska City)

UNLN; Richardon County (Rulo) UNLN; Sioux County (Monroe Canyon) UNLN. OHIO: Preble County (No locality

given) PUM. OKLAHOMA: Beckham County (No locality given) CAS; Cleveland County (No locality given) UONO; Grady

County (Chickasha) USNM; Payne County (Stillwater) OSUS; Woodward County (Woodward) CNHM.

The costipennis group

This group is characterized by the form of the virga, form of the median lobe of the male

genitalia, completely glabrous elytra, sulcate mentum surrounded by setae (shared with B.

mobilis and some Old World species), and abbreviated submentum. One species, B. costi-

pennis Motschulsky, is included.

Brachinus costipennis Motschulsky

(Figs. 18, 174, 186, 187, 188, 196, 197)

Brachinus costipennis Motschulsky, 1859: 138. Lectotype, here selected, a female, MCZ

red type label number 8329. Further labelled with a green square and “37.” As I pointed

out (1965: 5), “this specimen is very likely a cotype from Motschulsky, with whom

LeConte is known to have corresponded.” Type locality. — California, as given originally

by Motschulsky.

Brachinus carinulatus Motschulsky, 1859: 139. Lectotype, here selected, a male, MMM,

labelled with a green square and ‘Brachynus carinulatus Motsch. California.” Type local-

ity. — California, as originally given by Motschulsky. Erwin 1965: 4.

Brachynus cognatus Chaudoir, 1876: 74. Lectotype, here selected, a female, MHNP, lab-

elled “cognatus m. Mexico” and “Orizaba” on green paper and “Ex Museo Chaudoir” on

white paper. Type locality. — Orizaba, Mexico, as given on Chaudoir’s label. NEW SYN-

ONYMY.

Brachinus cancellatus Bates, 1891: 269. Lectotype, here selected, a male, BMNH, labelled

“Chihuahua City, Mexico,” and “Hoge.” It is placed in the series labelled “B. cognatus

Chaudoir.” Type locality. — Chihuahua City, Mexico, as originally given by Bates. NEW

SYNONYMY.

Diagnostic combination. — The lack of elytral pubescence immediately separates mem-

bers of this species from all others of the genus.

Description. — Small-sized beetles, 5.0 to 8.0 mm.

Color. Ferrugineous, except antennal article 4 occasionally infuscated. Dorsal surface and

epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows rugose and punctate. Surface of pronotum microrugose,

with an occasional setiferous puncture, but usually glabrous.

Head. Frontal furrows moderately impressed. Antennal scape cylindrical. Ligula with

sclerotized center area ellipsoid-convex with two paramedian rows of three setae per row.

Mentum (fig. 18) sulcate medially, sulcus surrounded by setae. Submentum (fig. 18) shor-

tened and with accessory setae.

Prothorax. Pronotum (fig. 174) convex, flattened along center line, sides slightly reflexed.

Proepipleura and proepisterna usually glabrous. Anterior tibia with anterior edge strigose.

Pterothorax. Elytra short and convex with quite arcuate sides. Humeral angles square,

costae elevated and smooth, depressions between costae slightly rugose. Wings fully deve-

loped.

Abdomen. As described for genus.

I — (

Figs. 192-196. Right stylus of female ovipositor, ventral aspect. 192. Brachinus rugipennis Chaudoir, Hope, Arkansas.

193. Brachinus viridipennis Dejean, Mobile, Alabama. 194. Brachinus altemans Dejean, Seabrook, Texas. \95. Brach-

inus kansanus LeConte, Scandia, Kansas. 196. Brachinus costipennis Motschulsky, 26.1 miles north of Glenwood,

New Mexico. Figs. 1 97-20 1 . Geographical distribution maps. 197. Brachinus costipennis Motschulsky. 198 .Brachinus

kansanus LeConte. 199. Brachinus viridipennis Dejean. 200. Brachinus rugipennis Chaudoir. 201. Brachinus altemans

Dejean. Accompanying scale line equals 1 .0 mm.

Bombardier Beetles

87

Genitalia. Male (figs. 186, 187, 188). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend short. Median lobe slightly swollen medially, just be-

fore bend. Apex of shaft narrowed, blunt and deflected ventrally. Ligule short, broad, and

truncate. Virga (figs. 186, 187). Female (fig. 196). Stylus short, narrow, tapering to acute

apex.

Variation. — Besides the intrapopulational variation in body size and shape of the pron-

otum, these beetles are rather constant throughout the range of the species.

Flight. — These beetles have been recorded coming to lights in Mexico.

Etymology . — Latin, costa, rib; pennis, wing; referring to the raised costae of the elytra.

Collecting notes. — I collected these beetles along an intermittent stream on the eastern

slope of the Mount Hamilton Range near San Jose, California. The sides of the stream were

lined with specimens of Salix and Quercus species and an occasional specimen of Platanus

species. The beetles were beneath small and large stones near the water.

Life history . — Members of this species have been collected in all months, except January.

Teneral adults were collected in March in Oaxaca and in July in California.

Distribution. — (Fig. 197). The range of this species extends from Utah and northern Cali-

fornia to Chiapas, Mexico, and east to the Big Bend area of Texas. The specimens labelled

“Arkansas,” “Kansas,” and “Yukon,” must be suspected of being mislabelled. I have seen

1,810 specimens from the following localities:

CANADA

YUKON: (No locality given) ISNH, WSUP.

CENTRAL AMERICA

GUATEMALA: (Agua Caliente) MCZ; (Los Amates) MCZ.

MEXICO

BAJA CALIFORNIA: (Hamilton Ranch) CAS; (San Vicente) CAS; (Tijuana) CNHM. DURANGO: (Durango City) MCZ;

(12.2 miles south of El Banco) UASM. JALISCO: (Atenquique) CAS. OAXACA: (25.0 miles south of Mitla) ISUA;

(Oaxaca City) CAS; (22.5 miles west of Oaxaca) UASM; (Paderon, Rio Tehuantepec) AMNH; (Rio Atoyac, near Jucha-

tengo) UASM; (Rio Malatengo, 11.1 miles north of Matias Romero) UASM; (72.5 miles south of Valle Nacional) UASM.

PUEBLA: (near Petlalcingo) UASM; (Puente Tepexco, near Tepexco) UASM. SAN LUIS POTOSI: (El Pujal) CPBo;(2.7

miles west of Santa Catarina) UASM; (Tamazunchale) AMNH, CAS, MCZ. SINALOA: (Roserio) CAS. SONORA: (7.2

miles southeast of Alamos) GRNo; (Hermosillo) CAS; (Sonoyta) AMNH. VERACRUZ: (Fortin de las Flores) UASM,

CPBo; (Orizaba) UNLN. ZACATECAS: (Juchipila 0.9 miles north of Jalpa) UASM.

UNITED STATES

ARIZONA: Cochise County (Chiricahua Mountains) CAS, (17.0 miles east of Douglas) UCR, (Dragoon Mountains, Texas

Pass) MCZ, (Portal) GRNo, RCGr, (San Pedro River, near Palominas) UASM, (San Pedro River, east of Sierra Vista) UATA,

(Tombstone) SDNHM; Coconino County (Grand Canyon, mile 52.0) UATA, (Grand Canyon, Havasupai Indian Reser-

vation) UMSP, (Oak Creek Canyon, near Flagstaff) CAS; Gila County (Carrizo Creek, near Carrizo) DRWh, (Gila Valley)

CAS, (near Globe) UATA, (Payson) UATA, (Pinal Mountains) SDNHM, (San Carlos Lake) UATA, (Winkelman) UATA;

Graham County (Aravaipa) CAS, (Aravaipa Creek, between Klondyke and Aravaipa) DRWh, (Geronimo) CAS, UATA;

Maricopa County (Phoenix) CNC, MCZ, OUCO; Navajo County (8-15.0 miles northeast of Whiteriver) AMNH; Pima County

(Saint Xavier Mountains) CAS, (Tucson) AMNH, CAS, MCZ, USNM; Pinal County (Aravaipa Canyon) CUNY, (Florence)

ANSP, CMPP, (Sycamore Camp, 9.0 miles northwest of Payson) CAS; Santa Cruz County (Lewis Springs, San Pedro River)

UASM, (Patagonia) CAS, CNHM, CUNY, UATA, UCD, UCR, (5.0 miles southwest of Patagonia) AMNH, (Pena Blanca)

UASM; Yavapai County (Bumble Bee) CAS, (Camp Verde) CAS, (Congress) UATA, (Prescott) AMNH, MCZ, (Verde River)

USNM, (5.0 miles north of Wickenburg) UMAH; Yuma County (Fort Yuma) USNM, (Yuma) LACM, USNM; County

unknown (Gila River) CNC. ARKANSAS: Hempstead County (Hope) MCZ. CALIFORNIA: Alameda County (Oakland

Hills) CAS; Calaveras County (Mokelumne Hill) CAS; Fresno County (Camp Greeley) CAS, (Le Fevre Creek) CAS, (Sanger)

CAS, (Trimmer) UMSP; Humboldt County (Garberville) CAS, LACM; Imperial County MCZ, (Carrizo) SDNHM, (Castiac)

UIMI; Kern County AMNH, USNM, (Caliente) BMNH; Los Angeles County PSUU (Big Dalton Dam) UCD, (Big Tujunga)

LACM, (Burbank) CNHM, ISNH, (Frenchman’s Flats) GRNo, LACM, (Lake Arrowhead) CAS, (Los Angeles) CNHM,

UATA, UNLN, USNM, (Los Angeles River) LACM, UCR, (Newhall) CUNY, USNM, (Pasadena) ANSP, CMPP, CNHM,

CUNY, LACM, MCZ, (Rio Hondo) LACM, (Tujunga Pass) UCD, (San Francisquito Canyon) LACM, UCD, (San Gabriel

Canyon) GRNo, TCBa, (San Gabriel Mountains, Camp Bonita) LACM, (Santa Monica) UMSP; Madera County (Course-

gold) CAS, UIMI; Mendocino County (Navarro River, 2.0 miles northwest of Philo) CAS, (Ukiah) WBa; Monterey County

(Salinas) USNM, (Stone Canyon) CAS; Orange County (Costa Mesa) UCD, UCR, (Huntington Beach) USNM, (Olive)

TAMU, (Santa Ana) CMPP, MCZ, TAMU; Placer County (No locality given) CAS ; Riverside County (Corona) USNM, (El-

sinore) CMPP, (Elsinore Lake) CAS, (Gilman Hot Springs) UCD, (Hemet) VVBa, (Palm Canyon) LACM, (Riverside) ANSP,

CAS, CUNY, UMAH, USNM, (Simond’s) LACM, (Temecula) CAS, (White Water Canyon) SJSC, UCR; Sacramento County

(Folsom) USNM; San Bernardino County OUCO, (Afton Canyon) USNM, (San Bernardino) CAS, ISNH, (Cajon) CNHM,

(Cajon Pass) UCD, (Cajon Wash) LACM, (Colton) CAS, CUNY, MCZ, UATA, UCR, UMAH, (Redlands) MCZ; San Diego

88

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County CCha, UMAH, (Banner) SDNHM, (Dehesa) SDNHM, (Descanso) CAS, (El Monte Oaks) SDNHM, (Mission Dam)

SDNHM, (Mission Valley) SDNHM, (Mountains of San Diego County) USNM, (Oceanside) CAS, (Pine Valley) CNC, MCZ,

SDNHM, UMSP, (Poway) CAS, USNM, (Rincoln) SDNHM, (San Diego) ANSP, CNHM, CUNY, MCZ, SDNHM, USNM, (San

Pasqual) UCR, (Valley Center) SDNHM, (Warner’s Hot Springs) PUM, (Warner’s Ranch) SDNHM; San Francisco County

(San Francisco) CMPP; San Luis Obispo County (Arroyo Grande) CAS; Santa Barbara County (Santa Barbara) MCZ, (Santa

Cruz Island) CAS, (Santa Cruz Island, Christie Beach) TLEr, (Santa Cruz Island, Christie Ranch) UCR, (Shephard’s Inn)

CAS; Santa Clara County (Adobe Creek) CAS; Sonoma County (Dry Creek, 9.0 miles northwest of Healdsburg) SJSC,

(Duncan Mills) CAS, (2.0 miles east of Healdsburg) CAS, (Rio Nido) CAS; Stanislaus County (15.0 miles west of Patterson)

TLEr; Tehama County (western hills of Tehama County) CAS; Tulare County (Sequoia National Park) WBa; Ventura

County (Santa Paula) ANSP, CAS; Yolo County (Davis) UCD; Counties unknown (Aliso Creek) UWSW, (Colorado Desert)

MCZ, (Sylvania) MCZ. KANSAS: Douglas County (No locality given) CUNY. NEW MEXICO: Catron County (near Ara-

gon) AMNH, (San Francisco Creek, 26.1 miles north of Glenwood) DRWh, UASM; Grant County (18.0 miles north of

Mimbres, Roberts Lake) TLEr. TEXAS: Brewster County (Big Bend National Park, Hot Springs) CNC, (Rio Grande) CAS.

UTAH: Garfield County (Boulder) ISUA; Grand County OUCO, (Moab) CAS, Washington County SDNHM, (3.0 miles

south of Gunlock) GRNo, (Saint George) AMNH, KSU, MCZ, (Santa Clara Creek) UCD, (Zion National Park) CAS.

The alternans group

This small group of three species is characterized by the tripartite virga of the endophallus

with elongate median apex, and plurisetose ligula. The group is divided into two subgroups.

The alternans subgroup

This subgroup is characterized by the accessory setae of the mentum and submentum, and

the narrow ligule of the male median lobe. Two closely related species, B. alternans Dejean

and B. viridipennis Dejean, are included

Brachinus alternans Dejean

(Figs. 172, 173, 189, 190, 191, 194, 201)

Brachinus alternans Dejean, 1825: 316. Lectotype, here selected, a female, MHNP, labelled

“alternans m. in Amer. bor.” on green paper, “Georgia” on green paper, “D. Escheri” on

green paper, and “Ex Museo Chaudoir” on white paper. Type locality. — Georgia, as orig-

inally given by Dejean.

Brachinus librator Dejean, 1831: 425. Lectotype, here selected, a male, MHNP, labelled

“librator m. in Amer. bor.” and “LeConte.” on green paper, and “Ex Museo Chaudoir”on

white paper. Type locality. — North America, as originally given by Dejean. NEW SYNO-

NYMY.

Brachinus deyrollei Laferte, 1841: 42. Lectotype, here selected, a male, MHNP, labelled

“Missouri, Reiche” and “Ex Museo Chaudoir.” Type locality. — Missouri, as given on the

label. NEW SYNONYMY.

Brachinus strennus LeConte, 1844: 48. Lectotype, here selected, a female, MCZ red type

label number 5844. Further labelled with an orange disc and “76.” Type locality. —

Georgia, as originally given by LeConte. NEW SYNONYMY.

Brachinus tormentarius LeConte, 1848: 200. Lectotype, here selected, a female, MCZ red

type label number 5845. Further labelled with a yellow disc and “77.” Type locality . —

Western States, as originally given by LeConte. NEW SYNONYMY.

Brachinus distinguendus Chaudoir, 1868: 287. Lectotype, here selected, a male, MHNP,

labelled “fumans h. in Amer. bor.” on green paper, and “Ex Museo Chaudoir” on white

paper. Type locality. - United States, as originally given by Chaudoir. NEW SYNONYMY.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Large-sized beetles, 1 1.5 to 16.5 mm.

Color. Antennae and venter various. Elytra blue, rarely with greenish luster.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows, head behind eyes, and surface of pronotum moderately

punctate, punctures moderately impressed.

Head. Frontal furrows moderately impressed. Antennal scape robust, widened apically.

Bombardier Beetles

89

Ligula with sclerotized center area ellipsoid-convex, plurisetose. Mentum and submentum

with numerous accessory setae.

Prothorax. Pronotum (fig. 172) convex, flattened along center line, sides barely reflexed.

Surface with punctures moderately impressed. Proepipleura and proepisterna pubescent.

Anterior tibia with anterior margin weakly strigose.

Pterothorax. Elytra elongate, broad, moderately costate. Humeral angle square. Costae

smooth, glabrous, depressions between costae pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 189, 190, 191). Median lobe with plane of shaft hardly rotated from

plane of basal bend. Basal bend short. Median lobe arcuate, slightly swollen at middle, apex

variable. Ligule short, narrow and truncate. Virga (figs. 189, 190). Female (fig. 194). Stylus

broad, arcuate, and elongate, apex narrowly rounded.

Variation. — The members of this species are among the most variable of the genus in

North America. Intrapopulational variation in the shape of the pronotum is illustrated in

figures 172 and 173. Antennal articles 3 and 4 are or are not infuscated. The color of the

ventor involves four combinations of ferrugineous and infuscated areas, as follows: sides of

abdomen infuscated, remainder ferrugineous; the preceding, plus the mesepisterna and

metepisterna, infuscated; both the preceding, plus the metasternum at sides, infuscated; and

finally, the preceding, plus all the abdomen, infuscated (except ferrugineous dimples). The

pale condition is commonest in the midwest, and rare toward the south and east. However,

one or more of these conditions is common to single populations. Variation occurs in the

apex of the shaft of the male genitalia also. The bent or unbent condition is independent of

the color cline described above, and (as color variation) is common to local populations.

Flight. — C. W. O’Brien has recorded these beetles flying to “blacklights” in Florida.

Etymology . — Latin, alternus, alternate; referring to the costate elytra.

Life history. — Members of this species have been collected during all months of the year.

Teneral adults were collected in April in Illinois, in August in Tennessee, in September in

Nebraska and Illinois, and in November in North Carolina. Overwintering probably takes

place as an adult.

Distribution. — (Fig. 201). The range of this species extends from New Mexico north to

Minnesota, east to Connecticut, and south to the Florida Keys. I have seen 1,141 specimens

from the following localities:

UNITED STATES

ALABAMA: Clarke County (Salt Mountain, 6.0 miles south of Jackson) UMAH; Lee County (Auburn) AUAA; Mobile

County (Magazine Point) CAS, (Mobile) ANSP, CAS, MCZ, (Mount Vernon) AUAA, OUCO; Tuscaloosa County (Tus-

caloosa) UASM. ARKANSAS: (No locality given) AMNH; Conway County (No locality given) UAFA; Jefferson County

(Pine Blinf) ISNH; Lawrence County (Imboden) CAS, USNM; Mississippi County UAFA, (Osceola) UMAH. CONNECTI-

CUT: New Haven County (Hamden) CAS. DISTRICT OF COLUMBIA: (Washington) UMSP. FLORIDA: Alachua County

(Gainesville) FDAG, RFre, UMAH, (Grant’s Cave) FDAG, (Poe Springs) UMAH; Baker County (Glen Saint Mary) FDAG;

Brevard County (Indian River) ISNH, USNM; Broward County (Fort Lauderdale) UMAH; Charlotte County (Punta Gorda)

CNC, CNHM; Citrus County (No locality given) CAS; Collier County (Everglades) OUCO, UMAH, USNM; Dade County

UATA, (18.0 miles northwest of Hialeah) TLEr, (Homestead) AMNH, FDAG, TLEr, (Long Pine Key) MCZ, (Miami)

UCD, (Paradise Key) AMNH, CNC, USNM, (Royal Palm State Park) AMNH, PUM, UCR, UMAH; Duval County (Arlington)

AMNH, (Jacksonville) AMNH, ANSP, USNM; Glades County (Moore Haven) USNM, (Palmdale) AMNH; Hendry County

(Clewiston) RCGr, (La Belle) PUM; Hernando County (Brooksville) CAS; Highlands County (Archbold Biology Station)

CMPP, CUNY, PSUU, (Highlands Hammock State Park) TLEr, (4.5 miles west of Venus) TCBa; Hillsborough County

(Plant City) UMAH, (Tampa) CAS; Jefferson County (Monticello) AMNH; Lake County UMAH, (Leesburg) CAS; Lee

County (Fort Myers) PUM, (Sanibel Island) CAS; Leon County (Lake Jackson) UMAH; Levy County (No locality given)

ANSP; Madison County (Greenville) FDAG; Manatee County (Oneco) UASM; Marion County (No locality given) MCZ;

Orange County (Orlando) OUCO, (Winter Park) MCZ; Osceola County (Kissimmee) AMNH, ANSP, CUNY, PUM; Palm

Beach County (Canal Point) CUNY, (Lake Worth) CMPP, (Stewart) UMAH; Pinellas County (Dunedin) CAS, PUM,

UWMW, (Saint Petersburg) CAS, PUM; Putnam County (Crescent City) USNM, (Welaka) UMAH; Saint Johns County

(Saint Augustine) AMNH, CAS; Sarasota County (Myakka State Park) CUNY, UASM, (Sarasota) PUM; Seminole County

(Sanford) MCZ, PUM; Volusia County (Enterprise) ANSP, CAS, OUCO, USNM, (Ormond Beach) PUM; Counties unknown

(Capron) USNM, (Cutier) USNM, (Detroit) USNM, (Hardkinsville) MCZ, (Lake Apopka) AMNH, (Lake Okeechobee) PUM,

(Lake Poinsett) USNM, (Port Sewall) AMNH, (Sand Point) USNM. GEORGIA: Charlton County (Okefenokee Swamp,

90

Erwin

Billy Island) CUNY, USNM; Thomas County (ThomasviUe) AMNH, USNM. ILLINOIS: Alexander County (Olive Branch)

CAS, CMPP, CNHM; Cook County (Chicago) CAS, (Palos Park) UMAH; Jackson County (Carbondale) ISNH, (Fountain

Bluff) ISNH; Johnson County (South of Vienna) RTBe; Richland and Lawrence Counties (Wabash) MCZ; Saint Clair

County (No locality given) CAS, UWMW; Union County (Alto Pass) CNHM; Counties unknown (Pike) ISNH. INDIANA:

Knox County (No locality given) PUM; Perry County (No locality given) PUM; Posey County (Hovey Lake) CEWh, PUM,

(Mount Vemon) CEWh; Spencer County (No locality given) CAS, PUM; Vigo County (No locality given) PUM. KANSAS:

Coffey County (No locality given) ULLK; Douglas County (Baldwin) OSUC, (5.0 miles north of Baldwin City) RFre,

(Lawrence) CAS, PUM, UCD, UMAH, USNM, (3.0 miles northwest of Lawrence) UNLN; Franklin County (No locality

given) KSU; Montgomery County (Independence) CAS; Riley County (Manhattan) KSU; Shawnee County (Topeka) CMPP,

KSU; County unknown (Fort Hays) MCZ. KENTUCKY: Bell County (Pineville) UAFA; Jefferon County (Louisville)

UAFA; Jessamine County (Indian Falls) TCBa; Mercer County (Dix Dam) TCBa; Rockcastle County (Crooked Creek)

TCBa. LOUISIANA: (No locality given)iSNH, UMSP; Cameron Parish USNM, (Grand Chenier) CNC; Iberia Parish (Avery

Island) ANSP; Madison Parish (Tallulah) TAMU, UMAH; Natchitoches Parish (Natchitoches) UMAH; Orleans Parish (New

Orleans) CAS, USNM, ZMLS; Ouachita Parish (Calhoun) UAFA; Plaquemines Parish (Naim) MCZ; Vermilion Parish

(Gueydan) USNM; Vemon Parish (Rosepine) UAFA. MARYLAND: Dorchester County (near Lloyds) USNM; Harford

County (Edgewood) CUNY. MICHIGAN: Allegan County (Allegan) CAS. MINNESOTA: Olmsted County (No locality

given) UMSP. MISSISSIPPI: Attala County (Cole Creek, Natchez Trace) RCGr; Carrol County (Avalon) UMAH; George

County (Lucedale) CUNY; Greene County (Leakesville) CUNY; Hinds County (Jackson) UMAH; Lamar County (Lumber-

ton) CUNY; Lauderdale County (Meridian) UMAH; Leflore County (Greenwood) UMAH; Perry County (New Augusta)

CUNY; Sharkey County (Rolling Fork) USNM; Tunica County (Dundee) UMAH; Washington County (Leroy Percy State

Park) RCGr; County unknown (Moon) AMNH. MISSOURI: Buchanan County (Saint Joseph) USNM; Marion County

(Hannibal) CAS; Saint Charles County (Saint Charles) MCZ; Vemon County (Nevada area) TLEr; County unknown (Big

Oak State Park) RTBe. NEBRASKA: Dakota County (South Sioux City) UNLN; Nemaha County (Peru) CNHM; Sarpy

County (Bellevue, Childs’ Point) CAS, UNLN. NEW JERSEY: Bergen County (Ramsey) USNM; Cape May County (Five

Mile Beach) USNM; Gloucester County (Westville) MCZ; Ocean County (Lakehurst) AMNH; County unknown (Split Rock

Lake) USNM. NEW YORK: New York County (New York City) MCZ; Westchester County (Peekskill) CAS, MCZ. NORTH

CAROLINA: Catawba County (Newton) UCR; Dame County (Cape Hatteras) USNM; Duplin County (Wallace) UCR;

Haywood County (Crestmont) UMAH; (Mount Sterling) CUNY; Robeson County (Boardman) USNM; Wake County

(Raleigh) UCR, UNCR; County unknown (Beauford) MCZ. OHIO: Clinton County (No locality given) OUCO; Hamilton

County (Cincinnati) UMAH; Scioto County (No locality given) OUCO. OKLAHOMA: Alfalfa County (No locality given)

OSUS; Choctaw County (No locality given) OSUS, UONO; Cleveland County OUCO, UONO, (Norman) CAS; Dewey

County (Seiling) OSUS; Le Flore County (Poteau) OSUS; McCurtain County RCGr, (Eagletown) OSUS, (Idabel) OSUS;

Mazes County (No locality given) UONO; Nowata County (13.0 miles west of Vinita) RFre, UASM; Oklahoma County

(Oklahoma) CEWh; Tillman County (No locality given) CAS; Tulsa County (Collinsville) CAS. PENNSYLVANIA: Alle-

gheny County (Pittsburgh) CMPP. SOUTH CAROLINA: Florence County (Scranton) UMAH; Jasper County (Savannah

River Refuge) UASM; Oconee County (Clemson) UASM; Sumter County (Poinsett State Park) VMKi. TENNESSEE: Knox

County (Knoxville) MCZ; Morgan County (Sunbright) CMPP; Obion County (Reelfoot Lake) UMAH; Overton County

(Cleeks Mill) TCBa; Putnam County (Cookeville) TCBa; Sevier County (Great Smoky Mountains National Park) CMPP;

Smith County (Peyton Creek) TCBa. TEXAS: Brazos County (bottoms) ISNH, (College Station) MCZ, TAMU; Cameron

County (Brownsville) OUCO, (Brownsville, Esperanza Ranch) USNM; Colorado County (Columbus) MCZ, UMSP; Dallas

County CAS, ISNH, (Dallas) CMPP, MCZ; Dimmit County (Corrizo Springs) ISUA; El Paso County (El Paso) CMPP;

Grayson County (Juniper Point, Lake Texoma, 12.0 miles north of Whitesboro) RCGr; Harris County (Houston) UCD,

(Seabrook) CAS; Hunt County (Commerce) OUCO; Jeff Davis County (Davis Mountains) OUCO; Kendall County (Com-

fort) CMPP; Liberty County (Devers) UMAH; Runnels County (Ballinger) USNM; Travis County (S. F. Austin State Park)

CNHM; Victoria County (Victoria) USNM; County unknown (Fuller) USNM. VIRGINIA: Alexandria County (No locality

given) USNM; Fairfax County (Mount Vemon) USNM; Nansemond County (Cypress Chapel) ISUA; Norfork County

(Dismal Swamp) AMNH, CAS, USNM; Spotsylvania County (Fredericksburg) CAS. WEST VIRGINIA: Berkeley County

(Leetown) RTBe; Greenbrier County (White Sulphur Springs) MCZ.

Brachinus viridipennis Dejean

(Figs. 179, 183, 184, 185, 193, 199)

Brachinus viridipennis Dejean, 1831: 426. Lectotype, here selected, a female, MHNP, lab-

elled “v. viridipennis m. in Amer. bor.”, “LeConte” on green paper, and “Ex Museo Chau-

doir” on white paper. Type locality. — “l’Amerique septentrionale” as originally given

by Dejean, but herewith restricted to Mobile, Alabama.

Brachinus viridis LeConte, 1844: 49. Lectotype, here selected, a male, MCZ red type label

number 5840. Further labelled with a gold disc and “85.” Type locality. - Georgia, as

originally given by LeConte. LeConte 1862: 524.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. - Large-sized beetles, 8.9 to 15.0 mm.

Color. Antennal article 4, sides of mesosternum and metasternum, mesepisterna, mete-

pisterna, and abdominal sterna and terga infuscated. Dorsal surface and epipleura of elytra

greenish to bluish.

Microsculpture. As described for genus.

Macrosculpture. As in alternans.

Bombardier Beetles

91

Head. As in alternans.

Prothorax. As in alternans, except anterior tibia with anterior edge punctate, with punc-

tures occasionally forming elongate grooves, but not strigae.

Pterothorax. Elytra elongate, narrow, weakly costate. Humeral angle sloped. Metasternum

short, its length behind middle coxa less than diameter of middle coxa (fig. 26). Wings re-

duced outside stigma.

Abdomen. As described for genus.

Genitalia. Male (figs. 183, 184, 185). Median lobe with plane of shaft rotated slightly

from plane of basal bend. Basal bend short. Median lobe straight, swollen medially, apex

narrow, elongate. Ligule long, narrow, slightly widened at apex. Virga (figs. 183, 184). Fe-

male (fig. 193). Stylus long, narrow, arcuate, narrowly rounded apically.

Variation. — Besides the intrapopulational variation in body size and shape of the pro-

notum, these beetles may have greenish or bluish colored elytra. Those with bluish elytra

are few and occur among the greenish populations.

Flight. — The flight of these beetles has never been recorded, and it is probable that they

cannot fly.

Etymology . — Latin, viridis, green; pennis, wing; referring to the greenish elytra of

these beetles.

Life history. — Members of this species have been collected in all months, except January,

May, and August, but no teneral adults were seen.

Distribution. — (Fig. 199). The range of this species extends from eastern Texas to Flor-

ida. I have seen 81 specimens from the following localities:

UNITED STATES

ALABAMA: Clay County (Ashland) AUAA; Mobile County (Alabama Port) CAS, (Coden) AMNH, (Mobile) ANSP, CAS,

CUNY, OUCO, UASM, UMAH, USNM. ARKANSAS: Hempstead County (Hope) MCZ. FLORIDA: Baker County (Glen

Saint Mary) FDAG; Duval County (Jacksonville) MCZ, USNM. GEORGIA: Chatham County (Savannah) CAS; County

unknown (Fort Stewart) TLEr. SOUTH CAROLINA: Beaufort County (Hardeeville) UMAH. TEXAS: Colorado County

(Rock Island, Skull Creek) UMSP.

The rugipennis subgroup

This subgroup is characterized by the wide ligule of the male genitalia, absence of acces-

sory setae, and narrow prothorax. One species, B. rugipennis Chaudoir, is included.

Brachinus rugipennis Chaudoir

(Figs. 171, 180, 181, 182, 192, 200)

Brachynus rugipennis Chaudoir, 1868: 297. Lectotype, here selected, a female, MHNP,

labelled “Etas Unis Guex” in the box and “Ex Museo Chaudoir” on the specimen Type

locality. — United States, as given originally by Chaudoir’s label, but herewith restricted

to Texas.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Small to medium-sized beetles, 7.1 to 9.4 mm.

Color. Antennal articles 3 and 4, abdominal terga, and sides of abdominal sterna infus-

cated, the latter two usually very dark. Remainder of abdominal sterna and usually some of

metepisterna lightly infuscated, otherwise ferrugineous. Dorsal surface and epipleura of ely-

tra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and surface of pronotum punctate, punctures shallowly

impressed.

Head. Frontal furrows shallowly impressed. Antennal scape narrow, almost cylindrical.

Ligula as in alternans. Mentum and submentum without accessory setae.

Prothorax. Pronotum (fig. 171) slightly convex, flattened along center line, sides narrowly

92

Erwin

reflexed. Surface with punctures moderately impressed. Proepipleura and proepisterna pub-

escent anteriorly and posteriorly, glabrous medially. Anterior tibia with anterior surface

strigose.

Pterothorax. Elytra elongate, narrow, weakly costate. Humeral angle slightly sloped. Cos-

tae rugose, as well as depressions between costae.

Abdomen. As described for genus.

Genitalia. Male (figs. 180, 181, 182). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend moderately long. Median lobe arcuate, broadened to

apex, and with apex broadly rounded. Ligule short, widened apically, truncate. Virga (figs.

180, 181). Female (fig. 192). Stylus short, narrow, parallel-sided, narrowly rounded api-

cally.

Variation. — This species is rather constant throughout its range, even in body size.

Flight. — These beetles have been collected repeatedly at lights throughout the range of

the species.

Etymology. — Latin, ruga, wrinkle or fold; pennis, wing; referring to the rugose costae.

Collecting notes. — C. Armin collected these beetles along margins of lakes and irrigation

ditches, and at the edges of small streams in Colorado.

Life history. — Members of this species have been collected from December to July, and

in October. Teneral adults have been collected in March and December in Florida, and in

October in Massachusetts.

Distribution. — (Fig. 200). The range of this species extends from western Colorado to

Massachusetts, and south to Florida. There appears to be discontinuity between the New

England populations and the Floridian one, and between these populations and those west

of the Mississippi. I have seen 327 specimens from the following localities:

UNITED STATES

ARKANSAS: Garland County (Hot Springs National Park) SJSC; Hempstead County (Hope) CUNY, MCZ, UMAH; Pike

County (Delight) CMPP; Washington County (No locality given) UAFA; White County (Searcy) UWSW. COLORADO:

Boulder County (El Dorado Springs) CArm, (Teller Lake) CArm; Montrose County (Montrose) MCZ. FLORIDA: Alachua

County (Gainesville) UMAH; Brevard County (Eau Gallie) MCZ, (Indian River) USNM; Charlotte County (Charlotte Har-

bor) AMNH, (Punta Gorda) AMNH, UMAH; Collier County (Collier Seminole State Park) TLEr, (Naples) CUNY; Dade

County (Royal Palm State Park) PUM; Hendry County (La Belle) OUCO; Highlands County (Archbold Biology Station)

CUNY, PSUU, (Childs) RCGr, (Lake Placid) AMNH; Hillsborough County (Tampa) USNM; Lee County (Fort Myers)

CNC; Lake County USNM, (Fruitland Park) UMAH; Manatee County (Bradenton) CAS, (Oneco) UASM; Marion County

MCZ (Silver Springs) CAS; Orange County (Orlando) MCZ, (Pinecastle) FDAG, (Winter Park) MCZ; Osceola County

(Kissimmee) AMNH; Pinellas County (Dunedin) CAS, PUM, TAMU, (Tarpon Springs) CNC; Sarasota County (Englewood)

AMNH, (Sarasota) PUM; Taylor County (Steinhatchee) USNM; Volusia County (Enterprise) CAS, USNM; Counties un-

known (Port Sewall) AMNH, (Suwannee River) CAS. GEORGIA: Lowndes County (Valdosta) UMAH. KANSAS: Staf-

ford County (Salt Flats Area) UASM. MASSACHUSETTS: Hampden County (Chicopee) KSU, MCZ, (Wilbraham) MCZ;

Plymouth County (Plymouth) AUAA. OKLAHOMA: Marshall County (Lake Texoma, Willis) RCGr, (Madill) RCGr.

NEBRASKA: Cherry County (Hackberry Lake) UNLN. NEW JERSEY: Cape May County (Seven Mile Beach) OUCO;

Mercer County (Trenton) CAS; Warren County (Phillipsburg) CAS; County unknown (Anglesea) CAS. NEW MEXICO:

Bernalillo County (Albuquerque) ANSP, CAS, MCZ, USNM; Sandoval County (Los Alamos) CNC; Taos County (Rio

Grande River, near Taos) CAS. PENNSYLVANIA: Montgomery County (Areola) ANSP; Montour County (Danville) CAS;

Northhampton County (Easton) CAS, (Watergap) AMNH, USNM. TENNESSEE: Lake County (Parnell Point) RTBe.

TEXAS: Anderson County (Elkhart) TAMU; Blanco County (2.0 miles south of Round Mountain) UASM; Brazos County

(College Station) MCZ, TAMU; Dallas County (Dallas) MCZ, UASM; Deaf Smith County (Hereford) TAMU; Montgomery

County (Willis) USNM; Nueces County (Corpus Christi) MCZ; County unknown (Bathage) CAS. VIRGINIA: (No locality

given) UMSP. WYOMING: (western Wyoming) USNM.

The hirsutus group

The members of this group are characterized by the form of the virga, compressed median

lobe with a ventral sulcus, lack of lateral pronotal setae, and strongly costate elytra. Two

species, B. hirsutus Bates and B. pallidus Erwin, are included.

Bombardier Beetles

93

Brachinus hirsutus Bates

(Figs. 205, 212, 213, 214, 220, 224)

Brachinus hirsutus Bates, 1884: 295. Lectotype, here selected, a male, BMNH, labelled

“Pinos Altos, Chihuahua, Mexico, Buchan-Hepburn,” “B. C. A. Col. I. 1. Brachinus hir-

sutus Bates,” “Type, H. T.” and “Brachinus hirsutus Bates” (handwritten). Type local-

ity. — Pinos Altos, Chihuahua, as originally given by Bates.

Brachinus puncticollis LeConte, 1858: 28. NOMEN NUDUM. Erwin, 1965: 13.

Notes. — In 1965 I wrongly placed B. puncticollis as a synonym of B. tschernikhi Manner-

heim. After seeing LeConte’s specimen, I now place the name here.

Diagnostic combination. — The densely pubescent cordiform pronotum, together with

the strongly costate elytra, lack of lateral pronotal setae, and pale venter, separates members

of this species from any others in the American Southwest and Mexico.

Description. — Medium-sized beetles, 7.6 to 10.3 mm.

Color. Ferrugineous, sides of abdomen usually slightly infuscated. Dorsal surface and epi-

pleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows, head behind eyes, and surface of pronotum densely

punctate and rugose. Punctures moderately impressed.

Head. Frontal furrows moderately impressed. Antennal scape robust, widest at middle.

Ligula with sclerotized center area ellipsoid-convex with two lateral rows of three setae per

row. Mentum and submentum with accessory setae.

Prothorax. Pronotum (fig. 205) slightly convex, flattened along center line, sides moder-

ately reflexed. Lateral setae absent. Proepipleura and proepisterna totally pubescent. Anter-

ior tibia with anterior surface strigose.

Pterothorax. Elytra moderately long, narrow, strongly costate. Humeral angle square.

Costae smooth, glabrous, depressions pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 212, 213, 214). Median lobe with plane of shaft rotated slightly

from plane of basal bend. Basal bend long. Median lobe nearly straight, swollen just before

compressed shaft. Apex of shaft narrow and acute, ventral surface sulcate. Ligule short,

narrow, truncate. Virga (figs. 212, 213). Female (fig. 224). Stylus short, parallel-sided,

rounded apically.

Variation. — The shape of the pronotum is much more constant in the members of this

species than in other North and Middle American species. Intrapopulational variation occurs

in the total size and in the color of the elytra (blue to bluish green).

Flight. - The flight of these beetles has been recorded by G. R. Noonan in Upper Sabino

Canyon, Arizona.

Etymology . — Latin, hirsutus, hair; referring to the dense pubescence of the pronotum

of these beetles.

Collecting notes. — G. E. Ball and I collected these beetles in coarse gravel at the edges of

an intermittent stream near El Banco, Durango, Mexico.

Life history. — Members of this species have been collected in all months of the year ex-

cept February and November. Many teneral adults have been collected in June and July in

Arizona, and in January in Jalisco, Mexico. The life cycle is probably much the same as in

B. pallidus (Erwin, 1967).

Distribution . — (Fig. 220). The range of this species extends from southern Utah to the

Mexican High Plateau. I have seen 456 specimens from the following localities:

MEXICO

DURANGO: (12.2 miles south of El Banco) UASM; (Nombre de Dios) AMNH; (Rio Chico, 15.7 miles west of Durango)

UASM; (Rio Florido, near Las Nieves) UASM; (Rio Nazas, near Rodeo) UASM. HIDALGO: (Rio Tula, near Tasquillo)

UASM. JALISCO (0.4 miles west of Coculo) UASM. SAN LUIS POTOSI: (Puente La Parada, 7.5 miles northwest of Mex-

Erwin

Figs. 202-205. Pronotum, right half, dorsal aspect. 202. Brachinus cibolensis new species, near Paradise, Arizona.

203. Brachinus cinctipennis Chevrolat, 13.0 miles southeast of Lagos de Moreno, Jalisco, Mexico. 204. Brachinus

pallidus Erwin, Kings Canyon, California. 205. Brachinus hirsutus Bates, Sonoyta, Sonora, Mexico. Figs. 206-217.

Male genitalia. 206. Brachinus cinctipennis Chevrolat, 23.0 miles, southeast of Lagos de Moreno, Jalisco, Mexico,

ventral aspect. 207 & 208. Lateral & dorsal aspects of same. 209. Brachinus cibolensis new species, Douglas, Arizona,

ventral aspect. 210 & 211. Lateral & dorsal aspects of same. 212. Brachinus hirsutus Bates, Sonoyta, Sonora, Mexico,

ventral aspect. 213 & 2 1 4. Lateral & dorsal aspects of same. 215. Brachinus pallidus Erwin, Kings Canyon, California,

ventral aspect. 216 & 217. Lateral & dorsal aspects of same. Accompanying scale lines equal 1.0 mm.

Bombardier Beetles

95

quitic) UASM; (2.7 miles west of Santa Catarina) UASM. SONORA: (10.0 miles south of Alamos) UCD; (Sonoyta)

AMNH. ZACATECAS: (Rio Juchipila, 0.9 miles north of Jalpa) UASM; (1.3 miles southeast of Sain Alto) UASM.

UNITED STATES

ARIZONA: Cochise County (Chiricahua Mountains) USNM, (14.0 miles northeast of Douglas) UCR, (Bear Creek, Montezu-

ma Pass, Huachuca Mountains) CNC, (Portal) RCGr, (San Pedro River, east of Sierra Vista) UATA, (San Pedro River, 10.0

miles east of Sierra Vista) OSUC, (Tombstone) SDNHM; Coconino County (Bill Williams Fork) MCZ, (Grand Canyon, mile

llb.5) UaTA; Gila County (East Verde River, 6.0 miles north of Payson) LACM, (Globe) KSU, (Payson) UATA, (Sierra

Ancha Mountains) UMAH; Graham County (Aravaipa Creek, between Klondyke and Aravaipa) DRWh, (San Carlos Reser-

voir) UATA; Maricopa County (Phoenix) CNHM, MCZ, OUCO, UATA, (Wickenburg) RCGr; Mohave County (Beaver Dam)

LACM, (16.0 miles north of Wikieup) LACM; Pima County (Cienega Creek, Pantano) CUNY, (Organ Pipe Cactus National

Monument, Quito Baquito) GRNo, (Redington) UATA, (Sabino Canyon) AMNH, (Upper Sabino Canyon) GRNo, (Saint

Xavier Mountains) CAS, (Santa Catalina Mountains) CAS, USNM, (Santa Catalina Mountains, Bear Canyon) CAS, UATA,

(Tucson) AMNH, CAS, CUNY, MCZ, UMAH, USNM; Pinal County (Aravaipa) CUNY, (Sycamore Camp, 9.0 miles north-

west of Payson) CAS, (Near Sombrero Butte) USNM; Santa Cruz County (Bear Canyon Bridge, Lochiel-Bisbee Road) CAS,

(Canelo Hills) UATA, (Nogales) CAS, USNM, (Patagonia) CNHM, CUNY, UATA, UCD, UCR, (Pena Blanca Lake, 16.0

miles northwest of Nogales) OSUC, (Santa Cruz River, near Nogales) CAS, (Yanks Spring, 4.0 miles southeast of Ruby)

AMNH; Yavapai County AMNH (Bumble Bee) CAS, (Camp Verde) CAS, PSUU, (29.0 miles northwest of Congress) UATA,

(Mayer) GRNo, (Prescott) CAS; Counties unknown (Hot Springs) CAS, (Palmerlee) CMPP, (Superstition Mountains)

UATA. NEW MEXICO: Catron County (Glenwood) DRWh, (San Francisco Creek, 26.1 miles north of Glenwood) UASM;

Grant County (near Gila) UASM. TEXAS: Jeff Davis County (Davis Mountains) CAS, UCD, USNM, (Fort Davis) AMNH,

CNC, (6-10.0 miles west of Fort Davis) UASM, (Limpia Canyon) CNHM, DRWh, TLEr, SJSC, UASM; Reeves County

(Balmorhea Lake) UASM; Presidio County (Presidio) TAMU. UTAH: Washington County (3.0 miles south of Gunlock)

GRNo, (Saint George) USNM, (Santa Clara) UWSW.

Brachinus pallidus Erwin

(Figs. 204, 215, 216, 217, 218, 222)

Brachinus pallidus Erwin, 1965: 8. The holotype male and allotype female are in CAS.

Type locality. — Mad River, 5.0 miles east of Mad River Post Office, Trinity County,

California.

Diagnostic combination — The diagnostic characteristics are given in the key.

Description. — Medium-sized beetles, 7.5 to 9.3 mm.

Color. As in hirsutus.

Microsculpture. As described for genus.

Macrosculpture. Head as in hirsutus , pronotum not as densely punctate.

Head. As in hirsutus , but ligula without constant setae.

Prothorax. As in hirsutus , but less densely setiferous. Pronotum (fig. 204).

Pterothorax. As in hirsutus , but only elytral depressions 6, 7, and 8 pubescent, except in

apical third.

Abdomen. As described for genus.

Genitalia. Male (figs. 215, 216, 217). As in hirsutus , except more compressed and without

ventral sulcus. Virga (figs. 215, 216). Female (fig. 222). Stylus straighter than in hirsutus ,

more acute apically.

Variation. — The members of this species vary within local populations in body size and

the shape of the pronotum. Otherwise, they are quite constant throughout the range of the

species.

Flight. — I have watched these beetles fly in captivity.

Etymology . — Latin, pallidus , pale; referring to the pale venter of these beetles.

Collecting notes. — These beetles occur along intermittent streams and permanent rivers

in coarse gravel.

Life history. — See p. 166 and Erwin 1967.

Distribution . — (Fig. 218). The range of this species extends from Los Angeles County in

southern California north to eastern Washington. I have seen 835 specimens from the follow-

ing localities:

UNITED STATES

CALIFORNIA: Alameda County (Alameda Creek) CAS, (Arroyo Mocho) TLEr, (Berkeley) CUNY, (Livermore) CAS,

(Los Moches Canyon, Livermore hills) CEWh, (Niles Canyon) ANSP, CAS, SDNHM, UASM, (Sunol) CAS; Amador County

Erwin

222

223

224

225

Figs. 218-221. Geographical distribution maps. 218. Brachinus pallidus Erwin. 219. Brachinus cibolensis new species.

220. Brachinus hirsutus Bates. 22 1 . Brachinus cinctipennis Chevrolat. Figs. 222-225. Right stylus of female ovipositor,

ventral aspect. 222. Brachinus pallidus Erwin, Kings Canyon, California. 223. Brachinus cibolensis new species, near

Paradise, Arizona. 224. Brachinus hirsutus Bates, Rio Chico, Durango, Mexico. 225. Brachinus cinctipennis Chevrolat,

13.0 miles southeast of Lagos de Moreno, Jalisco, Mexico. Accompanying scale line equals 1.0 mm.

Bombardier Beetles

97

(5.0 miles west of Sutter Creek on Horse Creek) TLEr; Butte County (Oroville) CAS; Calaveras County (Mokelumne Hill)

CAS, USNM; Contra Costa County (Marsh Creek) TLEr; Fresno County (Camp Greeley) CAS, (Kings Canyon) CAS, (La

Fevre Creek) ANSP, UASM; Glenn County (Elk Creek) CAS; Humboldt County AMNH, (6.0 miles east of Bridgeville)

GRNo, (Eel River, Fernbridge) UASM, (Fort Seward) CAS, UIMI, (Garberville) CAS, UIMI, (5.0 miles south of Garberville)

TLEr, (Schively) UCD; Lake County (Lower Lake) CAS, (Middletown) CAS, JSch, UIMI, (North Fork Cache Creek, High-

way 20) UCD, (Putah Creek) CAS; Los Angeles County USNM, (Los Angeles) CNHM; Madera County (Coursegold) CAS,

UIMI; Marin County USNM, (Fairfax) CUNY, (Point Reyes) CAS, UCD; Mariposa County (Jolon) CAS, (3.0 miles south-

east of Jolon) CAS; Mendocino County CNHM, (Eel River) CAS, (Long Valley Creek, 6.7 miles south of Laytonville) CAS,

UWSW; Merced County (Merced Falls) UCD; Napa County CMPP, (Monticello) UCD, (Pope Valley) CAS, (Rutherford)

TLEr, (Saint Helena) AMNH, ROM, (Saint Helena Creek) CAS, (10.0 miles east of Shell Peak) CAS; Placer County (Aub-

urn) UCD; Sacramento County (Folsom) JSch; San Joaquin County (No locality given) SJSC; San Luis Obispo County

(Atascadero) CAS, (Nacimiento River, Camp Roberts) PSUU, (Paso Robles Creek) CAS; Santa Clara County CNHM, (Ar-

royo Bayo) SJSC, (Gilroy Hot Springs) TLEr, (Hecker Pass) CAS, (Isabel Creek) TLEr, (Los Gatos) CAS, (Morgan Hill)

SJSC, (Mount Hamilton) CAS, JSch, (San Jose) CAS, (Santa Clara) CAS, (Uvas Creek) TLEr; Santa Cruz County (Santa

Cruz) USNM; Shasta County CNHM, (Anderson) CAS, (Redding) CAS, WBa; Siskiyou County (Klamath River) USNM,

(10.0 miles west of Montague) JSch, (south of Shasta River) CAS, (Yreka) AMNH; Sonoma County UNHM, (Cloverdale)

CUNY, (Del Puerto Creek) TLEr, (Dry Creek, 9.0 miles northwest of Healdsburg) SJSC, (Guerneville) CAS, (2.0 miles

east of Healdsburg) CAS, (Rio Nido) CAS, (Santa Rosa) CUNY, MCZ, OUCO; Tehama County (Red Bluff) CAS; Trinity

County CMPP, (Mad River) TLEr, (0.7 miles northwest of Ruth Dam) GRNo, (Weaverville) UWSW; Yolo County (Davis)

CBak, UCD; Counties unknown (Aliso) UWSW, (Latrobe) CAS, (San Antonio Mission) CUNY, (Sylvania) CAS. OREGON:

Douglas County (North Umqua River, near Winchester) JSch, (7.0 miles northwest of Roseburg) JSch, (Winchester) JSch;

Jackson County (Eagle Point) CAS, (Medford) CAS, UCD, (8-14.0 miles south of Ruch) JSch, (10.0 miles south of Ruch)

JSch, (Talent) UCD, (Trail) JSch, LRus; Josephine County (Applegate River) OSUC, (Illinois River) JSch, (Selma) JSch;

Umatilla County (Umatilla) MCZ. WASHINGTON: Spokane County (Spokane Falls) MCZ; Walla Walla County (Walla

Walla) OSUC.

The fumans group

This group is characterized by the virga, the sides of which are curled over ventrally from

base to apex, forming a central trough. The diversity of this group warrants its division into

twelve subgroups.

The cinctipennis subgroup

The members of this subgroup are characterized by an elytral ferrugineous sutural stripe

and a long narrow median lobe. Two species, B. cinctipennis Chevrolat and B. cibolensis new

species, are included.

Brachinus cinctipennis Chevrolat

(Figs. 203, 206, 207, 208, 221, 225)

Brachinus cinctipennis Chevrolat, 1835: 163. Lectotype, here selected, a male, HMO, Type

number Col. 113 1/3, further labelled “Brachinus cinctipennis Chev. Col de M. Z cent no

163 Mexico plaine de Mexico Aout sous des pierres Salle 59.” Type locality. — The high-

plain of Mexico, as originally given by Cheviolat, but herewith restricted to the State of

Mexico, Mexico.

Diagnostic combination. — The ferrugineous sutural stripe on the elytra, together with the

pale palpi, and extensively pale legs, separate these beetles from all others of the genus.

Description. — Medium-sized beetles, 6.6 to 9.4 mm.

Color. Antennal articles 3-11, apex of femur, mesepisterna, metepisterna, metasternum

at sides, and abdominal sterna and terga infuscated, otherwise ferrugineous. Dorsal surface

of elytra greenish-blue with ferrugineous sutural stripe, epipleura testaceous.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and surface of pronotum slightly rugose and punctate,

punctures barely impressed.

Head. Frontal furrows moderately impressed. Antennal scape cylindrical. Ligula with

sclerotized center area ellipsoid-convex with two lateral rows of three setae per row.

Prothorax. Pronotum (fig. 203) convex, sides barely reflexed. Proepipleura glabrous. Pro-

episterna with a few scattered setae both anteriorly and posteriorly. Anterior tibia with an-

terior surface strigose.

98

Erwin

Pterothorax. Elytra elongate, narrow, barely costate. Humeral angle square. Costae and

depressions pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 206, 207, 208). Median lobe with plane of shaft rotated 45° from

plane of basal bend. Basal bend long. Apex of shaft narrow, rounded. Ligule short, narrow,

narrowly rounded apically. Virga (figs. 206, 207). Female (fig. 225). Stylus short, broad,

narrowly rounded apically.

Variation. — Intrapopulational variation occurs in the following characteristics: presence

or absence of accessory setae on the submentum; number of setae on the ligula; extent of

infuscation on the tibia; width of the sutural stripe; and the color of the elytra.

Flight. - The flight of these beetles has not been recorded.

Etymology. - Latin, cinctus, belt , pennis, wing; referring to the broad, sutural stripeon

the elytra of these beetles.

Collecting notes. - G. E. Ball and I collected these beetles from beneath stones at the

edge of an artificial pond in Jalisco, Mexico.

Life history. — Members of this species have been collected in April, June to September,

November, and December. Teneral adults were collected in April in Puebla, Mexico.

Distribution. — (Fig. 221). The range of this species extends from San Luis Potosi to

Puebla. I have seen 105 specimens from the following localities:

MEXICO

AGUASCALIENTES: (Aguascalientes) AMNH;(E1 Rotono, 10.0 miles east of Aguascalientes) AMNH. DISTRITO FEDER-

AL: CPBo, (Mexico City) AMNH, BMNH, CUNY, TMBH. JALISCO: (Encarnacion de Diaz) UASM; (5.0 miles west of

Lagos) UATA; (13.0 miles southeast of Lagos de Moreno) USNM; (12.0 miles west of Ojuelos) CAS. MEXICO: (Lago

Zumpango, near San Juan Zitlaltepec) UASM. PUEBLA: (Lago Totolcingo, near Tlaxcala line) UASM. SAN LUIS POTOSI:

(San Luis Potosi) AMNH. STATE UNKNOWN: (Presa de Angulo) JHen.

Brachinus cibolensis new species

(Figs. 202, 209, 210, 211,219, 223)

Type locality. — Five miles west of Portal, Southwest Research Station, 5,400 feet, Cochise

County, Arizona.

Type specimens. - The holotype male and allotype female are in the entomological museum

at AMNH. The holotype was collected by E. Ordway, and the allotype was collected by

M. Statham at the type locality on July 27, 1956 and May 8, 1958, respectively. Twenty-

five paratypes collected in various localities on various dates are in AMNH, CAS, MCZ,

TLEr, and UASM.

Diagnostic combination. — The ferrugineous sutural stripe on the elytra, together with

the entirely black legs and palpi, separate these beetles from all others in Mexico and United

States.

Description. — Medium-sized beetles, 7.5 to 9.8 mm.

Color. Palpi, antennal articles 2-11, metepisterna, metasternum at sides, legs, and abdom-

inal terga and sterna infuscated to black. Dorsal surface of elytra blue with ferrugineous

sutural stripe, epipleura testaceous.

Microsculpture. As described for genus.

Macrosculpture. As in cinctipennis .

Head. As in cinctipennis.

Prothorax. As in cinctipennis. Pronotum (fig. 202).

Pterothorax. As in cinctipennis.

Abdomen. As described for genus.

Genitalia. Male (figs. 209, 210, 211). Median lobe with plane of shaft rotated 45° from

plane of basal bend. Basal bend long. Shaft long and narrow, apex rounded, slightly notched.

Ligule short, tapering to narrowly rounded apex. Virga (figs. 209, 210). Female (fig. 223).

Bombardier Beetles

99

Stylus short, broad, tapering to narrowly rounded apex.

Variation. — Intrapopulational variation occurs in the shape of the pronotum, width of

the sutural stripe on the elytra, and in the color of the elytra and antennal articles 2-1 1.

Flight. - These beetles have been collected at lights at Douglas, Arizona, Lordsburg, New

Mexico, and Durango City, Mexico.

Etymology . — Cibola, from the legendary Seven Cities of Cibola that Coronado searched

for in the American Southwest; Latin, ensis, denoting place, locality, or country.

Collecting notes. — G. E. Ball collected these beetles from under stones at the margin of a

dirt water tank in the Chiricahua Mountains of Arizona.

Life history. — Members of this species have been collected from June to September, but

no teneral adults were seen.

Distribution. — (Fig. 219). The range of this species extends from northern Arizona to

Durango City, Mexico. I have seen 78 specimens from the following localities:

MEXICO

DURANGO: (Durango) AMNH, UASM; (15.0 miles west of Durango) CNC.

UNITED STATES

ARIZONA: Cochise County (Benson) UATA, (Douglas) CUNY, UASM, UCR, (near Paradise) UASM, (Portal) GRNo, (San

Bernardino Ranch) ZMLS, (South West Research Station, 5.0 miles west of Portal) AMNH, UCD, (Tombstone) SDNHM,

(Willcox) AMNH, UATA; Coconino County (Bill Williams Fork) AMNH; Santa Cruz County (Nogales) CAS, CNHM, UCR.

NEW MEXICO: Bernalillo County (Albuquerque) CMPP, USNM; Hidalgo County (Animas) AMNH, (Lordsburg) CNC,

UCR, (Rodeo) CUNY, UCD; Socorro County (Socorro) CAS.

The quadripennis subgroup

The members of this subgroup are characterized by the presence of a ridge on the ventral

surface of the male genitalia. Five species, B. quadripennis Dejean, B. mexicanus Dejean,

B. neglectus LeConte, B. javalinopsis new species, and B. kavanaughi Erwin, are included,

eluded.

Brachinus quadripennis Dejean

(Figs. 230, 240, 241, 242, 247, 251)

Brachinus quadripennis Dejean, 1825: 316. Lectotype, here selected, a female, MHNP,

labelled “quadripennis m. in Amer. bor.” and “Lherminier” on green paper, and “Ex

Museo Chaudoir” on white paper. Type locality. — North America, as originally given by

Dejean, but herewith restricted to Florida.

Brachinus stygicornis Say, 1834: 415. Neotype designated by me, a male, in MCZ. Type

locality. - South Bend, Nebraska. NEW SYNONYMY.

Brachinus tschernikhi Mannerheim, 1843: 184. Types presumed lost (see Erwin, 1965: 13).

Type locality. — California, as originally given by Mannerheim. NEW SYNONYMY.

Notes. — The neotype designated by Lindroth for B. stygicornis was selected by me

from the Nebraska University material I had on loan, and sent to Lindroth. He subsequently

deposited all Say’s neotypes in MCZ. The specimen was labelled “South Bend, Neb.” “5/8/

09” and “R. W. Dawson Collection.” This locality is the nearest to Say’s original area from

which we had specimens. The original area was “crevices of rocks ... Engineer Cantonment,

near Council Bluff.”

The name B. sejungenius Chaudoir is found on some of Chaudoir’s specimens in MHNP.

This name was never published, and is therefore not really in existence, but I add this note

to prevent confusion in the future.

Diagnostic combination. — The diagnostic characters are given in the key, but in most

cases the infuscated tarsi and tibiae will separate these beetles from all others of the genus in

the United States, except individuals of B. phaeocerus and B. azureipennis . The members of

phaeocerus are smaller and have bright blue elytra, while the members of azureipennis are

Figs. 226, 230. Pronotum, right half, dorsal aspect. 226. Brachinus kavanaughi Erwin, Superior, Colorado. 230.

Brachinus quadripennis Dejean, Archbold Research Station, Florida. Figs. 227-229, 231-242. Male genitalia. 227.

Brachinus javalinopsis new species, 32.0 miles east of Douglas, Arizona, ventral aspect. 228. Lateral aspect of same.

229. Dorsal aspect of same. 23 1 . Brachinus kavanaughi Erwin, 6.9 miles north of Golden, Colorado, ventral aspect.

232. Lateral aspect of same. 233. Dorsal aspect of same. 234. Brachinus mexicanus Dejean, 0.9 miles northeast

of Cedar Springs, California, ventral aspect. 235. Lateral aspect of same. 236. Dorsal aspect of same. 237. Brachinus

neglectus LeConte, Guntown, Florida, ventral aspect. 238. Lateral aspect of same. 239. Dorsal aspect of same. 240.

Brachinus quadripennis Dejean, Grand Coulee, Washington, ventral aspect. 241. Lateral aspect of same. 242. Dorsal

aspect of same. Accompanying scale lines equal 1.0 mm.

Bombardier Beetles

101

larger, have black palpi and antennae, and have metallic blue luster on the infuscated ab-

domen.

Description. — Medium-sized beetles, 7.8 to 1 1.0 mm.

Color. Palpi, antennal articles 3 and 4, mesepimera, tibiae, and tarsi usually infuscated.

Abdominal sterna, terga, metepisterna, and metasternum at sides infuscated to black, other-

wise ferrugineous. Dorsal surface and epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows, and sometimes disc of pronotum rugose, sparsely punc-

tate, punctures barely impressed.

Head. Frontal furrows moderately impressed. Antennal scape cylindrical. Ligula with

sclerotized center area ellipsoid-convex with two or three setae apically. Mentum and sub-

mentum without accessory setae.

Prothorax. Pronotum (fig. 230) slightly convex, flattened along center line, sides slightly

reflexed. Proepipleura glabrous. Proepisterna with a few setae anteriorly and posteriorly,

glabrous medially. Anterior tibia with anterior margin strigose.

Pterothorax. Elytra elongate, narrow, moderately costate. Humeral angle various, but

usually prominent or at least square. Costae smooth on disc, depressions pubescent. Wings

entire.

Abdomen. As described for genus.

Genitalia. Male (figs. 240, 241, 242). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend short. Apex of shaft narrowed, ridged ventrally. Ligule

short, broad, narrowed apically. Virga (figs. 240, 241) elongated. Female (fig. 247). Stylus

broad, parallel-sided, rounded apically.

Variation. — The color of the tibiae, tarsi, and palpi vary from almost black to ferrugin-

eous. The pale form occurs infrequently in the northwestern populations, but is common in

the midwestern, eastern, and Floridian populations. Variation also occurs in the shape of the

pronotum and humeral angle within single population samples.

Flight. — These beetles have been collected repeatedly at lights throughout the range of

the species, especially Florida.

Etymology . — Latin, quadratus, square, pennis, wing; referring to the square shape of

the elytra.

Collecting notes. — L. Russell collected these beetles on the floodplain of the Flathead

River near Perma, Montana. They were beneath short grass, matted with algae, near an eph-

emeral pond formed by river floodwaters.

Life history. — Members of this species have been collected from January to September,

and teneral adults have been collected in March in Florida, and in August in Washington and

Michigan. These beetles probably overwinter as adults.

Distribution. — (Fig. 251). There are six areas from which these beetles have been com-

monly collected. These are the northwestern United States and adjacent Canada; the Great

Salt Lake area; the areas drained by the Mississippi, and its northern tributaries east of

longitude 105°, including the Great Lakes Region, New England, Florida and the eastern

part of the Gulf Coast, and Brownsville, Texas. I have seen 2,405 specimens from the

following localities:

CANADA

ALBERTA: (Medicine Hat) UASM. BRITISH COLUMBIA: (Kamloops) MCZ; (Oliver) CNC; (Osoyoos) CAS, UATA.

MANITOBA: (Aweme) OUNO. ONTARIO: (Saint Williams, southwest of Simcoe) ZMLS. SASKATCHEWAN: (Kenosee)

UNSS.

UNITED STATES

ALABAMA: Mobile County (Magazine Point) CAS. ARIZONA: (No locality given) USNM. ARKANSAS: Bradley County

(8.0 miles south of Warren) JSch; Mississippi County (Osceola) UMAH; Washington County (No locality given) UAFA.

102

Erwin

Figs. 243-245. Pronotum, right half, dorsal aspect. 243. Brachinus neglectus LeConte, Southern Pines, North Carolina.

244. Brachinus mexicanus Dejean, Temecula, California. 245. Brachinus javalinopsis new species, Victoria, Texas.

Figs. 246-250. Right stylus of female ovipositor, ventral aspect. 246. Brachinus javalinopsis new species, 15.0 miles

north of Las Cruces, New Mexico. 147. Brachinus quadripennis Dejean, Smyrna, Washington. 248. Brachinus mexi-

canus Dejean, Temecula, California. 249. Brachinus kavanaughi Erwin, Superior, Colorado. 250. Brachinus neglectus

LeConte, Jacksonville, Florida. Fig. 251. Geographical distribution map of Brachinus quadripennis Dejean. Accom-

panying scale lines equal 1.0 mm.

Bombardier Beetles

103

CALIFORNIA: Fresno County (Friant) CAS; Kings County (Hanford) USNM; Lake County (Cache Creek) UCR, (Clear

Lake) CAS, (Lower Lake) CAS; Los Angeles County USNM, (Pasadena) CMPP; Merced County (Los Banos) CAS, TLEr;

Sacramento County (Sacramento) TLEr, UCD, UIMI; Yolo County (Causeway) UCD, (Clarksburg) UCD, (Davis) CAS,

TLEr, UCD, UIMI. SOUTH DAKOTA: Beadle County (Huron) VMKi; Brookings County (Brookings) VMKi, (White)

VMKi; Lawrence County (Spearfish) VMKi; Union County (Elk Point) VMKi; Yankton County (Yankton) VMKi.

FLORIDA: Alachua County (Gainesville) CNC, FDAG, TLEr, UMAH, USNM, (Newnans Lake, 5.0 miles east of Gaines-

ville) RFre, UASM; Broward County (Fort Lauderdale) UMAH; Charlotte County (Punta Gorda) CNC, PSUU; Collier

County (Everglades) USNM; Columbia County (O’Leno State Park) CUNY; Dade County FDAG, (3.0 miles south of

Florida City) AMNH, (Homestead) CNC, TLEr, (Long Pine Key) MCZ, (Miami) CAS, (Royal Palm State Park) PUM; De

Soto County (Arcadia) CUNY; Duval County (Jacksonville) ISNH ; Glades County (Palmdale) AMNH; Hendry County

(Clewiston) OUCO; Highlands County (Archbold Biology Station) AMNH, CEWh, CMPP, CUNY, PSUU, RCGr, (Brighton)

UASM, (Fish Eating Creek, 4.0 miles west of Venus) RCGr, (Highlands Hammock State Park) TLEr; Hillsborough County

(Tampa) USNM; Lee County (Fort Myers) AMNH, CUNY; Levy County FDAG, (Manatee Springs State Park) RFre,

UASM, (Williston) UWSW; Liberty County (Camp Torreya) UMAH; Manatee County (Bradenton) FDAG, (Oneco) CNC,

CUNY, UASM; Marion County (No locality given) MCZ; Okeechobee County (Lake Okeechobee, 6.0 miles south of

Okeechobee) UASM; Orange County (Winterpark) MCZ; Palm Beach County (Belle Glade) CUNY, (North New River

Canal) UASM; Pinellas County (Clearwater) CNHM, (Dunedin) CAS, CUNY, PUM, (Tarpon Springs) CNC, CNHM; Polk

County (Lakeland) UMAH, (Lake Wales) FDAG, USNM, (Winter Haven) FDAG; Saint Johns County (Ponte Vedra Beach)

AMNH, (Saint Augustine) AMNH; Sarasota County (Myakka River State Park) CUNY, UASM, (Sarasota) PUM; Seminole

County (Sanford) CUNY, FDAG, JSch, PUM, USNM; Volusia County (Enterprise) CAS, USNM, (Ormond) AMNH; Coun-

ties unknown (Belleair) AMNH, (Fort Capron) ISNH, (Lake Harney) USNM, (Sand Point) USNM. GEORGIA: Lowndes

County (No locality given) OUCO; Thomas County (Thomasville) CNHM, USNM. IDAHO: Bonner County (Sagle) UWSW;

Latah County (Moscow) CMPP. ILLINOIS: (No locality given) ISNH. INDIANA: Vigo County (No locality given) PUM,

UWMW; Posey County (Hovey Lake) PUM; County unknown (Mineral Springs) CMPP. IOWA: Boone County (Ledges

State Park) ISUA; Dickinson County (Lake Okoboji) USNM, (Milford) USNM; Emmet County (No locality given) CAS;

Henry County (Mount Pleasant) RTBe; Johnson County (Iowa City) MCZ, USNM; Palo Alto County (Ruthven) ISUA,

(Silver Lake) USNM; Sioux County (Howarden) VMKi; Story County (Ames) ISUA, MSUM, OSUC, USNM, (Soper’s Mill

Dam, 3.0 miles east of Gilbert) ISUA; Woodbury County (Sioux City) UMSP. KANSAS: Atchison County (Atchison)

CMPP; Douglas County CUNY, (Lawrence) UWMW; Pottawatomie County (Onaga) CAS; Riley County (Manhattan) KSU;

Rooks County (No locality given) KSU; Saline County (Salina) CAS, CMPP, MCZ, PUM, ZMLS; Sheridan County (State

Lake, near SlUdley) RFre. LOUISIANA: Orleans Parish (New Orleans) ANSP. MASSACHUSETTS: Hampden County

(Longmeadow) USNM; Hampshire County (Mount Tom) MCZ; Middlesex County (Arlington) MCZ, (Concord) MCZ,

(Newton) MCZ, (Waltham) MCZ; Suffolk County (Dorchester) MCZ; County unknown (Forest Hills) MCZ. MICHIGAN:

Huron County (Point aux Barques) UMAH; Ingham County (East Lansing) UATA; Macomb County (Mount Clemens)

CNHM; Marquette County (Marquette) CAS; Oakland County (Milford) UMAH; County unknown (Aurelius) JSch.

MINNESOTA: Big Stone County UMSP, (Ortonville) ISNH; Clearwater County (Lake Itasca) UMSP; Houston County

(Mississippi Bluff, 1-2.0 miles north of State Line) UMSP; Jackson County (Jackson) USNM; Lincoln County (Lake Ben-

ton) VMKi; Lyon County (No locality given) UMSP; Norman County (No locality given) UMSP; Olmsted County (No

locality given) UMSP; Polk County (Crookston) UMSP; Ramsey County (Saint Paul) UMSP; Saint Louis County (Duluth)

ISNH; Steams County (Koronis Lake, Paynesville) USNM; Washington County (No locality given) UMSP; Wright County

(No locality given) UMSP. MISSOURI: Atchison County (Langdon) AMNH; Callaway County (Readsville) MCZ; Jackson

County (Kansas City) KSU; Saint Louis County (Ranken) JSch; Wayne County (William sville) CNC; Wright County (Gull

Creek, west of Mount Grove) TCBa. MONTANA: Cascade County (Great Falls) CAS; Flathead County (Kalispell) USNM;

Sanders County (Perma) LRus. NEBRASKA: Cass County (South Bend) UNLN; Cuming County (West Point) UNLN;

Dakota County (South Sioux) UNLN; Dodge County (Fremont) UNLN; Douglas County (Childs’ Point, Omaha) CAS,

UNLN; Lancaster County (Lincoln) UNLN, (Malcolm) USNM, (Roca) UNLN; Otoe County (Nebraska City) UNLN;

Sarpy County (Bellevue) UNLN; Saunders County (Ashland) SDSU, UNLN, (Cedar Bluffs) UNLN, USNM; Thomas Coun-

ty (Halsey) UMAH; York County (Bradshaw) UNLN, (York) UNLN. NEVADA: ANSP, USNM, MCZ, ISNH, Humboldt

County (Golconda) CBak; Washoe County (Reno) RTBe. NEW JERSEY: Cape May County (Five Mile Beach) OUCO;

Gloucester County (Woodbury) USNM. NEW YORK: Suffolk County (Montauk) CNHM. NORTH DAKOTA: Benson

County (11.9 miles west of York) UASM; McKenzie County (North Roosevelt National Park) AMNH; Morton County

(Heart River, 5.0 miles west of Mandan) UASM; Ramsey County (Devils Lake) MCZ, USNM, ZMLS. OHIO: Ashtabula

County (Jefferson) USNM; Wood County (No locality given) PUM. OREGON: Baker County UMAH, (Wallowa Mountains)

CAS; Curry County (8.0 miles east of Gold Beach) OSUC; Lake County (Hart Lake) JSch; Wasco County (The Dalles)

MCZ, USNM; Umatilla County (Umatilla) MCZ; Union County (Alicel) RESt, UIMI; County unknown (Stein Mountains)

CAS. PENNSYLVANIA: Allegheny County (Pittsburgh) CMPP. SOUTH DAKOTA: Bon Homme County (Springfield)

VMKi; Brookings County (Brookings) CMPP, SDSU, VMKi, (Volga) CAS, MCZ, USNM, VMKi, (White) VMKi; Brown

County (Stratford) LACM; Brule County (Chamberlain) SDSU; Buffalo County (Fort Thompson) SDSU; Clay County

(Vermillion) SDSU; Haakon County (Philip) SDSU; Ifaghes County (Pierre) VMKi; Jones County (Murdo) SDSU; Kings-

burg County (Erwin) USNM; Turner County (Centerville) VMKi; Union County (Elk Point) SDSU, VMKi; Yankton Coun-

ty (Yankton) VMKi; County unknown (Darwood Lake) VMKi. TENNESSEE: Lake County (Gray’s Landing) RTBe.

TEXAS: Cameron County (Brownsville) CNC, CNHM, CUNY, OUCO, USNM. UTAH: Cache County (Logan) USUL,

(Smith field) USUL, (Wellsville) USUL; Salt Lake County (Salt Lice City) UIMI, USNM, UWSW; Utah County (Provo)

MCZ, UASM, USNM, (Provo Canyon, 1.0 mile south of Springdale) UMSP, (Utah Lake, Provo) UASM, UMAH.

WASHINGTON: Adams County (Lake McElroy) CAS, USNM, UWSW, (Othello) UWSW, (Ritzville) CAS, CMPP, MCZ,

PUM, UMAH, USNM; Franklin County (Kahlotus) UWSW; Grant County (Coulee City) CAS, UWSW, (Crab Creek) UWSW,

(Grand Coulee, Dry Falls) WSUP, UWSW, ZMLS, (Grand Coulee, Meadow Creek) WSUP, (Moses Lake) JSch, UWSW,

(Smyrna) UWSW, (Soap Lake) UWSW, (Steamboat Rock) UWSW, (Stratford) CAS; Kittitas County (Vantage) UWSW;

Lincoln County (Sprague) CAS, CMPP, PUM, UWSW, (Sprague Lake) CAS, (Wilbur) UASM; Okanogen County (5.0 miles

south of Tonasket) CNHM; Spokane County (Cheney) JSch, UWSW, (Medical Lake) UWSW; Yakima County (Toppenish)

104

Erwin

UWSW. WISCONSIN: Dane County (No locality given) UWMW.

Brachinus mexicanus Dejean

(Figs. 234, 235, 236, 244, 248, 252)

Brachinus mexicanus Dejean, 1831: 428. Lectotype, here selected, a male, MHNP, labelled

“mexicanus m. in Mexico” “Hopfner” on green paper, and “Ex Museo Chaudoir” on

white paper. Type locality. — Mexico, as originally given by Dejean, but herewith res-

tricted to Baja California, Mexico.

Brachinus fidelis LeConte, 1862: 524. Lectotype, here selected, a female, MCZ red type

label number 5852 Further labelled with a gold disc and “B. fidelis Kern. LeC.” Type

locality. — Kern County, California, here designated, based on LeConte’s labels. NEW

SYNONYMY.

Brachinus convexus Chaudoir, 1837: 7. Lectotype, here selected, a female, MHNP, labelled

“convexus Mex. Chaud.” and “Ex Museo Chaudoir.” This specimen is pinned beneath a

specimen of B. mexicanus Dejean, and placed in that series. Type locality. — Mexico, as

given on Chaudoir’s label. NEW SYNONYMY.

Brachinus lecontei Motschulsky, 1859: 139. Primary homonym of B. lecontei LeConte,

1844: 49 (see Erwin, 1965: 10).

Diagnostic combination. — The diagnostic characters are given in the key, but these

beetles can be separated from all others in the study area by elytral pubescence restricted to

depressions 6, 7, and 8, dark venter, blue elytra, and lack of accessory setae on the mentum.

Description. — Medium-sized beetles, 7.5 to 9.6 mm.

Color. See under Variation, below.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and disc of pronotum rugose, sparsely punctate, punc-

tures weakly impressed.

Head. Frontal furrows moderately impressed. Antennal scape robust, widest at middle.

Ligula with sclerotized center area ellipsoid-convex with two lateral rows of three setae

each. Mentum and submentum without accessory setae.

Prothorax. As in quadripennis , except proepipleura variable. Pronotum (fig. 244).

Pterothorax. Elytra as in quadripennis , except costae variable and pubescence confined

to depressions 6, 7, 8, scutellar region, and across the apical sixth of the elytra. Wings

fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 234, 235, 236). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend short. Apex of shaft various, ridged ventrally. Ligule

short, broad, narrowed apically. Virga (figs. 234, 235). Female (fig. 248). Stylus broad,

tapering to acute apex.

Variation. — The composition of this species seems to be more complex than most Brach-

inus species in the study area. Two forms are clearly recognizable in most of the range, ex-

cept California. I refer to these forms as the “high-costae morph” and the “low-costae

morph.” In California, these two forms merge in all characteristics, with local populations

having both the extreme and intermediate forms. In Mexico, the high-costae morph is known

only in northern Baja California (Catavina). Other populations are in Phoenix, Arizona,

southern New Mexico, Hope, Arkansas, Chicago, Illinois, and Brookings, South Dakota. In

Phoenix, there appears to be a mixed population, but the specimens seen were collected at

different times. Except for the last three mentioned peripheral localities, the range of the

low-costae morph overlaps that of the high-costae morph.

The high-costae morph generally has pale antennae, pale abdominal center, well devel-

oped costae, and a few setae on the proepipleura. The low-costae morph is quite dark be-

Bombardier Beetles

105

Figs. 252-255. Geographical distribution maps. 252. Brachinus mexicanus Dejean. 253. Brachinus kavanaughi Erwin.

254. Brachinus javalinopsis new species. 255. Brachinus neglectus LeConte.

106

Erwin

neath, the abdominal center is infuscated, the third and fourth antennal articles are at least

clouded, the outer articles are infuscated rather than being dusky, and finally the proepip-

leura are usually glabrous.

Further variation occurs in the male genitalia. In the low-costae morph the shaft is nar-

rowed to apex, and the virga tapers evenly to the apex. The high-costae morph has the apex

of the shaft almost truncate, and the virga is pinched medially. The flattened rim of the

female stylus is more pronounced in the low-costae morph than in the high-costae morph,

but the general shape is the same.

Flight. — These beetles have been collected repeatedly at lights throughout the species range.

Collecting notes. — I have collected these beetles on numerous occasions in California,

New Mexico, and Mexico. They are always near water, either standing or flowing. At Rancho

Nuevo, Chiapas they were beneath logs in grassy soil. In Santa Clara County, California they

occur under stones in gravel covered with algae near stream margins.

Life history. — Members of this species have been collected in all months of the year. The

life cycle is much the same as in B. pallidus (Erwin, 1967).

Distribution. — (Fig. 252). The range of this species extends from Washington south to at

least Guatemala, and east to Arkansas and Illinois. I have seen 2,603 specimens from the

following localities:

CENTRAL AMERICA

GUATEMALA: (No locality given) MCZ.

MEXICO

AGUASCALIENTES: (Aguascalientes) CNHM; (11.0 miles west of Aguascalientes) UASM; (15.0 miles west of Pabellon)

UMAH. BAJA CALIFORNIA: (Arroyo de Purisima) CAS; (Arroyo del Rosario) CAS; (Catavina) CAS; (south of El Sauzal)

CAS; (1.3 miles northwest of El Truinfo) CAS; (Hamilton Ranch) CAS; (La Mision) GRNo; (12.4 miles east of La Paz)

CAS; (Mira Flores) CAS; (Rancho Stacion Salsipuedes) UMSP; (San Ignacio) CAS; (3.0 miles east of San Isidro) CMPP;

(San Vicente) CAS; (Tijuana) CNHM; (65.0 kilometers south of Tijuana) LACM; (Triunfo) CAS. CHIAPAS: (15.6 miles

west of Comitan) UASM; (Rancho Nuevo, 8.6 miles east of San Cristobal) RTBe, UASM; (1.0 miles north of San Cristobal)

RTBe; (8.6 miles east of San Cristobal) UASM. CHIHUAHUA: (Carta Blanca, 16.0 miles west of Matachic) AMNH;

(Catarinas) AMNH; (Chihuahua City) AMNH; (25.0 miles northwest of Chihuahua City) CNC; (Primavera) AMNH; (San

Jose Babicora) AMNH. COAHUILA: (Arroyo Palo Blanco, 15.0 miles north of Saltillo) DRWh; (Saltillo) AMNH, MCZ.

DISTRITO FEDERAL: (Penon Viejo) MCZ. DURANGO: (12.2 miles south of El Banco) UASM; (Encino) AMNH; (Nom-

bre de Dios) AMNH; (Rio Chico, 15.7 miles west of Durango) UASM. GUANAJUATO: (Rio Guanajuato, 9.8 miles

south of Sialo) UASM. GUERRERO: (13.9 miles west of Chilpancingo) UASM. HIDALGO: (Guadalupe) MCZ; (Huicha-

pan) LACM; (Rio Tula, near Tasquillo) UASM; (Tula) JHen. JALISCO: (Ajijic) JHen, UATA; (4.0 miles west of Ajijic)

AMNH; (21.4 miles south of Encamacion de Diaz) UASM; (Cocula) USNM; (9.1 miles northwest of Cautla) UASM; (Guad-

alajara) AMNH, MCZ; (9.0 miles east of Guadalajara) AMNH; (13.0 miles southeast of Lagos de Moreno) UASM; (17.9

miles west of Magdalena) UASM; (12.0 miles west of Ojuelos de Jalisco) CAS; (Puente Caquixtle, 9.7 miles east of

Encamacion de Diaz) UASM; (Rio Grande de Santiago, 12.5 miles west of Ixtlahuacan del Rio) UASM; (21.0 miles

northeast of Tepatitlan) CAS; (Valle de Guadaloupe) CAS. MEXICO: (Lago Zumpango, near San Juan Zitlaltepec) UASM;

(Tempascaltepec) CAS; (7.0 miles north of Tenancingo) UASM; (Tonatico) JHen; (Valle de Bravo) JHen; (Villa Carbon)

JHen. MICHOACAN: (7.0 miles south Ario de Rosales) UASM; (10.0 miles west of Jiquilpan) DRWh; (Morelia) UASM;

(Tuxpan) JHen; (50.0 miles west of Zitacuaro) MCZ. MORELOS: (Cuernavaca) ANSP; (Progreso) WSUP. NAYARIT: (5.1

miles north of Chapalilla) UASM; (Tepic) UATA; (19.0 miles southeast of Tepic) CAS; (19.3 miles southeast of Tepic)

UASM. NUEVO LEON: (3.2 miles south of Galeana) UASM; (1.1 miles east of Iturbide) UASM; (1.3 miles east of Iturbide)

UASM; (14.8 miles west of Linares) UASM; (Monterrey) AMNH; (6.0 miles south of Monterrey) FDAG; (Rio Linares,

20.0 miles west of Linares) CAS; (Rio Sabinas Hidalgo, 4.8 miles east of Sabinas Hidalgo) UASM; (Santa Rosa Canon,

14.8 miles west of Linares) UASM. OAXACA: (Mitla) CPBo; (25.0 miles south of Mitla) ISUA; (Oaxaca) AMNH, CAS,

MCZ; (Rio Atoyac, near Juchatengo) UASM; (1.4 miles west of Tamazulapan) UASM; (4.3 miles west of Tamazulapan)

UASM; (72.5 miles south of Valle Nacional) UASM. PUEBLA: (9.0 miles north of Amatitlan) CAS; (near Petlalcingo)

UASM; (Tehuacan) CAS; (near Tehuitzingo) UASM; (1.3 miles south of Tlatlauqui) UASM. QUERETARO: (33.0 kilo-

meters north of Acambay) UASM; (near Palmillas) UASM. SAN LUIS POTOSI: (Puente de la Parada, 7.5 miles northwest

of Mexquitic) UASM; (2.0 miles south of San Luis Potosi) CAS; (2.7 miles west of Santa Catarina) UASM. SINALOA:

(Culiacan) AMNH; (Real de Piaxtla) AMNH. SONORA: BMNH; (7.2 miles southeast of Alamos) GRNo; (10.0 miles

west of Alamos) AMNH; (Cocospera Canon, 8.0 miles east of Imeris) AMNH; (16.0 miles northeast of Ciudad Obregon)

CNC; (Pesqueria) CAS; (Rancho Atascosa, 42.0 kilometers south of Nogales) CNHM; (Rio Mayo, San Bernardo) CAS;

(Sonoyta) AMNH. VERACRUZ: (Jalapa) ANSP; (Orizaba) ISUA, UNLN. ZACATECAS: (Rio Juchipila, 0.9 miles north

of Jalpa) UASM; (1.3 miles southeast of Sain Alto) UASM. STATE UNKNOWN: (Tujipilco) CAS.

UNITED STATES

ARIZONA: Apache County (White Mountains) CAS, UATA; Cochise County (Benson) CAS, (Bisbee) CAS, (Cave Creek

Canyon) PSUU, TCBa, (Cave Creek Ranch) UASM, (Chiricahua Mountains) CAS, OUCO, UASM, UATA, USNM, (Chiri-

Bombardier Beetles

107

cahua Mountains, Rucker Canyon) CAS, (Chiricahua Mountains, Rucker Lake) UASM, (Chiricahua Mountains, Rustler’s

Park) CNC, (Douglas) CMPP, (Guadalupe Canyon, 32.0 miles east of Douglas) CUNY, (15.0 miles southeast of Hookers

Hot Springs) DRWh, (Huachuca Mountains) OUCO, USNM, (Portal) AMNH, CAS, SJSC, (San Bernardino Ranch) KSU,

LA CM, (San Pedro River, near Palominas) UASM, (San Pedro River, 10.0 miles east of Sierra Vista) OSUC, UATA, (South

West Research Station, 5.0 miles west of Portal) AMNH, FDAG, GRNo, UCD, UCR, (Tombstone) SDNHM, (Willcox)

AMNH; Coconino County (Grand Canyon National Park) CAS, (Havasupai Indian Reservation, Supai Grand Canyon)

UMSP; Gila County (Coolidge Dam) SJSC, (Globe) UASM, UATA, (14.0 miles south of Globe) UASM, (Payson) UATA,

(Winkelman) UATA; Graham County (Aravaipa) CAS, (Aravaipa Creek, between Klondyke and Aravaipa) DRWh,

(Galiuro Mountains) UASM, (Gila River,) near Geronimo) UASM, (San Carlos Reservoir) SJSC, UATA; Maricopa

County (Phoenix) MCZ, OUCO, UASM, USNM, (Salt River, Phoenix) MCZ, UATA; Mohave County (Beaver Dam)

ISUA, (Hubbard Ranch) USNM, (Littlefield) UCD, (16.0 miles north of Wikieup) LACM; Navajo County (Show

Low) LACM; Pima County (Baboquivari Mountains, Browns Canyon) AMNH, CAS, (Cienega Creek, Pantano) CUNY,

(Coyote Mountain, 0.5 miles north of Mendoza Canyon) UATA, (Saint Xavier Mountains) CAS, (Santa Catalina

Mountains) CAS, UATA, USNM, (Santa Catalina Mountains, Bear Canyon) CAS, (Santa Catalina Mountains, Sabino

Canyon) CAS, TLEr, UASM, UATA, (Santa Rita Mountains) CAS, (Tucson) CAS, CUNY, UASM, UATA, UIMI,

UMAH, USNM; Pinal County (Aravaipa Canyon) CUNY, (Sycamore Camp, 9.0 miles northwest of Payson) CAS;

Santa Cruz County (Canelo) UATA, CUNY, (Lewis Springs, San Pedro River) UASM, (Nogales) CAS, CNHM, UASM,

UCD, USNM, (Pajarita Mountains) CAS, (Patagonia) CAS, CNHM, CUNY, UATA, (5.0 miles southwest of Patagonia)

AMNH, (Pena Blanca) CUNY, OSUC, UASM, (Santa Rita Mountains, Madera Canyon) UASM, UCD, WHTy, (Tumacacori

Mountains, Sycamore Canyon) CUNY, (Yanks Spring, 4.0 miles southeast of Ruby) AMNH; Yavapai County (Bumble

Bee) CAS, (Camp Verde) PSUU, (Congress) UATA, (Cottonwood) UIMI, (Granite Mountain) CAS, (Prescott) CAS, UASM,

USNM, (Wickenburg) ISUA, (5.0 miles north of Wickenburg) UMAH; Counties unknown (Atasco Mountains, Sycamore

Canyon) UATA, (Cayetano Mountains) UMAH, (Kohl’s Ranch) UATA, (Santa Cruz River) UATA, (Senator) AMNH,

(Superstition Mountains) UATA, (Texas Pass) CUNY, (Tortolita Mountains) CAS. ARKANSAS: Garland County (Hot

Springs) CAS; Hempstead County (Hope) MCZ. CALIFORNIA: Alameda County (Livermore) CAS, (Niles Canyon) UCD,

(Oakland Hills) CAS, (San Leandro) UIMI; Amador County (5.0 miles west of Sutter Creek) TLEr; Colusa County (High-

way 20 and 16) UCD, (Rumsey Canyon) UCD; El Dorado County (No locality given) CAS; Fresno County (La Fevre

Creek) CAS; Glenn County (Elk Creek) CAS; Imperial County (Calpatria) CAS; Inyo County (Big Pine) CAS, (Deep

Springs Lake) CAS, (Diaz Lake) CAS, (Freeman) CAS, (Independence) CAS, (Little Lake) CAS, (Lone Pine) CAS, (Olan-

cha) CAS, (Owens Lake) CAS, (Westgard Pass Plateau) CAS; Kern County (Caliente) ZMLS; Los Angeles County

(Alhambra) CAS, (Arroyo Seco Canyon) WBa, (Big Dalton Dam) UCR, (Frenchman Flats) CAS, (Los Angeles)

CAS, (Pasadena) CAS, (San Dimas) CAS, (Tapica County Park) GRNo; Madera County (Coarsegold) CAS, (O’

Neals) UCD; Mariposa County (Mariposa) CAS; Merced County (Merced) CAS; Monterey County (Carmel) UASM, (Jblon)

CAS, (3.0 miles southeast of Jolon) CAS, (Stone Canyon) CAS; Napa County (Monticello) UCD; Orange County (Black

Star Canyon) UCD, (Laguna Beach) CAS; Placer County (No locality given) CAS; Riverside County (Colton) CAS, (Hemet)

WBa, (Palm Canyon) CAS, (Palm Springs) CAS, (Riverside) CAS, (San Jacinto Mountains) CAS, (Temecula) CAS; San

Benito County (Panoche Valley) CAS; San Bernardino County (Afton Canyon) USNM, (Cajon Pass) UCD, (0.9 miles north-

east of Cedar Springs) GRNo; San Diego County (Carrizo) UIMI, (Chicken Creek) CAS, (3.0 miles south of Dehesa) TLEr,

(Guatay) UIMI, (Jacumba) CAS, (Knaus) CAS, (Mission Valley) UCD, (Mount Palomar) CAS, (9.0 miles east of Pine

Valley) UCD, (Poway) CAS, (San Juan Capistrano) UIMI, (Sweetwater River) RESt; San Joaquin County (Corral Hollow)

TLEr; San Luis Obispo County (Atascadero) CAS, (Cambria) CAS, (San Luis Obispo) CAS, (Santa Margarita) CAS; Santa

Barbara County (Bluff Camp, San Rafael Mountains) UCD, (Canada del Venadito) UCD, (Cuyama River) CAS, (Gaviota)

CAS, (Santa Cruz Island) CAS, (Santa Ynez River, San Lucas) CAS; Santa Clara County (Alum Rock Park) CAS, (Arroyo

Bayo) SJSC, (Gilroy Hot Springs) TLEr, (Pacheco Pass) UIMI, (Uvas Creek) TLEr; Santa Cruz County (Santa Cruz)

CAS; Stanislaus County (Del Puerto Canyon) UIMI, (Del Puerto Creek) TLEr; Tehama County (Hills west of Tehama

County) CAS; Tulare County (Kaweah) CAS; Ventura County (Fillmore) CAS, (Foster Park) UCD, (Ojai) ZMLS, (Santa

Paula) CAS, UCD, (Ventura) CAS, (Wheeler Hot Springs) CAS; Yolo County (Davis) UCD, (Putah Canyon) UCD, (Putah

Creek) TLEr. ILLINOIS: Cook County (Riverside) UMAH. NEW MEXICO: Catron County (near Argon) AMNH, (Cooney

Canyon, 10.0 miles east of Alma ) UASM, (2.0 miles west of Luna) CCha, (San Francisco Creek, 26.1 miles north of

Glenwood) UASM; Dona Ana County (Mesquite) LACM; Grant County (Gila River, near Gila) UASM, (Sapillo Creek,

26.0 miles north of Silver City) TLEr, (Silver City) USNM, (16.0 miles west of Silver City) CAS; Hidalgo County (Pelon-

cillo Mountains) GRNo, (Post Office Canyon, 12.0 miles southeast of Rodeo) SJSC, (Rodeo) UCR; Quay County (Tucum-

cari) MCZ; San Miguel County (Sapello Canyon) USNM. NEVADA: Humboldt County (Soldier Meadows) NS DA; Kye

County (Beatty) OSUC, UWSW; Washoe County (Reno) RTBe, UCD, USNM. OREGON: (No locality given) UMSP.

SOUTH DAKOTA: Brookings County (Brookings) ANSP. TEXAS: Blanco County (2.0 miles south of Round Mountain)

UASM; Brewster County (Alpine) CAS, CUNY, MCZ, (Big Bend National Park) CNC, MCZ, UASM, (Glenn Springs)

UMAH, (Green Valley) CAS, (Saint Helena Canyon) CAS; Cameron County (Brownsville) CMPP, DHKa; Culberson County

(2.5 miles east of Nickel Creek Station) CNHM; Fayette County (Flatonia) ANSP; Hudspeth County (9.0 miles west of

Sierra Blanca) OUCO; Jeff Davis County (Barrel Springs Creek, 22.0 miles west of Fort Davis) DRWh, (Davis Mountains)

CAS, OUCO, UCD, USNM, (Fort Davis) AMNH, CNC, MCZ, (6-10.0 miles west of Fort Davis) UASM, (Limpia Canyon,

2.0 miles northwest of Fort Davis) CNC, CNHM, DRWh, TLEr, UASM; Presidio County (Presidio) TAMU; Reeves County

(Balmorhea Lake) UASM, (Pecos) JSch; San Saba County (Can^ San Saba) MCZ; Taylor County (25.0 miles southwest

of Abilene) CMPP; Travis County (Austin) CAS; Val Verde County (9.0 miles southeast of Del Rio) DRWh. UTAH: Cache

County (Logan) USUL; Utah County (Wasatch Mountains, Provo Canyon) CAS; Washington County (3.0 miles south of

Gunlock) GRNo, (12.0 miles north of La Verkin) UMSP, (Santa Clara Creek) CAS, UCD, UWSW, (Saint George) AMNH,

ISUA, MCZ, USNM, (Zion National Park) CAS. WASHINGTON: Spokane County (Spokane) CAS.

108

Erwin

Brachinus kavanaughi Erwin

(Figs. 226, 231, 232, 233, 249, 253)

Type locality. — Superior, Boulder County, Colorado, along Coal Creek.

Type specimens. - The holotype male and allotype female are in CAS. Both were collected

by D. H. Kavanaugh at the type locality, on June 4, 1968. Twenty-five paratypes col-

lected on various dates and at various localities are in CAS, DHKa, MCZ, TLEr, and

UASM.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. - Medium-sized beetles, 8.0 to 12.0 mm.

Color. Antennal article 3 at apex and all of 4, metepisterna, metasternum at sides, abdom-

inal sterna and terga infuscated to black, otherwise ferrugineous. Dorsal surface and epip-

leura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and disc of pronotum rugose and punctate, punctures

moderately impressed.

Head. As in quadripennis, except antennal scape robust, widest at middle.

Prothorax. As in quadripennis , except proepipleura with setae both anteriorly and pos-

teriorly, glabrous medially. Pronotum (fig. 226).

Pterothorax. As in quadripennis.

Abdomen. As described for genus.

Genitalia. Male (figs. 231, 232, 233). Median lobe with plane of shaft as in quadripennis.

Basal bend longer, more arcuate. Apex not so flattened. Ligule short, narrowed apically.

Virga (figs. 231 , 232). Female (fig. 249). Stylus broad, but narrowed toward apex.

Variation. — This is a fairly constant species throughout its range. The total size and the

shape of the pronotum vary within populations.

Flight. - These beetles have been collected at lights in Del Rio, Texas.

Etymology . - The latinized form of Kavanaugh, named for D. H. Kavanaugh, an excel-

lent collector of carabid beetles who collected the types.

Collecting notes. - Kavanaugh, my wife, and I collected these beetles near the type local-

ity at an elevation of 5,400 feet. They were found beneath stones during the day along a

clear cold stream whose edge was composed of stones embedded in clay and gravel, alter-

nating with gravel bars. Beetles of this species also have been collected on the shores of

lakes in Colorado by C. Armin.

Life history. — Members of this species have been collected from February to September.

Teneral adults were collected in May in Texas and in September in Illinois. Overwintering

probably takes place as an adult.

Distribution. - (Fig. 253). The range of this species extends from western New York

through the Great Lakes region, south to Nuevo Leon, Mexico, and west to central Colorado

and New Mexico. I have seen 508 specimens from the following localities:

MEXICO

NUEVO LEON: (Monterrey) AMNH; (Rio Blanquillo, 7.0 miles north of Montemorelos) UASM; (Rio Salinas at Cienaga

de Flores) UASM. TAMAULIPAS: (20.0 miles north of Ciudad Victoria) MCZ.

UNITED STATES

COLORADO: Boulder County (Coal Creek, 6.9 miles north of Golden) DHKa, (Coal Creek, Marshall Lake Area)

CArm, (Coal Creek, Superior) CArm, DHKa, (Four Mile Creek) CArm, (Hayden) CArm, (Left Hand Creek) CArm,

(McCall Lake, Lyons) CArm, (North Saint Vrain, Lyons) CArm, (Red Gulch, Lyons) CArm, (Teller Lake) CArm;

Yuma County (Wray) KSU; County unknown (Regnier) AMNH. KANSAS: Comanche County (No locality given)

KSU; Montgomery County (Independence) CAS; Riley County (Manhattan) KSU; Sheridan County (State Lake,

near Studley) RFre, UASM. ILLINOIS: Alexander County (Horseshow Lake) RTBe; Cook County (Chicago) CMPP;

Knox County (Galesburg) MCZ; La Salle County (Ottawa) RTBe; Vermilion County (Kickapoo) RTBe, (Oakwood)

ISNH; County unknown (30.0 miles south of Grape Creek) RTBe; MISSOURI: Gasconade County (Gasconade

River) UWSW; Jefferson County (Kimmswick) UMAH; Saint Louis County (Saint Louis) CAS, SDNHM; Wright

County (Gull Creek, west of Mount Grove) TCBa. NEW MEXICO: Bernalillo County (Cedro Canyon) AMNH;

Bombardier Beetles

109

Eddy County (Blue Spring, 10.0 miles east of Carlsbad) USNM; San Miguel County (Sapello Creek, Sapello) UASM. NEW

YORK: Tompkins County (Ithaca) UASM. NEBRASKA: Banner County (Glen Rock Canyon) UNLN; Glen Sioux County

(No locality given) AMNH. OHIO: Franklin County (Columbus) OUCO; Hamilton County (Cincinnati) UMAH; Ottawa

County (Put-in-Bay, South Bass Island) UMAH. OKLAHOMA: Comanche County (Wichita National Forest) CAS, OSUS.

SOUTH DAKOTA: Lawrence County (Spearfish) VMKi. TEX^lS: Blanco County UMAH, USNM, (Round Mountain)

OUCO, USNM, (2.0 miles south of Round Mountain) UASM; Brazos County (College Station) TAMU; Burnet County

(Inks Lake State Park) UMSP; Cameron County (Brownsville) CNC, USNM; Comal County (New Braunfels) USNM;

Cornell County (No locality given) MCZ; Culberson County (2.5 miles east of Nickel Creek Station) CNHM; Eastland

County (No locality given) UMSP; Gillespie County (No locality given) OUCO; Gray County (McClellan) UMSP; Hays

County (San Marcos) CAS; Kimble County (Roosevelt) CAS; Lampasas County (Lampasas River) CUNY; Lee County

(Fedor) CMPP; McLennan County (Waco) MCZ, USNM, ZMLS; Pecos County (Sheffield) CAS; San Saba County (Camp

San Saba) MCZ; Shelby County (Kerrville) CNC; Taylor County (25.0 miles southwest of Abilene) AMNH, CMPP, CNHM;

Travis County UMAH, (Austin) AMNH, MCZ, WSUP; Val Verde County (Del Rio) CAS, CNC, (9.0 miles southeast of

Del Rio) DRWh; Victoria County (Victoria) USNM; Counties unknown (Fuller) USNM, (Tiger Mills) USNM. WYOMING:

Platte County (Glendo Reservoir, near Glendo) DRWh.

Brachinus javalinopsis new species

(Figs. 227, 228, 229, 245, 246, 254)

Type locality. - Willcox, Cochise County, Arizona.

Type specimens. — The holotype male and allotype female are in AMNH. Both were collec-

ted at the type locality by T. Cohn, P. Boone, and M. Cazier on September 7, 1950.

Fifteen paratypes collected on the same day at the same locality by the same collectors

are in AMNH, CAS, MCZ, TLEr, and UASM.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Large-sized beetles, 12.3 to 13.5 mm.

Color. As in quadripennis, except antennal articles 3 and 4 infuscated only apically, and

tibiae, tarsi, and palpi ferrugineous.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows rugose and sparsely punctate, disc of pronotum densely

rugose and punctate, punctures moderately impressed.

Head. Frontal furrows moderately to deeply impressed. Antennal scape robust, widest

about middle. Ligula with center area ellipsoid-convex and plurisetose. Mentum and sub-

mentum with accessory setae.

Prothorax. Pronotum (fig. 245) convex, flattened along center line, sides widely reflexed.

Proepipleura and proepisterna as in quadripennis. Anterior tibia with anterior margin punc-

tate, punctures occasionally forming shallow strigae.

Pterothorax. As in quadripennis , except elytra broader, costae sharper, and pubescence

denser and shorter. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 227, 228, 229). Median lobe with plane of shaft rotated about 45°

from plane of basal bend. Basal bend short. Apex of shaft narrowed, ridged ventrally. Ligule

short, broad, truncate apically. Virga (figs. 227, 228). Female (fig. 246). Stylus broad, nar-

rowed apically, rounded at apex.

Variation. — Intrapopulational variation occurs in the shape of the pronotum, the color

of the antennal articles, and in the total size. The color of antennal articles 3 and 4 is paler

in some individuals.

Flight. — These beetles have been collected repeatedly at lights throughout the range of

the species.

Etymology. - Spanish, javili, wild pig; Latin, opsis, likeness; referring to the very large,

broad habitus of these beetles.

Collecting notes. — D. Larson and W. Sharp have collected these beetles at margins of

ponds in Texas.

Life history. — Members of this species have been collected from May to September, and

in January. Teneral adults were collected in July in New Mexico. Overwintering probably

110

Erwin

takes place as an adult.

Distribution. — (Fig. 254). The range of this species extends over much of the American

Southwest, from western Arizona to the Texas panhandle, and south to Brownsville. I have

seen 1 18 specimens from the following localities:

UNITED STATES

ARIZONA: Cochise County (Douglas) UCD, UCR, (14.0 miles northeast of Douglas) UCR, (17.0 miles east of Douglas)

UCR, (Guadalupe Cknyon) CUNY, (3.0 miles east of Johnson) RCGr, (10.0 miles north of Paradise) TLEr, (Portal) GRNo,

UCD, (South West Research Station, 5.0 miles west of Portal) AMNH, UCD, (Willcox) AMNH, UATA; Gila County (Globe)

UATA; Graham County (Thatcher) UCD; Maricopa County (Gila Bend) CAS. MISSOURI: (No locality given) ISNH. NEW

MEXICO: Dona Ana County (Las Cruces) CNHM, (15.0 miles north of Las Cruces) RCGr, (State College) USNM; Eddy

County AMNH, (Black River, near Whites City) UWSW, (Carlsbad) UWSW, (Whites City) UWSW; Hidalgo County (Lords-

burg) CNC, (Post Office Canyon, Peloncillo Mountains) UCR, (Rodeo) GRNo, UCD, UCR, (13.0 miles north of Rodeo)

AMNH; Luna County (Deming) AMNH; Quay County (Tucumcari) SDSU; San Miguel County (Las Vegas) UIMI. TEXAS:

Brewster County (9.0 miles north of Alpine) OSUC, (Hot Spring, Big Bend National Park) CNC; Frio County (5.0 miles

north of Dilley) UASM; Hudspeth County (9.0 miles southwest of Del City) AMNH; Kleburg County (Kingsburg) USNM;

Presidio County (7.0 miles north of Marfa) RCGr; Randall County (Palo Duro State Park) UMSP; Scurry County (Snyder)

TCBa; Val Verde County (Del Rio) UASM; Victoria County (Victoria) USNM.

Brachinus neglectus LeConte

(Figs. 237, 238, 239, 243, 250, 255)

Brachinus neglectus LeConte, 1844: 49. Lectotype, here selected, a male, MCZ red type

label number 31,775. This specimen is unlabelled, but stands third in a series of four

specimens behind label “B. quadripennis Dejean.” Type locality. — Georgia, as given

originally by LeConte.

Diagnostic combination. — The diagnostic characters are given in the key.

Description. - Medium-sized beetles, 8.4 to 12.3 mm.

Color. Metasternum at sides, metepisterna, and abdominal sterna and terga infuscated,

otherwise ferrugineous. Dorsal surface and epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and sometimes disc of pronotum rugose and sparsely

punctured, punctures moderately impressed.

Head. As in quadripennis , except antennal scape robust, widest at middle, mentum and

submentum with accessory setae.

Prothorax. As in quadripennis, except anterior margin of anterior tibia punctate, punc-

tures rarely forming strigae. Pronotum (fig. 243).

Pterothorax. As in quadripennis .

Abdomen. As described for genus.

Genitalia. Male (figs. 237, 238, 239). Median lobe with plane of shaft rotated slightly

from plane of basal bend. Basal bend long. Apex of shaft narrowed, ridged ventrally. Ligule

short, truncate. Virga (figs. 237, 238). Female (fig. 250). Stylus broad basally, narrowing

to almost acute apex.

Variation. — The few specimens available representing this species exhibit the usual var-

iation in the shape of the pronotum and its macrosculpture.

Flight. - These beetles have been collected repeatedly at lights in Georgia and Florida.

Etymology . — Latin, neglectus, forgotten.

Life history. — Members of this species have been collected from March to September.

Teneral adults have been collected in May in Georgia. These beetles probably overwinter as

adults.

Distribution. — (Fig. 255). The range of this species extends from North Carolina to

southern Florida, and west to southern Alabama. I have seen 32 specimens from the follow-

ing localities:

Bombardier Beetles

111

UNITED STATES

ALABAMA: Mobile County (Mobile) CAS, (Saraland) CAS. FLORIDA: Alachua County FDAG, (Gainesville) TLEr,

UMAH; Columbia County (O’Leno State Park) CUNY; Duval County (Jacksonville) CAS; Highlands County (Archbold

Biology Station) PSUU; Levy County (Manatee Springs State Park) RFre; Orange County (Winter Park) MCZ; Pinellas

County (Dunedin) CAS; Polk County (Lakeland) UMAH; County unknown (Guntown) ANSP. GEORGIA: Charlton

County (Okefenokee Swamp, Billy’s Island) CUNY; Chatham County (Savannah) CAS; Thomas County (Thomasville)

ANSP, MCZ. NORTH CAROLINA: Moore County (Southern Pines) CAS. SOUTH CAROLINA: (No locality given)

MCZ.

The phaeocerus subgroup

This subgroup is characterized as follows: median lobe with long narrow apex, elytra

bright blue with a contrasting black suture, and outer antennal articles black. Four species,

B. phaeocerus Chaudoir, B. azureipennis Chaudoir, B. consanguineus Chaudoir, and B.

imporcitis new species, are included.

Brachinus phaeocerus Chaudoir

(Figs. 257, 258, 259, 265, 275, 279)

Brachinus phaeocerus Chaudoir, 1868: 300. Lectotype, here selected, a male, MHNP,

labelled “Tejas” and “Ex Museo Chaudoir.” Type locality. — Texas, as originally given

by Chaudoir’s label.

Diagnostic combination. - The diagnostic characteristics are given in the key.

Description. — Small-sized beetles, 7.2 to 8.1 mm.

Color. Antennal articles 3 and 4, mesepimera, metepistema, metasternum at sides, ab-

dominal sterna and terga, tibiae and tarsi infuscated to black. Antennal articles 5-11, and

metasternal process (between middle coxae), usually darkly infuscated. Female somites 7

and 8, and male somites 8 and 9, and remainder of beetle, ferrugineous. Dorsal surface and

epipleura of elytra bright blue with black sutural margins.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and surface of pronotum densely rugose and punctate,

punctures moderately deeply impressed.

Head. Frontal furrows moderately deeply impressed. Antennal scape robust, but nearly

cylindrical. Ligula with sclerotized center area with two lateral rows of three setae per row.

Mentum and submentum without accessory setae.

Pro thorax. Pronotum (fig. 265) slightly convex, flattened along center line, sides slightly

re flexed. Proepipleura glabrous. Proepisterna with a few setae both anteriorly and poster-

iorly, glabrous medially. Anterior tibia with anterior surface strigose.

Pterothorax. Elytra moderately long, narrow, moderately costate. Humeral angle square

to prominent. Costae and depressions pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 257, 258, 259). Median lobe with plane of shaft rotated slightly

from plane of basal bend. Basal bend moderately long. Apex of shaft narrowed to apex,

elongate. Ligule short, narrow, truncate. Virga (figs. 257, 258). Female (fig. 275). Stylus

narrow, tapering to acute apex.

Variation. — Intrapopulational variation occurs in the shape of the pronotum, and degree

of infuscation of the outer antennal articles.

Flight. — G. E. Ball collected these beetles at lights in Big Bend National Park, Texas.

Etymology . — Greek, phaios, dusky brown; keros , horn; referring to the infuscated

antennae of these beetles.

Collecting notes. — G. E. Ball collected these beetles in Carex marshes at Colorado

Springs; and my wife and I collected them with B. kavanaughi specimens along streams near

Golden, Colorado.

Life history. — Members of this species have been collected from February to September.

Figs. 256, 260, 261, 265. Pronotum, right half, dorsal aspect. 256. Brachinus consanguineus Chaudoir, Rio Papagayo,

Guerrero, Mexico. 260. Brachinus azureipennis Chaudoir, 20 miles north of El Mogote, Guerrero, Mexico. 261. Bra-

chinus imporcitis new species, Globe, Arizona. 265. Brachinus phaeocerus Chaudoir, 25.0 miles southwest of Abilene,

Texas. Figs. 257-259, 262-264, 266-271. Male genitalia. 257. Brachinus phaeocerus Chaudoir, 25.0 miles southwest of

Abilene, Texas, ventral aspect. 258. Lateral aspect of same. 259. Dorsal aspect of same. 262. Brachinus imporcitis

new species, Globe, Arizona, ventral aspect. 263. Lateral aspect of same. 264. Dorsal aspect of same. 266. Brachinus

consanguineus Chaudoir, Rio Mayo, Sonora, Mexico, ventral aspect. 267. Lateral aspect of same. 268. Dorsal aspect

of same. 269. Brachinus azureipennis Chaudoir, 2.0 miles north of El Mogote, Guerrero, Mexico, ventral aspect.

270. Lateral aspect of same. 271. Dorsal aspect of same. Accompanying scale lines equal 1.0 mm.

Bombardier Beetles

113

Figs. 272-275. Right stylus of female ovipositor, ventral aspect. 272. Brachinus consanguineus Chaudoir, 41.4 miles

north of Acapulco, Guerrero, Mexico. 273. Brachinus azureipennis Chaudoir, 2.0 miles north of El Mogote, Guerrero,

Mexico. 274. Brachinus imporcitis new species, Pinal Creek, Arizona. 275. Brachinus phaeocerus Chaudoir, 25.0 miles

south of Abilene, Texas. Figs. 276-279. Geographical distribution maps. 276. Brachinus azureipennis Chaudoir.

277. Brachinus consanguineus Chaudoir. 278. Brachinus imporcitis new species. 279. Brachinus phaeocerus Chau-

doir. Accompanying scale line equals 1.0 mm.

114

Erwin

Teneral adults were collected in May in Texas and in June in Oklahoma. Overwintering

probably takes place as an adult.

Distribution. — (Fig. 279). The range of this species extends from Nebraska south to

Chihuahua, Mexico. The distribution seems to be continuous to eastern Kansas, but a great

discontinuity lies between Kansas and G. E. Ball’s New York record. Ball recorded the

capture of these specimens in his collecting notes, so there is no doubt that he took them

alive near Ithaca, New York. To the west, these beetles extend to the east flank of the Chiri-

cahua Mountains and the White Mountains of Arizona. They also range down the Gila River

system into Arizona where they apparently hybridize with B. imporcitis new species. I have

seen 213 specimens from the following localities:

MEXICO

CHIHUAHUA: (San Rafael) AMNH.

UNITED STATES

ARIZONA: Cochise County (Cave Creek Canyon, Portal) TCBa. COLORADO: Boulder County (Bear Creek canyon)

CArm, (Coal Creek, Marshall Lake Area) CArm, (Coal Creek, Plainview) CArm, (Coal Creek, east of Superior) CArm,

(Four Mile Creek) CArm, (Left Hand Creek, Lyons) CArm, (Red Gulch, Lyons) CArm, (Rocky Flats Reservoir) CArm,

(South Saint Vrain, east of Lyons) CArm; El Paso County (near Colorado Springs) UASM; Jefferson County (2.0 miles

north of Golden) DHKa, TLEr; Weld County (Greeley) USNM. KANSAS: Douglas County (Lawrence) MCZ; Riley

County (Manhattan) KSU. NEBRASKA: Banner County (Glen Rock Canyon) UNLN. NEW MEXICO: Catron County

(Cooney Canyon, 10.0 miles east of Alma) UASM, (San Francisco Creek, 26.1 miles north of Glenwood) UASM; Grant

County (18.0 miles north of Mimbres) TLEr, (Silver City) MCZ; Taos County (Rio Grande River, near Velarde) UASM.

NEW YORK: Tompkins County (Ithaca) UASM. OKLAHOMA: Alfalfa County (No locality given) OSUS, Comanche

County (Wichita National Forest) CAS; Jackson County (No locality given) CAS. TEXAS: Blanco County (Round Moun-

tain) OUCO, (2.0 miles south of Round Mountain) UASM; Brewster County (Alpine) CAS, (Big Bend National Park)

UASM; Culberson County (2.5 miles east of Nickel Creek Station) CNHM; Eastland County (No locality given) UMSP;

Hemphill County (Canadian) USNM; Kerr County (10.0 miles north of Kerrville) UMSP; Taylor County (25.0 miles

southwest of Abilene) CNHM; Travis County (Austin) WSUP.

Brachinus imporcitis new species

(Figs. 32, 261, 262, 263, 264, 274, 278)

Type locality. - Pinal Creek, Globe, Gila County, Arizona.

Type specimens. - The holotype male and allotype female are in CUNY. Both were collect-

ed at the type locality by A. and H. Dietrich on May 8, 1953 at an elevation of 5,500

feet. Fifteen paratypes collected at various localities on various dates are in AMNH, CAS,

MCZ, TLEr, and UASM.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. - Small-sized beetles, 7.0 to 8.9 mm.

Color. Apex of antennal articles 3 and 4, metepisterna, and abdominal sterna and terga

infuscated, otherwise ferrugineous. Outer antennal articles usually dark. Dorsal surface and

epipleura of elytra bright blue, with black sutural margins.

Microsculpture. As described for genus.

Macrosculpture. As in phaeocerus.

Head. As in phaeocerus, except antennal scape widest about middle.

Prothorax. As in phaeocerus, except proepisterna glabrous. Pronotum (fig. 261).

Pterothorax. As in phaeocerus, except humeri narrow, sloped and wings reduced outside

stigma (fig. 32).

Abdomen. As described for genus.

Genitalia. Male (figs. 262, 263, 264). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend short. Apex of shaft narrowed to apex, slightly curved

dorsally. Ligule short, broad, rounded apically. Virga (figs. 262, 263). Female (fig. 274).

Stylus narrow, tapering to narrowly rounded apex.

Variation. — Intrapopulational variation occurs in the shape of the pronotum, and total

size, otherwise the characters are constant.

Bombardier Beetles

115

Flight. — These beetles have never been recorded in flight and it is doubtful that they can

fly.

Etymology. - Latin, Imporcitis, god of plowing; referring to the furrow-like costae of

the elytra.

Life history. — Members of this species have been collected from May to September, and

in January. Teneral adults have been collected in May at Bumble Bee, Arizona. Overwinter-

ing probably takes place as an adult.

Distribution. - (Fig. 278). The range of this species is confined to Arizona. In the San

Simon Valley these beetles occur along the Gila River system where they apparently hybrid-

ize with B. phaeocerus. I have seen 81 specimens from the following localities:

UNITED STATES

ARIZONA: Cochise County (Chiricahua Mountains) UATA, USNM, (Chiricahua National Monument) CAS, (Southwest

Research Station, 5.0 miles west of Portal) AMNH; Gila County (Globe) KSU, UMAH, (Middle Pioneer Camp, Pinal Moun-

tains) UMNH, (Pinal Creek, Globe) CUNY, (base of Pinal Mountains) CAS, UATA, (Roosevelt Lake) UMAH; Greenlee

County (Clifton) CAS; Pima County (Tucson) AMNH; Yavapai County AMNH, (Bumble Bee) CAS, (Crown King) CAS,

(Monte Crypta Mine) CUNY, (Potter Creek, Prescott) CAS, (Prescott) CAS, MCZ, USNM; County unknown (Superstition

Mountains) UATA.

Brachinus azureipennis Chaudoir

(Figs. 260, 269, 270, 271, 273, 276)

Brachynus azureipennis Chaudoir, 1876: 75. Lectotype, here selected, a male, MHNP,

labelled “Matamoros,” and “Ex Museo Chaudoir.” Type locality. — Matamoros, Puebla,

Mexico, as originally given by Chaudoir.

Diagnostic combination. — The black palpi, antennae, tibiae and tarsi, plus the metallic

blue luster of the venter will separate these beetles from all others of the genus in North and

Middle America.

Description. - Medium-sized beetles, 7.8 to 10.3 mm.

Color. Palpi, antennal articles 2-11, tibiae, tarsi, and elytra near suture, black. Metepister-

na, metasternum at sides, and abdominal sterna and terga black with metallic blue luster,

otherwise ferrugineous. Dorsal surface and epipleura of elytra bright blue with black sutural

margins.

Microsculpture. As described for genus.

Macrosculpture. As in phaeocerus.

Head. As in phaeocerus, except ligula with only two or three setae apically.

Prothorax. As in phaeocerus , except proepipleura and proepisterna completely pubescent.

Pterothorax. As in phaeocerus.

Abdomen. As described for genus.

Genitalia. Male (figs. 269, 270, 271). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend short. Apex of shaft narrow and elongate. Ligule mod-

erately long, narrow, tapering to apex. Virga (figs. 269, 270). Female (fig. 273). Stylus

broad at base, narrowed to acute apex.

Variation. — Intrapopulational variation occurs in the shape of the pronotum, body size

and in the brightness of blue color of the elytra.

Flight. — The flight of these beetles has been recorded by F. G. Werner in Jalisco, Mexico.

Etymology . — French, azur, blue; Latin, pennis, wing; referring to the bright blue color

of the elytra of these beetles.

Collecting notes. — G. E. Ball and D. R. Whitehead collected these beetles in a wet pasture

beneath stones near El Mogote, Guerrero, Mexico. There was no water in the near vicinity.

Life history. — Members of this species have been collected in January and February,

July and August, and October, but no teneral adults were seen.

Distribution. - (Fig. 276). The range of this species extends from southern Arizona south

116

Erwin

to Guerrero, Mexico. I have seen 107 specimens from the following localities:

MEXICO

AGUASCALIENTES: (4.0 miles southwest of Aguascalientes) AMNH; (15.0 miles west of Pabellon) UMAH. CHIHUA-

HUA: (Madera) AMNH; (Primavera) AMNH; (Salaices) AMNH; (San Jose Babicora) AMNH. DISTRITO FEDERAL:

(Temascaltepec) CAS. DURANGO: (Durango) AMNH; (15.0 miles west of Durango) CNC; (6.0 miles northeast of El Salto)

AMNH; (Otinapa) AMNH. GUERRERO: (2.0 miles north of El Mogote) UASM. JALISCO: (Guadalajara) AMNH; (16.0

kilometers west of Jalostotitlan) MCZ. MEXICO: (4.3 miles north of Ixtapan de La Sal) UASM. MICHOACAN: (Tuxpan)

RCGr. SAN LUIS POTOSI: (Puente La Parada, 7.5 miles northwest of Mexquitic) UASM. ZACATECAS: (Presa Choquen)

JHen.

UNITED STATES

ARIZONA: Cochise County (San Pedro River, near Palominas) UASM, (South West Research Station, 5.0 miles west of

Portal) AMNH; Santa Cruz County (Canelo) UATA.

Brachinus consanguineus Chaudoir

(Figs. 256, 266, 267, 268, 272, 277)

Brachynus consanguineus Chaudoir, 1876: 76. Lectotype, here selected, a male, MHNP,

labelled “Ex Museo Chaudoir” and standing first of two specimens in front of box label

“consanguineus Chaudoir, Mexique, Toluca Boucardi.” Type locality. — Toluca, Mexico,

as originally given by Chaudoir.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Medium-sized beetles, 10.2 mm.

Color. Antennal articles 3-1 1, metepisterna, and abdominal sterna and terga infuscated,

otherwise ferrugineous. Dorsal surface and epipleura of elytra bright blue with black sutural

margins.

Microsculpture. As described for genus.

Macrosculpture. As in phaeocerus, except punctures finer.

Head. As in phaeocerus, except frontal furrow more deeply impressed, antennal scape

widest apically, and ligula plurisetose.

Prothorax. As in phaeocerus, except sides widely reflexed, and proepipleura and proepis-

terna pubescent. Pronotum (fig. 256).

Pterothorax. As in phaeocerus.

Abdomen. As described for genus.

Genitalia. Male (figs. 266, 267, 268). Median lobe with plane of shaft rotated slightly

from plane of basal bend. Basal bend short. Apex of shaft elongate, tubular. Ligule very

short, truncate. Virga (figs. 266, 267). Female (fig. 272). Stylus narrow, longer than in

phaeocerus, narrowed apically.

Variation. — Too few specimens are known to evaluate geographic variation.

Flight. — The flight of these beetles has not been recorded.

Etymology . - Latin, consanguineus, related by blood; probably referring to the similar-

ity of these beetles to other members of the genus.

Life history. — Members of this species have been collected in August and December, but

no teneral adults were seen.

Distribution. — (Fig. 277). The range of this species extends from Sonora to Guerrero

along the western side of Mexico. I have seen four specimens from the following localities:

MEXICO

GUERRERO: (Rio Papagayo, 41.4 miles north of Acapulco) UASM. SINALOA: (28.0 miles east of Villa Union) CNC.

SONORA: (Rio Mayo, Caramechi) CAS.

The oaxacensis subgroup

This subgroup is characterized as follows: virga elongate and strongly sclerotized, lateral

pronotal setae lacking, and highly raised elytral costae. One species, B. oaxacensis new

species, is included.

Bombardier Beetles

117

Brachinus oaxacensis new species

(Figs. 280, 290, 291, 292, 297, 300)

Type locality. — Twenty-five miles south of Mitla, Oaxaca, Mexico.

Type specimens. — The holotype male is in ISUA. The allotype female is in UASM. The

male was collected at the type locality on January 4, 1956, by J. C. Schaffner. The female

was collected by G. E. Ball and D. R. Whitehead on March 3, 1966 along the Rio Atoyac,

near Juchatengo, Oaxaca. Three paratypes are in JHen, TLEr, and UCD.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Medium-sized beetles, 10.2 to 10.6 mm.

Color. Sides of abdominal sterna slightly infuscated, otherwise ferrugineous. Dorsal sur-

face and epipleura of elytra bright bluish-green.

Macrosculpture. Frontal furrows and surface of pronotum finely rugose and punctate,

punctures moderately impressed.

Head. Frontal furrows moderately impressed. Antennal scape cylindrical. Ligula with

sclerotized center area ellipsoid-convex with two lateral rows of three setae per row. Men-

tum and submentum with accessory setae.

Prothorax. Pronotum (fig. 280) slightly convex, flattened along center line, sides slightly

reflexed. Lateral setae absent. Proepipleura and proepisterna with numerous setae throughout

their length. Anterior tibia with anterior surface strigose.

Pterothorax. Elytra elongate, broad, strongly costate. Humeral angle square. Depressions

with numerous setae, costae smooth. Depression 1 with erect depression setae twice as long

as elytral pubescence. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 290, 291, 292). Median lobe with plane of shaft rotated from plane

of basal bend about 45°. Basal bend short. Apex of shaft broadly rounded. Ligule short,

broad, rounded apically. Virga (figs. 290, 291). Female (fig. 297). Stylus narrow, parallel-

sided, rounded apically.

Variation. — Too few specimens are known to evaluate geographic variation.

Flight. — The flight of these beetles has not been recorded.

Etymology. - Oaxaca, the place where the types are from; Latin, ensis, place, locality

or county.

Collecting notes. — G. E. Ball and D. R. Whitehead collected one specimen of this species

from under a rock at the edge of the Rio Atoyac, Oaxaca, Mexico.

Life history. — Members of this species have been collected in March, May, and Decem-

ber, but no teneral adults have been seen.

Distribution. — (Fig. 300). The range of this species extends from Sonora to Oaxaca. I

have seen five specimens from the following localities:

MEXICO

GUERRERO: (Cacahuamilpa) JHen. OAXACA: (Rio Atoyac, near Juchatengo) UASM; (25.0 miles south of Mitla) ISUA.

SONORA: (10.0 miles southeast of Alamos) UCD.

The patruelis subgroup

The members of this subgroup are characterized by the short median lobe, with three

ligules, and their narrow sloped humeri. One species, B. patruelis LeConte, is included.

Brachinus patruelis LeConte

(Figs. 282, 284, 285, 286, 296, 301)

Brachinus patruelis LeConte, 1844: 50. Lectotype, here selected, a female, MCZ red type

label number 5842, further labelled with a pink disc and “B. conformis Dej. patruelis

LeC.” Type locality. — New York, as originally given by LeConte.

Figs. 280-283. Pronotum, right half, dorsal aspect. 280. Brachinus oaxacensis new species, Rio Atoyac, Oacaca, Mex-

ico. 281. Brachinus ovipennis LeConte, Charlotte, Vermont. 282. Brachinus patruelis LeConte, Towaco, New Jersey.

283. Brachinus conformis Dejean, Archbold Research Station, Florida. Figs. 284-295. Male genitalia. 284. Brachinus

patruelis LeConte, Fall River, Massachusetts, ventral aspect. 285. Lateral aspect of same. 286. Dorsal aspect of same.

287. Brachinus conformis Dejean, Archbold Research Station, Florida, ventral aspect. 288. Lateral aspect of same.

289. Dorsal aspect of same. 290. Brachinus oaxacensis new species, Cacahuamilpa, Guerrero, Mexico, ventral aspect.

291. Lateral aspect of same. 292. Dorsal aspect of same. 293. Brachinus ovipennis LeConte, Prince Edward County,

Ontario, Canada, ventral aspect. 294. Lateral aspect of same. 295. Dorsal aspect of same. Accompanying scale lines

equal 1 .0 mm.

Bombardier Beetles

119

Diagnostic combination. - The diagnostic characteristics are given in the key.

Description. — Small-sized beetles, 5.7 to 8.3 mm.

Color. Antennal articles 3 and 4, mesepisterna, metepisterna, metasternum at sides, and

abdominal sterna and terga infuscated, otherwise ferrugineous. Dorsal surface and epipleura

of elytra blue.

Micro sculpture. As described for genus.

Macrosculpture. Frontal furrows rugose, sparsely punctate. Surface of pronotum sparsely

punctate, punctures barely impressed.

Head. Frontal furrows moderately impressed. Antennal scape robust, widest at middle.

Ligula with sclerotized center area ellipsoid-convex with two apical setae. Mentum and sub-

mentum without accessory setae.

Prothorax. Pronotum (fig. 282) convex, slightly flattened along center line, sides moder-

ately reflexed. Proepipleura glabrous. Proepisterna with a few setae both anteriorly and

posteriorly, glabrous medially. Anterior tibia with anterior surface strigose.

Pterothorax. Elytra short, weakly costate. Humeral angle narrow, sloped. Costae and

depressions pubescent. Metasternum short, its length less than diameter of middle coxa.

Wings reduced outside stigma, as in figure 32.

Abdomen. As described for genus.

Genitalia. Male (figs. 284, 285, 286). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend short. Median lobe very short and robust. Apex of

shaft broadly rounded. Ligule short, broad, with two lateral accessory ligules. Virga (figs.

284, 285). Female (fig. 296). Stylus short, narrow, narrowly rounded apically.

Variation. — Intrapopulational variation occurs in the shape of the pronotum, body size,

and in the prominence of the humeri.

Flight. — The flight of these beetles has not been recorded, and it is probable that they

cannot fly.

Etymology . — Latin, patruelis, cousin, kin; referring probably to the similar habitus of

these beetles to other Brachinus species.

Life history. — Members of this species have been collected from April to September.

Teneral adults were collected in May in Massachusetts.

Distribution. — (Fig. 301). The range of this species extends from Massachusetts south to

New Jersey. Western populations are in Michigan and Illinois. I have seen 61 specimens

from the following localities:

UNITED STATES

CONNECTICUT: New London County (Old Lyme) CAS. ILLINOIS: (No locality given) ISNH. MASSACHUSETTS:

Bristol County (Fall River) CAS; Middlesex County (Framingham) CAS, UNLN, (Newton) MCZ; Norfolk County (Brook-

line) MCZ; Plymouth County (Marion) MCZ; Suffolk County (Boston) CAS, (Dorchester) MCZ, UMAH; County unknown

(Blue Hills) ISUA, USNM, (Forest Hills) USNM, (Freetown) CAS. MICHIGAN: Wayne County (Detroit) CAS. NEW JER-

SEY: Bergen County (Fort Lee) CAS; Burlington County (Atrion) CAS; Morris County (Towaco) USNM; Warren County

(Great Piece Meadows) AMNH. NEW YORK: New York County (Staten Island) CAS, (Yonkers) CNHM; Suffolk County

(Southold) CUNY ; County unknown (Newtown, Long Island) CAS.

The conformis subgroup

This subgroup is characterized as follows: median lobe small, narrow, and chisel-shaped,

and stylus of female ovipositor elongate and narrow. One species, B. conformis Dejean, is

included.

Brachinus conformis Dejean (Figs. 283, 287, 288, 289, 298, 302)

Brachinus conformis Dejean, 1831: 427. Lectotype, here selected, a female, MHNP, lab-

elled “conformis m. in Amer. bor.” on green paper, “LeConte” and “Ex Museo Chau-

doir” on white paper. Type locality. — North America, as originally given by Dejean, but

herewith restricted to Florida.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Small-sized beetles, 5.0 to 8.0 mm.

Color. Terminal palpal articles, antennal articles 2, 3, and 4, metepisterna, and abdominal

120

Erwin

296

297

298 299

H

Figs. 296-299. Right stylus of female ovipositor, ventral aspect. 296. Brachinus patruelis LeConte, Fall River, Massa-

chusetts. 297. Brachinus oaxacensis new species, Rio Atoyac, Guerrero, Mexico. 298. Brachinus conformis Dejean,

Archbold Research Station, Florida. 299. Brachinus ovipennis LeConte, Ithaca, New York. Figs. 300-303. Geograph-

ical distribution maps. 300. Brachinus oaxacensis new species. 301. Brachinus patruelis LeConte. 302. Brachinus con-

formis Dejean. 303. Brachinus ovipennis LeConte. Accompanying scale line equals 1.0 mm.

Bombardier Beetles

121

sterna and terga infuscated, otherwise ferrugineous. Dorsal surface and epipleura of elytra

blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and surface of pronotum punctate and microrugose,

punctures barely impressed.

Head. Frontal furrows shallowly impressed. Antennal scape robust, widest at middle.

Ligula with sclerotized center area ellipsoid-convex with two lateral rows of three setae per

row. Mentum and submentum without accessory setae.

Prothorax. Pronotum (fig. 283) convex, slightly flattened along center line, sides barely

reflexed. Proepipleura and proepisterna pubescent both anteriorly and posteriorly, glabrous

medially. Anterior tibia with anterior surface strigose.

Pterothorax. Elytra elongate, narrow, moderately costate. Humeral angle square. Costae

smooth, glabrous on disc, depressions between costae pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 287, 288, 289). Median lobe with plane of shaft barely rotated

from plane of basal bend. Basal bend short. Shaft straight, narrow, flattened, apex narrowly

rounded. Ligule short, narrowed apically, rounded at apex. Virga (figs. 287, 288). Female

(fig. 298). Stylus elongate, narrow, rounded apically.

Variation. — Too few specimens are known to evaluate geographic variation.

Flight. - These beetles have been collected at lights repeatedly in Florida.

Etymology . - Latin, conformis, similar; referring to the similarity between these beetles

and those of other species in the genus.

Life history. — Members of this species have been collected in May, June, and November,

and no teneral adults have been seen.

Distribution. - (Fig. 302). The range of this species extends from middle Florida to

northern Florida. I have seen 34 specimens from the following localities:

UNITED STATES

FLORIDA: Alachua County (Gainesville) USNM; Duval County (Jacksonville) OUCO; Highlands County (Aichbold

Biology Station) PSUU, (Highlands Hammock State Park) TLEr; Osceola County (Kissimmee) USNM; Pinellas County

(Dunedin) PUM; Putnam County (Welaka) CUNY.

The ovipennis subgroup

This subgroup is characterized by compressed and collapsed median lobe, the orientation

of the virga in the median lobe, and the ovate elytra. One species, B. ovipennis LeConte, is

included.

Brachinus ovipennis LeConte

(Figs. 281, 293, 294, 295, 299, 303)

Brachinus ovipennis LeConte, 1862: 525. Lectotype, here selected, a male, MCZ red type

label number 31,774, further labelled with a pink, disc, “89” and “B. cephalotes LeC,

perplexus Dej, cephalotes LeC, ovipennis LeC.” Type locality. - Middle states, as indi-

cated by LeConte’s pink disc, but herewith restricted to Vermont.

Brachinus cephalotes LeConte, 1862: 525. Lapsus calami.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Medium-sized beetles, 7.4 to 10.7 mm.

Color. Metepisterna and sides of abdomen infuscated, otherwise ferrugineous. Dorsal sur-

face and epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and surface of pronotum rugose and punctate, punctures

barely impressed.

122

Erwin

Head. Frontal furrows moderately impressed. Antennal articles robust, widest about

middle. Ligula with sclerotized center area ellipsoid-convex with two lateral rows of three

setae per row. Mentum with one to four accessory setae at middle. Submentum with acces-

sory setae.

Prothorax. Pronotum (fig. 281) slightly convex, flattened along center line, sides slightly

reflexed. Proepipleura and proepisterna pubescent both anteriorly and posteriorly, glabrous

medially. Anterior tibia with anterior surface punctate, punctures in some specimens fused

to form weak strigae.

Pterothorax. Elytra elongate, narrow, weakly costate. Humeral angle sloped. Costae and

depressions pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 293, 294, 295). Median lobe with plane of shaft rotated 45° from

plane of basal bend. Basal bend short. Apex of shaft narrowly rounded. Dorsal surface of

median lobe .usually collapsed, ligule ill-defined. Virga (figs. 293, 294), oriented sideways in

median lobe. Female (fig. 299). Stylus short, narrow, narrowly rounded at apex.

Variation. — Intrapopulational variation occurs in body size, shape of the pronotum,

shape of antennal scape, and amount of area infuscated on venter.

Flight. - The flight of these beetles has not been recorded.

Etymology . — Latin, ovum, egg; pennis, wing; referring to the ovoid-shaped elytra of

these beetles.

Collecting notes. — R. T. Bell collected these beetles at the edge of ponds in Vermont,

and V. M. Kirk collected them in pitfall traps in cornfields in South Dakota.

Life history. — Members of this species have been collected from March to October. Ten-

eral adults were collected in April in New York, and in September in Indiana, Kansas, and

New York. Overwintering probably takes place as an adult.

Distribution. — (Fig. 303). The range of this species extends from New England and

Quebec, west to Kansas and South Dakota, south to Texas. I have seen 528 specimens

from the following localities:

CANADA

ONTARIO: (Belleville) CNC; (Brimley) ZMLS; (Cedaivale) CAS; (De Cew Falls) CNC; (Erindale) CAS; (Forest Hill

Village) CAS; (Point Pelee) ZMLS; (Prince Edward County) CAS, CUNY, MCZ; (Sarnia) UMAH; (Toronto) CAS, CNC,

MCZ, ZMLS.

UNITED STATES

CONNECTICUT: Litchfield County (Cornwall) CAS; New Haven County (New Haven) CAS. ILLINOIS: Alexander Coun-

ty (Cairo) ISNH; Cook County (Chicago) CAS, (Evanston) UMAH, (Glencoe) UMAH, (La Grange) CAS, (Riverside)

UMAH, (Schiller Park) CNHM; Lake County (Lake Zurich) RTBe, (Waukegan) USNM; Madison County (Mitchell) USNM;

Mason County (Havana) ISNH; Rock Island County (Rock Island) UMAH. INDIANA: Knox County (No locality given)

PUM; Posey County (Hovey) CEWh, PUM, (Mount Vernon) CEWh. IOWA: Dickenson County (Lake Okoboji) USNM;

Johnson County (Iowa City) USNM. KANSAS: Douglas County (Lawrence) PUM, UMAH; Geary County (No locality

given) KSU; Hamilton County (No locality given) CAS; McPherson County (McPherson) CMPP; Sedgwick County USNM,

(Wichita) CAS; Seward County (No locality given) KSU. MASSACHUSETTS: Hampden County (Chicopee) MCZ, USNM.

MICHIGAN: Huron County (Charity Island) UMAH; Kent County (Grand Rapids) UNLN; Macomb County (Mount

Clemens) CNHM; Saint Clair County (Port Huron) USNM; Washtenaw County USNM, (Ann Arbor) JSch, UMAH, (Salem)

UMAH; Wayne County (Detroit) USNM; County unknown (Pentwater) CNHM. MINNESOTA: Goodhue County (Lake

Pepin, east of Frontenac) UMSP, Houston County (No locality given) UMSP; County unknown (Cook Creek) UMSP.

MISSOURI: Saint Louis County (Webster Groves) USNM. NEBRASKA: Douglas County (Omaha) UNLN; Lancaster

County (Lincoln) UNLN; Otoe County (Nebraska City) UNLN. NEW YORK: Cayuga County (Montezuma marsh) UASM;

Chautauqua County (Dunkirk) CAS; Columbia County (Copake Falls) CNHM; Dutchess County (Red Hook) UMSP;

Erie County (Buffalo) ANSP, CAS, CMPP, ISNH, USNM, (Hamburg) CAS; Genesee County (East Bethany) FDAG; Monroe

County (Honeoye Falls) CUNY, (Rochester) LACM, MCZ; New York County (Bronx Park) CAS, (Staten Island) MCZ;

Niagara County (Olcott) CUNY; Ontario County (Canandaigua) MCZ; Oswego County (Minetto, Oswego) CUNY; Tomp-

kins County (Groton) UCD, (Ithaca) CAS, CUNY, GRNo, ISNH, KSU, MCZ, OUCO, PSUU, UASM, UCR, UNLN, (Six

Mile Creek, Ithaca) TLEr; Schuyler County (Watkins Glen) VMKi; Wyoming County (Gainesville) CUNY; County unknown

(Atwaters) USNM, (Danby) UNLN. OHIO: Ashtabula County (Conneaut) PUM, (Rock Creek) PUM, (Saybrook) PUM;

Cuyahoga County (Berea) USNM; Hamilton County (Cincinnati) UMAH; Lake County (Perry) PUM; Lucas County (No

locality given) PUM; Ottawa County (Gypsum) OUCO; Sandusky County (Winous Point) SJSC. OKLAHOMA: Cleveland

County (No locality given) UONO; Comanche County (Wichita National Forest) CAS. PENNSYLVANIA: Allegheny

County (Pittsburgh) CMPP; Crawford County (Conneaut Lake) CMPP; Erie County (Fort Erie) CAS; Tioga County (Rut-

Bombardier Beetles

123

land) ANSP. RHODE ISLAND: Washington County (Watch Hill) USNM. SOUTH DAKOTA: Bennett County (Martin)

VMKi; Brookings County (Brookings) SDSU, VMKi, (Volga) VMKi, (White) VMKi; Butte County (Belle Fourche) VMKi;

Harding County (Buffalo) VMKi; Hutchinson County (Menno) VMKi; Jones County (Murdo) SDSU; Lawrence County

(Spearfish) VMKi; Meade County (Bear Butte) VMKi. TEXAS: Dallas County (Dallas) MCZ. VERMONT: Addison County

(Ferrisburg) UMAH; Chittenden County (Burlington) UATA, (Home Creek Delta, Charlotte) RTBe, (Shelburne) CAS,

(Shelburne Pond, Shelburne) RTBe; Franklin County (La Moille River, East Georgia) RTBe; Grand Isle County (Alburg)

ISNH. WISCONSIN: Dane County (Madison) UWMW; Green County (Brodhead) UMAH; Milwaukee County (No locality

given) UWMW.

The tenuicollis subgroup

This subgroup is characterized as follows: median lobe elongate and narrow, elytra large,

square, and strongly costate, and mesepisterna infuscated. One species, B. tenuicollis Le-

Conte, is included.

Brachinus tenuicollis LeConte

(Figs. 307, 312, 313, 314, 324, 329)

Brachinus tenuicollis LeConte, 1844: 49. Lectotype, here selected, a female, MCZ red type

label number 5849, further labelled with a pink disc and “v. librator Dej., similis LeC.,

79 = tenuicollis - cat., similis.” Type locality. — New York, as originally given by LeConte.

Brachinus ballistarius LeConte, 1848: 199. Lectotype, here selected, a male, MCZ red type

label number 5846, further labelled with a pink disc. Type locality. — New York, as

originally given by LeConte. NEW SYNONYMY.

Brachinus similis LeConte, 1848: 199. Lectotype, here selected, a female, MCZ red type

label number 5849, further labelled as above. Type locality. — New York, as originally

given by LeConte. NEW SYNONYMY.

Diagnostic combination. - The large size, strongly costate elytra, and infuscated mese-

pisterna separate these beetles from all others in North America.

Description. — Large-sized beetles, 11.9 to 14.5 mm.

Color. Mesepisterna, mesepimera, metepisterna, metasternum at sides, and abdominal

sterna and terga infuscated to black. Antennal articles 3 and 4 infuscated at apex in some

specimens, otherwise beetles ferrugineous. Dorsal surface and epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and disc of pronotum rugose and punctate, punctures

barely impressed.

Head. Frontal furrows moderately impressed. Antennal scape cylindrical, or almost so.

Ligula with sclerotized center area ellipsoid-convex with two lateral rows of three setae per

row. Mentum and submentum without accessory setae.

Prothorax. Pronotum (fig. 307) slightly convex, flattened along center line, sides slightly

reflexed. Proepipleura and proepistema with or without setae. Anterior tibia with anterior

surface punctate.

Pterothorax. Elytra elongate, broad, strongly costate. Humeral angle square. Depressions

pubescent, costae at least on disc glabrous, smooth. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 312, 313, 314). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend short. Apex of shaft narrow, almost acute. Ligule

short, narrow, truncate. Virga (figs. 312, 313). Female (fig. 324). Stylus broad, rounded

apically.

Variation. — Intrapopulational variation occurs in the shape of the pronotum and in body

size. The proepipleura and proepistema are glabrous or setose, the setae either spread uni-

formly over the surface, or restricted anteriorly and posteriorly.

Flight. — These beetles have been collected repeatedly at lights throughout the range of

the species.

124

318 319 320 321 322 323

Figs. 304-307, 311. Pronotum, right half, dorsal aspect. 304. Brachinus cyanipennis Say, Milton, Vermont. 305. Bra-

chirtus galactoderus new species, Rio Papagayo, Guerrero, Mexico. 306. Brachinus gebhardis Erwin, Horse Creek,

California. 307. Brachinus tenuicollis LeConte, Olcott, New York. 311. Brachinus medius Harris, Ritzville, Washing-

ton. Figs. 308-310, 312-323. Male genitalia. 308. Brachinus cyanipennis Say, Milton, Vermont, ventral aspect. 309 &

310. Lateral & dorsal aspects of same. 312. Brachinus tenuicollis LeConte, Olcott, New York, ventral aspect. 313 &

314. Lateral & dorsal aspects of same. 315. Brachinus medius Harris, Atbara, British Columbia, Canada, ventral

aspect. 316 & 317. Lateral & dorsal aspects of same. 318. Brachinus galactoderus new species, 12.0 miles south of

Gusave, Sinaloa, Mexico, ventral aspect. 319 & 320. Lateral & dorsal aspects of same. 321. Brachinus gebhardis

Erwin, Gilroy Hot Springs, California, ventral aspect. 322 & 323. Lateral & dorsal aspects of same. Accompanying

scale lines equal 1 .0 mm.

Bombardier Beetles

125

Etymology . — Latin, tenuis, thin; collis, neck; referring to the narrow pronotum of

these beetles.

Life history. - C. H. Lindroth informs me that H. G. James (Belleville, Ontario) reared

at least two specimens of this species from pupal cells of Hydrophilus obtusus Say (Chatter-

ton, Ontario, 2. VIII. 56). These beetles have been collected from March to September. It is

probable that adults overwinter much the same as in B. pallidus.

Distribution. — (Fig. 329). The range of this species extends from Colorado and western

Texas to the east coast from Massachusetts to Maryland and in Florida. The California re-

cord is doubtful. I have seen 649 specimens from the following localities:

CANADA

ONTARIO: (Belleville) RTBe; (Chatterton) ZMLS; (De Cew Falls) CNC; (Point Pelee) ROM; (Prince Edward County)

CUNY, UATA, UCD; (Toronto) RTBe; (Trenton) CAS, CNC. QUEBEC: (Hull) USNM; (Montreal) CAS; (Outrem’t)

CAS; (Rigaud) CAS; (Saint Eustache) CAS; (Saint Rose) CAS.

UNITED STATES

ARKANSAS: Benton County (Rogers) KSU; Conway County (No locality given) UAFA; Hempstead County (Hope)

MCZ; Lawrence County (Imboden) MCZ; Mississippi County (Osceola) JSch, UMAH; Washington County (No locality

given) UAFA. CALIFORNIA: (No locality given) UMSP. COLORADO: Boulder County (Reservoir, Four Mile Mesa)

CArm. CONNECTICUT: Hartford County (Windsor) MCZ; New London County (Old Lyme) CAS. FLORIDA: Columbia

County (O’Leno State Park) CUNY; Seminole County (Sanford) PUM. ILLINOIS: Cook County (Chicago) UMAH,

WSUP, (Elk Grove) CMPP, (La Grange) CAS, JSch, MCZ, UMAH, USNM, ZMLS, (Palos Park) RCGr, (Riverside) ANSP;

Gallatin County (Shawneetown) ISNH; Lake County (Fort Sheridan) UMAH; La Salle County (No locality given) RTBe;

Rock Island County (Moline) UMSP; Union County (Alto Pass) CNHM; Counties unknown (Falling Spring) UMAH, (Foun-

tain Bluff) ISNH. INDIANA: Knox County (No locality given) PUM; Marion County (No locality given) PUM; Porter

County (Dunes Beach) CEWh; Posey County (Hovey Lake) CEWh, PUM; Tippecanoe County (Lafayette) PUM; Vigo

County (No locality given) PUM. IOWA: Buchanan County (Independence) MCZ; Johnson County (Iowa City) USNM.

KANSAS: Ellis County (No locality given) KSU; Douglas County (No locality given) CAS; Riley County (No locality

given) KSU; Shawnee County (Topeka) CMPP; County unknown (Williston) MCZ. KENTUCKY: Edmonson County

(Mammoth Cave Nat Pk) TCBa. LOUISIANA: Calcasieu Parish (Sam Houston State Park) CUNY; East Baton Rouge Parish

(Baton Rouge) LSUB. MARYLAND: Charles County (No locality given) MCZ. MASSACHUSETTS: Middlesex County

(Billerica) MCZ; (Boston) CAS, (Concord) MCZ, (Sudbury ) MCZ, (Tyngsboro) CAS, (Waltham) MCZ, (Wayland) MCZ;

Norfolk County (Dover) MCZ, (Newton) MCZ; Suffolk County (West Roxbury) MCZ; County unknown (Forest Hills)

USNM. MICHIGAN: Allegan County (Allegan) CAS; Clinton County UMAH (Bath) JSch; Eaton County (Grand Ledge)

USNM; Genesee County (Flint) RCGr; Gratiot County (No locality given) JSch, UMAH; Huron County (Charity Island)

UMAH; Ingham County (Haslett) OSUS; Iosco County (No locality given) UMAH; Kent County (Grand Rapids) CNHM,

OSUS; Lapeer County (Sawdel Lake) RCGr, UMAH; Livingston County (E. S. George Reserve) OSUS, UMAH; Macomb

County (Mount Clemens) CNHM; Midland County (No locality given) JSch; Oakland County (No locality given) OSUS,

UMAH; Saginaw County (No locality given) UMAH; Van Buren County (Paw Paw Lake) UMAH; Washtenaw County

OSUS, (Ann Arbor) UMAH, (Whitmore Lake) CCha, JSch; Wayne County (Rockwood) UMAH; Wayne County (Detroit)

MCZ, UMAH, USNM, (Eight Mile Road) UMAH; Counties unknown (Aurelius) OSUS, (Dun Scotus) PSUU. MINNESOTA:

Dakota County (No locality given) UMSP; Hennepin County (No locality given) UMSP; Houston County (3.0 miles north

of Hokah) UMSP; LeSeuer County (Lake Madison) UMSP; Nicollet County (Saint Peter) UMSP; Pine County (mouth of

Snake River) UMSP, (4.0 miles east of Pine City) USNM; Ramsey County (Saint Paul) ISNH; Washington County (Saint

Croix River) UMSP; Watonwan County (No locality given) UMSP. MISSISSIPPI: George County (Lucedale) CUNY. MIS-

SOURI: Carter County (Van Buren) UMAH; Vernon County (Nevada Area) TLEr. NEBRASKA: Lancaster County

(Lincoln) UNLN; Nemaha County (Peru) UNLN. NEW JERSEY: Essex County (Cedar Grove) USNM; Morris County

(Boonton) USNM, (Lincoln Park) CAS, CNHM, (Towaco) USNM. NEW MEXICO: San Miguel County (Las Vegas) CMPP.

NEW YORK: Cayuga County (Montezuma Marsh) UASM; Erie County (Buffalo) ISNH, MCZ; Genesee County (Batavia)

CUNY; Monroe County (Rochester) LACM, MCZ; New York County (Cypress Hills) AMNH; Niagara County (Olcott)

CUNY; Ontario County (Canandaigua) MCZ; Oswego County (Oswego) ANSP, CUNY; Saint Lawrence County (Rossie)

JSch; Tompkins County (Ithaca) CAS, CUNY, MCZ; County unknown (Meadowdale) CUNY. OHIO: Ashtabula County

(Rock Creek) PUM. OKLAHOMA: (No locality given) CCha. TENNESSEE: Lake County (Gray’s Landing) RTBe. TEXAS:

Brazos County (College Station) TAMU; Brewster County (Alpine) MCZ; Dallas County (Dallas) MCZ; Hudspeth County

(9.0 miles southwest of Del City) AMNH; San Patricio County (Welder Wildlife Refuge, near Sin ton) CNC; Victoria County

(Victoria) USNM; County unknown (Fuller) USNM. VERMONT: Chittenden County (Burlington) RTBe, (Shelburne

Pond, Shelburne) RTBe. WISCONSIN: Calumet County (No locality given) PUM; Crawford County (Wauzeka) UWMW;

Dane County (No locality given) UWMW; Wood County (Griffith Street Nursery) UWMW, (Port Edwards) UWMW.

The cyanipennis subgroup

This subgroup is characterized by: median lobe with broad ligule, pronotum strongly

cordate, and erect depression setae two to three times as long as the elytral pubescence. One

species, B. cyanipennis Say, is included.

126

Erwin

v328

Figs. 324-328. Right stylus of female ovipositor, ventral aspect. 324. Brachinus tenuicollis LeConte, La Grange, Illin-

ois. 325. Brachinus galactoderus new species, Lake Tehuantepec, Oaxaca, Mexico. 326. Brachinus cyanipennis Say,

East Georgia, Vermont. 327. Brachinus gebhardis Erwin, Uvas Creek, California. 328. Brachinus medius Harris,

McLeans Bogs, New York. Figs. 329-333. Geographical distribution maps. 329. Brachinus tenuicollis LeConte. 330.

Brachinus cyanipennis Say. 331. Brachinus medius Harris. 332. Brachinus gebhardis Erwin. 333. Brachinus galacto-

derus new species. Accompanying scale line equals 1 .0 mm.

Bombardier Beetles

127

Brachinus cyanipennis Say (Figs. 304, 308, 309, 310, 326, 330)

Brachinus cyanipennis Say, 1823: 143. Neotype designated by me, a male, in MCZ. The spe-

cimen was selected from University of Iowa material, and sent to Lindroth. He subse-

quently deposited all Say’s neotypes in MCZ. Type locality. — Ames, Iowa. This locality

is the nearest to Say’s original area from which we had specimens. The original area was

“fissures of the rocks . . . near Engineer Cantonment” near Council Bluffs, Iowa.

Brachinus cephalotes Dejean, 1825: 317. Lectotype, here selected, a male, MHNP, labelled

“cephalotes mihi in Amer. Bor.” “Latreille” and “Ex Museo Chaudoir” and standing first,

in first of two rows below box label “cyanipennis Say.” Type locality. — “Amerique sep-

tentrionale” as originally given by Dejean. NEW SYNONYMY.

Brachinus cordicollis LeConte, 1862: 525. Lapsus calami.

Brachinus rejectus LeConte, 1862: 525. Lectotype, here selected, a female, MCZ red type

number 5843, further labelled with a pink disc and “B. cordicollis LeC. cyanipennis Say,

cordicollis LeC., rejectus LeC.” Type locality. — Kansas, as originally given by LeConte.

NEW SYNONYMY.

Diagnostic combination. - The erect depression setae of the elytra standing 2 to 3 times

higher than the ely tral pubescence separate these beetles from all others in the United States.

Description. — Medium-sized beetles, 8.0 to 12.0 mm.

Color. Metepisterna usually infuscated, otherwise ferrugineous. Dorsal surface and epi-

pleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and surface of pronotum rugose and punctate, punc-

tures moderately impressed.

Head. Frontal furrows moderately impressed. Antennal scape robust, widened apically.

Ligula with sclerotized center area ellipsoid-convex with two lateral rows of three setae per

row. Mentum of some specimens with one or two small setae at center, submentum without

accessory setae.

Prothorax. Pronotum (fig. 304) convex, strongly cordiform, sides moderately reflexed.

Proepipleura and proepisterna with a few scattered setae both anteriorly and posteriorly,

glabrous medially. Anterior tibia with anterior surface strigose.

Pterothorax. Elytra moderately long, narrow, moderately costate. Humeral angle square.

Costae smooth on disc, otherwise pubescent, depressions pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 308, 309, 310). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend long. Apex of shaft blunt. Ligule short, widened api-

cally, truncate. Virga (figs. 308, 309). Female (fig. 326). Stylus long, narrow, almost acute

apically.

Variation. — Intrapopulational variation occurs in the following characteristics: the center

of the mentum bears or lacks a few accessory setae; the antennal articles 3 and 4, metepis-

terna, and sides of the abdomen are or are not infuscated; the apex of the shaft is or is not

notched; and the virga is narrowly or broadly rounded apically.

Flight. — These beetles have been recorded at lights in South Dakota.

Etymology . — Greek, kyanos, blue; Latin, pennis, wing; referring to the blue elytra of

these beetles.

Collecting notes. — C. Chantal collected these beetles on river terraces above the Becan-

cour River in Quebec. The beetles were beneath stones on sandy clay.

Life history. - Members of this species have been collected from March to October. Ten-

eral adults were collected in May in Kansas. Overwintering probably takes place as an adult.

Distribution. - (Fig. 330). The range of this species extends from New Brunswick through

New England, west to New Mexico, and south to Texas and Alabama. I have seen 2,095

specimens from the following localities:

128

Erwin

CANADA

MANITOBA: (Lake Jessica, northeast of Winnipeg) UASM. NEW BRUNSWICK: (Saint John) USNM. ONTARIO: (Belle-

ville) ZMLS; (Go Home Bay) CNC; (Kenora) ZMLS; (2.0 miles north of Little Current) UASM; (Marmora) CNC; (Oliphant,

Bruce Peninsula Lake Huron) ZMLS; (Otter Lake, 9.0 miles south of Smith Falls) CCh; (Presque Isle) UASM; (Sever)

CMPP; (Toronto) CAS, RTBe; (Walsingham, southwest of Simcoe) ZMLS. QUEBEC: (Becancour) CCh, OSUC; (Como)

CNC, CUNY, MCZ; (Montreal) MCZ; (Outrem’t) CAS; (Rigaud) CAS, CCh; (Saint Ours) CCh; (Terreb’ne) CAS.

UNITED STATES

ALABAMA: Baldwin County (No locality given) UASM; Mobile County (Calvert) ANSP, UASM, (Mobile) CAS. ARKAN-

SAS: Pulaski County (Little Rock) AMNH. CONNECTICUT: Fairfield County (Canbury) AMNH; Hartford County (Bur-

lington) ISUA; Litchfield County AMNH, (Cornwall) CAS, CUNY; New Haven County (Hamden) CAS, (New Haven)

CAS, (South Meriden) CAS, MCZ; Tolland County (Union) ISUA; County unknown (Yoshen) AMNH. DISTRICT OF

COLUMBIA: (Black Pond) USNM, (Rock Creek, Washington) USNM. ILLINOIS: Champaign County (Urbana) UMPP;

Hancock County (Hamilton) ISUA; Knox County (Galesburg) MCZ; La Salle County (No locality given) RTBe; McHenry

County (McHenry) WSUP; Vermilion County (Camp Robert Drake, Fairmount) RTBe. INDIANA: Fulton County (No

locality given) PUM; Kosciusko County (Winona Lake) UMAH; Knox County (No locality given) PUM; Lake County

(Miller) UMAH; Montgomery County (Shades) PUM, RTBe. IOWA: Boone County (Ledges State Park) ISUA; Clayton

County (Guttenberg) ISUA, USNM; Dickinson County (Spirit Lake) USNM, UWMW; Johnson County (Iowa City) MCZ,

USNM; Lee County (Fort Madison) ANSP, CMPP, MCZ, ZMLS; Palo Alto County (Ruthven) ISUA; Story County (Ames)

ISUA; County unknown (Herrold) CAS. KANSAS: Douglas County (Lawrence) ANSP, CMPP, UWMW, (Lone Star) CNC;

Reno County (Medora) MCZ; Riley County (Marlatt) KSU. KENTUCKY: Fayette County (Lexington) TCBa; Rowan

County (Morehead) CNC. MAINE: Cumberland County (Sebago Lake, South Casco) AMNH; Oxford County (Norway)

MCZ, (Paris) CUNY; Penobscot County (Orono) UATA, UMSP. MARYLAND: Baltimore County (Baltimore) CAS,

MSUM; Montgomery County (No locality given) USNM; County unknown (Plummers Islands) USNM. MASSACHUSETTS:

Barnstable County (Barnstable) MCZ, (Hyannis) ISUA, (Woods Hole) WSUP; Bristol County (Attleboro) ZMLS, (Fall

River) CAS, (Swansea) MCZ; Franklin County (Northfield) MCZ; Hampden County (Chicopee) MCZ; Hampshire County

(Amherst) CEWh, MCZ, (Mount Tom) CMPP, USNM; Middlesex County (Arlington) CAS, UASM, (Billerica) CAS, UASM,

ZMLS, (Concord) MCZ, UASM, ZMLS, (Lincoln) MCZ, (Pepperill) UATA, (Sherbom) MCZ, (Sudbury) MCZ, (Tyngsboro)

MCZ, (Wayland) MCZ, (Waltham) MCZ, (Woburn) MCZ; Nantucket County (Nantucket) ISNH, MCZ; Norfolk County

(Brookline) MCZ, (Dedham) MCZ, (Dorchester) MCZ, (Dover) MCZ, (Sharon) CUNY, (Stoughton) USNM, (Wellesley)

MCZ, (Westwood) MCZ; Worcester County (Ashbumham) CEWh; County unknown (Mount Toby) MCZ. MICHIGAN:

Allegan County (Allegan) CAS; Charlevoix County (Beaver Island) UMAH; Eaton County (Grand Ledge) USNM; Grand

Traverse County (Marion Island) UMAH; Huron County (SandjPoint) UMAH; Kalamazoo County OUCO, (Gull Lake

Biology Station) JSch; Kent County (Grand Rapids) UNLN; Lake County (Loon Lake) UMAH; Oakland County (No

locality given) UMAH; Oceana County (Crystal Valley) CNHM; Otsego County (Lake Manuka) UMAH; Ottawa County

(No locality given) KSU, PUM; Washtenaw County (Ann Arbor) UMAH, (Whitmore Lake) JSch. MINNESOTA: Benton

County (No locality given) UMSP; Big Stone County (No locality given) UMSP; Clearwater County (Itasca State Park)

UMSP, (Itasca State Park, BohallLake) UMSP, (Itasca State Park, De Soto Lake) UMSP; Crow Wing County (Mille Lacs

Lake, near Garrison) UMSP; Dakota County (No locality given) UMSP; Douglas County (Alexandria) PSUU, UMSP; Hen-

nepin County (Minneapolis) UMSP; Houston County (Mississippi Bluff, 1-2.0 miles north of State Line) UMSP; Lac qui

Parle County (Madison) UMSP; Lake County (Basswood Lake) UMSP; Norman County (No locality given) UMSP; Otter

Tail County (Battle Lake) UMSP; Pine County (Saint Croix River, 10.0 miles east of Pine City) UMSP, (Snake River, 6.0

miles east of Pine City) UMSP; Ramsey County UMSP, (Saint Paul) ISNH, WSUP; Traverse County (No locality given)

UMSP; Washington County (Saint Croix River, 3.0 miles north of Stillwater) UMSP; County unknown (Cliff) UMSP. MIS-

SISSIPPI: George County (Lucedale) CUNY. MISSOURI: Jefferson County (Kimmswick) USNM; Saint Louis County (No

locality given) LACM. NEBRASKA: Dakota County (Hubbard) UNLN, (South Sioux City) UNLN; Lancaster County

(Lincoln) UNLN; Saunders County (Cedar Bluffs) UNLN. NEW HAMPSHIRE: Cheshire County (Swanzey Pond) MCZ;

County unknown (Three Mile Island) MCZ. NEW JERSEY: Bergen County (Bear Swamp, Ramsey) AMNH, (Emerson)

CAS, (Oakland) CAS; Cumberland County (Bridgeton) CAS; Essex County (Cedar Grove) USNM; Hunterdon County

(Hampton) AMNH; Morris County (Riverdale) MCZ; Passaic County (Great Notch) USNM; Somerset County (North

Branch) DRWh; Union County (Berkeley Heights) AMNH; Warren County (Phillipsburg) CAS; County unknown (Chelcea)

CMPP, (Split Rock Lake) CAS, USNM. NEW MEXICO: (No locality given) CMPP. NEW YORK: Clinton County (Platts-

burg) CAS; Columbia County (Hudson) CAS; Cortland County (McLean Bogs) CUNY; Dutchess County (Red Hook)

UMCP; Essex Cty (Fort Ticonderoga) PUM; New York Cty (Broad Channel) CAS, (Bronx Park) CAS, (Brooklyn) LACM,

USNM, (New York City) CAS, MCZ, (Rockaway Beach) MCZ, (Staten Island) CAS, USNM, (Yonkers) CAS, CHNM, MCZ;

Niagara County (Olcott) CUNY; Orange County (West Point) UMAH, USNM; Otsego County (Unadilla) MCZ; Putnam

County (2.0 miles northwest of Brewster) PUM; Queens County (Cunningham Park) CAS; Rockland County (Bear Moun-

tains) CNHM; Saint Lawrence County (Canton) OSUS; Suffolk County (Babylon) AMNH, (Montauk) CNHM, MCZ,

(Southold) CUNY, (Wildwood State Park) CUNY; Tompkins County (Ithaca) CAS, CUNY, UASM, UIMI, (McLean) UMSP,

(Tomkins Cove) AMNH, (Varna) UASM; Ulster County (Ashokan) AMNH, (Phoenicia) CAS; Washington County (Salem)

CAS; Westchester County (Bedford) CAS, (Peekskill) MCZ; Wyoming County (Silver Lake) CAS; Counties unknown (Cats-

kill Mountains) AMNH, (Hebron) AMNH, USNM, (Miller’s Port) CUNY. OHIO: Coshocton County (Mohican River) PUM.

OKLAHOMA: Cleveland County (No locality given) OUCO; Payne County (Stillwater) OSUS. PENNSYLVANIA: Alleg-

heny County (Pittsburgh) CAS; Bradford County (Susquehanna River, Wyalusing) CAS, UASM; Burks County (No locality

given) MCZ; Chester County (West Chester) UWMW; Cumberland County (New Cumberland) UASM, VMKi; Dauphin

County (Harrisburg) CUNY, VMKi; Franklin County (Chambersburg) USNM; Monroe County (No locality given) USNM;

Montgomery County (Areola) OUCO; Northampton County (Easton) CAS, UASM, (Delaware Water Gap) AMNH; Pike

County (Camp Colang) CNHM; Philadelphia County (Chestnut Hill) USNM; Venango County (French Lick Creek, south of

Venango) PUM; Counties unknown (Belfast) CAS, (Bethlehem) CNHM, (Edge Hill) USNM, (Lehigh Water Gap) USNM,

Bombardier Beetles

129

(North Cumberland) CAS. RHODE ISLAND: Kent County (Quonset Point) CAS; Newport County (Newport) CNHM;

Providence County (Providence) CMPP; County unknown (Moswansicut Lake) CMPP. SOUTH DAKOTA: Brookings

County (Brookings) CMPP, SDSU, (Volga) VMKi; Hyde County (Highmore) SDSU. TENNESSEE: Lake County (No local-

ity given) RTBe. TEXAS: Coryell County (No locality given) MCZ; Frio County (5.0 miles north of Dilley) UASM; Kerr

County (Kerrville) CNC; McLennan County (Waco) MCZ; Travis County (Austin) WSUP. VERMONT: Addison County

(Lewis Creek, North Ferrisburg) RTBe; Chittenden County (Burlington) RTBe, (Gilette Pond, Richmond) RTBe, (La

Moille River, Milton) RTBe, (Sandbar State Park) RTBe, (Shelburne) CAS; Franklin County (LaMoille River, East Georgia)

RTBe; Grand Isle County (Alburg) RTBe; Lamoille County (Ithiel Falls, Johnson) RTBe; Orange County (Wells River)

MCZ; Rutland County (West Haven, Lake Champlain) RTBe; Windham County (Brattleboro) USNM, (West River, Brook-

line) RTBe, (West River, Newfane) RTBe, (West River, Townshend) RTBe. VIRGINIA: Fairfax County (Great Falls)

ISUA, USNM; Loudoun County ANSP, (Harpers Ferry) USNM. WISCONSIN: Bayfield County (Lake Namekagon)

UWMW; Dane County (No locality given) UWMW; Dodge County (Beaver Dam) UMAH; Milwaukee County (No locality

given) CAS; Portage County (Stevens Point) UWMW; Sauk County (Praire du Sac) CNHM, (Victory) USNM.

The medius subgroup

This subgroup is characterized as follows: size small, venter infuscated, female stylus

small, arcuate, and median lobe small, generalized. One species,#, medius Harris, is included.

Brachinus medius Harris

(Figs. 31 1, 315, 316, 317, 328, 331)

Brachinus medius Harris, 1828: 117. Lectotype, here selected, a male MCZ red type label

number 2641 1. Type locality. — Boston, Massachusetts. Here designated, because Harris

neither labelled his specimens, nor gave a locality in his descriptions, but be collected in

the Boston area.

Brachinus minutus Harris, 1828: 1 17. Lectotype, here selected, a male, MCZ red type label

number 26412, further labelled “471 female, 104.” Harris inisidentified the sex of the

beetle. Type locality. — Boston, Massachusetts (see above). NEW SYNONYMY.

Diagnostic combination. - The small size, glabrous proepisterna, and infuscated abdomen

separate these beetles from all others in the study area.

Description. — Small-sized beetles, 5.2 to 7.3 mm.

Color. Antennal articles 3-11, metepisterna, and abdominal sterna and terga infuscated.

Antennal articles 1 and 2, metepisternum at sides, apex of tibia, and tarsi infuscated in some

specimens, in others these sclerites are ferrugineous. Dorsal surface and epipleura of elytra

blue, with greenish luster in some specimens.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and surface of pronotum rugose and punctate, punctures

barely impressed.

Head. Frontal furrows moderately impressed. Antennal scape robust, widened apically.

Ligule with sclerotized center area ellipsoid-convex with two lateral rows of three setae per

row. Mentum and submentum without accessory setae.

Prothorax. Pronotum (fig. 311) slightly convex, flattened along center line, sides slightly

reflexed. Proepipleura, and usually proepisterna glabrous, the latter sometimes with 1-3

setae near anterior edge. Anterior tibia with anterior surface strigose.

Pterothorax. Elytra short, narrow, slightly costate. Humeral angle square. Depressions

and costae pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 315, 316, 317). Median lobe with plane of shaft rotated 45° from

plane of basal bend. Basal bend long. Apex of shaft blunt, slightly turned dorsally. Ligule

short, narrowed toward apex. Virga (figs. 315, 316). Female (fig. 328). Stylus small, short,

narrow, slightly arcuate, narrowly rounded apically.

Variation. — Intrapopulational variation occurs in the following characteristics: shape of

pronotum; body size; proepipleura with or without setae; and color as indicated above. The

elytra are greenish only in the Brownsville, Texas area.

Flight. — These beetles have been collected at lights in South Dakota, Minnesota, and

130

Erwin

Texas.

Etymology. - Latin, medius, middle. Harris described this species in 1828, using two

names because he had two very small specimens and two medium-sized specimens, in com-

parison with other New England Brachinus species. He called the small ones minutus and the

others medius.

Collecting notes. — D. R. Whitehead collected these beetles from under stones along an

intermittent stream in Texas. G. E. Ball collected them on the shores of a Texas lake. Other

records indicate they also inhabit bogs, edges of reservoirs, and marshes.

Life history. - Members of this species have been collected in all months, except Decem-

ber. Teneral adults were collected in September in Michigan. Overwintering probably takes

place as an adult.

Distribution. — (Fig. 331). The range of this species is disjunct. The general pattern is

much the same as in B. quadripennis . I have seen 979 specimens from the following local-

ities:

CANADA

BRITISH COLUMBIA: (Atbara, near Creston) UASM, ZMLS; (Osoyoos) CAS, ISNH, ISUA, ZMLS; (Salmon Arm) CNHM,

CUNY, MCZ, UATA; (Shuswap Lake) ZMLS. ONTARIO: (Belleville) CNC; (Britannia) CNC; (Constance Bay) CNC;

(Hamilton) CNHM; (Osgoode) CNC; (Toronto) CAS; (Trenton) MCZ. QUEBEC: (Becancour) CCha; (Choisypr Rigaud)

ZMLS; (Fort Coulonge) CAS, CNC; (Montreal) CAS; (Rigaud) CAS; (Saint Eustache) CAS; (Saint Ours) CCha; (Saint Rose)

CAS.

MEXICO

TAMAULIPAS: (9.9 miles west of Pesca) UASM.

UNITED STATES

ALABAMA: Mobile County (Mobile) ANSP, CAS, UMAH, USNM, (Orchard) CAS, KSU, (Spring Hill) USNM. CALI-

FORNIA: ANSP, UMSP, USNM, UWMW, San Diego County (San Diego) MCZ. COLORADO: Boulder County (Boulder

Creek) CArm, (Crystal Lake) CArm, (Erie) CArm, (Longmont, Bellmire Reservoir) CArm, (Longmont, Divide Reservoir)

CArm, (Rod and Gun Club Lake) CArm, (Teller Lake) CArm, (Viele Lake) CArm; Weld County (Greeley) USNM.

CONNECTICUT: (No locality given) USNM. FLORIDA: Alachua County (Gainesville) FDAG, (4.0 miles north of High

Springs) CNC, (New nans Lake, 5.0 miles east of Gainesville) RFre; Duval County (Jacksonville) CAS, OUCO; Highlands

County (Archbold Biology Station) PSUU; Hillsborough County (Tampa) USNM; Manatee County (Oneco) UASM; Marion

County (3.0 miles southwest of Lake Marion) CNC; Osceola County (No locality given) FDAG; Pinellas County (Dunedin)

PUM, UMAH; Sarasota County (Myakka River State Park) CUNY, UASM. GEORGIA: Baker County (Newton) CNC;

Charlton County (Okefenokee Swamp) TLEr. KANSAS: Sheridan County (State Lake, near Studley) RFre, UASM.

IDAHO: Bonner County (Sagle) UWSW; County unknown (Sand Point) UWSW. ILLINOIS: Cook County (Chicago)

UWSP; McHenry County (Algonquin) ISNH; Rock Island County (Moline) UMSP. INDIANA: Vigo County (No locality

given) PUM. IOWA: Buchanan County (Independence) MCZ, USNM; Clayton County (Guttenberg) USNM; Des Moines

County (Burlington) ANSP; Dickenson County (Lake Okoboji) USNM, (Spirit Lake) UWMW; Johnson County (Iowa City)

MCZ, USNM; Lee County (Fort Madison) CAS, (Keokuk) USNM; Story County (Ames) ISUA. MARYLAND: (No locality

given) UMSP. MASSACHUSETTS: Bristol County (Fall River) ISUA, MCZ; Hampshire County (Mount Tom) CMPP,

USNM; Middlesex County (Acton) CNC, (Arlington) CNC, MCZ, (Billerica) CNC, CUNY, (Boston) CAS, USNM, (Cam-

bridge) ANSP, (Concord) MCZ, (Sudbury) CEWh, MCZ, (Waltham) MCZ, (Wayland) CNC, CNHM, MCZ, UASM; Norfolk

County (Newton) MCZ, (Sharon) CUNY; Plymouth County (Marion) MCZ; Suffolk County (Dorchester) MCZ; Worces-

ter County (Fitchburg) USNM; County unknown (Forest Hills) USNM. MICHIGAN: Alcona County UMSP; Charlevoix

County (Beaver Island) UMAH; Cheboygan County (No locality given) CUNY; Huron County (Charity Island) UMAH;

Kalamazoo County (Gull Lake Biology Station) TFH1; Lapeer County UMAH, (Lapeer State Game Area) RCGr; Livingston

County (E. S. George Reserve) UMAH; Menominee County (Menominee) CEWh; Oakland County (No locality given)

UMAH; Wayne County (Detroit) MCZ, UMAH; Washtenaw County (Ann Arbor) UMAH. MINNESOTA: Crow Wing County

(Nisswa) UMSP; Hennepin County (Bloomington) ISNH; Olmsted County (No locality given) UMSP; Polk County (Crook-

ston) UMSP; Ramsey County (Saint Paul) UMSP. MISSOURI: (No locality given) ANSP, USNM. MONTANA: Sanders

County (Perma) LRus. NEBRASKA: (No locality given) ANSP. NEW HAMPSHIRE: Strafford County (Milton) MCZ.

NEW JERSEY: Bergen County (Emerson) CAS; Cape May County (Cape May) CAS; Morris County (Boonton) USNM.

NEW YORK: Cortland County (McLean Bogs) CAS, CUNY; Erie County (Buffalo) AMNH, ANSP; Essex County (Fort

Ticonderoga) PUM; Monroe County (Rochester) LACM; Nassau County (Hewlett) USNM, (Roslyn) CAS; New York Coun-

ty (New York City) MCZ, (Yonkers) CAS, MCZ; Niagara County (Olcott) CUNY; Orange County (West Point) USNM;

Queens County (Queens) CAS; Suffolk County (Brookhaven) UASM, VMKi, (Montauk Point) CNHM, CUNY, (Riverhead)

VMKi; Tompkins County (Ithaca) CAS; Westchester County (Bedford) CAS, (Mount Vernon) CNHM; County unknown

(Chicago Bog) UMSP. NORTH DAKOTA: Benson County (11.9 miles west of York) UASM; Grand Forks County (Univer-

sity) USNM. OKLAHOMA: Marshall County (Lake Texoma, Willis) RCGr. OHIO: Adams County (No locality given)

OUCO; Cuyahoga County (Cleveland) MCZ; Lawrence County (Miller) UMAH; Tuscarawas County (No locality given)

OUCO. OREGON: Clackamas County (Oregon City) CAS; Multnomah County (Portland) CAS, USNM; County unknown

(Sauvie Island) JSch. PENNSYLVANIA: Lackawanna County (Scranton) UASM, USNM; Westmoreland County (Jeannette)

Bombardier Beetles

131

CMPP. SOUTH CAROLINA: Florence County (Florence) VMKi; Sumter County (Poinsett State Park) VMKi. SOUTH

DAKOTA: Beadle County (Huron) VMKi; Brookings County (Brookings) SDSU, VMKi; Brown County (Hecla) SDSU;

Clay County (Vermillion) SDSU; Lawrence County (Spearfish) VMKi; Yankton County (Y ankton) VMKi. TEXAS: Blanco

County (Cypress Mill) USNM; Cameron County (Brownsville) CAS, CNC, OUCO, SJSC, TLEr, USNM; Hidalgo County

(Hidalgo) CMPP, (McAllen) UMAH, (Weslaco) TAMU; Reeves County (Balmorhea Lake) UASM, (Pecos) CNC; Sutton

County (Sonora) TAMU; Terrell County (Chandler Ranch) UASM, (16.0 miles north of Dryden) UASM, (Lozier Canyon)

MCZ; Val Verde County (9.0 miles southeast of Del Rio) DRWh; Victoria County (Victoria) USNM. UTAH: Salt Lake

County (Salt Lake City) USNM; Utah County (Provo) MCZ, USNM. VERMONT: Addison County (Dead Creek, Addison)

RTBe, (Lewis Creek, North Ferrisburg) RTBe; Chittenden County (Shelburne) CAS. WASHINGTON: Adams County

(Ritzville) CAS, PUM, USNM, UWSW; Lincoln County (Sprague Lake) CAS; County unknown (Yakima River, Morgan’s

Ferry) MCZ. WISCONSIN: Bayfield County (No locality given) UWMW; Dane County (No locality given) ANSP; Wood

County (Griffith Street Nursery) UWMW.

The gebhardis subgroup

This subgroup is characterized by elytra with pubescence restricted to depression 8, and

by the form of the median lobe. The two species, B. gebhardis Erwin, and B. galactoderus

new species, included here are very similar in their external form.

Brachinus gebhardis Erwin

(Figs. 306, 321, 322, 323, 327, 332)

Brachinus gebhardis Erwin, 1965: 6. Holotype male and allotype female are in CAS. Type

locality. — Uvas Creek, 5.0 miles west of Morgan Hill, Santa Clara County, California.

Diagnostic combination. — The presence of pubescence in the eighth depression only,

the pale center of the venter, and the accessory setae of the mentum separate these beetles

from all others in North and Middle America.

Description. — Medium-sized beetles, 7.0 to 12.0 mm.

Color. Metepisterna and sides of abdomen infuscated, otherwise ferrugineous. Dorsal

surface and epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and surface of pronotum rugose and punctate, punctures

moderately impressed.

Head. Frontal furrows moderately impressed. Antennal scape cylindrical. Ligula with

sclerotized center area ellipsoid-convex with two lateral rows of three setae per row. Men-

tum and submentum with accessory setae.

Prothorax. Pronotum (fig. 306) slightly convex, flattened along center line, sides narrowly

reflexed. Proepipleura and proepisterna with scattered setae both anteriorly and posteriorly,

glabrous medially. Anterior tibia with anterior surface strigose.

Pterothorax. Elytra elongate, narrow, moderately costate. Humeral angle square. Depres-

sion 8 pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 321, 322, 323). Median lobe with plane of shaft moderately rotated

from plane of basal bend. Basal bend short. Apex of shaft narrowed, almost acute. Ligule

moderately long, broad, and rounded apically. Virga (figs. 321, 322). Female (fig. 327).

Stylus narrow, arcuate, almost acute apically.

Variation. — Intrapopulational variation occurs in the following characteristics: width

of the pronotal explanation at the anterior angles; presence or absence of the accessory setae

on the mentum; sides of abdominal sterna infuscated or not; occasionally the metasternum

is infuscated at sides. On the whole, the Arizona populations have larger members without

mental accessory setae, but with wider pronotal explanations, and wider shafts of the

median lobe.

Flight. — These beetles have been collected at lights by G. E. Ball in Arizona.

Etymology . — Greek, geb, born; Old French, hardi, shovel-shaped; referring to the.

dorsal outline of these beetles.

132

Erwin

Collecting notes. - In Santa Clara County, California, these beetles occur along the mar-

gins of intermittent streams. The sides of the streams are generally rocky with underlying

gravel, and very little vegetation near the actual stream, although emergent specimens of

Salix species (at high water) are found in some areas.

Life history. — Members of this species have been collected in all months, except Dec-

ember. Teneral adults were collected in September in Santa Clara County, California. Over-

wintering probably takes place in the adult stage.

Distribution. - (Fig. 332). The range of this species extends from northern California,

to the San Jacinto Mountains of southern California. Other aggregates of populations occur

in southern Arizona, and in southern Baja California. I have seen 151 specimens from the

following localities:

MEXICO

BAJA CALIFORNIA: (1.3 miles northwest of El Truinfo) CAS.

UNITED STATES

ARIZONA: Graham County (10.0 miles southwest of Safford) UCR; Santa Cruz County (Madera Canyon) UCD, CUNY,

(Pena Blanca) UASM, (Yanks Spring, 4.0 miles southeast of Ruby) AMNH; Pima County (Box Canyon, Santa Rita

Mountains) CNHM, (west side of Baboquivari Mountains) CAS, (Browns Canyon, Baboquivari Mountains) AMNH, (Santa

Catalina Mountains) CAS, (Santa Catalina Mountains, Sabino Canyon) CAS, TCBa, UASM, (Santa Rita Mountains) CAS,

(Tanque Verde) UATA, (Tucson) CAS. CALIFORNIA: Alameda County (Alameda Creek) CAS, (Arroyo Mocho) TLEr,

(Berkeley) USUL; Amador County (Horse Creek) TLEr; Fresno County (Le Fevre) ANSP; Los Angeles County (2.7 miles

south of Little Rock Ranger Station) GRNo, (Pasadena) CAS, MCZ, (San Francisquito Canyon) LACM, (San Gabriel

Canyon) CAS, TCBa, (Soledad Canyon) LACM, (Tanbark Flat) UCR; Monterey County (Bryson) CAS; Orange County

(Lower San Juan Campground) LACM; Riverside County (San Jacinto Mountains) CAS; San Diego County (Valley Center)

SDNHM; San Luis Obispo County (Atascadero) CAS, (San Luis Obispo) CAS; Santa Barbara County (Buellton) CAS,

(Cuyama River) CAS, (4.0 miles east of Los Prietos) UCD, (Oso Canyon) UCD, (Santa Barbara) MCZ, (Santa Cruz Island)

CAS, (West Santa Ynez River) UCD; Santa Clara County CNHM, (Gilroy Hot Springs) TLEr, (Mount Hamilton) JSch,

(Pacheco Pass) UIMI, (Uvas Creek) TLEr; Stanislaus County (Del Puerto Creek) TLEr, (20.0 miles west of Patterson) TLEr;

Ventura County (Foster Park) UCD, (Ojai) MCZ, (Santa Clara River, Santa Paula) CUNY; County unknown (Cachuma

Reservoir) CAS, (Shephards Inn) MCZ.

Brachinus galactoderus new species

(Figs. 305, 318, 319, 320, 325, 333)

Type locality. - Rio Papagayo, 41.4 miles north of Acapulco, Route 95, 700 feet, Guerrero,

Mexico.

Type specimens. — The holotype male and allotype female are in MCZ. Both were collected

at the type locality by G. E. Ball and D. R. Whitehead on December 20, 1965. Sixteen

paratypes collected at various localities and on various dates are in CAS, MCZ, TLEr, and

UASM.

Diagnostic combination. — The milky appearance of the surface of the pronotum will

separate these beetles from all others in Mexico.

Description. - Medium-sized beetles, 9.3 to 10.6 mm.

Color. Metepisterna, metasternum at sides, abdominal sterna and terga infuscated, other-

wise ferrugineous. Dorsal surfaces and epipleura of elytra slate-black without blue luster.

Microsculpture. As described for genus, except surface of pronotum with isodiametric

meshes raised into beads which apparently scatter the reflected light producing a milky

appearance.

Macrosculpture. As in gebhardis.

Head. Frontal furrows moderately impressed. Antennal scape robust, widest about mid-

dle. Ligula with sclerotized center area ellipsoid-convex with two apical setae. Mentum and

submentum without accessory setae.

Prothorax. As in gebhardis, except proepipleura glabrous.

Pterothorax. As in gebhardis.

Abdomen. As described for genus.

Bombardier Beetles

133

334

336

340 341 342

343 344 345

Figs. 334-339. Pronotum, right half, dorsal aspect. 334. Brachinus fumans Fabricius, Kahlotus, Washington. 335.

Brachinus perplexus Dejean, Dundee, Mississippi. 336. Brachinus favicollis Erwin, Jamul, California. 337. Brachinus

puberulus Chaudoir, Victoria, Texas. 338. Brachinus imperialensis Erwin, Douglas, Arizona. 339. Brachinus velutinus

Erwin, Davis, California. Figs. 340-345. Male genitalia. 340. Brachinus favicollis Erwin, Jamul, California, ventral

aspect. 341 & 342. Lateral & dorsal aspects of same. 343. Brachinus perplexus Dejean, Dundee, Mississippi, ventral

aspect. 344 & 345. Lateral & dorsal aspects of same. Accompanying scale lines equal 1.0 mm.

134

Erwin

Genitalia. Male (figs. 318, 319, 320). Median lobe with plane of shaft rotated about 30°

from plane of basal bend. Basal bend short. Apex of shaft narrowed, almost acute. Ligule

long, broad, tapering to narrow apex. Virga (figs. 318, 319). Female (fig. 325). Stylus short,

broad, tapering to narrowly rounded apex.

Variation. - Intrapopulational variation occurs in the shape of the pronotum, and in

total size. A cline may occur in the color of the knees. Those populations in the north have

darkened knees, while those in Oaxaca and Guerrero have pale knees. No specimens are

available from Jalisco and Michoacan to see where this trend develops.

Flight. — The flight of these beetles has not been recorded.

Etymology . — Greek, galaktos, milky; derus, neck; referring to the milky appearance of

the pronotum of these beetles.

Collecting notes. — G. E. Ball and D. R. Whitehead collected these beetles from under

rocks embedded in gravel at the margins of rivers in Mexico.

Life history. — Members of this species have been collected in all months, except Feb-

ruary and October, but no teneral adults have been seen.

Distribution. — (Fig. 333). The range of this species extends from Sonora, south to Oax-

aca on the west side of Mexico. I have seen 66 specimens from the following localities:

MEXICO

GUERRERO: (Rio Mezcala, 23.7 miles north of Zumpango) UASM; (Rio Papagayo, 41.4 miles north of Acapulco) UASM.

NAYARIT: (Rio Acaponeta, 2.4 miles south of Acaponeta) UASM; (Rio de las Canyas) CAS; (8.7 miles east of San Bias)

GRNo; (19.0 miles southeast of Tepic) CAS. OAXACA: (Lago Tehuantepec, Benito Juarez Dam) UASM; (Rio Niltepec,

18.4 miles west of Zanatepec) UASM. SINALOA: (3.0 miles east of Culiacan) GRNo; (3.4 miles west, 5.0 miles south of

Culiacan) GRNo; (12.0 miles south of Guasave) UASM; (Rio Panuco, 11.2 miles northeast of Concordia) UASM;(Vene-

dillo) CAS. SONORA: (7.2 miles southeast of Alamos) GRNo; (10.0 miles southeast of Alamos) UCD; (Rio Mayo, San

Bernardo) CAS.

The fumans subgroup

This subgroup is characterized by swollen median lobe, pale venter with infuscated sides,

coarsely punctate pronotum, and generally similar habitus. Six species, B. fumans (Fabri-

cius), B. perplexus Dejean, B. puberulus Chaudoir, B. velutinus Erwin, B. favicollis Erwin,

and B. imperialensis Erwin, are included.

Brachinus fumans (Fabricius)

(Figs. 334, 346, 347, 348, 358, 369)

Carabus fumans Fabricius, 1781: 307. Lectotype, designated by Lindroth in Kiel Museum,

labelled “in America, D. Blackburn.”

Brachinus cyanopterus LeConte, 1844: 49. Lectotype, here selected, a female, MCZ red

type label number 5847, further labelled with a pink disc and “B. fumans Fab., Dej.,

cyanopterus Say.” Type locality. - New York, as originally given by LeConte. LeConte

1848: 203, LeConte 1862: 524.

Brachinus sufflans LeConte, 1848: 204. Lectotype, here selected, a female, MCZ red type

label number 5648, further labelled with a pink disc and “86 v. sufflans LeC.” Type

locality. — New York, as originally given by LeConte. LeConte 1862: 524.

Brachinus affinis LeConte, 1848: 204. Lectotype, here selected, a male, MCZ red type label

number 31881, further labelled with a yellow disc and “2747” and standing ninth in a

series of 16 specimens behind box labelled “B. fumans Dej.” Type locality . — Indiana, as

originally given by LeConte. NEW SYNONYMY.

Brachinus perplexus LeConte, 1862: 524. This name must be considered a lapsus calami

because LeConte (1848) correctly sites Dejean as author of this name. However, Le-

Conte’s concept of perplexus was different than that of Dejean, according to their lab-

elled specimens.

Bombardier Beetles

135

Figs. 346-357. Male genitalia. 346. Brachinus fumans Fabricius, Vantage, Washington, ventral aspect. 347 & 348.

Lateral & dorsal aspects of same. 349. Brachinus imperialensis Erwin, Douglas, Arizona, ventral aspect. 350 & 351.

Lateral & dorsal aspects of same. 352. Brachinus velutinus Erwin, Davis, California, ventral aspect. 353 & 354. Lateral

& dorsal aspects of same. 355. Brachinus puberulus Chaudoir, Hidalgo County, Texas, ventral aspect. 356 & 357.

Lateral & dorsal aspects of same. Figs. 358-363. Right stylus of female ovipositor, ventral aspect. 358. Brachinus

fumans Fabricius, Kahlotus, Washington. 359. Brachinus favicollis Erwin, El Sauzal, Baja California, Mexico. 360.

Brachinus puberulus Chaudoir, Texas. 361 . Brachinus imperialensis Erwin, Douglas, Arizona. 362. Brachinus perplexus

Dejean, Dundee, Mississippi. 363. Brachinus velutinus Erwin, Davis, California. Accompanying scale lines equal 1 .0 mm.

136

Erwin

Brachinus tabasconus Bates, 1891: 268. Lectotype, here selected, a male, BMNH, labelled

“San Juan Bautista, Tabasco” and “Hoge” and standing first behind label “tabasconus

Bates.” Type locality. — San Juan Bautista, Tabasco, Mexico, as originally given by Bates.

NEW SYNONYMY.

Brachinus amplipennis Bates, 1891: 268. Lectotype, here selected, a female, BMNH, lab-

elled “Paso del Norte, Chihuahua, Hoge” “Tr. Ent. S. L. 1891. Brachinus amplipennis

Bates” “Syntype” and “1891-64.” Type locality. - Villa Lerdo, Durango, Mexico, as

originally given by Bates. NEW SYNONYMY.

Brachinus atbarae Stehr, 1950: 102. Holotype, male, at OUCO, labelled “Atbara, B. C.

Canada, 24-IV-45, G. Stace Smith Coll.” Allotype, a female, at OUCO, labelled as holo-

type, except 7-V-46. Thirteen paratypes in OUCO and UBC. Type locality. - Atbara, Bri-

tish Columbia, Canada. NEW SYNONYMY.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. - Medium-sized beetles, 9.0 to 14.0 mm.

Color. Metepisterna, sides of abdominal sterna, and terga infuscated, otherwise ferrugin-

eous. Dorsal surface and epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows, head behind eyes, and surface of pronotum rugose and

punctate, punctures moderately impressed.

Head. Frontal furrows moderately impressed. Antennal scape robust, widened apically.

Ligula with sclerotized center area ellipsoid-convex with two lateral rows of three setae per

row. Mentum and submentum without accessory setae.

Prothorax. Pronotum (fig. 334) slightly convex, flattened along center line, sides slightly

reflexed. Proepipleura and proepisterna with a few setae both anteriorly and posteriorly,

glabrous medially. Anterior tibia with anterior surface punctate, punctures elongate, some-

times merged.

Pterothorax. Elytra elongate, narrow, moderately costate. Humeral angle square or pro-

jecting. Costae smooth on disc, depressions pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 346, 347, 348). Median lobe with plane of shaft rotated about 45°

from plane of basal bend. Basal bend short. Shaft swollen medially, very robust, apex blunt

and slightly turned dorsally. Ligule short, broad, narrowed toward apex. Virga (figs. 346,

347). Female (fig. 358). Stylus narrow, parallel-sided, rounded apically.

Variation. - Intrapopulational variation occurs in the following characteristics: body size;

shape of the pronotum; height of costae; length to width ratio of elytra; and color of terga.

No dines are apparent.

Flight. — These beetles have been repeatedly collected at lights in Arizona and Texas.

Etymology . - Latin, fumans , smoke; referring to the crepitating ability of these beetles.

Collecting notes. — D. R. Whitehead collected these beetles in gravel from under stones at

the edges of an intermittent stream in Texas. In Colorado, C. Armin collected these beetles

at lake and reservoir edges and along irrigation canals. In Arizona, these beetles were col-

lected in wet meadows.

Life history. - Members of this species have been collected from March to October. Ten-

eral adults were collected in British Columbia and Quebec in September. Overwintering

probably takes place as an adult.

Distribution . — (Fig. 369). The range of this species extends from Maine to Washington,

south to Tabasco, Mexico. It is the most widespread species in North America. I have seen

2,055 specimens from the following localities:

Bombardier Beetles

1

Figs. 364-369. Geographical distribution maps. 364. Brachinus imperialensis Erwin. 365. Brachinus favicollis Erwin.

366. Brachinus perplexus Dejean. 367. Brachinus velutinus Erwin. 368. Brachinus puberulus Chaudoir. 369. Brachinus

fumans Fabricius.

138

Erwin

CANADA

ALBERTA: (5.0 miles south of Etzikom) DRWh; (Lethbridge) UASM. BRITISH COLUMBIA: (Atbara, Creston) CUNY,

UASM, ZMLS; (Rykerts, south of Creston) ZMLS. MANITOBA: (Aweme) UASM. ONTARIO: (Belleville) UASM; (Cedar-

dale) CAS; (Grand Bend, Lake Huron) ZMLS; (Ottawa) CNC; (Pelee Island) CNC, CUNY, PUM; (Port Hope) CAS; (Prince

Edward County) CAS; (South Cajuga, southwest of Dunnville, Lake Erie) ZMLS; (Toronto) CAS, CUNY, USNM, ZMLS.

QUEBEC: (Becancour) CCh; (Fort Coulonge) CNC; (Hull) CAS, USNM; (lie Saint Helene) UASM; (Lachine) CAS; (Mont-

real) CAS, UWMW; (Outrem’t) CAS; (Rigaud) HGou, ZMLS; (Saint Eustache) CAS; (Saint Foy) CCh; (Saint Jean) CAS;

(Saint Rose) CAS; (Vaudreuil) CAS.

MEXICO

CHIHUAHUA: (12.0 miles north of Escalon) CNC. COAHUILA: (Torreon) MCZ. NUEVO LEON: (Montemorelos) CAS;

(Monterrey) AMNH; (5.0 miles south of Monterrey) CNC; (Rio Sabina^ Hidalgo, 7.9 miles east of Sabinas Hidalgo) UASM.

SAN LUIS POTOSI: (Presa de Guadalupe, 55.3 miles west of Ciudad del Maiz) UASM. SINALOA: (12.0 miles south of

Guasave) UASM; (5.0 miles north of Mazatlan) GRNo; (Real de Piaxtla) AMNH; (Venedillo) CAS. SONORA: (Bahia

San Francisquito) GRNo; (Hermosillo) CNHM; (Rio Mayo, San Bernardo) CAS. TAMAULIPAS: (Ciudad Mante) AMNH;

(Ciudad Victoria) AMNH, CNHM; (Gomez Farias) UASM; (73.1 miles north of Manuel) UASM.

UNITED STATES

ALABAMA: Mobile County (Mobile) CAS; Tuscaloosa County (Tuscaloosa) UASM. ARIZONA: Cochise County (Cave

Creek Ranch) UASM, (Douglas) UASM, (2.0 miles northeast of Douglas) OUCO, (South West Research Station, 5.0 miles

west of Portal) CUNY, FDAG; Pima County (Tucson) AMNH, CAS, USNM; Pinal County (No locality given) JSch; Santa

Cruz County (6.0 miles north of Nogales) UASM; County unknown (East Bridge) UMSP. ARKANSAS: Bradley County

(No locality given) UAFA; Conway County (No locality given) UAFA; Garland County (Hotsprings) CAS; Greene County

(No locality given) OSUC; Hempstead County (Hope) CUNY, MCZ, UMAH; Izard County (Franklin) ISUA; Lawrence

County (Imboden) CAS, LACM, MCZ; Pulaski County (8.0 miles north of Camp Robinson) CNHM, (Little Rock) AMNH,

MCZ; Pike County (Delight) CNHM; Washington County (No locality given) ISNH, UAFA. CALIFORNIA: San Diego

County (No locality given) USNM; San Francisco County (San Francisco) CMPP. COLORADO: Boulder County (Arvada)

CArm, (Baseline Lake) CArm, (Bellmire Reservoir, Longmont) CArm, (Boulder Creek, near Boulder) CArm, (Hillcrest

Lake) CArm, (Hodgson-Harris Reservoir, Louisville) CArm, (Lyons) CArm, (McIntosh, Longmont) CArm, (Prince Lake,

Erie) CArm, (Reservoir Four Mile Mesa) CArm, (Valmont) CArm; Denver County (Denver) CAS, USNM; La Plata County

(Durango) MCZ; Otero County (Rocky Ford) USNM; Weld County (Greeley) USNM. CONNECTICUT: Fairfield County

(Danbury) AMNH, (Redding) UCR, (Stamford) USNM; New London County (Norwich) MCZ, (Old Lyme) CAS, USNM;

Windham County (Pomfret) CAS. DISTRICT OF COLUMBIA: (Black Pond) USNM, (Piney Branch) USNM, (Rock Creek)

USNM, (Woodridge) USNM. FLORIDA: Lake County (No locality given) UMAH; Orange County (Orlando) LACM.

GEORGIA: Thomas County (Boston) TAMU. IDAHO: Boise County (2.0 miles west of Boise) CNHM, (Boise River,

Boise) CNHM, UMAH; Bonner County (Sagle) UWSW; Canyon County (Homedale) CNHM, (Parma) UIMI, UMAH; Idaho

County (Clearwater) WSUP; Latah County (Kendrick) UIMI, (Moscow) UIMI; Nez Perce County (Lewiston) OUCO, UIMI,

(Myrtle) UIMI, (Snake River, 4.0 miles south of Lewiston) UIMI, (Spalding) UIMI. ILLINOIS: Alexander County (Olive

Branch) CAS; Bureau County (Bureau) MCZ; Champaign County (Urbana) CNHM; Clark County (No locality given) PUM;

Cook County (Chicago) CNHM, MCZ, WSUP, (La Grange) CAS, (Willow Springs) CAS; Greene County (Hillview) UCD;

Kendall County (Oswego) CNHM; Lake County (Fort Sheridan) UMAH, (Highland Park) UMAH; La Salle County (Ottawa)

RTBe; McHenry County (McHenry) JSch, WSUP; McLean County (Heyworth) USNM; Monroe County (Bloomington)

UMAH; Pike County (Florence) UCD; Putnam County (No locality given) ISNH; Richland and Lawrence Counties (Wab-

ash) AMNH; Rock Island County (M)line) UMSP; Saint Clair County (No locality given) CAS; Sangamon County (Spring-

field) CNHM; Tazewell County (Tremont) CNHM; Vermilion County (Kickapoo State Park) RTBe, (Muncie) ISj$H;

Warren County (Pine) CMPP; Counties unknown (Falling Springs) LACM, (Saint Claire) LACM. INDIANA: Harrison Coun-

ty (No locality given) PUM; Lake County (Miller) CNHM, PUM; Porter County (Beverly Shores) CNHM; Posey County

(Hovey Lake) CEWh, PUM, (Mount Vernon) CEWh; Steuben County (No locality given) PUM; Tippecanoe County

(Lafayette) PUM. IOWA: Dickinson County ( Lake Okoboji) USNM, (Spirit Lake) USNM; Guthrie County (8.0 miles south-

west of Bayard) ISUA; Henry County (Mount Pleasant) CAS, ISUA, MCZ, RTBe, UASM, UMAH; Howard County (Elma)

AMNH; Johnson County (Iowa City) CUNY, LACM, MCZ, USNM, (Solon) USNM; Iowa County (No locality given) MCZ;

Palo Alto County (Ruthven) ISUA; Story County (Ames) CAS, GRNo, ISUA, MSUM, RTBe, USUL; Van Buren County

(Lacey-Keosauqua State Park) ISUA; Woodbury County (Sioux City) UMSP; County unknown (Herrold) CAS. KANSAS:

Atchison County (Atchison) CMPP; Clay County (No locality given) ANSP, USNM; Coffey County (No locality given)

ULLK; Douglas County (Lawrence) UCD, UMAH, USNM, UWMW, (12.0 miles south of Lawrence) UCD; Ellis County (No

locality given) KSU; Ellsworth County (Kanopolis Kam State Park) RCGr; Franklin County (No locality given) KSU,

UMAH; Hamilton County (No locality given) CAS; Leavenworth County (Tonganoxie) MCZ; Montgomery County (Indep-

endence) CAS; Pottawatomie County (Onaga) CAS, KSU, OSUC; Riley County UIMI, (Manhattan) KSU; Rooks County

(No locality given) KSU; Saline County (Salina) CMPP, KSU, MCZ; Shawnee County (Topeka) CMPP; Wilson County (No

locality given) LACM; Counties unknown (Fort Hays) MCZ, (WiUiston) MCZ. KENTUCKY: Jefferson County (Louisville)

ULLK. MAINE: (No locality given) CNHM. MARYLAND: Montgomery County (Cabin John) USNM; Prince Georges

County (College Park) UMCP; County unknown (Plummers Islands) USNM. MASSACHUSETTS: Barnstable County

(Barnstable) MCZ, (Woods Hole) JSch, USNM; Bristol County (Fall River) CAS, (Somerset) CAS; Dukes County (Martha’s

Vineyard) USNM; Essex County (Manchester) MCZ, (Topsfield) UASM; Franklin County (Northfield) MCZ, ZMLS;

Hampden County (Chicopee) MCZ, (Springfield) MCZ, ZMLS; Hampshire County (Mount Tom) CMPP; Middlesex

County (Billerica) CAS, UATA, (Cambridge) CAS, MCZ, (Concord) MCZ, (Tyngsboro) CAS, MCZ; Nantucket

County (Nantucket) ISNH; Norfolk County (Brookline) AMNH, (Newton) MCZ, (Wellesley) MCZ; Plymouth County

(Duxbury) MCZ, (Marion) MCZ; Suffolk County (Dorchester) MCZ, ZMLS; County unknown (Forest Hills) USNM.

MICHIGAN: Huron County (Charity Island) UMAH; Livingston County (E.S. George Reserve) UMAH; Oceania

County (Pentwater) UMAH; Wayne County (Detroit) USNM; Washtenaw County (Ann Arbor) UMAH; (Sharon)

UMAH. MISSOURI: Atchison County (Langdon) AMNH; Buchanan County (Saint Joseph) USNM: Greene County

Bombardier Beetles

139

(Willard) ANSP, UASM; Jackson County (Buckner) TAMU; New Madrid County (Big Oak State Park) RTBe; Pettis

County (Sedalia) CNHM; Platte County (Parkville) ISUA; Saint Charles County (Saint Charles) MCZ; Saint Louis

County (Saint Louis) CAS, (Valley Park) UMAH; Vernon County FDAG, (Nevada) TLEr. MINNESOTA: Benton

County (No locality given) UMSP; Big Stone County (No locality given) UMSP; Clearwater County (Bohall, Lake Itasca

State Park) UMSP, (De Soto Lake, Itasca State Park) UMSP; Douglas County (Alexandria) PSUU; Goodhue County (Lake

Pepin, east of Frontenac) UMSP; Hennepin County (Minneapolis) UMSP; Houston County (near Brownsville) UMSP; Olm-

stead County (No locality given) UMSP; Norman County UMSP, (Ada) USNM; Pine County UMSP, (Snake River, 4.0 miles

east of Pine City) USNM; Ramsey County (Saint Paul) ISNH, UMSP; Saint Louis County ( Park Point, Duluth) AMNH;

Steams County (Lake Koronis, Paynesville) USNM; Traverse County (No locality given) UMSP; Washington County (3.0

miles south of Afton) UMSP, (Saint Croix River) UMSP; County unknown (Cliff) UMSP. MONTANA: Lake County

(Poison) UWSW; Liberty County (Chester) MSUM; Mineral County (6.0 miles southeast of Saint Regis) LRus; Sanders

County (Paradise) LRus; Stillwater County (Columbus) MSUM. NEBRASKA: Cass County (South Bend) UNLN, USNM;

Dakota County (South Sioux City) UNLN; Lancaster County (Lincoln) CAS, UNLN, (Malcolm) CAS, USNM, (Roca)

UNLN; Nemaha County (Peru) LSUB; Nuckolls County (Superior) OSUC; Otoe County (Nebraska City) UNLN; Saunders

County (Ashland) SDSU, (Cedar Bluffs) UNLN; Thomas County (Halsey) UMAH, UNLN. NEVADA: Humboldt County

(Golconda) CBak. NEW HAMPSHIRE: Cheshire County (Jaffrey) MCZ; Grafton County (H anover) USNM; Strafford

County (Durham) ISNH. NEW JERSEY: Bergen County (Demarest) USNM, (Emerson) CAS, (Fort Lee) AMNH, (Palisade)

MCZ, (Ramsey) AMNH; Camden County (Clementon) USNM; Cape May County (Seven Mile Beach) OUCO; Essex County

(South Orange) CAS ; Gloucester County (Glassboro) USNM, (Grenloch) USNM, (Malaga) ANSP; Morris County (Boonton)

USNM, (Chester) AMNH; Middlesex County (South Amboy) AMNH; Passaic County (Clifton) USNM, (Oak Ridge) CNHM,

(Passaic) AMNH, (Paterson) AMNH; Somerset County (No locality given) USNM; Sussex County (Hopatcong) AMNH;

Warren County (Phillipsburg) CAS; Counties unknown (Estling Lake) USNM, (Gugmard) AMNH, (Midvale) USNM, (Snake

Hill) USNM. NEW MEXICO: Sandoval County (Los Alamos) CNC. NEW YORK: Albany County (Altamont) CUNY;

Columbia County (Copake Falls) CNHM; Cortland County (McLean Bogs) CAS, CUNY, ISNH, UMCP; Delaware County

(Cooks Falls) CAS; Dutchess County (Poughkeepsie) UASM; Erie County (Buffalo) ISNH; Niagara County (Olcott) AUAA;

New York County (Bronx Park) AMNH, (Flatbush) AMNH, (Mosholu) AMNH, (Nepera Park) CAS, (New York City)

CAS, MCZ, (Parkville) AMNH, (Rattlesnake Creek) CAS; Orange County (Huguenot) AMNH, ( Pine Island) CUNY, (West

Point) USNM; Richmond County (Staten Island) AMNH, CAS, USNM; Queens County ( Jamaica) AMNH, MCZ; Rockland

County (Suffem) USNM; Saint Lawrence County (Rossie) JSch; Tompkins County (Ithaca) CAS, CUNY, KSU, OUCO,

PSUU, UASM, UMCP, UNLN, (McLean) UMSP; Ulster County (Ashokan) AMNH (Oliverea) USNM; Wayne County (Sod-

us) UASM; Westchester County (New Rochelle) CAS, (Peekskill) MCZ, (White Plains) CAS, USNM, (Yonkers) MCZ; Coun-

ty unknown (Catskill) CAS, (Danby) UNLN. NORTH CAROLINA: Beaufort County (Washington) MCZ; Haywood

County (Mount Sterling) CAS; Moore County (Southern Pines) USNM; New Hanover County (Wilmington) CAS; Pasquo-

tank County (Elizabeth City) MCZ; Polk County (Tryon) USNM; Robeson County (Lumberton) CNC; County unknown

(Faison) CNC. OHIO: Ashtabula County (Chestnut Grove) PUM, (Jefferson) PUM, (Rock Creek) PUM, (Saybrook) PUM;

Hamilton County (Cincinnati) UMAH; Ottawa County (Bass Island) PUM, (Put-in-Bay) UMAH; Putnam County (Sugar

Creek) PUM; Summit County (Hudson) MCZ. OKLAHOMA: Cleveland County (Norman) OSUS, (Uono) CAS; Delaware

County (Jay) OSUS; McCurtain County (Idabel) OSUS; Oklahoma County (No locality given) OSUS; Payne County

(Stillwater) OSUS; Rogers County (Catoosa) CNHM; Tulsa County (Tulsa) CAS. OREGON: Baker County (Robinette)

UWSW, (Snake River, Farewell Bend) UWSW; Douglas County (Roseburg) MCZ; Gilliam County (5.0 miles west of Arling-

ton) JSch; Malheur County (Ontario) UWSW; Umatilla County (Camp Umatilla) MCZ; Union County (Alicel) RESt, UIMI;

Wallaowa County (Wallaowa) OSUC; Wasco County (The Dalles) MCZ; County unknown (Olds Ferry) UIMI. PENNSYL-

VANIA: Allegheny County CAS, (Pittsburgh) CMPP; Bradford County (Susquehanna River, Wyalusing) UASM; Centre

County (State College) PSUU; Chester County (Unionville) CMPP; Dauphin County (Harrisburg) UASM, (Hemmelstown)

ANSP; Delaware County (Castlerock) USNM; Fayette County (No locality given) CMPP; Luzern County (Wyoming)

USNM; Monroe County (Echo Lake) CAS; Northampton County (Easton) CAS, (Delaware Water Gap) AMNH, UASM;

Philadelphia County (Olney) USNM, (Philadelphia) USNM; Pike County (Milford) CNHM;Tioga County UMAH, (Rutland)

ANSP; Wayne County (White Mills) CAS; Westmoreland County (Jeannette) CMPP; Counties unknown (Conk Forest)

PSUU, (Edge Hill) USNM, (Rockville) CAS, (The Rock) PSUU. RHODE ISLAND: Kent County (Quonset Point) CAS,

(Warwick) UMAH; Newport County (Newport) USNM, (Portsmouth) CAS; Providence County (Providence) CMPP; Wash-

ington County (Watch Hill) USNM; County unknown (Touisset) UMAH. SOUTH CAROLINA: Oconee County (Clemson)

WSUP. SOUTH DAKOTA: Brookings County (Brookings) CMPP, SDSU, VMKi, (Volga) CAS; Hughes County (Canning)

SDSU, (Pierre) SDSU, VMKi; Tripp County (Winner) VMKi. TENNESSEE: Lake County (Gray’s Landing) RTBe;Monroe

County (Unaka Mountains) ANSD; Morgan County (Burrville) CNHM; Counties unknown (Lookout Mountain) MCZ,

(Reelfoot Lake) CUNY, ISNH. TEXAS: Bexar County (San Antonio) CUNY, (10.0 miles northwest of San Antonio) CAS;

Blanco County (Round Mountain) CAS; Brazos County ISNH, (College Station) TAMU; Brewster County (Castolon)

TAMU, (Glenn Springs) UMAH; Cameron County (Brownsville) CAS, CUNY, DHKa, MCZ, USNM, (Esperanza Ranch,

Brownsville) USNM; Comal County (New Braunfels) UASM; Dallas County (Dallas) MCZ, UASM; Dimmit County

(No locality given) TAMU; Hidalgo County (Mission) UASM, (Weslaco) TAMU; Kleburg County (Kingsville) CUNY;

Lamar County (Paris) USNM; Lee County (Fedor) CMPP; Reeves County (Balmorhea Lake) UASM, (Pecos) ISUA;

Taylor County (Abilene) CAS; Terrell County (Lozier Canyon) MCZ; Travis County (Austin) UASM; Uvalde County

(Uvalde) CAS; Val Verde County (Del Rio) UASM, (Devil’s River, Del Rio) CNC, (9.0 miles southeast of Del Rio)

DRWh; Victoria County (Victoria) USNM; Webb County (Laredo) CAS. UTAH: Cache County (Logan) USUL,

(Wellsville) USUL; Salt Lake County (Fort Douglas) MCZ; Uinta County (No locality given) CMPP; Utah County

(Provo) CAS; Weber County (Ogden) CNHM, USNM. VERMONT: Bennington County (No locality given) CAS,

MCZ, USNM; Chittenden County (Burlington) RTBe; Franklin County (La Moille River, East Georgia) RTBe;

Lamoille County (Stowe) AMNH; Windsor County (White River Junction) CAS. VIRGINIA: Fairfax County MCZ,

(Alexandria) USNM, (Great Falls) USNM; Loudon County (Bluemont) USNM, (Harpers Ferry) USNM; Montgomery Coun-

ty (No locality given) USNM; Nelson County (No locality given) USNM; Spotsylvania County (Fredericksburg) CAS.

WASHINGTON: Adams County (Lake McElroy) CAS, PUM, USNM, (Lind) CAS, ( Ritzville) USNM; Asotin County

140

Erwin

(Grande Ronde River, Anatone) UWSW; Benton County (Hanford) UWSW; Franklin County (Kahlotus) CAS, UWSW,

(Pasco) UWSW, WSUP; Grant County (Coulee City) CAS, UWSW, (Dry Falls, Grand Coulee) UWSW, WSUP, (Stratford)

CAS, UIMI, USNM, UWSW; Kittisas County (Vantage) CAS, UWSW; Lincoln County (Sprague) USNM, (Sprague Lake)

CAS; Pend Oreille County (Usk) USNM; Spokane County (Cheney) WSUP; Walla Walla County (Burbank) UWSW, (College

Place) JSch; Whitman County (Almota) WSUP, (Wawawai) USNM, UWSW, WSUP; Counties unknown (Vila) UWSW, (Yak-

ima River, Morgan’s Ferry) MCZ. WEST VIRGINIA: USNM, Tucker County (No locality given) SJSC. WISCONSIN:

Bayfield County (No locality given) UWMW; Dane County (No locality given) UWMW, UWSW; Dodge County (Beaver

Dam) CAS, UMAH; Green County (Albany) CAS, (Brodhead) UMAH; Milwaukee County (Milwaukee) MCZ; Sauk Coun-

ty (Praire du Sac) CNHM. WYOMING: (No locality given) CMPP.

Brachinus puberulus Chaudoir

(Figs. 337, 355, 356, 357, 360, 368)

Brachinus puberulus Chaudoir, 1868: 294. Lectotype, here selected, a male, MHNP, lab-

elled “stygicornis sec. LeConte Say” and “Ex Museo Chaudoir.” Type locality. — Texas,

as originally given by Chaudoir.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Medium-sized beetles, 9.5 to 12.4 mm.

Color. As in fumans.

Microsculpture. As described for genus.

Macrosculpture. As in fumans, except punctures finer.

Head. As in fumans, except antennal scope widest at middle.

Prothorax. As in fumans, except anterior tibia with anterior surface strigose. Pronotum

(fig. 337).

Pterothorax. As in fumans, except costae weaker.

Abdomen. As described for genus.

Genitalia. Male (figs. 355, 356, 357). Median lobe with plane of shaft barely rotated

from plane of basal bend. Basal bend short. Median lobe swollen medially. Apex of shaft

narrow, slightly turned dorsally. Ligule short, parallel-sided, rounded apically. Virga (figs.

355, 356). Female (fig. 360). Stylus long, narrow, rounded apically.

Variation. — Too few specimens are known to evaluate geographic variation.

Flight. - The flight of these beetles has not been recorded.

Etymology. - Latin, pubis, down or hair; ulus, diminutive; referring to the short pubes-

cence covering these beetles.

Life history. - I have seen one adult collected in May, but it was not teneral.

Distribution. — (Fig. 368). This species is known only from Texas. I have seen six speci-

mens from the following localities.

UNITED STATES

TEXAS:MCZ, MHNP, Cameron County (Brownsville) WHTy; Hidalgo County (No locality given) TAMU; Victoria County

(Victoria) USNM.

Brachinus favicollis Erwin

(Figs. 336, 340, 341, 342, 359, 365)

Brachinus favicollis Erwin, 1965: 1 1 . Holotype male and allotype female both in CAS.

Type locality. - Jamul, San Diego County, California.

Diagnostic combination. — The very deep punctures of the pronotum separate these

beetles from others in the study area.

Description. — Medium-sized beetles, 9.5 to 10.5 mm.

Color. As in fumans, except terga ferrugineous, and sterna more narrowly infuscated at

side.

Microsculpture. As described for genus.

Macrosculpture. As in fumans, except pronotal punctures much deeper.

Head. As in fumans, except mentum and submentum with accessory setae.

Prothorax. Pronotum (fig. 336) similar to fumans. Proepipleura and proepisterna with

Bombardier Beetles

141

numerous scattered setae. Anterior tibia with anterior surface strigose.

Pterothorax. As in fumans, except costae more highly elevated.

Abdomen. As described for genus.

Genitalia. Male (figs. 340, 341, 342). Median lobe with plane of shaft rotated about 45°

from plane of basal bend. Basal bend short. Shaft not as swollen as in fumans, apex more

narrowed and slightly twisted. Ligule short, broad, truncate. Virga (figs. 340, 341). Female

(fig. 359). Stylus narrow, widened apically and rounded.

Variation. — Intrapopulational variation occurs in the shape of the pronotum, thecolor of

the elytra, and the degree to which the metepistema are infuscated.

Flight. — These beetles have been collected at lights at Camp Verde, Arizona.

Etymology . — Latin, favus, honeycomb; collis, neck; referring to the strongly punc-

tured pronotum of these beetles.

Collecting notes. — D. R. Whitehead collected these beetles at the edge of a permanent

stream near Hooker’s Hot Springs, Arizona. G. R. Noonan collected specimens from under

stones in an intermittent stream bed in southern California at an elevation of 3,250 feet.

The stream side was sandy and with small stones. The common trees in the bottom of the

gully, through which the stream flowed, were specimens of Platanus, Salix, Alnus, Populus,

and Fraxinus species.

Life history. — Members of this species have been collected from March to August, and in

January and October. Teneral adults were collected in May in California and Arizona, and

in July in Arizona. Overwintering probably takes place in the adult stage.

Distribution. — (Fig. 365). The range of this species extends from eastern Arizona into

California, and south to Baja California. I have seen 152 specimens from the following

localities:

MEXICO

BAJA CALIFORNIA: (Catavina) CAS; (San Vicente) LACM; (South of El Sauzal) CAS.

UNITED STATES

ARIZONA: Cochise County (Bass Canyon, Tenney’s Mule Shoe Ranch, near Hooker’s Hot Springs) DRWh; Gila County

(East Verde River, 6.0 miles north of Payson) LACM, (10.0 miles south of Globe) AMNH, (Base of Pinal Mountains) PUM,

UATA; Graham County (Geronimo) UATA; Navajo County (8-15.0 miles northeast of Whiteriver) AMNH; Pima County

(Santa Catalina Mountains) CAS, (Tucson) CAS, UATA; Pinal County (Aravaipa Creek) CUNY; Yavapai County (Camp

Verde) CAS, (Cottonwood) UIMI; County unknown (Carrizo) UATA, (Kohl’s Ranch) UATA. CALIFORNIA: Imperial

County (Carrizo) UMAH; Los Angeles County CMPP, (Azusa) LACM, (Los Angeles) LACM, (Pasadena) CAS, (San Francis-

quito Canyon) LACM, (San Gabriel Canyon) GRNo, (Tujunga Creek) LACM; Orange County (Black Star Canyon) UCD,

(Trabuco) GRNo; Riverside County (Hemet) VVBa, (Palm Canyon) CNC, LACM, MCZ, (Palm Springs) UCD, (Temecula)

CAS; San Bernardino County (Mojave River) CAS; San Diego County UCD, (Jumul) CAS, (Mission Valley) SDNHM, (Pamo

Valley) LACM, (Poway Groove) GRNo; Santa Barbara County (4.0 miles east of Los Prietos) UCD; County unknown (San

Juan) UWSW.

Brachinus perplexus Dejean

(Figs. 335, 343, 344, 345, 362, 366)

Brachinus perplexus Dejean, 1831: 426. Lectotype, here selected, a male, MHNP, labelled

“perplexus ? var. in Amer. Bor.” on green paper, and “Ex Museo Chaudoir” on white

paper. This specimen stands first in second row of nine specimens. Type locality . — North

America, as originally given by Dejean, but herewith restricted to Florida.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. - Medium-sized beetles, 7.9 to 1 1.0 mm.

Color. As in fumans, except terga usually ferrugineous.

Microsculpture. As described for genus.

Macrosculpture. As in fumans.

Head. As in fumans, except antennal scape widest about middle, and mentum and sub-

mentum with accessory setae.

142

Erwin

Prothorax. As in fumans , except anterior tibia with anterior surface strigose. Pronotum

(fig. 335).

Pterothorax. As in fumans, except costae weaker and more densely punctate, and humeral

angle not so prominent.

Abdomen. As described for genus.

Genitalia. Male (figs. 343, 344, 345). Median lobe with plane of shaft rotated 45° from

plane of basal bend. Basal bend long. Apex of shaft narrow, slightly twisted. Ligule moder-

ately long, broad, truncate. Virga (figs. 343, 344). Female (fig. 362). Stylus short, broad,

slightly bent at apical third, narrowly rounded apically.

Variation. - Intrapopulational variation occurs in the body size and in the shape of the

pronotum.

Flight. - These beetles have been collected at lights in Alabama and Louisiana.

Etymology . — Latin, perplexus, intricate, puzzling; referring to the similarity of these

beetles to the other species in the genus.

Life history. - Members of this species have been collected in February, and May to

August, but no teneral adults have been seen.

Distribution. - (Fig. 366). The range of this species extends from Florida to Texas, and

north to southern Iowa. I have seen 71 specimens from the following localities:

UNITED STATES

ALABAMA: Mobile County (Mobile) CAS; Tallapoosa County (Smith Mountain Tower) AUAA. ARKANSAS: Conway

County (No locality given) UAFA; Desha County (No locality given) UAFA; Hempstead County (Hope) MCZ; Lawrence

County (No locality given) CAS. FLORIDA: Dade County (Royal Palm State Park) PUM; Pinellas County (Tarpon Springs)

CAS. LOUISIANA: Ouachita Parish (Calhoun) UAFA; Vernon Parish (Rosepine) UAFA. MISSISSIPPI: Carrol County

(Avalon) UMAH; George County (Lucedale) CUNY; Pike County (McComb) UWSW; Tunica County (Dundee) UMAH.

NORTH CAROLINA: Moore County (Southern Pines) CNC. OKLAHOMA: Le Flore County (Summerfield) OSUS.

TENNESSEE: Lake County (Gray’s Landing) RTBe; Shelby County (Memphis) CAS. TEXAS: Dallas County (Dallas) MCZ;

Kleberg County (Kingsville) CUNY; Morris County (Daingerfield State Park) UASM; Victoria County (Victoria) USNM;

County unknown (Fuller) USNM.

Brachinus velutinus Erwin

(Figs. 339, 352, 353, 354, 363, 367)

Brachinus velutinus Erwin, 1965: 17. Holotype male and allotype female both in UCD.

Type locality. — Davis, Yolo County, California.

Diagnostic combination . - The diagnostic characteristics are given in the key.

Description. - Small-sized beetles, 7.0 to 8.2 mm.

Color. As in fumans, except terga and metepisterna usually ferrugineous.

Microsculpture. As described for genus.

Macrosculpture. As in fumans, except punctures very fine.

Head. As in fumans, except antennal scape almost cylindrical, andmentum with acces-

sory setae.

Prothorax. As in fumans, except much narrower, proepipleura and proepisterna pubes-

cent throughout, and anterior tibia with anterior edge strigose. Pronotum (fig. 339).

Pterothorax. Elytra narrower than in fumans, more sloped at humeri, costae hardly pres-

ent, and with denser pubescence.

Abdomen. As described for genus.

Genitalia. Male (figs. 352, 353, 354). Median lobe with plane of shaft rotated about 45°

from plane of basal bend. Basal bend short. Median lobe slightly swollen medially. Apex of

shaft narrowed, slightly twisted, and slightly bent dorsally. Ligule short, narrow, and round-

ed apically. Virga (figs. 352, 353). Female (fig. 363). Stylus short, broad, arcuate, narrowly

rounded apically.

Variation. — Too few specimens are known to evaluate geographic variation.

Flight. — The flight of these beetles has not been recorded.

Bombardier Beetles

143

Etymology . — Latin, velutinus, velvety; referring to the appearance of the elytral pub-

escence on these beetles.

Collecting notes. — W. H. Tyson collected members of this species in a Typha-Scirpus

marsh near Los Banos when the water was near maximum height.

Life history. — Members of this species have been collected in April, May, and September,

but no teneral adults were seen.

Distribution. — (Fig. 367). The range of this species extends along the Central Valley of

California in the Sacramento and San Joaquin drainage systems. I have seen 19 specimens

from the following localities:

UNITED STATES

CALIFORNIA: PUM, USNM, Contra Costa County (Brentwood) CAS; Stanislaus County (Newman) UCD; Tulare County

(Visalia) UCD; Yolo County (Davis) CAS, UCD.

Brachinus imperialensis Erwin

(Figs. 338, 349, 350, 351, 361, 364)

Brachinus imperialensis Erwin, 1965: 17. Holotype and allotype both in CAS. Type local-

ity. — Potholes, Imperial County, California.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. — Medium-sized beetles, 7.0 to 12.1 mm.

Color. Metepisterna and sides of abdominal sterna infuscated, otherwise ferrugineous.

Dorsal surface and epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. As in fumans.

Head. As in fumans , except antennal scape widest at middle, and mentum and submentum

with accessory setae.

Prothorax. As in fumans, except anterior tibia with anterior surface strigose.

Pterothorax. As in fumans, except costae weaker and elytra totally pubescent.

Abdomen. As described for genus.

Genitalia. Male (figs. 349, 350, 351). Median lobe with plane of shaft rotated about 45°

from plane of basal bend. Basal bend short. Apex of shaft broad, slightly twisted. Ligule

moderately long, broad, rounded apically. Virga (figs. 349, 350). Female (fig. 361). Stylus

short, arcuate, rounded apically.

Variation. — Intrapopulational variation occurs in body size and in the shape of the pro-

notum.

Flight. - These beetles have been collected repeatedly at lights throughout the range of

the species.

Etymology. - Imperial, English, from Imperial County, the place where the types were

collected; ensis, place, locality, or county.

Life history. — Members of this species have been collected in March and April, June to

September, and November, but teneral adults were not seen.

Distribution. — (Fig. 364). The range of this species extends from eastern Colorado,

south to the Mexican Highplain, and from California east to Texas. I have seen 218 speci-

mens from the following localities:

MEXICO

DURANGO: (Durango) AMNH; (Tlahualilo) USNM. SAN LUIS POTOSI: (San Luis Potosi) AMNH. SINALOA: (Los

Mochis) CAS. SONORA: (Ciudad Obregon) AMNH; (Hermosillo) CAS.

UNITED STATES

ARIZONA: Cochise County (Cave Creek Ranch) SJSC, UASM; (Douglas) CNHM, CUNY, FDAG, UASM, UCD, UCR;

(2.0 miles northeast of Douglas) OUCO; (5.0 miles west of Portal) UCD; (South West Research Station, 5.0 miles west of

Portal) AMNH, CUNY; Coconino County (Cameron Trading Post) UASM; Graham County (Thatcher) UCD; Pima County

(San Simon) LACM; (Tucson) MCZ, USNM; Yuma County (Fort Yuma) USNM, (Yuma) CAS, MCZ, TLEr, TMBH, UATA,

USNM; County unknown (East Bridge) ISUA, MCZ. CALIFORNIA: Imperial County (Brawley) CAS, CBak, (Calexico)

144

Erwin

UCD, UWSW, (Calipatria) CAS, (El Centro) CAS, CNHM, TLEr, (HoltviUe) CAS, (Imperial Valley) UATA, (Needles) TLEr,

(Palo Verde) UIMI, (Potholes) CAS; Los Angeles County (No locality given) CAS, UWSW; Riverside County (Blythe) CAS,

(Colorado River, Blythe) TLEr, (Lake Elsinore) USNM, (Palm Springs) USNM. COLORADO: Yuma County (Wray) KSU.

NEVADA: Clark County (Logandale) NSDA, (Overton) UASM, (Overton Boat Landing) JSch. NEW MEXICO: Curry

County (Clovis) USNM; Hidalgo County (Animas) AMNH, (Rodeo) CUNY; Luna County (Deming) MCZ. TEXAS: Brew-

ster County (Alpine) CAS, (Castolon) TAMU; Cameron County (Brownsville) CNC, CUNY, ISNH; El Paso County (El

Paso) CMPP; Hidalgo County TAMU, (McAllen) UMAH; Jeff Davis County (Fort Davis) CNC; Kleberg County (Kingsville)

CUNY; Randall County (Canyon) TAMU, (Palo Duro State Park) UMSP; Reeves County (Pecos) ISUA; San Patricio Coun-

ty (Welder Wildlife Refuge, near Sin ton) CNC; Travis County (Austin) UASM; Victoria County (Victoria) UASM.

The cordicollis group

The single characteristic shared by the species of this group is the form of the virga of the

endophallus. This structure is characteristically H-shaped with a dorsal fin on the midline or

some modification of the basic H-shape. The more darkly pigmented areas extend from the

median fin to the lateral lobes. The eight species belonging to this group are arrayed in four

subgroups.

The cordicollis subgroup

The species of this subgroup are characterized by the robust median lobe, usually with a

ventral depression on the shaft, and the true H-shaped virga. Four species, B. cordicollis

Dejean, B. sublaevis Chaudoir, B. cyanochroaticus Erwin, and B. ichabodopsis new species,

are included.

Brachinus cordicollis Dejean

(Figs. 376, 377, 378, 379, 387, 391)

Brachinus cordicollis Dejean, 1826: 466. Lectotype, here selected, a male, MHNP, labelled

“cordicollis m. in Amer. Bor.” and “D. LeConte.” Type locality. — North America, as

originally given by Dejean, but herewith restricted to Fairfax County, Virginia.

Brachinus velox LeConte, 1848: 206. Lectotype, here selected, a male, MCZ red type label

number 5850, further labelled with a pink disc, “90” and “v. velox LeC.” Type locality. —

New York, as originally given by LeConte. LeConte, 1862: 524.

Brachinus leptocerus Chaudoir, 1868: 296. Lectotype, here selected, a female, MHNP,

labelled “Ex Museo Chaudoir” and standing fifth in a series of six specimens behind box

label “leptocerus Chaud. Etas Unis Guex.” Type locality. — United States, as originally

given by Chaudoir, but herewith restricted to New York. NEW SYNONYMY.

Brachynus gracilis Blatchley, 1910: 160. Lectotype, previously selected (Blatchley, 1930:

33), a male, PUM, labelled “Marshall Co., Ind. W. S. B. 10-14-03” “L-l 1/2-36” red type

label, and a black bordered label “Holotype Brachynus gracilis Blatchley, 1910.” Type

locality. — Marshall County, Indiana, as originally given by Blatchley. NEW SYNONYMY.

Notes. — The specimen labelled gracilis holotype cannot be a holotype because Blatchley

did not originally designate it so.

Diagnostic combination. - The diagnostic characteristics are given in the key.

Description. — Medium-sized beetles, 7.0 to 10.2 mm.

Color. Antennal articles 3 and 4 various. Metepisterna, and abdominal sterna and terga

infuscated, otherwise ferrugineous. Dorsal surface and epipleura of elytra blue.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows and disc of pronotum rugose and punctate, punctures

moderately impressed.

Head. Frontal furrows moderately impressed. Antennal scape cylindrical or almost so.

Ligula with center area ellipsoid-convex with two lateral rows of three setae per row. Men-

tum and submentum without accessory setae.

Prothorax. Pronotum (fig. 379) slightly convex, flattened along center line, sides narrowly

Figs. 370, 374, 375, 379. Pronotum, right half, dorsal aspect. 370. Brachinus ichabodopsis new species, Hardkinsville,

Florida. 374. Brachinus sublaevis Chaudoir, Archbold Biology Station, Florida. 375. Brachinus cyanochroaticus

Erwin, 11.9 miles west of York, North Dakota. 379. Brachinus cordicollis Dejean, Toronto, Canada. Figs. 371-373,

376-378, 380-385. Male genitalia. 371. Brachinus sublaevis Chaudoir, Archbold Biology Station, Florida, ventral as-

pect. 372 & 373. Lateral & dorsal aspects of same. 376. Brachinus cordicollis Dejean, Ithaca, New York, ventral

aspect. 377 & 378. Lateral & dorsal aspects of same. 380. Brachinus cyanochroaticus Erwin, 1 1 .9 miles west of York,

Dakota, ventral aspect. 381 & 382. Lateral & dorsal aspects of same. 383. Brachinus ichabodopsis new species, Hard-

kinsville, Florida, ventral aspect. 384 & 385. Lateral & dorsal aspects of same. Accompanying scale lines equal 1.0

mm.

146

Erwin

reflexed. Proepisterna pubescent throughout, proepipleura various. Anterior tibia with an-

terior surface strigose.

Pterothorax. Elytra elongate, narrow, costae weakly elevated. Humeral angle square.

Costae and depressions pubescent. Wings fully developed.

Abdomen. As described for genus.

Genitalia. Male (figs. 376, 377, 378). Median lobe with plane of shaft rotated 45° from

basal bend. Basal bend long. Apex of shaft broad and blunt, with slight notch at middle.

Venter of shaft with elongate longitudinal depression. Ligule elongate, narrow, rounded

apically. Virga (figs. 376, 377). Female (fig. 387). Stylus narrow, parallel-sided, rounded

apically.

Variation. — Besides the intrapopulational variation in the shape of the pronotum and the

body size, setae of the proepipleura are present in some individuals, absent from others.

Flight. — The flight of these beetles has not been recorded.

Etymology . — Latin, cordis , heart; collis, neck; referring to the cordiform outline of

the pronotum of these beetles.

Collecting notes. — C. Chantal collected these beetles from under stones on sandy clay

terraces above the Becancour River in Quebec.

Life history. — Members of this species have been collected from April to October. Tener-

al adults were collected in April in New York, in May in Ontario, in July in Pennsylvania,

and in September in Wisconsin. Overwintering is probably in the adult stage.

Distribution. — (Fig. 391). The range of this species extends from New Brunswick and

Virginia west to Utah, Colorado, and New Mexico. I have seen 864 specimens from the

following localities:

CANADA

NEW BRUNSWICK: (Saint John) UWMW. ONTARIO: (Belleville) UASM; (Erindale) CAS; (Lake Abitibi, Low Bush) CAS;

(Rogue Hills) CAS; (Summerville) CAS; (Toronto) CAS; (Willowdale) CAS. QUEBEC: (Becancour) CCh; (Como) MCZ;

(Perrot Isle) CMPP; (Vaudreuil) CAS.

UNITED STATES

ARKANSAS: Washington County (No locality given) UAFA. COLORADO: Boulder County (Viele Lake) CArm; Denver

County (Denver) ISNH. ILLINOIS: Cook County (Chicago) WSUP, (Des Plaines) CNHM, (Willow Springs) UMAH; Kendall

County (Oswego) CNHM; Lake County (Volo) RTBe; La Salle County (No locality given) RTBe; McHenry County (Algon-

quin) ISNH; Putnam County (No locality given) ISNH. INDIANA: CNHM, Lake County (Hessville) CMPP. IOWA: Buch-

anan County (Independence) MCZ; Henry County (Mount Pleasant) CAS; Howard County (Elma) AMNH; Johnson County

(Iowa City) MCZ, USNM, (Solon) USNM; Story County (Ames) ISUA. KANSAS: Riley County (No locality given) KSU;

Shawnee County (Topeka) KSU. MARYLAND: (No locality given) MCZ. MASSACHUSETTS: Hampden County (Chico-

pee) MCZ; Middlesex County (Cambridge) MCZ, (Concord) MCZ, (Sudbury) MCZ, (Wayland) MCZ. MICHIGAN: Barry

County (Otis Lake) TFH1; Charlevoix County (Garden Island) UMAH; Cheboygan County (No locality given) UMAH;

Gratiot County (No locality given) UMAH; Kent County (Grand Rapids) UNLN; Saint Claire County (Port Huron)

USNM; Washtenaw County (Ann Arbor) UMAH, (Whitmore Lake) JSch; Wayne County (Detroit) MCZ; County unknown

(Aurelius) OSUS. MINNESOTA: Clearwater County (De Soto Lake, Itasca State Park) UM*SP; Douglas County (Alexandria)

PSUU; Hennepin County (No locality given) UMSP; Houston County (No locality given) UMSP; Mille Lacs County (Gar-

rison) UMSP, (Mille Lacs Lake, near Garrison) UMSP; Olmstead County (No locality given) UMSP; Otter Tail County

(Battle Lake) UMSP; Pine County (Snake River, 4.0 miles east of Pine City) UMSP; Ramsey County (No locality given)

UMSP; Red Lake County (Plummer) UMSP; Washington County (No locality given) UMSP; County unknown (Vineland)

UMSP. MISSOURI: (No locality given) ISNH. NEBRASKA: Dodge County (Fremont) UNLN; Lancaster County (Lincoln)

UNLN; Saunders County (Ashland) UNLN. NEW JERSEY: Gloucester County (Woodbury) USNM; Hunterdon County

(Hampton) AMNH; Middlesex County (New Brunswick) AMNH; Passaic County (Passaic) AMNH; Somerset County (No

locality given) UMAH; Sussex County (Hopatcong) AMNH; Union County (Elizabeth) AMNH; County unknown (Snake

Hill) AMNH. NEW MEXICO: Bernalillo County (Albuquerque) USNM. NEW YORK: Albany County (Altamont) CNHM,

CUNY ; Cayuga County (Cayuga) MCZ, (Fair Haven) LACM; Clinton County (Plattsburg) WSUP; Erie County (East Aurora)

CUNY, (Hamburg) CAS; Greene County (Catskill) USNM; Jefferson County (Cape Vincent) CUNY; Monroe County

(Rochester) LACM, MCZ; New York County (Bronx Park) CAS, (New York City) MCZ; Niagara County (Olcott) AUAA,

CUNY; Ontario County (Fish Creek, near Victor) UASM, (Geneva) UMAH; Suffolk County (Babylon) AMNH; Schuyler

County (Cayuta Lake) UASM, (Watkins Glen) AMNH, MCZ, VMKi; Tompkins County (Buttermilk Falls) PUM, (Ithaca)

AUAA, CAS, CUNY, ISNH, KSU, LACM, MCZ, OUCO, UASM, UCR, UIMI, UNLN, (Ludlowville) CUNY, (Varna) UASM;

Westchester County (Peekskill) MCZ; Wayne County (No locality given) UASM; County unknown (Enfield Falls)

CNC, (Taughannock Falls) CUNY. OHIO: Delaware County (No locality given) PUM; Franklin County (Columbus)

OUCO, PUM; Pickaway County (No locality given) SJSC; Putnam County (Auglaize River) PUM; Ottawa County

(Bass Island) PUM; Williams County (Saint Joseph River) PUM. PENNSYLVANIA: Allegheny County (Pittsburgh)

Bombardier Beetles

147

CMPP, OSUS; Bradford County (Susquehanna River, Wyalusing) GRNo; Crawford County (Conneaut Lake) CMPP;

Cumberland County (Camp Hill) CUNY; Dauphin County (Harrisburg) VMKi; Erie County CMPP, (Erie) CUNY;

Franklin County (Chambersburg) USNM; Montgomery County (Areola) ANSP, OUCO; Northampton County (Easton)

CAS, (Delaware Water Gap) AMNH; Pike County (Milford) SDNHM; Westmoreland County (Jeanette) CMPP; County

unknown (Edge Hill) USNM, (Lehigh Water Gap) USNM. UTAH: (No locality given) ISUA. VERMONT: Bennington

County (No locality given) MCZ; Chittenden County (East Georgia) RTBe, (Grand Isle) MCZ. VIRGINIA: Fairfax County

(Great Falls) USNM; Loudoun County (No locality given) ANSP, MCZ. WEST VIRGINIA: County unknown (Brush Creek)

CUNY. WISCONSIN: Dane County (No locality given) UWMW; Green County (Brodhead) UMAH; Winnebago County

(Oshkosh) CAS.

Brachinus cyanochroaticus Erwin

(Figs. 375, 380, 381, 382, 388, 392)

Brachinus cyanochroaticus Erwin, 1969: 283 . The holotype male and allotype female are

in MCZ. Type locality. - Eleven miles west of York, Benson County, North Dakota.

Diagnostic combination. — The diagnostic characteristics are given in the key.

Description. - Medium-sized beetles, 7.0 to 1 1.3 mm.

Color. Antennal articles 3 and 4 apically, metepisterna, metasternum at sides, and abdom-

inal sterna and terga infuscated, otherwise ferrugineous. Dorsal surface and epipleura of

elytra blue, usually with metallic luster.

Microsculpture. As described for genus.

Macrosculpture. Frontal furrows rugose and punctate. Disc of pronotum punctate, punc-

tures barely impressed.

Head. Frontal furrows moderately impressed. Antennal scape robust, widest at middle,

otherwise head as in cordicollis.

Prothorax. As in cordicollis , except sides of pronotum (fig. 375) more widely reflexed

and proepipleura glabrous.

Pterothorax. As in cordicollis.

Abdomen. As described for genus.

Genitalia. Male (figs. 380, 381, 382). Median lobe with plane of shaft slightly rotated

from plane of basal bend. Basal bend long. Shaft as in cordicollis. Entire median lobe more

robust than in cordicollis. Virga (figs. 380, 381). Female (fig. 388).

Variation. - The members of this species vary considerably in the shape of the pronotum

and in the color of the elytra. These are both local variations, but generally the color has a

metallic luster in the north and western portions of the range, while in the south and eastern

parts of the range, the color becomes dull blue.

Flight. - The flight of these beetles has not been recorded.

Etymology . — Greek, kyanos, blue; chroaticus, color of the skin; referring to the blue

color of the elytra of these beetles.

Collecting notes. — H. Goulet collected these beetles near old beaver houses in the com-

pany of Platypatrobus lacustris Darlington, and G. E. Ball and R. T. Bell collected them at

the edges of ponds.

Life history. - Members of this species have been collected from April to October. Ten-

eral adults were collected in May in Quebec. At least some adults overwinter.

Distribution. — (Fig. 392). The range of this species extends from British Columbia and

Idaho, east to Massachusetts, and south to Kansas. I have seen 342 specimens from the fol-

lowing localities:

CANADA

BRITISH COLUMBIA: (Salmon Arm) CAS. MANITOBA: (Boissevain, south of Brandon) ZMLS; (Whitewater Lake)

UASM, ZMLS. ONTARIO: (Belleville) CUNY; (Britannia) CAS; (Craigleith, West Collingwood, Georgian Bay) ZMLS;

(De Cew Falls) CNC; (Grand Bend, Lake Huron) ZMLS; (Gravenhurst) AMNH; (Hyde Park Comer) RTBe; (Lake of the

Woods, Clearwater Bay) CNC; (Point Pelee) ZMLS; (Port Credit) ZMLS; (Prince Edward County) CUNY ; (Toronto) CAS;

(Turkey Point, Lake Erie) ZMLS. SASKATCHEWAN: (Regina) ZMLS. QUEEEC: (Dorval) CAS; (Gatineau, 1.0 mile south-

west of Meach Lake) CNC; (Lachine) CAS; (Lac Saint Jean) CAS; (La Trappe) ZMLS; (Montreal) CAS; (Outrem’t) CAS;

(Rigaud) CAS, HGou; (Saint Eustache) CAS; (Saint Rose) CAS; (Venise) CCh.