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HARVARD UNIVERSITY

LIBRARY

OF THE

Museum of Comparative Zoology

C ■ 0. - Q f 7y

Mus. COMP. ZOOL.

. library

LJuaestiones may20,P74

entomological

Harvard

Rsity

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

VOLUME IX

1973

11

CONTENTS

Editorial — On Finality 1

Griffiths - Studies on boreal Agromyzidae (Diptera). III. Phytomyza miners

on Cnidium and Conioselinum (Umbelliferae) 3

Steiner - Solitary wasps from subarctic North America - II. Sphecidae from

the Yukon and Northwest Territories, Canada: Distribution and

ecology 13

Perrault - A taxonomic review of the eastern Nearctic species complex

Pterostichus (Haplocoelus) Adoxus (Coleoptera: Carabidae) 35

Book review 41

Book review 44

Book review 47

Editorial - For Love or Money? 51

Shorthouse — The insect community associated with Rose Galls of

Diplolepis polita (Cynipidae, Hymenoptera) 55

Larson — An annotated list of the Hydroadephaga (Coleoptera: Insecta) of

Manitoba and Minnesota 99

Richards — Biology of Bombus polaris Curtis and B. hyperboreus Schonherr

at Lake Hazen, Northwest Territories (Hymenoptera: Bombini) 115

Editorial — Guess Whose Universe? 159

Thomas — The deer flies (Diptera: Tabanidae: Chrysops ) of Alberta 161

Whitehead - Annotated Key to Platynus, including Mexisphodrus and most

“Colpodes”, so far described from North America including

Mexico (Coleoptera: Carabidae: Agonini) 173

Griffiths - Studies on boreal Agromyzidae (Diptera). IV. Phytomyza miners on

Angelica, Heracleum, Laserpitium and Pas tinaca (Umbelliferae) 219

Book review 254

Book review 255

Book review 257

Announcement 259

Announcement 260

Book review 261

Book review 263

Noonan — The Anisodactylines (Insecta: Coleoptera: Carabidae: Harpalini):

Classification, Evolution, and Zoogeography

267

INDEX

m

Acacia, 313, 368, 371

Acciavatti, R. E. (see Kurczewski, F. E.),

28, 33

Achillea, 26

millefolium, 16, 20, 29

Acilius, 110

fraternus, 110

mediatus, 110

semisulcatus , 110

Aconitum, 232

septentrionale, 143

acridids, 20, 23, 24

Acroceridae, 254

Acrogeniodon, 345

bedeli, 345

Aculeata, 33, 142, 145

Adelges abietis, 97

lariciatus, 96

Adephaga, 423

Agabini, 1 1 3

Agabus, 99, 106-108, 113

ajax, 108

ambiguus, 107

anthracinus, 108

arcticus, 108

bicolor, 107

browni, 1 1 3

canadensis, 107

clavatus (= antennatus), 108

colymbus, 106, 1 13

confinis, 107

congener, 107

discolor, 107

disintegratus, 107

hudsonicus, 113

inf us cat us, 107

inscrip tus, 107

kenaiensis, 108

minnesotensis, 108

nigroaeneus ( = erichsonii) , 1 08

ontarionis, 108

phaeopterus, 107

pseudoconfertus, 108

punctulatus, 106

semipunctatus , 107

seriatus, 106

sharpi (= falli), 107

subfuscatus, 107

triton, 106

Agabus (continued)

velox, 1 13

verus (= clavicornis), 108

Agaporus, 1 13

Agassiz, J. L. R., 321, 419

Agelena labyrinthica, 258

Agoni, 174, 175

Agonini, 173-214, 215

Agonum, 173, 174, 175

baroni, 189

(Platynus) bilime ki, 213, 215, 216

cavatum, 191

chihuahuae, 191

(Hemiplatynus) chihuahae, 191

consulare, 192

curtipenne, 201

decentis group, 174, 175

dominicense, 195

euprepes, 196

harfordi, 204

hypolithos group, 174, 175

infidum, 203

larvale group, 174, 175

leptodes, 198

logicum, 199

lymphaticum, 203

morelosense, 203

ovatulum, 205

perleve, 206

pinalicum, 200

puncticeps group, 174

umbripenne, 213

Agromyza, 240

heraclei, 240

Agromyzid, 220, 242, 243

miners, 3

Agromyzidae, 3-8, 219-253, 254

Agromyziden, 8, 241, 242, 243

Agromyzider, 241

Agromyzinen, 8

Alder, H., 66, 95

Allen, P., 6, 8, 225, 229, 233, 234, 235, 241

Allocinopus, 277, 280, 284-285, 389, 405,

440, 443, 472

angustulus, 285

castaneus, 285

latitarsis, 285

ocularis, 285

sculp ticollis , 284, 285

IV

Allocinopus (continued)

smithi, 285

Alluaud, C., 344, 345

Alnus, 16, 21, 24, 28

tenuifolia, 58

Alpinobombus, 1 15, 1 17, 121, 124, 126,

132, 142

Alysiinae, 242

Alysson, 21, 27

triangulifer , 15, 21, 27

Alyssonini, 27

Amara, 283, 348, 354, 375

Amathusiidae, 47

Amelanchier alni folia, 58

amino acids, 140, 145

Ammophila, 13, 20, 25-26, 30, 32

azteca, 14, 22, 25-26, 31, 33

mediata, 15, 26

strenua, 15, 26

Ammophilini 25-26, 3 1

Amphasia, 281-282, 378-379, 380, 394, 395,

396, 397, 398, 399, 406, 441, 444, 473

fulvicollis, 378, 380

interstitialis, 378, 380, 467, 468, 471

sericeus, 379, 467, 471

Anacolpodes recticollis, 208

Anadaptus, 269, 282, 283, 284, 373-374,

395, 396, 398, 406, 435, 441, 444, 473

Anchomenus, 192, 200, 204

(Platynella) baroni, 189

brullei, 190

cavatus, 191

chevrolati, 203

concisus, 192

consularis, 192

(Plantyus) curtipennis , 201

dominicensis, 195

( Plocodes) guerrerensis, 2 1 4

harfordi, 204

( Platynella) infidus, 203

(Platynella) logicus, 199

(Plocodes) longiceps, 201

lymphaticus, 203

(Platynella) morelosensis, 203

nugax, 203

ovatulus, 205

pinalicus, 200

simplicior, 203

suffectus, 192

Ander, von Kjell, 116, 140

Anderson, R. D., 113

Andrews, H. E., 287, 344, 345, 346, 347, 419

Angelica , 8, 219, 220, 221, 222, 226, 229, 231,

232, 234, 235, 238, 240, 243

archangelica, 220, 231, 234, 240

arguta, 240

atropurpurea, 226

decursiva, 235, 240, 248

genuflexa, 231, 234, 252

kiusiana, 236

lucida, 231, 234, 253

miqueliana, 232

palustris, 221, 239, 240

polyclada, 232, 236

razulii, 235

sylvestris, 229, 231, 234, 235, 239, 240, 252

Anisochirus , 278

alluaudi , 278

Anisodactyli, 276

Anisodactylidae, 276

Anisodactylides, 276

Anisodactylina, 264, 267, 268, 269, 271, 275,

276, 278, 279, 344, 378, 379, 384, 385, 401,

403, 404, 405, 406, 407, 435, 440, 441, 442,

443, 444, 472, 473, 476

contemporary zoogeography of the subtribe,

403-407

evolutionary trends and convergences,

397-399

key to the genera, 279-284

key to the subgenera, 279-284

phylogeny of the genera and subgenera,

388-397

Anisodactylinae, 276

Anisodactylines,

classification, evolution and zoogeography,

267-480

Anisodactylini, 276, 426

Anisodactylitae, 276

Anisodactylites, 276

Anisodactyloid, 388, 389, 390, 392, 393, 394,

395, 397, 399, 435, 466, 472, 473, 476, 478,

479

Anisodactylus , 267, 268, 271, 274, 278, 283,

284, 295, 309, 341, 348-349, 349-351, 352,

355, 356, 374, 375, 376, 377, 382, 396, 398,

401, 402, 404, 405, 406, 424, 425, 426, 435,

441, 443, 473, 475, 480

V

Anisodactylus (continued)

abaculus, 349

aethiops, 369

agricola, 350

alternans, 374, 398

amaroides, 375, 376, 468, 470

amplicollis, 349

anthracinus, 356, 357, 358, 366-368,

369, 398, 403, 414, 416, 417, 446,

449, 450, 464, 475, 480

antoinei, 35 1

arizonae, 320

atricornis, 351

beryllus , 361

binotatus, 277, 278, 350, 351, 468, 470,

471

breviceps, 364

caenus, 375, 376, 470

calif ornicus , 348, 350

carbonarius, 349, 350, 351, 356

consobrinus, 350, 396

convexus, 367

crassus, 368, 369

darlingtoni, 267, 356, 357, 358, 367,

370-371, 403, 414, 415, 416, 446,

450, 456, 463, 475, 480

dejeani, 267, 268, 352, 353, 468, 469,

470

depressus , 307

dilatatus, 366, 367

discoideus, 373, 374, 396, 398

dulcicollis, 355, 356, 357, 359, 360,

361, 362-363, 372, 398, 402, 413,

414, 415, 417, 446, 449, 450, 451,

457, 465, 475, 480

ellipticus , 362, 363

elongatus, 362, 363

furvus, 348, 350

gravidus, 368, 369

haplomus, 357, 363-364, 367, 370,

372, 403, 414, 415, 416, 417, 438,

446, 449, 456, 466, 475, 480

harpaloides, 354, 355, 356, 357, 359

360-361, 363, 372, 398, 402, 414,

415, 416, 417, 446, 449, 450, 457,

458, 464, 475, 480

harrisi, 350

hauseri, 350, 351

heros, 352, 353

Anisodactylus (continued)

hispanus, 351

intermedius , 354, 470

karennius , 350, 351

kempi, 369

kirbyi, 348, 350

laetus, 266, 267, 268, 376, 377, 468, 470

limbatus, 342

loedingi, 349, 350, 351, 468, 470

longicollis, 364

mandschuricus , 349, 354

marginatus, 369

melanopus, 350

men/fa, 356, 358, 365, 367, 368-370, 371,

372, 402, 403, 413, 414, 416, 417, 446,

449, 457, 464, 475, 480

metallescens, 349

modicus, 363

nemorivagus, 350, 351

nigerrimus, 350

nigricornis, 351

nigrita, 350

nivalis, 374, 467

oblongus, 365

obscuripes, 349

obtusicollis, 349

ochropus, 343

opaculus, 354, 355, 356, 357, 358-360,

402, 413, 414, 415, 417, 446, 449, 450,

456, 464,475,480

overlaeti, 343

ovularis, 356, 358, 361, 363, 366, 372, 398,

403, 415, 416, 417, 446, 449, 457, 465,

475.480

paulus, 359

peropacus, 364

picinus, 337

pinguis, 368, 369

pitychrous, 374

poeciloides, 353, 468, 470

porosus, 374, 398

propinquus, 351

pueli, 350

punctatipennis, 267 , 268, 351, 352, 470

rotundangulus , 373, 374, 467, 468, 469

rufus, 292

rusticus, 356, 357, 358, 364-366, 367, 372,

398, 403, 414, 416, 417, 446, 449, 450.

456.465.475.480

VI

Anisodactylus (continued)

sadoensis, 351, 352

sanctaecrucis, 374

sayi, 309

schaubergi, 349

semirubidus, 372

shibatai, 351

signatus, 350, 351

similis, 349, 350

sjostedti, 346

sulcipennis , 369

texanus, 355, 356, 357, 361-362, 402,

414, 415, 416, 446, 449, 450, 465,

475, 480

tricuspidatus , 349, 350, 351

tristis, 365

verticalis, 374, 375, 467, 471

virens, 353

viridescens, 313,314

wolcotti, 369

xanthopus, 340

zabroides, 344

Anisostichus, 282, 338-339, 388, 391, 392,

397, 406, 435, 440, 443, 472

amoenus, 338, 339, 391

laevis, 339, 391,467

octopunctatus , 339, 391

posticus, 339, 391

Anisotarsus, 267, 268, 269, 273, 274, 281,

289, 290, 292, 293, 294, 295-300, 304,

338, 339, 374, 375, 390, 393, 399, 400,

403, 405, 406, 407, 408, 409, 410, 412,

413, 414, 422, 435, 440, 443, 445, 447,

448, 450, 452, 453, 454, 458, 459, 460,

461,472,474, 479

contemporary zoogeography of the

subgenus, 407-409

historical zoogeography of the subgenus,

409-413

key to the north american species,

297-300

phylogeny of the new world species,

399-402

Anthomyiidae, 254

Antoine, M., 419

Aoki, K., 32

Apatelus, 339

Apator (= Agabus) bifarius , 108

aphid, 24, 96

Aphididae, 92

Aphroteniinae, 387, 420

Apidae, 134, 140, 142, 143, 144, 145

Apis arctica, 1 15

mellifera, 257, 258

mellifica (= A. mellifera), 258

Aplocentrus, 283, 375-376, 377, 396, 406,

441, 444, 473

Arachnid, 140

Arachnida, 144

Araneae, 141, 145

Arctagrostis latifolia, 1 17

arctic insects, 32, 33, 140, 143

Arnica alpina, 136

arthropod visual systems, 257-258

Asahina, E., 32

Ashlock, P. D., 382, 419

Ashmead, W. H. 58, 59, 95

Asilidae, 254

Askew, R. R., 62, 63, 71, 83, 90, 95

aspen, 16, 25

Astata, 22-23, 32

nubecula, 14, 20, 21, 22-23, 31

Astatinae, 22-23, 31, 34

Auclair, J. L. 134, 140

Auffenberg, W., 409, 419

Axelrod, D. I. 301, 321, 408, 409, 411, 415,

416,419

Ayres, K. D., (see Kinsey, A. C.), 58, 65, 97

Azetecarpalus, 419

Baker, H. G., 31,32

Ball, G. E., 41-43, 263-264, 270, 355, 378,

387,409,414,419,420

Ballion, E., 351

Barbula icmadophila, 120

Barr, T. C., Jr., 174, 176, 195, 202, 207, 212,

213, 214, 215

Barypina, 387

Basilewsky, P., 278, 287, 337, 338, 341, 342,

343,344, 345,350,405,420

Bassett, H. F., 65, 95

Bates, H. W 188, 189, 190, 191, 192, 193, 194,

195, 196, 197, 198, 199, 200, 201, 202, 203,

204, 205, 206, 207, 208, 209, 210, 21 1, 212,

213, 214, 215, 276, 285, 286, 289, 296, 298,

299, 300, 302, 308, 310, 311,314, 318, 322,

323, 324, 325, 327, 329, 330, 331, 332, 333,

334, 335, 336, 344, 345, 346, 349, 350, 351,

374, 420

Vll

Batrachion, 321, 322

chalconatum, 322

rana , 322

rufipalpum, 322

Batrachium, 321

Bedel, L., 420

bees, 27, 33, 140, 143, 145

beetles, 20

Beiger, M., 229, 231, 234, 235, 237, 238,

241

Belicek, J., 47-48, 261-262

Bembidion, 386

Berland, L., 66, 95

Bertram, G. C. L. 131, 140

Besbicus mirabilis, 88, 96

Be tula, 16, 21

papyri fern, 58

Beutenmuller, W. 59, 65, 95

Bidessini, 1 1 4

Bidessus, 101-102

af finis, 101

flavicollis, 101

granarius, 102

Bird, R. D., 99, 113

Blackburn, T., 289, 290

Blackwelder, R. E., 175, 188, 189, 190,

191, 192, 193, 194, 195, 196, 197, 198,

199, 200, 201, 202, 203, 204, 205, 206,

207, 208, 209, 210, 211, 212, 213, 214,

215

Blair, K. G., 63, 65, 67, 68, 71, 73, 75, 88,

95

Blair, W. F., 409, 420

Blair, W. G.,412, 420

Blanchard, C. E., 203, 215

Blatchley, W. S. 295, 306, 309, 318, 420

blattids, 23

Boheman, C. H., 342

Bolivar y Pieltain, C., 174, 178, 189, 190,

200, 209, 213, 215, 216

(see Barr, T. C., Jr.), 174, 213, 215

Bolivaridius, 173

ovatellus, 195

Bombias, 126, 142

Bombidae, 140

Bombinae, 145

Bombini, 115-157

Bombus, 115, 116, 126, 137, 138, 141,

142, 143, 144

Bombus (continued)

agrorum, 115, 126, 140, 141, 142

alpinus, 116, 144

arcticus, 1 1 5

balteatus, 116, 124, 125, 138

(Alpinobombus) balteatus, 132

fervidus, 144

hyperboreus, 115-157

(Alpinobombus) hyp erb or eus, 146

(Alpinobombus) hyperboreus clydensis, 145

hypnorum, 144

inexpectus, 138, 145

(Bombus) lucorum, 119

medius, 126

(Pyrobombus) melanopygus, 132

(Pyrobombus) mixtus, 132

polaris, 115-157

(Alpinobombus) polaris, 146

(Pyrobombus) sit kensis, 132

(Pyrobombus) sylvicola, 126, 132

Bouche, P. F., 240, 241

Brachinida, 422

Brachinus, 382, 409

Brachycera, 242

Braconidae, 232, 242

Bradycellina, 388

Braendegaard, J., 116, 140

Braschnikow, W. C„ 235, 238, 239, 240, 241

Breed, W. J. (see Elliot, D. H.), 388, 422

Bremidae, 142

Brennan, J. M., 161, 163, 164, 166

Brian, A. D., 121, 127, 132, 134, 137, 140, 141

Brinck, P., 116, 117, 121, 134, 138, 141

Brischke, C. G. A., 233, 235, 241

Britton, E. B., 387, 420

Bromeliads, 313, 334

Broscidae, 419

Broscina, 387

Broscine, 387, 390

Broscini, 386, 387, 419

Broun, T., 284, 420

Brown, W. J.. 99, 113

Brucella abortus, 255

Bruggeman, P. F.,.116, 127, 134, 138, 141

Brulle, G. A., 322, 420

Brundin, L., 270, 382, 383, 386, 387, 420

Bryum, 117, 120

Bugbee, R. E., 59, 68, 69, 95

bugs, 20

Vlll

Buhr, H., 3, 8, 56, 95, 229, 231, 235,

238, 239, 241

Bukatsch, F., 134, 141

bumblebee, 34, 141, 142, 143, 144, 145

honey, 143

Buquet. J. B. L., 352

Burdick, N. A. (see Kurczewski, F. E.), 28,

33

Burgeon, L., 338, 342, 343, 344

Burks, B. D. (see Krombein, K. V.), 24, 26,

33, 59, 97

Burton, J. J. S. (see Pechuman, L. L.),

162, 166

Butler, C. G. (see Free, J. B.), 132, 135,

138, 142

butterflies, 32, 33

Calcidoidea, 75, 97

Callan, E. McC., 65, 66, 67, 75, 95, 96

Callidina, 423

Callier gon giganteum, 1 20

Callimome, 96

Callimomidae, 96

Callirhytis quercussuttoni, 97

Calosoma, 386

Caltagirone, L., 70, 71, 96

Cameron, R. S. (see Johnson, N. E.), 318,

366, 423

Camnula, 24

Campylium arcticum, 117, 120

Carabici, 40, 425

Carabidae, 35-39, 40, 173-214, 215, 216,

217, 263, 267-480

Carabiques, 216

Caraboidea, 40

Carabus binotatus, 348, 349

etruscus, 380

germanus, 381

Carboniferous, 386

Cardamine pratensis , 1 1 7

Carex, 58, 318

aquatilis , 120

aquatilis var. stans, 1 1 7

Carpenter, G. D., 116, 141

Carret, A., 380, 381, 420

Casey, T. L., 40, 189, 192, 199, 201, 203,

216, 295, 305, 306, 307, 311, 312, 314,

317, 320, 349, 350, 355, 358, 359, 360,

362, 363, 364, 365, 366, 367, 369, 372,

373,374,379, 420, 421

Cassiope tetragona, 136, 156

Castelnau (de Laporte), F. L. N. C., 286, 287,

288,289, 290, 296, 421

Cataglyphis bicolor, 257, 258

Caucalis, 238

cecidology, 55, 96

Cecidomyiidae, 56

Celtis, 98, 321

Cenogmus, 280, 287-288, 389, 390, 405, 435,

440, 443, 472

castelnaui, 287, 288, 466, 467, 468, 471,

472

interioris, 288

opacipennis, 288

Cephalogyna, 349, 350

loedingi, 350

Cerastium, 136

beeringianum , 117

Ceratina, 34

Cercerini, 27-28, 32, 34

Cerceris, 20, 27-28, 34

nigrescens, 20, 28, 31

nigrescens nigrescens, 15, 21, 27-28

Ceroptres, 62, 88

chalcid-flies, 96

chalcidoid Hymenoptera, 94, 98

chalcis fly, 98

Char a, 104

Chaudoir, M. de., 188, 189, 190, 191, 192, 193,

194, 195, 196, 197, 198, 199, 200, 201, 202,

203, 204, 205, 206, 207, 208, 209, 210, 211,

212, 213, 214, 216, 288, 289, 290, 295, 296,

298, 299, 300, 307, 321, 322, 337, 338, 342,

345,346,357, 362,363,421

Chernov, Y. I., 116, 141

Chevrolat, L. A. A., 216, 289, 290, 321, 322, 421

Chironomid, 387

Chironomidae, 254

Chorthippus curtipennis, 24

Chlorochroa uhleri, 22

chrysidid wasps, 21, 22

Chrysopa vulgaris, 257

Chrysops, 161-171

aestuans, 161, 165, 168, 170

ater, 161-162, 164, 165, 170

callidus, 161

carbonarius, 161, 162

carbonarius nubiapex, 161

discalis, 161, 162, 165, 168, 170

IX

X

XI

Colpodes (continued)

teter, 2 1 2

transfuga, 212

transversicollis, 213

tristis, 201

trujilloi, 196, 197

unilobatus, 213

valens, 213

validus, 213

variabilis, 214

versicolor , 194

violaceipennis, 214

Colymbetes, 109-110, 114

dahuricus, 110

dolobratus, 110

longulus, 109, 110

rugipennis, 1 1 0

sculptilis, 110

Colymbetinae, 106-110

Compositae, 8, 242

Conioselinum, 3-8, 242

chinense, 3, 4, 6, 7, 8, 1 1

tataricum, 4

Coptotomus, 109

interrogatus, 109

Coquerel, J. C, 191, 216

Corbet, P. S., 31,32, 117, 118, 121,

127, 128, 131, 133, 141, 149

(see Oliver, D. R.), 117, 144

Cornus stolonifera, 58

Cosens, A., 55, 61, 66, 96

Crabro, 15,28,30, 33

advenus, 33

latipes, 15, 21, 28, 33

Crabroninae, 13, 20, 21, 28-30, 31, 33

crabronine, 20, 21, 22, 31

Crabronini, 28-30

Crasodactylus , 280, 286-287, 389, 390,

404, 405, 435, 440, 443, 472, 476

indicus, 286, 287, 467

punctatus, 286, 287, 405

Creobina, 387

Cretaceous, 387, 388, 389, 391, 392,

393, 394, 395, 396, 397, 402, 413,

415,476, 477

Criniventer, 280, 292-293, 390, 399,

406, 435, 440, 443, 472

rufus, 292

Crossocerus, 15, 22, 28

Crowson, R. A., 388, 421

Cruciferae, 137

Cry sis (Pent aery sis) shanghaiensis, 32

Csiki, E., 188, 189, 190, 191, 192, 193, 194,

195, 196, 197, 198, 199, 200, 201, 202,

203, 204, 205, 206, 207, 208, 209, 210,

211, 212, 213, 214, 216, 274, 285, 286,

287, 288, 289, 290, 296, 322, 350, 353,

374, 421

Cullumanobombus , 126, 142

Cumber, R. A., 1 19, 122, 126, 132, 133, 141

Cumming, M. E. P., 56, 96

Curculionidae , 318

Curtis, J., 338, 339, 421

Cybister, 1 1 1

fimbriolatus , 1 1 1

Cybisterinae, 1 1 1

Cychrini, 419

Cyclorrhapha, 382, 422

cynipid, 56, 95, 96

galls, 93, 94, 98

wasps, 98

Cynipidae, 55-94, 95, 96, 97, 98

Cynipoidea, 74, 95, 96, 98

Cynips divisa, 92

kollari, 68, 88, 95

Cyrtolaus, 175

D’Abrera, B., 47

Dacnusa fuscipes, 232

Dalla Torre, W. K., 58, 96

Danaidae, 47

Daptini, 378

Daptus incrassatus, 377

Darlington, P. J. Jr., 196, 216, 263, 270, 290,

296, 346, 382, 383, 384, 386, 388, 403,

404, 405,409,413,421

Darwin, C., 384, 421

Daucus car ota, 29

Day, J. H., 117, 141

deer flies, 161-171

Dejean, P. F. M. A., 40, 191, 199, 201, 202,

203, 216, 289, 290, 295, 296, 298, 299,

300, 308, 313, 320, 322, 339, 340, 342,

343, 347, 348, 349, 350, 351, 353, 354,

358, 365, 366, 373, 374, 377, 378, 380,

381,382, 422

delphacid, 25

Delphinium, 232

Desmopachria, 101

Xll

Desmopachria (continued)

convexa, 101

Diabasis, 166

Diachromus, 277, 279, 380, 381, 397,

398, 404, 405, 435, 441, 444, 473

germanus, 381, 468, 471

Diaphoromerus, 289, 295, 296, 393

iridipennis, 295

species group, 296

Diatypus, 280, 293, 294, 295, 337-338,

339, 391, 392, 397, 399, 435, 440,

443, 472

Dichaetochilus, 341, 342

jeanneli, 267 , 268, 342

Dicheirus, 269, 277, 281, 309, 381-382,

395, 396, 397, 398, 406, 425, 435,

441 , 444, 473

brunneus, 382

dilatatus, 382

dilatatus angulatus, 276, 277, 341, 382

obtusus, 382, 398

piceus, 277, 309, 382

strenuus, 382

Dichiropsis, 337, 338

Dicranoncus, 216

Dietz, R. S., 384, 388, 389, 392, 393,

394, 395, 409, 413, 422, 476, 477,

478, 479

Digby, P. S. B., 31, 32, 141

digger wasps, 32, 33

Dineutus, 1 1 1

assimilis, 1 1 1

discolor, 1 1 1

horni, 1 1 1

nigrior, 1 1 1

Dinkelman, M. G., (see Malfait, B. T.),

384, 424

Diodontus, 14, 22, 24

Diplolepis, 58, 61, 65, 66, 67, 68, 71, 74

75, 88, 93, 94, 95, 98

bicolor, 57, 59, 68

eglanteria, 61

eglanteriae, 61

ignota, 67

japonica, 65, 66, 87, 88, 92

mayri, 58

multispinosus, 57

occidentalis, 59

polita, 55-94

Diplolepis (continued)

rosae, 58, 61, 65, 66, 67, 71, 75

(= Rhodites) rosae, 97

rosarum, 61

spinosellus, 59

Diploplectron, 22, 32

peglowi, 14, 21, 22

Diptera, 3-8, 29, 144, 161-171, 219-253,

254, 255,257,382, 422

Dipteren, 8, 241

Dipteres, 241

Distichium, 120

capillaceum, 120

Ditrichum flexicoule, 120

Dorf, E., 409, 413, 415, 416, 421

Dorylinae, 255

Doutt, R. L., 63, 96

Downes, J. A., 31, 32, 127, 131, 132, 141

Drepanocladus aduncus, 120

brevifolius, 117

revolvens, 117, 120

Drosophila, 44, 257, 258

melanogaster, 257

Dryas, 117, 119, 120, 121

integrifolia, 136, 156

-Kobresia habitats, 119, 121

Dryophanta erinacei, 98

Dryudella, 21, 23, 31

picta, 14, 21, 23

Dubach, P., 32

Duftschmidt, C. E., 351

Dyscolus, 173

acuminatus, 187

( Ophryodactylus) aequinoctialis , 1 88

anchomenoides , 199

caeruleomarginatus , 190

chalcopterus , 207

cupripennis, 193

cyanea, 190

cyanipennis, 193

nebrioides, 193

nitidus, 202

(Stenocnemus) pallidipes, 205

purpuratus, 207

variabilis, 214

Dytiscidae, 99, 101-1 11, 113, 114

Dytiscus, 110, 114

anxius , 1 10

dauricus, 1 1 0

Xlll

Dytiscus (continued)

fasciventris, 110

harrisi, 1 1 0

hybridus, 1 1 0

parvulus, 1 1 0

sublimbatus (= cordieri), 1 10

verticalis, 110

Eady, R. D., 58, 96

Ectemnius, 15, 21, 29-30, 31

arcuatus, 15, 22, 29

dives, 15, 22, 29

lapidarius, 15, 29

nigrifrons, 15, 21, 29

trifasciatus, 15, 29-30

Egbert, D., (see McCracken, I.), 59, 65, 97

Ekstam, O., 134, 141

Elaphrini, 386

elder, 25

etiology, 97

Elliot, D. H., 388, 422

(see Kitching, J. W.), 388, 423

Elliptoleus, 175

van Emden, F., 290, 291, 292, 293,

295, 296, 301, 306, 308, 310, 312,

317, 320, 322, 338, 339, 374, 422

Empididae, 254

Eocene, 413

Epeiridae, 23

Ephestia kuhniella, 257

Epicauta, 42

Epilachna, 261

Epilobium, 16, 20, 29

angusti folium, 58

latifolium, 136, 137, 156

Equisetum, 120

arvense, 1 17

variegatum, 117

Erechites hieracifolia , 318

Ericaceae, 137

Erichson, W. F., 296, 322, 381, 422

Erigeron canadense, 3 1 0

Eriophorum, 120

scheutzeri, 1 17

triste, 1 1 7

Eriosoma lanigerum, 25

Erwin, T. L., 270, 382, 392, 409, 414,

422,

(see Ball, G. E.), 270,419

Eucalyptus, 301

Euceroptres, 62

Eudichirus, 279, 341, 342, 343, 394, 440, 443,

472

Eulophidae, 62

Eupelmella vesicularis, 62

Eupelmidae, 62

Europhilus, 174, 175

Euryderus, 419

Eurytoma, 68, 69, 70, 71, 95

curt a, 79

flavicrurensa, 68

incerta, 68

longavena, 55, 62, 63, 64, 65, 68-71, 72, 73,

74, 76, 77, 78, 79, 80, 81, 82, 84, 85, 94

monemae, 69

pachy neuron, 68

parva, 71 , 97

robusta , 71

rosae, 70, 71

terrea, 68

Eurytomidae, 55, 62

Eurytomids, 68, 71

Eurytrichus, 295

flebilis, 310, 311

nitidipennis, 305

piceus, 309

Eutrema edwardsii, 1 1 7

Evans, D., 85, 88, 96

Evans, H. E., 21, 22, 23, 24, 26, 27, 28, 29, 30,

32, 33

Evarthrus, 409, 414, 420

Fabricius, J. C., 348, 349, 350, 351, 352, 374

Fagales, 56

Fagan, M. M. (see Rohwer, S. A.), 58, 97

Fairmaire, L., 342

Fall, H. C., 99, 104, 113

Falls, D. F. (see Morgan, W. J.), 384, 425

Felt, E. P., 56, 58, 96

Feronia adoxa, 35, 39

funesta, 201

interfector, 39

lugens, 199

moesta, 201

monacha, 202

opaca, 201

terminata, 295, 313

tristis, 35, 39

Fervidobombus, 126, 142

fireweed, 16, 20, 21

XIV

Flagg, W. (see Scholander, P. F.), 32, 34

flies, 20

Fooden, J., 385, 388, 422

Forbes, S. A., 360, 422

Formica polyctena, 257

de Fourcroy, A. F., 58, 96, 322

Free, J. B., 127, 131, 132, 135, 137,

138, 142

Freitag, R. (see Ball, G. E.), 409, 414, 420

(see Lindroth, C. H.), 39, 40, 316,

317,367, 424

Freuchen, P., 117, 127, 134, 142

Frey, R., 230, 235, 241

Friese, H., 116, 117, 126, 127, 132, 134,

138, 142

Frison, T. H., 117, 121, 127, 134, 142

Frost, S. W., 241

Fullaway, D. T., 59, 67, 96

Fulmek, L., 71, 74, 75,96

Fye, R. E., 132, 142

gall aphids, 96

community, 61-63

flies, 95

makers, 96, 97

wasps, 97

galls, 95, 96, 97, 98

Gary, N. E., 127, 131, 142

Gaumer, G. C. (see Kurczewski, F. E.),

28, 33

Gavriliok, V. A., 127, 131, 134, 142

Geiger, R., 31, 33

Gelechiidae, 26

Geoffroy, E. T., 58, 96

Geometridae, 26

Geomys, 370

Geopinus, 270, 281, 377-378, 396, 406,

435,441,444, 473

incrassatus, 378, 396, 397, 398, 468, 471

Germar, E. F., 289, 296, 358, 368, 422

Gerstaecker, C. E. A., 349

Glossina, 1

glycerol, 32, 34

Glyphomerus , 71

stigma , 55, 62, 63, 64, 70, 71-73, 74,

75, 76, 77, 78, 79, 80, 81, 82, 84, 85,

94

Glyptolenus , 175

Gnathaphanus , 280, 289, 290, 390, 398,

404, 405, 435, 440, 443, 472

Gnathaphanus (continued)

aridus, 290

basilewski, 296

chinensis , 290

chujoi, 290

denisonensis , 290

froggatti, 290

glamorgani, 290

goryi, 290

herbaceus, 290

kansuensis, 290

latus, 290

licinoides, 290

melbournensis , 290

minutus, 290

papuensis, 290

parallelus, 290

philippensis , 290

picipes, 290

pulcher, 290

punctifer, 290

rectangulus, 290

riverinae, 290

sculpturalis , 290

subolivaceus, 290

upolensis, 290

vulneripennis , 289, 290

whitei, 290

Gory, H. L., 290

Gorytes, 27

albosignatus, 15,21,27

Gorytini, 27, 31

Goureau, C., 224, 241

Graham, A., 409, 422

Granovsky, A., (see Ignoffo, C. M.), 92, 96

Grant, V., 134, 142

Graphoderus, 99, 111, 113

fasciatocollis (= fascicollis), 1 1 1

liberus, 1 1 1

manitobensis, 1 1 1

occidentals, 1 1 1

perplexus, 1 1 1

grasshoppers, 20

Griffiths, G. C. D., 3-8, 219-253, 254, 382, 422

Groschke, F., 236, 237, 242

ground-beetles, 40

Guerin-Meneville, F. E., 286, 287, 422

Gymnaetron, 28

antirrhini, 28

XV

Gynandromorphus, 281, 380-381, 397,

398, 404, 435,441,444, 473

etruscus, 381, 468, 471

peyroni, 380, 381

Gynandrotarsus, 267, 268, 269, 270, 273,

283, 296, 351, 354-356, 360, 361, 362,

363, 364, 366, 367, 368, 370, 371, 396,

398, 402, 403, 406, 413, 414, 415, 416,

417, 435, 441, 444, 446, 449, 450, 456,

457, 464, 465, 473, 475, 480

contemporary zoogeography of the

subgenus, 413-415

historical zoogeography of the sub-

genus, 415-417

key to the species, 357-358

phylogeny of the species, 402-403

Gyrinidae, 99, 111-112

Gyrinus, 99, 111-112, 113

aeneolus, 1 1 1

affinis, 1 1 2

analis, 1 12

aquiris, 1 1 2

bifarius, 1 1 2

borealis, 1 1 2

con finis, 112

dichrous, 1 1 1

impressicollis, 112

latilimbus, 1 1 2

lugens, 112

maculiventris, 112

minutus, 1 1 1

opacus, 1 1 2

piceolus, 1 1 3

picipes, 112

pugionis, 1 1 2

ventralis, 1 1 1

wallisi, 112

Habrocytus, 55, 62, 63, 64, 70, 73,

75-76, 77, 78, 79, 80, 81, 82, 84, 85

bedeguaris, 75, 76, 96

medicaginis, 75, 98

periclisti, 75, 96

trypetae, 75

Habronema, 255

Habu, A., 263-264, 290, 345, 346, 350,

351,422

hackberry, 97

Haematobia irritans, 255

Haematopota, 166

Haldeman, S. S., 314, 350, 422

Haliplidae, 99, 100-101

Haliplus, 99, 100, 113

( Liaphlus) apostolicus, 1 00

(s. str.) blanchardi, 100

(Paraliaphlus) borealis, 100

(Liaphlus) canadensis, 100

(Liaphlus) connexus, 100

( Liaphlus) cribrarius, 1 00

(s. str. ) immaculicollis, 100

(s. str. ) longulus , 100

(Paraliaphlus) pantherinus, 100

(s. str. ) strigatus , 100

( Liaphlus) subguttatus, 100

(Paraliaphlus) triopsis, 100

Hallam, A. (see Smith, A. G.), 384, 389, 392,

410,413,426

Hamilton, W. I. (see Marler, P. R.), 127, 144

Haplocentrus, 375

Haplocoelus, 35

Harpali, 263

Harpalina, 264, 278, 388

Harpalinae, 216, 217, 421

Harpaline, 278

Harpalinen, 425, 426

Harpalini, 263, 264, 267-480

Harpalomimetes, 282, 346-347, 395, 404, 440,

443, 473

andrewesi, 346, 347

sjostedti, 346

Harpalus, 263, 264, 278, 287, 288, 381

agilis, 3 1 3

agitabilis, 305

anthracinus, 366

caenus, 375

conspectus, 305

dulcicollis, 362

laevis, 338

lateralis, 288

maculicomis, 307, 308

madagascariensis , 278

merula, 368

mexicanus, 295, 320-321

oblongius cuius , 339

ocreatus, 313-314

patronus, 307

poeciloides, 353

punctilabris , 344

rotundicollis , 287

XVI

Harpalus (continued)

rusticus, 354, 364

sericeus, 379

similis, 313, 317

testaceus, 314, 317

virescens, 308

viridellus, 322

viridulus, 322

wilkensi , 322

Harper, A. M., 57, 96

Harrell, B. E. (see Martin, P. S.), 395,

424

Harris, B. J. (see Kurczewski, F. E.), 24,

28, 33

Harrison, J. W. H., 61, 96

Hartig, F., 235, 236, 237, 242

Hartman, F., 23, 33

Hasselrot, T. B., 121, 126, 127, 132,

133, 134, 142

Hatch, M. H., 113, 175,216

Hayekius, 345

constrictus, 345

Heilprin, A., 403, 404, 422

Hellen, W., 116, 142

Heming, B., 44-46

Hemiplatynus, 192, 213

(Hemiplatynus) chihuahuae, 191

(Stenoplatynus) umbripennis, 213

Hemiptera, 22

Hendel, F., 224, 225, 226, 228, 233,

235, 238, 240, 242

Hendrichs, J. (see Barr, T. C., Jr.), 174,

213, 215

(see Bolivar y Pieltain, C.), 174, 178,

189, 190, 200, 209, 213, 215, 216

Hennig, W., 270, 382, 383, 387, 422

Henriksen, K. L., 116, 138, 142

(see Braendegaard, J.), 116, 140

Heptagyiae, 387, 420

Heracleum, 219, 220, 221, 222, 225,

226, 227, 228, 234, 238, 240

lanatum, 58, 225, 226, 228, 230, 234,

235, 236, 251

mantegazzianum, 238

sphondylium, 220, 224, 225, 226, 227,

238, 253

Hering, E. M., 3, 4, 8, 220, 225, 227, 230,

231, 232, 234, 236, 237, 238, 240, 242

(see Groschke, F.), 236, 237, 242

Hering, M., 3, 8, 228, 230, 231, 233, 236, 237,

238, 239, 242

Hesperiidae, 47

Hexatrichus, 283, 353-354, 396, 398, 404, 405,

435, 441 , 444, 473

Heyne, A., 190

Himmer, A., 132, 134, 142

Hippodamia, 42

Hobbs, G. A., 117, 121, 124, 126, 132, 138,

142, 143

Hock, R. J. (see Scholander, P. F.), 32, 34

Hocking, B., 1-2, 31, 33, 51-53, 127, 131,

134, 135, 137, 143, 159-160

Hodek, I., 261-262

H0eg, O. A., 134, 137, 143

Hoffmeyer, E. B., 71, 96

Holden, J. C. (see Dietz, R. S.), 384, 388, 389,

392, 393, 394, 395, 409, 413, 422, 476, 477,

478, 479

Holm, A. (see Carpenter, G. D.), 116, 141

Holmen, K., 134, 143

Homoptera, 74

honey bee, 142, 145

components of, 145

guides, 144

Hope, F. W., 290

Hopkins, D. M., 409, 422

Horn, G. H., 276, 374, 382, 423

horse flies, 1 66

Howden, H. F., 408, 409, 412, 414, 423

Huber, L. L., 74, 96

Hudson, J. E., 255-256

Hugel, M. F., 134, 143

Hull. D. L., 382, 423

Hulten, E., 3, 8, 220, 242

humble-bees, 141, 143, 144, 145

Hurd, P. D. (see Baker, H. G.), 31, 32

Hydaticinae, 110-111

Hydaticus, 99, 110

modes tus, 110, 113

piceus, 110

stagnalis, 1 1 3

Hydroadephaga, 99-1 13

Hydrocanthari, 40, 425

Hydroporinae, 101-106, 114

Hydroporus, 99, 103-106, 113

(s. str.) appalachius, 105

(s. str.) arcticus, 104

(s. str.) badiellus, 105

XVII

Hydroporus (continued)

(Heterosternus) clypealis, 103

(s. str.) columbianus, 104

consimilis, 103

(s. str.) dentellus, 104

( s . str.) despectus, 105

(s. str.) dichrous, 104

(Oreodytes) duodecimlineatus , 106

(s. str.) fuscipennis, 105

(s. str. ) glabriusculus, 105

( Deronectes) griseostriatus, 1 06

laevis, 106

lapponum, 113

(s. str.) melancephalus, 105

(s. str.) melsheimeri, 104

(s. str. ) niger, 104

(s. str.) notabilis, 104

(s. str.) obscurus, 105

occidentalism 105

(Heterosternus) paugus, 104

(s. str.) pervicinus, 105

(Heterosternus) planiusculus , 104

(s. str.) rectus, 105

(Deronectes) rotundatus (= elegans), 106

(s. str.) rufinasus, 106

( Oreodytes) scitulus, 1 06

(Heterosternus) sericeus (= superioris),

104

(s. str.) signatus, 105

( Heterosternus) solitarius, 1 04

(He ter os tern us) s tagnalis ,104

(Deronectes) striatellus, 106

(s. str.) striola, 105

(s. str.) tartaricus, 105

(s. str.) tenebrosus, 105

(s. str.) tristis, 106

(Heterosternus) undulatus, 103

( He ter os tern us) vi ttatus, 103

Hydrovatus, 101

pustulatus, 101

Hygrotus, 99, 102-103

acaroides, 102

canadensis, 102

compar, 102

den tiger, 103

dispar, 102

farctus, 102

impressopunctatus, 103

masculinus, 103

Hygrotus (continued)

nubilus, 102

patruelis, 102

punctatus (= sayi), 102

punctilineatus, 102, 103

salinarius, 103

sellatus, 102

suturalis, 102

tumidiventris, 103

turbidus, 102

unguicularis , 103

Hylemya cilicrura, 30

Hymenoptera, 32, 33, 34, 42, 55-94, 95, 96,

97, 98, 115-157, 232, 257

Hyperaspis, 42

Hyperodes, 3 1 8

delumbis, 28

Hypharpax, 281, 288-289, 389, 390, 404, 405,

435, 440, 443, 472

abstrusus, 289

aerus, 289

antarticus, 289

australis, 289

bostocki, 289

celeb ensis, 289

dampieri, 289

dentipes, 288, 289

deyrollei, 289

flavit arsis, 289

flindersi, 289

habitans, 289

inornatus, 289

interioris, 289

kingi, 289

krefti, 289

moestus, 289

nitens, 289

obsoletus, 289

opacipennis, 289

peroni, 289

puncticollis , 289

queenslandicus , 289

ranula, 289

rotundipennis, 289

sculp turalis , 289

simplicipes, 289

sloanei, 289

varus, 289

vilis, 289

XV111

Hypocrabro chrysargirus, 29

trifasciatus , 30

Hyponomer, 244

Ignoffo, C. M., 92, 96

Ilybius, 99, 108-109, 114

angustior , 108

bigut tulus, 109

discedens, 109

fraterculus, 109

pleuriticus, 108

subaeneus, 109

inquilines, 96

Insect-flower associations, 143

relations, 33

insect galls, 96

biology of, 96

morphology of, 96

Irving, L. (see Scholander, P. F.), 32, 34

Ishida, H. (see Nakane, T.), 290

Ithytolus , 175

Iwosiopelus masaudai, 290

jackpine, 16

Jackson, C. I., 128, 143

Jacobson, G. G., 1 17, 127, 134, 138,

143, 276, 423

Jacquelin du Val, P. N. C., 276, 423

Jamieson, C. A. (see Auclair, J. L.), 134,

140

Jeannel, R., 276, 278, 337, 340, 342, 343,

345, 350, 351, 353, 354, 386, 423

Jedlicka, A., 340, 344, 345, 346, 349,

423

Jensen, J. A. (see Elliot, D. H.), 388, 422

Johansen, F., 1 17, 121, 127, 134, 143

Johnson, N. E., 318, 366, 423

Juncus albescens, 1 1 7

biglumis, 117

castaneus, 1 17

Jurassic, 387, 388, 389, 392, 476, 477

Kalaplasmic galls, 56

Kaltenbach, J. H., 224, 233, 242

Kareya, 344

Karl, O., 235, 242

Kevan, P. G., 31, 33, 137. 143

Kieffer, J. J. (see Dalla Torre, W. K.), 58,

96

King, P. G. 409, 423

Kinsey, A. C., 56, 57, 58, 65, 66, 96, 97

Kitching, J. W., 388, 423

Klug, J. C. F., 338, 342

knapweed gall-fly, 79, 98

Knee, W. J., 135, 143

Knuth, P., 134, 143

Kock, A. (see Schwartz, I.), 134, 145

Kolbe, H. J., 342

Krombein, K. V., 23, 24, 26, 27, 30, 33, 59, 97

(see Muesebeck, C. F. W.), 21, 22, 23, 24, 25,

26, 27, 28, 29, 30, 34, 59, 62, 67, 97

Kubska, J., 229, 235, 242

Kuiken, K. A. (see Weaver, N.), 134, 145

Kuntzen, H., 343

Kurczewski, E. J. (see Kurczewski, F. E.), 24,

25, 29, 33

Kurczewski, F. E., 23, 24, 25, 28, 29, 33

Kuroda, M., 240, 243

Kuschel, G., 403, 404, 406, 423

Kiister, E., 56, 97

Kuster, J. E., 257-258

Kuznetzov-Ugamskij, N. N., 65, 66, 97

LaBerge, W. E. (see Michener, C. D.), 126, 144

Labyrinthodont amphibians, 388

Laccophilinae, 101

Laccophilus, 101

maculosus, 101

inconspicuus (= biguttatus), 101

Laccornis, 106

conoideus, 106

Lacordaire, J. T., 276, 423

La Ferte-Senectere, F. T., 349, 354, 355, 357,

360, 362, 423

Laporte, F. L. de., 193, 216

La Roi, G., 57, 97

Larrinae, 21 , 23, 33

Larson, D. J., 99-113,409,423

Larson, R. I., 51

Laser, 238

Laserpitium, 219, 220, 222, 234, 235, 238

latifolium234, 236, 237

leafhoppers, 20

Leclercq, J., 29, 30, 33

LeConte, J. L., 40, 192, 198,211,216, 298,

299, 305, 309, 310, 347, 350, 354, 355, 357,

358, 362, 368, 374, 375, 376, 377, 382, 423,

424

Ledum, 61

groenlandicum , 58

Leech, H. B., 99, 113

Leguminosae, 137

XIX

Leng, C. W., 200, 209, 216, 306, 424

Leopold, A. S., 413, 424

Leopold, E. B. (see Wolfe, J. A.), 410, 427

Lepidoptera, 42, 47, 145

lepidopterous caterpillars, 20

Lequillon, 290

Lestica, 30

producticollis, 15, 30

Lestiphorus cockerelli, 15, 21, 27

Lewis, W. H., 61,94, 97

Libanotis, 238

Libytheidae, 47

Lindner, E., 254

Lineus, 2

Linnaeus, C., 381

Linnaniemi, W. M., 235, 243

Liodessus, 101

Liquidamber forest, 334

Liriomyza, 220

lutea, 220

strigata, 220

wachtli , 220

L0ken, A., 117, 121, 127, 134, 143

Long, F. L. (see Clements, F. E.), 134,

141

Longstaff, T. G., 127, 134, 143

Loricerini, 4 1 9

Losina-Losinsky, L. K., 32, 33, 34

Louwerens, 290, 296

Lubliner-Mianowska, K., 134, 143

Lundqvist, A., 235, 238, 243

Lutshnik, V., 278, 354, 424

Lycaenidae, 47

lygaeids, 22

Lyneborg, L. (see Ryden, N.), 229,

240, 243

Lyon, R. J., 62, 97

Lystrosaurus, 423

MacGinitie, H. D., 409, 424

MacLeay, W. J., 289, 290, 296, 424

MacLeay, W. S., 288, 290, 344, 424

Malfait, B. T., 384, 424

Malyshev, S. I., 56, 62,71,97

Mani, M. S., 56, 57, 62, 65, 79, 92, 97

Mannerheim, C. G. von. 216, 381, 424

Manning, A., 135, 144

Manning, S. A., 229, 235, 243

mantids, 23

Marler, P. R., 129, 144

Martin, J. E. H. (see Mosquin, T.), 1 16, 134,

144

Martin, P. S., 395, 409, 411,412, 416, 424

Maslin, T. P., 383, 424

Matthew, W. D., 384, 424

Mayr, E., 270, 382, 425

Mecoptera, 257

Medler, J. T., 119, 122, 144

(see Fye, R. E.), 132, 142

(see Knee, W. J.), 135, 143

Megabombus hyperboreus, 144

Mehringer, P. J. (see Martin, P. S.), 409, 411,

416, 424

Meijere, J. C. H. de., 4, 8, 225, 228, 229, 230,

231, 233, 234, 235, 237, 238, 239, 240, 243

Melanagromyza, 220

Melandrium affine, 136

Melanoplus, 24

Meloe, 42

Menetries, E., 382

Mesembrina mystacea, 254

Mesozoic, 387, 388

Mexisphodrus , 173-214, 215

profundus, 207

tlamayensis, 207

veraecrucis, 214

Michalska, Z., 229, 235, 243

Michener, C. D., 126, 144

Microsarus, 289, 290

insularis, 289

melbournensis , 289

Microtus ochrogaster , 372

midges, 382, 387

Migadopini, 385, 386

Milliron, H. E., 117, 121, 127, 134, 138, 139,

144

Milstead, W. W. (see Auffenberg, W.), 409, 419

Mimesa, 21, 24-25

clypeata, 21, 25, 31

pauper, 14, 21, 24-25

Mimumesa, 14, 25

XX

Mimumesa (continued)

clypeata, 14

Miocene, 392, 411, 413

Miscophini, 23

Miscophus, 23, 33

americanus, 14, 21, 23, 31

mites, 145

Monroe, E., 31, 34

Moore, I., 313, 425

Morawitz, A. V., 351, 352

Mordwilkoja vagabunda, 92, 96

Moreau, R. E., 394, 406, 425

Morgan, W. J., 384, 425

Morphogynandrus , 380, 381

Moser, J. C., 71, 74, 97

Mosquin, T., 116, 134, 144

moths, 26

Motschoulsky, V. von., 194, 217, 374

Muesebeck, C. F. W., 21, 22, 23, 24, 25,

26, 27, 28, 29, 30, 34, 59, 62, 67, 97

Muller, G., 340, 342, 425

Mulsant, E., 339

Murray, A., 337, 338, 425

Musca autumnalis, 28

domestica, 28, 29, 30, 257

Muscidae, 254, 255

Mutchler, A. J. (see Leng, C. W.), 200,

209, 216

McAlpine, J. F., 116, 134, 137, 138,

144

McCracken, I., 59, 65, 97

McLachlan, R., 134, 144

M’Nary, J. (see Cockerell, T. D. A.),

134, 141

Nakane, T., 290

Napomyza carotae, 220

nectar, 141. 143, 144. 145

Nelson, G. L., 382, 425

Nemaglossa , 279

Neoscutopterus, 109

angustus, 109

horni, 109

Newman, E., 40, 378, 380, 425

Newport, G., 132, 144

Niblett, M., 61, 65, 70, 71, 76, 88, 89,

97

Nielsen, B. O. (see Ryden, N.), 229, 240,

243

Neilsen, E. T., 132, 134, 144

Neilsen, I. C. (see Johansen, F.), 117, 121, 127,

134, 143

Nitelopterus, 33

Noonan, G. R., 267-480

Norway spruce gall, 97

Nothiger, R. (see Ursprung, H.), 43

Nothofagus, 413

Notiobia, 267, 268, 269, 270, 273, 281, 291,

293-337, 338, 339, 389, 390, 391, 392, 393,

394, 403, 406, 409, 413, 417, 435, 440,

443, 455, 456, 462, 463, 472, 474, 476,

477, 479

aeneola, 322

aequata, 33 1

agilis, 3 1 7

agitabilis, 306

amethystina, 296, 401, 408, 410, 474

angusticollis , 314,317

angustula, 296

antarticus, 289

aulica, 322

australasiae, 296

bamboutensis, 338

basilewski, 296

bradytoides, 296, 400, 401, 474

brevicollis , 294, 295, 298, 300-301, 302,

368, 399, 401, 410, 41 1, 445, 448, 454,

460, 474, 479

calathoides, 311,312

castaneus, 310,311,312

cephalus, 305, 306

chalcites, 296, 401, 408, 410, 474

championi, 322

chiriquensis , 294, 322

chloroderus, 308

concinna, 322

concolor, 322

connivens, 314, 317

convexulus, 311,312

cooperi, 267, 268, 294, 325-326, 327, 448,

451,455,462

cupreola, 294

cupripennis, 294, 296, 400, 474

cyanippa, 293, 294, 295, 298, 300, 302, 399,

401, 410, 445, 447, 454, 458, 460, 474, 479

cyanippus, 302

delicatus, 305, 306

diffusus, 338

disaparilis, 294, 322

XXI

Notiobia (continued)

dohrni, 337, 338

dubia, 322

elata, 296

elongensis, 338

ewardsi, 296

ewarti, 267, 268, 294, 324, 326-327,

448, 450, 456, 462

extraneus, 311, 312, 313

feanus, 338

flavipalpis, 296

flebilis, 299, 310-311,312, 399, 400,

408, 410, 411, 445, 448, 452, 460,

474, 479

flebilis castaneus, 310

flebilis flebilis, 310, 311

flebilis purpurascens, 310, 311

flindersi , 289

floridanus , 314, 317

foveicollis, 314, 317, 318

fuscipennis, 314, 317

germari, 296

hebes, 314, 317

hilariola, 293, 295, 298, 302-303,

399, 410, 445, 447, 454, 460, 474,

479

inaequalipennis, 296

inaudax , 311, 312

incerta, 294, 322

innerans, 314, 317

iridipennis, 295, 296

jucunda, 322

key to the subgenus, 323-325

kivuensis, 338

laeviusculus, 300, 301

lamprota , 298, 303, 318-319, 401,

408, 410, 445, 447, 453, 459, 474,

479

lamprotus, 318-319

lapeyrousei , 296

laticollis, 296

latiusculus, 296

leiroides, 294, 324, 327-329, 334, 448,

451, 454, 462

leonensis, 338

limbipennis, 294, 321-322, 324, 329-

330, 448, 451,455,458, 463

longipennis, 322

ludicollis, 296

Notiobia (continued)

maculicornis , 299, 300, 307-308, 400, 413,

445, 447, 452, 459, 474, 479

margaretae, 296

melaena, 294, 325, 329, 331-332, 448, 451,

455,462

melanara, 296

mexicana, 298, 320-321, 400, 401, 407,

408, 410, 41 1 , 445, 447, 452, 458,

459, 474, 479

nebrioides, 293, 321, 322

nicki, 267, 268, 290, 291, 292

nigrans, 296

nitidipennis, 298, 299, 304, 305-306, 315,

399, 400, 410, 413, 445, 448, 450, 452,

459, 474, 479

oblongiuscula, 296

obscura, 294, 323, 325, 332-333, 448, 451,

455, 463

obscura virens, 332

ocreatus , 316,317

opaca, 296

ovata, 296

pallipes, 325, 333-334, 448, 451, 455,

463

pallipes subaurata, 333-334

papuella, 296

papuensis, 296

parallelus, 314, 317

parilis , 294, 324, 325, 334-335, 428, 463

patronus, 307

patrueloides, 296

perater, 296

peruviana, 296

picea, 298, 309-310, 312, 317, 400, 410,

413, 445, 447, 452, 459, 474, 479

picinus, 338, 469

planiuscula, 296

planoimpressa, 296

polita, 296

porcatula, 296

praeclara, 294, 296, 322

purpurascens, 291 , 299, 31 1-313, 399, 400,

408, 410, 411,413, 445, 447, 452, 474,

479

quadricollis , 296

queenslandica, 296

queenslandicus , 289

rectangula, 296

XXII

Notiobia (continued)

remarks on the phylogeny of the species of

the subgenus, 402

ruficrura, 322

rugosipennis, 296

ruwenzoricus, 338

schlingeri , 267, 268, 300, 303-305,

399, 400, 410, 445, 447, 452, 460,

474, 479

schnusei, 296

sculp tip ennis, 296

sericipennis, 296

similis, 313, 317, 322

sinuessa, 329

smithi, 338

stubeli, 296

subaurata, 322

subovalis, 314, 317

subvirens, 314, 317

tenuitarsis, 305, 306

terminata, 297, 298, 299, 300, 302,

305, 306, 307, 308, 309, 310, 311,

312, 313-318, 320, 399, 400, 407,

408, 412, 413, 445, 447, 448, 450,

452, 461, 474, 479

terminata subvirescens, 3 1 7

terminatus, 310, 313

transversicollis, 322

tucumana, 277, 291, 293, 295, 296,

399, 400, 401,435,474

tucumanus nicki, 29 1

umbrata, 293, 294, 322, 324, 325,

335- 336, 337, 448, 451, 456, 463

umbrifera, 293, 294, 322, 324, 325,

336- 337, 448, 451,455,462

vernicatus, 314, 317

virescens, 298, 299, 300, 308-309,

400, 410, 445, 447, 452, 460, 474,

479

viridellus, 322

viridipennis, 296

viridula, 322

viridulus, 322

wilkensi, 294, 322

wilkensi concolor, 322

wilkensi flavicincta, 322

wilkensi pallipes , 322

zoogeography of the subgenus, 413

Notiobioid, 388, 389, 390, 393, 394, 397,

Notiobioid (continued), 466, 472, 476, 477,

478, 479

Notman, H., 307, 425

Nowakowski, J. T., 222, 225, 228, 231, 232,

235, 238, 240, 243

Nummi, W. O. (see Hobbs, G. A.), 132, 143

Nunberg, M., 235, 240, 243

Nymphalidae, 47

Nysson, 20, 21, 27

lateralis, 15, 27

subtilis, 15, 27

Nyssoninae, 27, 31

Nyssonini, 27

oak galls, 95

marble gall, 95

Oligocene, 392,412,415,416

Oligoxemus, 341, 342

Oliver, D. R., 117, 144

(see Milliron, H. E.), 117, 121, 127, 134,

138, 139, 144

Onychopterygia , 216

Ony pterygia, 175

Ophiomyia, 220

Ophonus constrictus, 345

Ophryodactylus, 173, 214

Ophyra leucostoma, 30

Opuntia, 363

Ormyridae, 62

Ormyrus, 62

Orothecium chryseum, 1 17

Orthoptera, 258

orthopteroid, 23

Orthothecium chryseum , 120

Osmorhiza, 232

aristata, 232

Osten Sacken, C. R. (Baron), 62, 65, 67, 97,

161, 162, 166

Oxybelini, 30

Oxybelus, 20, 30

bipunctatus, 33

quadrinotatus, 30

uniglumis, 31

uniglumis 4-notatus, 1 5

uniglumis quadrinotatus, 21, 30

Pachauchenius , 289, 290

celtidivesicula, 74

laeviceps, 289

philipp ensis, 289

Pachypsylla galls, 98

XX111

Pagonia, 166

Paleocene, 402, 415

Palmen, E. (see Lindroth, C. H.), 275,

276, 424

Pangoniinae, 166

Pantamorus peregrinus, 370

Panzer, G. W. F.,351

Papaver radicatum, 136

Papilionidae, 47

Papp, C. S. (see Swan, L. A.), 41

Paradiatypus, 294, 338, 392, 393, 397,

399, 435

Paraphytomyza, 242

parasite, 96, 97, 98, 145

parasitic cynipoids, 97

Parastomoxys mossambica, 255

Park, O. W., 135, 144

Parry, D. A., 31, 34

Pastinaca, 219, 220, 221, 223, 225, 226,

238

saliva, 224, 225, 226, 240

pea-galls, 97

Pechuman, L. L., 162, 165, 166

Peck, O., 68, 71,74, 75, 97

Pedicularis arctica, 117, 136, 156

capitata, 136, 156

hirsuta, 117, 136

Pelmatellina, 279

Pelmatelline, 279

Pelmatellus, 279

Peltodytes, 100-101

edentulus, 100

tortulosus, 101

Pemphredon, 25

bipartior, 14, 21, 25, 31

montana, 14, 25

Pemphredoninae, 13, 20, 21, 24-25, 31

pemphredonine, 21, 22, 31

Pemphredonini, 25

pentatomid, 22

Periclistus, 62, 67, 71, 75, 88, 93

brand ti, 75

brand tii, 71

calif ornicus , 67

piceus, 67

pirata, 55, 62, 63, 64, 67-68, 69,

70, 71, 72, 73, 74, 75, 76, 77, 78,

79, 80, 81, 82, 84, 85, 86, 87, 88-90,

92, 93

Perigonini, 42 1

Peringuey, L., 276, 342, 425

Permian, 388

Perrault, G. G„ 35-40

Perty, J. A. M., 293, 321, 322, 425

Petasites, 8, 242

Peucedanum, 238

Phanagnathus , 343-344, 394, 435, 440, 443,

473

overlaeti, 277 , 344

Philanthinae, 27-28

Philip, C. B., 161, 162, 163, 166

Phillips, W. J., 70, 71, 97

Philophuga, 423

Phylloxeridae, 96

Phytagromyza, 242

Phytomyza, 3-8, 219-253

aconiti, 232, 233

albiceps group, 3, 4, 219, 222-232, 233

angelicae, 5, 8, 10, 11, 219, 221, 222, 232,

233-235, 236, 237, 238, 239, 248, 250,

253

angelicae group, 219, 232-240

angelicae kibunensis, 235, 236

angelicastri , 221, 222, 228-230, 240, 247,

250, 252

angelicella, 219, 226

angelicivora, 221, 232, 233, 238-240, 250

aralivora, 4, 220

archangelicae, 219, 220, 221, 222, 230-232,

247, 250, 252

arnaudi , 221, 222, 232

bifida, 235

cicutae, 222

cnidii, 3, 4, 5-6, 9

conii , 222

conioselini, 3, 4, 5, 7-8, 9, 10, 220

heracleana, 221, 222, 232, 233, 236, 237-

238, 249, 253

heraclei, 224, 225

heracleiphaga, 219, 225

key, 4-5. 220-222

kibunensis, 219, 221, 232, 233, 236, 248

lanati, 222, 225, 230, 244

laserpitii, 219, 233, 234, 235

latifolii, 222, 232, 233, 236-237, 249

mertensiae, 4

milii, 242

mylini, 4

XXIV

Phytomyza (continued)

nepetae, 4

nigra, 224

nilssoni, 230, 23 1

obscurella, 6, 239

oenanthes , 222

osmorhizae, 4, 220

pastinacae, 219, 220, 221, 222, 223,

225, 226, 240, 245

pauliloewi, 232, 233

polycladae, 221, 222, 232

Ringdahli, 243

riparia , 220

sehgali, 4

selirii, 232, 233, 240

sk, 222

ji/fli, 232, 233, 240

sitchensis, 3, 4, 5, 6, 7, 8, 9, 10, 11, 220

sphondyliivora, 221, 222, 227, 246

spondylii, 219, 222-223, 224, 226,

227, 228, 229, 230, 240

spondylii heracleiphaga, 220, 222, 224,

225, 230, 245, 251

spondylii spondylii, 221, 224-225

tlingitica, 219, 220, 221, 222, 227-

228, 246, 250, 251

Phytomyzinae, 243

Picea glauca, 58

Pieridae, 47

Pimpinella, 238

Pinus sylvestri, 366, 370

Plath, O. E., 132, 135, 137, 144

Platymetopus flavilabris, 264

Platynella, 173, 174, 189, 199, 203,

214, 215

Platynidius, 175

Platyninae, 216

Platynus, 173-214, 216

acuminatus, 176, 187-188, 191

acutulus , 187, 188

aeneicauda, 192, 198

(Trapezodera) aeneicauda, 182, 188

aequinoctialis , 186, 188, 193, 211

agilis, 173, 174, 176, 188, 197, 198

amplicollis, 184, 188-189

anchomenoides, 189

angulosus, 185, 189

aphaedrus, 177, 189

approximatus, 189

Platynus (continued)

araizai, 178

( Rhadine) araizi, 1 89

atratus, 173, 174, 189

baroni, 181, 189

bicolor, 173, 174, 189, 192

bilimeki, 189

biovatus, 181, 189-190, 210

boneti, 178

(Rhadine) boneti, 190

brachyderus, 187, 190

brullei, 179, 190

brunneomarginatus , 192

brunnipennis, 190, 199

caeruleomarginatus , 187, 190

caeruleus, 111, 190-191

cavatus, 176, 191

chalcopterus , 191

championi, 173, 174, 180, 191, 195, 207

chaudoiri, 183, 191, 210

chevrolati, 191

chihuahuae, 180, 191

(Hemiplatynus) chihuahuae, 191-192

chloreus , 173, 174, 192, 202

colibor , 173, 174, 183, 189, 192, 209

columbinus, 111, 178, 192

concisus, 173, 174, 186, 189, 192, 206,

211

conicicollis, 111, 182, 192

consularis, 185, 192-193, 195

convexulus, 111, 192

cordatus, 186, 193

crossomerus, 193

cupripennis, 173, 174, 184, 188, 193, 194,

202

curtipennis, 173, 174, 193

cyanides, 181, 194

cyanipennis, 173, 174, 194

cycloderus, 173, 174, 177, 194, 207, 212,

214

decentis group, 177, 179, 185, 192

delicatulus, 111, 194

deyrollei, 180, 194

dilutus, 187, 194-195

districtus, 205

(Platynella) districtus, 176, 195

dominicensis , 183, 195

durangensis, 180, 195, 213

ebeninus, 185, 195

XXV

Platynus (continued)

erythrocerus, 181, 195

euides, 173, 174, 195

euprepes, 195

(Rhadine) euprepes, 178, 195-196

evanescens, 196

falli, 179, 196, 210

femoralis, 177, 196

forreri, 180, 196

fragilis, 173, 174, 196

fratellus, 173, 174, 184, 196-197,

198,213

funestus, 197

gracilis, 187, 197

guatemalensis, 197

guerrerensis, 173, 174, 197

haptoderoides, 185, 197

harfordi, 173, 174, 197

harpaloides, 179, 197, 206

hondurae, 173, 174, 197

hypolithos group, 176, 178

ilagis, 173, 174, 187, 188, 197-198

incommodus, 173, 174, 198

infidus , 173, 174. 198

inops, 182, 188, 192, 198

iricolor, 181, 198, 208

jalapensis, 173, 174, 198

key, 176-187

laetiusculus, 187, 197, 198

larvalis group, 178

leptodes, 198

(Rhadine) leptodes, 178, 198

lifragis, 173, 174, 187, 196, 198

limbicollis, 186, 198-199

logicus, 179, 182, 199

longiceps, 173, 174, 199, 201

( Colpodes) longiceps, 200

longipes, 178, 199

lucilius, 177, 199

lugens, 180, 189, 190, 199, 202

lymphaticus, 173, 174, 200

lyratus, 173, 174, 185, 200, 206

lyrophorus, 184, 188, 200

macrons , 184, 200, 109

marginatus, 212

marginicollis , 179, 200

medellini, 178

(Rhadine) medillini, 200

megalops, 173, 174, 177, 199, 200-201

Platynus (continued)

melanocnemis , 177, 201

meridanus, 201

metallicus, 183, 191, 201

minimus, 180, 201

moestus, 173, 174, 180, 193, 197, 199,

201- 202, 204, 213

monachus, 173, 174, 179, 182, 192, 202

(Platynella) montezumae, 176, 202

morelosensis, 173, 174, 202

nebrioides, 173, 174, 202

neglectus, 202

niger, 183,202, 207

nitidus, 173, 174, 179, 182, 185, 198, 199,

202- 203, 206, 211

nugax, 173, 174, 185, 200, 202, 203, 210

nyctimus, 179, 203

obscurellus, 173, 174, 186, 189, 197, 198,

203- 204

obscurus, 185, 186, 204

olivaceus, 187, 204

omaseoides, 176, 204

opacus, 204

orbicollis, 181, 204-205

ovatellus, 205, 209

ovatulus, 183, 205

ovipennis, 176

ovipennis group, 176, 178

pallidipes, 176, 205

parviceps, 205

pectoralis, 184, 187, 205

pelaezi, 178

(Rhadine) perlevis, 178, 205-206

petilus, 173, 174, 206

phaeolomus, 181, 206

picicornis, 183, 206

pinalicus, 173, 174, 206

planicollis, 206

platysmoides, 179, 206

porrectus, 111, 182, 192, 206

pristonychoides, 180, 206, 208

procephalus, 173, 174, 181, 191. 193, 197,

207

profundus, 185, 207

prolongatus, 173, 174, 207

pterostichoides, 179, 185, 207

punctatostriatus, 202, 207

purpuratus, 186, 191, 207

purulensis, 1 84, 207-208

XXVI

Platynus (continued)

quadrilaterus, 181, 198, 208

recticollis, 173, 174, 208

rectilineus, 173, 174, 206, 208

(Anacolpodes) rectilineus, 180

reflexicollis, 186, 208

reflexus, 184, 208

robustus, 184, 208

rotgeri, 178

(Rhadine) rotgeri, 209

rubidus, 187, 209

ruficornis, 184, 209

rufiventris, 177, 209

rufulus, 209

scabricollis , 183, 209

segregatus, 179, 196, 209-210

semiopacus, 177, 181, 189, 190, 193,

210

severus, 181, 182, 210

sexfoveolatus , 186, 210

sexpunctatus , 210

simplicior , 173, 174, 210

sphodroides, 180, 210

spinifer, 181, 210

stenos, 173, 174, 211

steropoides, 180, 211

striatopunctatus, 184, 211

stricticollis , 178, 211

subauratus, 187, 21 1

subcyaneus, 186, 188, 211

suffectus, 173, 174, 211

tenuicollis, 173, 174, 175, 177, 189,

211-212

tenuicollis group, 177, 179

tenuicornis, 111, 194, 212

teter , 181, 182, 212

tinctipennis, 182

tlamayensis, 185, 212

( Platynella) tolucensis, 176, 212

transfuga, 179, 181, 189, 212

transversicollis, 173, 174, 184, 213

trifoveolatus, 185

trifoveolatus group, 179, 185

tristis, 213

trujilloi, 173, 174, 213

umbripennis, 189

( St eno platynus) umbripennis, 180, 213

unilobatus, 173, 174, 213

valens, 180, 195, 213

Platynus (continued)

validus, 186, 213

variabilis, 173, 174, 186, 193, 197, 213,

214

veraecrucis, 207

(Mexisphodrus) veraecrucis, 182, 214

versicolor, 173, 174, 214

violaceipennis, 186, 214

Pleistocene, 394, 395, 407, 408, 410, 41 1, 412,

413,414,415,416,417

Pliocene, 386, 395, 401, 410, 41 1, 412, 415,

416

Plumb, G. H., 55, 97

Poa annua, 318

Podalonia, 13, 20, 26, 30

luctuosa, 15, 26

robusta, 15, 21 , 26

Podonominae, 387, 420

pollen, 140, 141, 143, 145

pollination, 144

flower, 143

Polyderis, 392

Polygonum viviparum, 117, 136

pompilid, 13, 20

Pompilidae, 13, 34

Pontania, 70

pacifica, 96

pomum, 61

poplar, 96

Populus, 61 , 86

balsamifera, 58

deltoides, 96

tremuloides, 58

Porsild, A. E., 117, 144

Powell, J. M., 118, 131, 135, 144

Powell, J. S. (see Elliot, D. H.), 388, 422

Pratt, D. (see Dubach, P.), 32

Progonochaetus , 267 , 268, 274, 276, 277 , 279,

340-342, 393, 394, 405, 406, 435, 440, 443,

473, 478

aeruginosis, 342

angolanus, 342

approximatus , 342

arnoldi, 342

atrofuscus, 342

bamboutensis, 342

basilewski, 261, 268, 342

bicoloripes, 342

brittoni, 342

XXV11

Progonochaetus (continued)

caffer, 342, 467

chev alien, 342

colmanti, 342

cursorius, 342

decorsei, 343

dilatatus , 342

discrepans, 342

emarginatus , 342

inchoatus, 342

incrassatus, 342

jeanneli, 276, 277, 343, 467

kafakumbae, 342

kapangae, 342

laeticolor, 342

laevistriatus, 342, 394, 467

limbatus, 342

longesulcatus, 342

merus, 342

moestus , 342

nigricrus, 342

obtusus, 342

ochropus , 341, 343

piceus, 342

planicollis, 342

prolixus , 342

pseudo chr opus , 341, 343

mdebecki, 342

sakalava, 342

seyrigi, 342

straneoi, 342

subcupreus, 342

vagans, 342

xanthopus, 341, 342

Prosoplasmic galls, 56

Prostomoxys saegerae, 255

Protognathus, 344

perrieri, 344, 345

zabroides , 344, 345

psenine wasps, 34

Psenini, 24-25

Pseudamphasia, 282, 379, 397, 398, 406,

44 1, 444, 473

Pseudanisodactylus , 267, 268, 277, 284,

351-352, 396, 404, 441, 444, 473

Pseudaniso tarsus, 267 , 268, 276, 277 ,

280, 290-292, 390, 406, 435, 440,

443, 472

widfci, 291,467, 472

Pseudaplocentrus , 267 , 268, 282, 376, 377,

396, 406, 436, 441, 444, 473

Pseudhexatrichus , 267 , 268, 277, 282, 349,

352-353, 396, 398, 404, 405, 424, 441,

444, 473

Pseudo dichirus , 268, 269, 283, 351, 354, 396,

398, 404, 405, 435, 441 , 444, 473

intermedius, 351

Pseudognathaphanus , 282, 341, 342, 344-345,

394, 395, 405, 406, 435, 440, 443, 473

dekkanus, 344

exaratus, 344

festivus, 344

laevistriatus, 342

perrieri, 345

punctilabris, 344

ru fit act or, 344

rusticus, 344

zabroides, 345

Pseudomorphini, 421

Psilidae, 254

Psithyrus, 138

Psyllidae, 74

psyllids, 24

Pteromalidae, 55, 62, 75, 96

Pterostichine, 175

Pterostichini, 35, 40

Pterostichus, 35, 40, 216, 275, 466

adoxus, 35, 36, 37, 38, 39

(Haplocoelus) adoxus, 35-39

rejectus, 35, 36, 38, 39

subarcuatus, 35, 36, 39

sufflatus, 35, 36, 39

sustentus, 35, 36, 39

tetricula, 35, 36, 39

/mto, 35, 36, 37, 38, 39

zephyrus, 35, 36, 39

Puel, L., 350, 351, 352, 354, 425

Putzeys, J. A. A. H., 296, 322, 342, 349, 425

Pyrobombus, 126, 143

Quendenfeldt, G., 342

Quensel, C., 380, 381

Quercus, 56

Quinlan, J. (see Eady, R. D.), 58, 96

Ragodactylus , 321

Ranunculus sulphureus, 1 1 7

trichophyllus , 1 1 7

Rasnodactylus , 341, 342, 343

jeanneli, 267, 268, 341, 342, 343

XXV111

Raven, P. H., 409, 41 1, 425

red-clover, 141, 143

Redtenbacher, L., 288, 425

Reiche, L., 207, 217, 340

Rey, C. (see Mulsant, E.), 339

Rhadine, 173, 175, 192, 196, 198,

206, 214, 215

perlevis, 205

Rhagionidae, 254

Rhagodactylus, 322

brasiliensis , 321, 322

Rhantus, 109

notatus, 109

plebeius (= binotatus), 109

sinuatus, 109

suturellus (= wallisi), 109

tostus, 109

zimmer manni (= suturellus), 109

Rhodites, 58, 95, 97

rosae, 95, 96

Rhopalosiphum rhois, 25

Rhysopus, 276, 277, 282, 347, 394,

395, 398, 399, 404, 435, 441, 443,

473

klynstrai, 347

Ribes lacustre, 58

Richards, K. W., 31, 34, 115-157

Richards, O. W., 115, 116, 134, 137,

138, 144

Riodinidae, 47

Robineau-Desvoidy, J. B., 224, 226,

243

robustella group, 242

Rohdendorf-Holmanova, E. B., 235,

238, 243

Rohwer, S. A., 58, 97

Rosa, 16, 21, 56, 58, 61, 67, 68, 97

acicularis , 55, 58, 60, 61, 66, 86, 92,

94, 97

arkansana , 61

calif ornica, 58

woodsii, 58, 61, 94

Rosaceae, 55, 56

Rosales, 56

rose galls, 95

gall wasp, 97

Ross, H. H., 409, 425

royal jelly, 145

Rubus , 24

ruby-tailed wasp, 32

Ryden, N., 229, 230, 231, 235, 238, 240,

243

Sagraemerus , 288, 289

javanus, 288

Salix, 16,21,58,86, 120, 129,313

arctica, 117, 119, 122, 136, 137, 156

lasiolepis, 96

Salomonsen, F. (see Freuchen, P.), 1 17, 127,

134, 142

Salt, R. W., 32, 34

Sanicula, 232

elate var. chinensis, 232

Sarcophaga rapax, 30

Sasakawa, ML, 220, 232, 235, 236, 243, 244

Satophagidae, 254

Satyridae, 47

Savage, J. M., 408, 425

Savile, D. B. O., 115, 116, 117, 127, 134, 135,

145, 147

sawflies, 26

Saxifragaceae, 8, 242

Saxifraga , 129

hir cuius, 1 1 7

nivalis, 117

oppositi folia, 1 19, 122, 131, 136, 137,

156

rivularis, 1 17

tricuspidata, 136, 156

Say, T., 40, 295, 298, 299, 313, 350, 357,

358, 364, 366, 375, 376, 378, 379, 380,

425

Scaphinotus, 419

petersi, 4 1 9

Scaritini, 427

Schaeffer, C., 200, 217, 349, 350, 357, 361,

425

Schauberger, E., 290, 344, 346, 350, 351,

352,425,426

Schaum, H., 339, 426

Schizogenius, 382, 427

Scholander, P. F., 32, 34

Schroder, D., 61, 65, 66, 67, 97

Schwarz, I., 134, 145

Sclater, P. L„ 403, 426

Scullen, H. A., 21, 28, 34

Scutopterus, 109

Scybalicus, 276, 280, 339-340, 390, 393, 394,

398, 404, 405, 435, 440, 443, 472, 476

XXIX

Scybalicus (continued)

biroi, 340

hirtus, 277, 340, 467

kabylianus, 340

oblongius cuius , 340, 469, 470

Scymnus, 42

Sehgal, V. K., 4, 8, 220, 226, 240,

244

Selander, R. B., 329, 334, 335, 337,

426

Selenophori, 263, 278

Selenophorus aeruginosus, 342

lugubris, 347

Senecio, 8, 242

Senecioneae, 8, 242

Sericoda, 174, 175

Seseli, 238

Severin, H. C. (see Young, F. N.), 1 10,

114

Shamsuddin, M., 166

Sharplin, C. D. (see Hocking, B.), 31,

33, 127, 131, 143

Shorthouse, J. D., 55-94, 97

(see Kevan, P. G.), 31, 33

Shuel, R. W., 135, 145

Silene acaulis , 136, 137

Silvius, 166

Simpson, G. G., 270, 384, 426

Simulium, 255

Sitona hispidula, 28

Skorikov, A. S., 116, 145

Sladen, F. W. L., 116, 117, 121, 122,

127, 132, 134, 135, 145

Sloane, T. G., 289, 290, 296, 426

Smirnovia tristis, 278

Smith, A. G., 384, 389, 392, 410,

413,426

Smith, A. V., 32, 34

Smith, F. (see Dubach, P.), 32

Smith, H. S., 62, 97

Solenius producticollis, 30

Solidago, 29, 58

Solier, A. J. J., 339

solitary bee, 34

solitary wasps, 13-32, 33, 34

S^mme, L., 32, 34

S0nderup, H. P. S., 235, 240, 244

Sorensen, T., 135, 145

Sparck, R. (see Braendegaard, J.), 116, 140

Sparre-Schneider, J., 134, 145

Spencer, K. A., 4, 8, 220, 222, 223, 224, 225,

226, 227, 229, 230, 234, 238, 240, 244

(see Hering, E. M.), 225, 240, 242

Sphagnum, 165

sphecid fauna distribution patterns, 30-32

sphecid wasps, 13, 20, 21, 22, 27, 30-32

Sphecidae, 13-32, 33, 34

Sphecinae, 25-26

Sphingidae, 26

spider wasps, 22

Spongopus, 283, 374-375, 396, 406, 435, 441,

444, 473

Spooner, G. M., 24, 25, 34

spruce, 16

-jackpine, 22

Stary, B., 235, 244

Steiner, A. L., 13-32, 34

Stellaria longipes, 136, 137, 156

Stenocnemus, 173, 188, 214

chevrolati, 188

versicolor, 194

Stephens, J. F., 351, 353

Sternlicht, M., 62, 97

Stewart, C. M. (see Dubach, P.), 32

Stilbolidus, 295

aztecanus, 320, 321

stink bugs, 22

Stomoxyine, 255

Stomoxys, 255

calcitrans, 255

ochrosoma, 255

Strand, E„ 116, 138, 145

Straneo, S. L., 174, 217

Stratiomyidae, 254

Straton, C. R. (see Alder, H.), 66, 95

Strickland, E. H., 23, 24, 26, 27, 28, 29, 30,

34, 161, 163, 166

Sturm, J., 342

Subterraneobombus , 126, 142

sumac, 25

Swales, D. E., 116, 134, 145

Swan, L. A., 41

Symphoromyia, 30

Synergus, 62

reinhardi, 68, 88

Syrphidae, 254

Syrphus ribesii, 29

Tabanid, 166

XXX

Tabanidae, 161-171

Tachinidae, 254

tachyine, 392

Tachysphex, 13, 20, 23-24, 30, 33

aethiops, 14, 20, 21, 23, 31

quebecensis, 14, 24, 31

terminatus, 14, 20, 24, 31, 33

Tachytini, 23-24, 31

Taketani, A. (see Yasumatsu, K.), 65,

66, 87, 88, 92, 98

Tanaka, K., 345, 346, 347, 350, 351,

426

Tanner, V. M., 275, 426

Tanno, K., 32, 34

Taschenberg, O. (see Heyne, A.), 190

Tauber, C. A. (see Tauber, M. J.), 225,

230, 244

Tauber, M. J., 225, 230, 244

Tecnophilus, 423

Tenthredinidae, 61, 96

Tephritidae, 74

Tertiary, 387, 388, 389, 390, 392, 393,

395, 397,410,411,412,415

Tetrastichus rosae , 62

tetrigids, 20, 24

Tettigoniidae, 33

Thawley, A. R., 135, 145

thecodont reptiles, 388

Thenarellus, 279

therapsid reptiles, 388

Theridiidae, 23

Thermonectes, 1 1 1

ornaticollis , 1 1 1

Thomas, A. W., 161-171

Thyanta, 23

Tipulidae, 254

Tomenthyonum nitens , 120

Torre-Bueno, J. R., 274, 426

Torymidae, 55, 62

torymids, 74

Torymus, 63, 74

bedeguaris , 55, 62, 63, 65, 70, 73, 74-75,

76, 77, 78, 79, 80, 81, 82, 84, 85

cy animus, 74

vesiculus , 74

Townes, H. K. (see Muesebeck, C. F. W.),

21, 22, 23, 24, 25, 26, 27, 28, 29, 30,

34, 59, 62, 67, 97

Trapezodera, 188

Trapezodera (continued)

aeneicauda, 188

Trechini, 386

Triassic, 388

Trichotichnus , 264

Triggerson, C. J., 62, 98

Trimerotropis, 23

Triplectrus, 354, 355, 356

aethiops, 369

beryllus, 360

breviceps, 364

convexus, 366

kempi, 369

longicollis, 364

marginatus, 369

modicus, 362

oblongus, 365

ovularis, 372

paulus, 359

peropacus, 363

semirubidus, 372

sulcipennis, 369

wolcotti, 369

Triplosarus, 280, 285-286, 389, 405, 435,

440, 443, 472

fulvescens, 285, 286

novaezealandiae, 286

Trypanosoma evansi, 255

Tryxalus, 24

Tschitscherine, T., 276, 345, 346, 353, 426

Tussilago, 8, 242

Tutin, T. G., 220, 244

Umbelliferae, 3-8,219-253

United States Department of Agriculture, 426

Urarus, 102

Urbahns, T. D., 75, 98

Urophora jaceana, 74, 98

Ursprung, H., 43

Ushatinskaya, R. S., 32, 34

Uvarov, B. P.. 31, 34

Van Dyke, E. C., 209, 217

Varley, G. C., 71,74, 75,79, 98

Vaurie, P. (see Selander, R. B.), 329, 334, 335,

337, 426

vespid, 22

Viburnum edule, 58

Viereck, H. L., 71,98

vilis group, 1 13

Virostek, J. F. (see Hobbs, G. A.), 132, 143

Vogt, P. R. (see Morgan, W. J.), 384,

425

Voigt, G., 235, 244

Wagner, F. von. (see Friese, H.), 1 1 7,

126, 127, 138, 142

Wallace, A. R., 403, 404, 405, 406,

426

Wallis, J. B., 99-113, 114

wasps, 32, 33, 34, 145

Weaver, N., 134, 145

weevils, 21, 27, 403

Wehner, R., 257

Weld, L. H., 56, 58, 59, 65, 98

Wells, B. W., 56, 98

Westwood, J. O., 348, 426

Whitehead, D. R., 410, 426, 427

Whitehead, Donald Robert., 173-214,

270, 382, 427

Whittaker, R. H., 94, 98

Wiedemann, C. R. W., 288, 289, 427

Wildner, G. (see Bukatsch, F.), 134, 141

Williams, F. X., 22, 34

willow, 16, 21

Wilson, E. W., 390, 427

Wingstrand, K. G. (see Brinck, P.), 1 16,

117, 121, 134, 138, 141

Wojtowski, F., 132, 134, 145

Wold, J. L. (see Scullen, H. A.), 21, 28,

34

Wolfe, J. A., 410, 427

Wood well, G. M., 94, 98

worker caste, 1 4 1

Wykes, G. R., 135, 145

Xenophonus, 278

hirtus, 278

Xestonotus, 283, 347-348, 394, 395,

399, 406, 435,441,443,473

lugubris, 277, 348, 466, 469, 471

yarrow, 16, 20, 21

Yarrow, I. H. H., 116, 138, 145

Yasumatsu, K., 65, 66, 87, 88, 92, 98

Young, F. N., 101, 102, 110, 114

Zacotus, 4 1 9

Zoerner, H., 229, 235, 239, 244

Zumpt, F., 255

D.-

Quaestiones

entomologicae

MUS. COMP, zool:

LIBRARY

FEB 1 7 1973

HARVARD

UNIVERSITY

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

VOLUME IX

NUMBER 1

JANUARY 1973

QUAESTIONES ENTOMOLOGICAE

A periodical record of entomological investigation published at the Department of

Entomology, University of Alberta, Edmonton, Alberta.

Volume 9 Number 1 2 January 1973

CONTENTS

Editorial - On Finality 1

Griffiths - Studies on boreal Agromyzidae (Diptera). III. Phytomyza

miners on Cnidium and Conioselinum (Umbelliferae) 3

Steiner — Solitary wasps from subarctic North America — II. Sphecidae from the

Yukon and Northwest Territories, Canada: Distribution and ecology 13

Perrault — A taxonomic review of the eastern Nearctic species complex

Pterostichus (Haplocoelus) Adoxus (Coleoptera: Carabidae) 35

Book review 41

Book review 44

Book review 47

Editorial — On Finality

Education means, by derivation, “a leading out” (e - out, ducere - to lead), by implication

and usage it means a leading of a person out of the darkness of ignorance into the illumina-

tion of knowledge, understanding, and wisdom. Wisdom most importantly, for neither

knowledge nor understanding constitute education, though they are essential ingredients of

it. Phonograph discs and magnetic tape accumulate knowledge more efficiently than the

mind of man; they do not understand, and are wise only in that they start in response to

the right stimulus and stop, usually, when they come to the end.

An honest doctor once remarked that birth and death, the two temporal ends of a man,

had little finality to them; that while a good doctor was usually pretty certain of a birth

when he saw one, most of them were much less certain about the precise moment of death

and admitted that the signing of a death certificate brought out the gambler in them. But

birth, as the beginning of life, as witness the controversies over abortion, is also at best ques-

tionable. So it is with insects; more or less precisely so with such as Glossina. At first sight

the egg-layers seem to start their lives with more precision, but surely life begins at fertiliza-

tion, so what of parthenogenesis? Do we go back to the most recent sexually produced

individual for the beginning? If a generation runs from fertilization to fertilization, as, in

the context of evolution, it must, when we speak of alternation of generations we give it a

different meaning. Different by a factor of two. But insect deaths are more dubious. Despite

our vast investments in them, they still defy definition; ask any dabbler in the study of

median lethal doses. Cryptobiosis and durable diapauses add further doubts.

The ends of an animal in space present problems of a different nature. Animal bodies

seem reluctant both to begin and to end. The front end of an animal, that end which

(usually) arrives first on the scene in the normal progress of the beast, is analogous to the

beginning of life, and, like birth it is usually more clearly defined, more abrupt. But it is

rarely completely abrupt; an advance guard of feelers, tentacles, antennae, or pseudopodia

precedes the main bulk of the body. The essential quality of a tail is its taper, a reluctance,

2

as it were, to come to an end. Trailing appendages or receptors often keep it company in its

reluctance. Of course a blunt beginning and a tapered finish are the essence of streamlining:

some Crustacea, unable to face up to this blunt beginning fold their two tapered ends

together and allow them to trail behind the superbly rounded bluntness of their folded

middles. Even the tails of those of us as have withdrawn them inside in embarrassment, are

reluctant to face up to finality. Smaller vertebrae successively succeed each other in what

Goethe called a gesture towards infinity. Morphologists recognize the reluctance of seg-

mented animals to start and to finish by giving special names to the first and last pieces of

the body, the acron and the telson. These parts really only differ from the segments in

between them in having only one neighbour instead of two, and in doing their best to intro-

duce the body and to bring things to an end.

Many forms of life have sidestepped the problem of beginning and ending by adopting a

radial rather than a bilateral symmetry. But this only compounds the problem — they have

to end in all directions instead of only in two, as witness the tapered arms of starfish. A

solution to the problem for bilateral animals which does not appear to have been pursued is

to join the two ends together to yield what one might call a ring worm; perhaps dogs and

cheese-skippers which chase their tails are playing with this idea. Perhaps the incredible

length of some nemertine worms of the genus Lineus, the bootlace worms, arises from a

simple reluctance to face the problem of ending.

Plants have a masterly way of their own of coming to an end, best shown by trees. Both

upwards and downwards, the extremities of these remarkable organisms combine tapering

with branching, thus having more and more parts of less and less size until they wind up

with a multitude of nothings. As in space, so also in time, plants take on life and give it up

with becoming pause.

To return to education, leading out is a gradual process. So is graduation, at least by

derivation, though it has become something of a sudden affair.

“Creatures animate with gradual life

Of growth, sense, reason, all summed up in man/’

There are trends in education today divergent from Milton’s view; but education - natural

education - must be a gradual process, integrated with (and of course embodying the study

of) life itself. Despite the element of repetition in it, the term ‘continuing education’ is a

valid one, for knowledge grows continually faster and must continue to nourish wisdom.

The finality of a final examination is antagonistic to education; life itself is the final exami-

nation of wisdom. So also terminal courses; they have no place in education until it is all

over. The only truly terminal course is that from the funeral parlour to the graveyard or

crematorium. A terminal course with a final examination is the end of everything.

Even an editorial must eventually come to an end. You might think that this is the end.

Well, it is. Almost.

Brian Hocking

STUDIES ON BOREAL AGROMYZIDAE (DIPTERA). III.

PHYTOMYZA MINERS ON CNIDIUM AND CONIOSELINUM (UMBELLIFERAE)

GRAHAM C. D. GRIFFITHS

Department of Entomology

University of Alberta Quaestiones entomologicae

Edmonton, Alberta T6G 2E3 9 : 3-11 1973

Phytomyza sitchensis n. sp. (type-locality Sitka, Alaska) and P. conioselini n. sp. ( type-

locality Chilkat Peninsula, Alaska) are recorded as miners of Conioselinum chinense (L.);

and P. cnidii n. sp. (type-locality Atkinson Point, Northwest Territories) as miner of Cnidium

cnidiifolium (Turcz.). No agromyzid miners of these plant genera were previously described.

Phytomyza sitchensis n. sp. (localite-type Sitka, Alaska) et P. conioselini n. sp. (localite-

type Peninsule de Chilkat, Alaska) sont rapportees comme mineuses du Conioselinum chi-

nense ( L. ); et P. cnidii n. sp. (localite-type Atkinson Point, Territoires du nord-ouest)

comme mineuse du Cnidium cnidiifolium (Turcz.). Aucune Agromyzide mineuse de ces

genres de plant es n’a ete decrite auparavant.

Phytomyza sitchensis n. sp. (Fundort vom Typus Sitka, Alaska) und P. conioselini n. sp.

(Fundort vom Typus Chilkathalbinsel, Alaska) werden als Minierer von Conioselinum chi-

nense (L.) besprochen; P. cnidii n. sp. (Fundort vom Typus Atkinson Point, Northwest

Territories) als Minierer von Cnidium cnidiifolium (Turcz.). Agromyziden-Minierer dieser

Pflanzengattungen sind bisher nicht beschrieben worden.

In the present paper three new species of Phytomyza are described from Alaska and

northwest Canada. All belong to the Phytomyza albiceps group, in the sense explained in

my previous paper (Griffiths, 1972b). These are the first agromyzid species described as

miners of Cnidium and Conioselinum, although there are previous records for Europe of

miners which were not bred (see below). Both host-plants of the agromyzid species here

described, Conioselinum chinense (L.) and Cnidium cnidiifolium (Turcz.), occur in north-

east Asia, as well as in North America (Hulten, 1968). The former is mainly a coastal plant,

ranging on the Pacific coast of North America from the Bering Straits to Washington State;

elsewhere it occurs on the eastern seaboard of North America, and in Asia on Hokkaido,

Sakhalin, the Kuril Islands and Kamchatka. Cnidium cnidiifolium is an arctic plant, not

reaching below the 60th parallel in North America; it has an extensive distribution in eastern

Siberia, as well as in Alaska, Yukon and along the arctic coast of the Northwest Territories.

The terminology and abbreviations used in my descriptions were explained in the first

paper of this series (Griffiths, 1972a). My use of the above plant names follows Hulten

(1968). The holotypes of the new species will be deposited in the Canadian National Col-

lection (Ottawa).

PREVIOUS RECORDS

An unknown Phytomyza species produces linear mines on Cnidium dubium (Schkuhr)

(= venosum Koch) in Poland and Germany. Recorded localities are Crossen-an-Oder (Kros-

no), Poland (Hering, 1936), Blumerode, Silesia, Poland (viii-ix.1934; Buhr, 1941) and Boiz-

enberg-an-Elbe, Mecklenburg, Germany (Buhr, 1932). Hering (1957:31 1, no. 1523) describes

the mine as follows.

“Channel begins in leaflet-centre, follows first one, then the other leaflet margin, finally

filling the entire leaflet; likewise 1-2 further leaflets are mined out; the early channel is

finally no longer recognizable. Mine whitish green when fresh, but soon becomes brownish.

4

Griffiths

Faeces in fine black particles, which in places are linked in beaded fashion, deposited irregu-

larly or in two rows. Semicircular slit on upper surface.”

Hering suggests that similar mines in Berlin Botanical Gardens were caused by Phytomyza

mylini Hering, but this was not confirmed by breeding. Linear mines of the type described

are produced on Umbelliferae by many different species of the Phytomyza albiceps group.

In the absence of any morphological information on larvae or adults, the species concerned

cannot be determined.

De Meijere (1937:238) has described and figured a larva collected by H. Buhr from mines

on Conioselinum tataricum Fisch. at Leningrad (Russia). This larva had only five bulbs on

its anterior spiracles, but about 16, arranged more or less in a circle, on its posterior spi-

racles. This description is not appropriate to the third instar larva of any of the three species

found by me on Conioselinum in Alaska, for all have more numerous spiracular bulbs.

Hering’s (1957:315, no. 1546) statement that the mines from Leningrad are “large, taking

in a large part of a point of the leaf, only on upper-surface” suggests that they are blotch-

mines; but he does not state this explicitly.

DIAGNOSIS

Caught adults of the Phytomyza albiceps group can be reliably identified only by dissec-

tion of the male genitalia. Fortunately the form of the aedeagus is strongly differentiated

between species of this group, allowing confident identification of many species which are

inseparable on external characters. The three new species described in this paper may be

included in Spencer’s (1969) key to Phytomyza species of Canada and Alaska by the exten-

sions given below. The second of these extensions (to couplet 88) incorporates an extension

previously proposed by Sehgal (1971).

84. Tarsi yellow; aedeagus as Spencer’s Figs. 402, 403 aralivora Spencer

Tarsi dark 84a

84a. Aedeagus as Spencer’s Figs. 473, 474 osmorhizae Spencer

Aedeagus as Fig. 7 sitchensis n. sp.

Aedeagus as Figs. 4, 5 conioselini n. sp.

88. Third antennal segment distinctly enlarged; aedeagus as Spencer’s Fig. 468

nepetae Hendel

Third antennal segment not enlarged 88a

88a. Aedeagus as Spencer’s Figs. 504, 505 sehgali Spencer

Aedeagus as Sehgal’s Figs. 1 10, 1 1 1 mertensiae Sehgal

Aedeagus as Figs. 1,2 cnidii n. sp.

The following key will facilitate identification of mines and immature stages of Phyto-

myza species on Conioselinum. No other genera of Agromyzidae are known to attack this

plant genus.

Key to Phytomyza mines on Conioselinum

1. On C. tataricum Fisch. Anterior spiracles of third instar larva (and puparium) with

five bulbs; posterior spiracles with about 16 bulbs P. sp. (de Meijere, 1937:238)

— On C. chinense (L.). Spiracular bulbs of third instar larva and puparium more numer-

ous 2

Boreal Agromyzidae

5

2. Mine primary blotch (Fig. 14A). Puparium with prominent anal lobes; posterior spi-

racles of puparium and third instar larva with 21-22 bulbs in broad ellipse (nearly

circular) (Fig. 11) P. sp. (compare angelicae Kaltenbach)

— Mine basically linear, though portions of the channel may coalesce in narrow leaf lobes

(Fig. 14B). Puparium without prominent anal lobes; posterior spiracles of puparium

and third instar larva with bulbs in narrow ellipse (Fig. 12, 13) 3

3. Puparium smoothly rounded, with intersegmental boundaries scarcely impressed (Fig.

9) P. sitchensis n. sp.

— Puparium as Fig. 1 0, with intersegmental boundaries distinctly impressed

P. conioselini n. sp.

TREATMENT OF SPECIES

Phytomyza cnidii new species

Adult. — Head with orbits narrowly projecting above eye in lateral view; genae in middle

1/3 to 1/4 of eye height; eyes with only sparse fine pubescence. Frons at level of front

ocellus 2-2 Vi times width of eye. Ors directed posteriorly, ori directed inwardly; posterior

ors about 2/3 as long as anterior ors; anterior ori variably developed, ranging from very

short to 2/3 as long as posterior ori; orbital setulae few (3-5), in one row. Peristomal margin

with vibrissa and 2-3 upcurved peristomal setulae. Third antennal article rounded distally,

with short pubescence.

3 + 1 dc; acr in 4-5 rows; 5-10 presutural ia; 4-7 postsutural ia; inner pa about half as long

as outer pa.

Second cross-vein (m-m) absent; m1 + 2 weak, absent from centre of wing in two females

(although they retain its terminal portion at wing tip). Costal ratio mg2/mg4 2. 2-2. 4. Wing

length 1. 8-2.0 mm.

Colour largely dark. Centre of frons dark brown, only slightly paler than black ocellar

plate, vertex and orbits; genae brown. Antennae black. Palpi black; labella orange-yellow.

Thorax finely grey-dusted, weakly shining, largely black with pale coloration only along

notopleural and mesopleural sutures (and in one specimen also at corners of humeral calli);

wing base yellow; squamae pale or somewhat infuscated, with dark fringe. Legs largely

black, with tips of front femora not contrasting, yellow-brown or red-brown. Basal cone of

ovipositor (9) largely shining, grey-dusted on dorsal surface only narrowly at base.

Male postabdomen with 8th sternum fused with 6th tergum. Telomeres not clearly de-

limited from periandrium, bearing dense group of setulae. Pregonites inconspicuous (weakly

pigmented), extending ventrally, shielding base of aedeagus at rest. Aedeagal hood with two

pairs of lateral sclerites. Aedeagus as Fig. 1,2; basal section with group of very small spinules

on dorsal surface between basal sclerites; medial lobe with well-defined loop of sclerotiza-

tion; distal section largely unpigmented, with distiphallus represented by slender strip of

sclerotization. Ejaculatory apodeme as Fig. 3.

Puparium and third instar larva. — Mandibles with two alternating teeth; right mandible

longer than left. Anterior spiracles with 8-10 bulbs in irregular ellipse. Posterior spiracles on

short conical processes, with 14-15 bulbs in narrow ellipse. Puparia dark brown or black,

1.7-1. 8 mm long, strongly arched, with clearly impressed intersegmental boundaries; anal

lobes weakly developed.

Mine. — Larvae leaf-miners on Cnidium cnidiifolium (Turcz.), leaving leaf before pupa-

rium formation. A description of the mine cannot be given, as the leaves from which the

type series was bred decomposed while in transit.

6

Griffiths

Types. — Holotype 6, 3 99 paratypes from larvae 26.vii.70 on Cnidium cnidiifolium

(Turcz.), 4 miles S Atkinson Point (on pingo), Northwest Territories, Canada, emerged

2.V.71, leg. P. G. Kevan.

Phytomyza sitchensis new species

Adult. — Head with orbits not projecting above eye in lateral view; genae in middle 1/3

to 1/4 of eye height; eyes with only sparse fine pubescence. Frons at level of front ocellus

about twice width of eye. Ors directed posteriorly, ori directed inwardly; posterior ors 3/4

to almost as long as anterior ors; anterior ori short or absent; orbital setulae few (4-6), in

one row. Peristomal margin with vibrissa and 3-5 upcurved peristomal setulae. Third anten-

nal article rounded distally, with short pubescence.

3 + 1 dc; acr in 3-4 rows; 5-6 presutural ia; 6-7 postsutural ia; inner pa about half as long

as outer pa.

Second cross-vein (m-m) absent. Costal ratio mg2/mg4 3. 4-3. 5. Wing length 2. 4-2.6 mm.

Colour almost entirely dark. Centre of frons and genae dark brown, scarcely paler than

rest of head. Labella orange-yellow. Thorax grey-dusted over black ground colour, only

weakly shining, with pale coloration only along notopleural and mesopleural sutures. Wing

base and squamae yellowish white, latter with dark margin and fringe. Legs largely dark,

with tips of front femora contrastingly yellow; tips of other femora less contrasting, yellow-

brown or dark. Basal cone of ovipositor (9) grey-dusted on about basal third.

Male postabdomen with 8th sternum fused with 6th tergum. Telomeres partly delimited

from periandrium by suture on outer side, bearing numerous fine setulae. Pregonites ex-

tending ventrally (shielding base of aedeagus at rest), but inconspicuous (weakly pigmented).

Aedeagal hood with two pairs of lateral sclerites (the more dorsal pair rather ill-defined).

Aedeagus as Fig. 7; right basal sclerite expanded at base; both basal sclerites with row of

conspicuous spinules distally above their dorsal margins; medial lobe with pair of slender

well-defined sclerites; distal section long, at its base with well-defined sclerite (mesophallus)

enclosing ejaculatory duct, largely membranous distally with only weak traces of terminal

pigmentation (distiphallus). Ejaculatory apodeme as Fig. 8.

Puparium and third instar larva. — Mandibles with two alternating teeth; right man-

dible longer than left. Anterior spiracles with two short horns, with about 14 bulbs in

ellipse. Posterior spiracles on short broad processes, with 26-33 bulbs in long narrow el-

lipse with wide gap on inner side (Fig. 12). Puparia (Fig. 9) shining black, about 1.7

mm long, smoothly rounded with intersegmental boundaries scarcely impressed; anal lobes

absent.

Mine. — Larvae leaf-miners on Conioselinum chinense (L.). Mine (Fig. 14B) entirely

linear, mainly following sinuations of leaflet margins, about 5 cm long, I-IV2 mm wide

terminally; faeces deposited as fine particles, mostly close together or forming beaded

strips; mine entirely on upper surface of leaf, appearing whitish green in reflected light;

larvae leaving leaf through semicircular slit on upper surface before puparium formation.

Types. — Holotype <5, 1 9 paratype from larvae 20-30.viii.69 on Conioselinum chinense

(L.), Starrigavan (on beach), Sitka, Alaska, emerged 1 1-14.V.70, leg. G. C. D. Griffiths. 1 9

paratype from larva 27-30.vi.68 on Conioselinum chinense (L.), Chilkat Peninsula (near

Haines), Alaska, emerged 23.vii.68, leg. G. C. D. Griffiths.

Remarks. — The breeding data given above indicate that sitchensis is multivoltine. The

smoothly rounded puparia are of the type described by Allen (1957) for P. obscurella

Fallen and other European species. The other known miners of Conioselinum do not have

puparia of this type.

Boreal Agromyzidae

7

Phytomyza conioselini new species (<5)

Adult. — Head with orbits only very narrowly projecting above eye in lateral view; genae

in middle 1/4 of eye height; eyes with only sparse fine pubescence. Frons at level of front

ocellus about 2 Vi times width of eye. Ors directed posteriorly, ori directed inwardly; poste-

rior ors from 2/3 to fully as long as anterior ors; anterior ori 1/2 to 2/3 as long as posterior

ori; orbital setulae numerous (8-10), irregularly arranged (a few lying between main row

and level of orbital bristles). Peristomal margin with vibrissa and 5-8 upcurved peristomal

setulae. Third antennal article rounded distally, with short pubescence.

3 + 1 dc (except 2 + 1 on one side in para type); acr and ia long; acr in 3-4 rows; 5-9

presutural ia; 4-9 postsutural ia; inner pa about half as long as outer pa.

Second cross-vein (m-m) absent. Costal ratio mg2/mg4 3.3. Wing length 2.5 mm.

Colour largely dark. Centre of frons ochreous, somewhat contrasting with dark orbits,

ocellar plate and vertex; genae ochreous to yellow-brown. Antennae dark. Palpi black;

labella yellow. Thorax strongly grey-dusted, scarcely shining, largely dark with pale colora-

tion only along notopleural and mesopleural sutures and on postalar callus (below outer

pa). Wing base and squamae whitish, latter with dark margin and fringe. Legs largely dark,

with tips of front femora contrastingly yellow; tips of other femora less contrasting, yellow-

brown or reddish.

Male postabdomen with 8th sternum fused with 6th tergum. Telomeres partly delimited

from periandrium by suture on outer side, bearing dense group of fine setulae. Pregonites

large, extending ventrally (shielding base of aedeagus at rest), but inconspicuous (weakly

pigmented). Aedeagal hood with two pairs of lateral sclerites (the more dorsal pair rather

ill-defined). Aedeagus as Fig. 4, 5; basal section without spinules; medial lobe with pair of

asymmetrically developed sclerites, that on left side much expanded with projecting point;

distal section very short, with complex sclerotization. Ejaculatory apodeme as Fig. 6.

Puparium and third instar larva. — Mandibles with two alternating teeth; right mandible

longer than left. Anterior spiracles with about 1 2 bulbs in irregular ellipse. Posterior spira-

cles (Fig. 13) on short conical processes, with 22-29 bulbs in narrow ellipse. Puparia (Fig.

10) dark brown or black, shining, 2. 1-2.2 mm long, strongly arched, with intersegmental

boundaries distinctly impressed; anal lobes absent.

Mine. — Larvae leaf-miners on Conioselinum chinense (L.). Mines linear, similar to those

of sitchensis; larvae leaving leaf through semicircular slit before puparium formation.

Types. — Holotype d, 1 d paratype from larvae 27.vi-2.vii.68 on Conioselinum chi-

nense (L.), Chilkat Peninsula (near Haines), Alaska, emerged 1 2-13. x. 68, leg. G. C. D.

Griffiths.

Remarks. — Puparia of this species and of sitchensis were obtained from linear mines on

Conioselinum collected on the Chilkat Peninsula. Unfortunately my records do not enable

me to separate the pressed mines according to species. There seems no obvious basis for

dividing them into two groups. However, the puparia are readily separable, for those of

conioselini have impressed intersegmental boundaries (Fig. 10), while those of sitchensis are

smoothly rounded (Fig. 9).

I do not know whether the late emergence of the two specimens indicates that this spe-

cies has a second generation in autumn, or was a “forced” emergence caused by delay in my

obtaining outdoor storage facilities.

The paratype male has an abnormal abdomen, with incomplete hypopygial rotation. The

cause of this was evidently unsuccessful parasitoid attack, for in the abdomen was found

a large capsule (0.325 x 0.15 mm) containing a hymenopterous larva. The aedeagus of

the paratype agrees with that of the holotype except that the right (not left) sclerite of

8

Griffiths

the medial lobe is expanded. I interpret its condition as abnormal in this respect, since

disturbances of the rotation process in cyclorrhaphous Diptera are often associated with

anomalous development of asymmetrical structures.

Phytomyza sp. (compare angelicae Kaltenbach)

In addition to larvae of sitchensis and conioselini, I also collected on the Chilkat Penin-

sula larvae of a third species of Phytomyza on Conioselinum chinense (L.), these producing

primary blotch-mines (Fig. 11, 14A). The mines and puparia of this species are similar to

those of P. angelicae Kaltenbach (on Angelica). Unfortunately I have so far obtained only

female flies. These are similar to angelicae, differing from the other Conioselinum-mmers

in having a bright yellow frons. I cannot determine whether they represent a distinct species

until males are obtained for critical comparison.

ACKNOWLEDGEMENTS

I am most grateful to P. G. Kevan for collecting and sending me the material from

Cnidium. My material from Conioselinum was collected on field trips supported by the

Boreal Institute of the University of Alberta. My wife Deirdre kindly prepared the illus-

tration of leaf mines (Fig. 14). I am grateful to M. von Tschirnhaus and K. A. Spencer

for confirming that none of the species described in this paper was previously known to

them.

REFERENCES

Allen, P. 1957. Larval morphology of some species of Phytomyza Fallen (Diptera: Agro-

myzidae). Proc. R. ent. Soc. Lond. (A) 32:171-181.

Buhr, H. 1932. Mecklenburgische Minen. I. Agromyziden-Minen. Stettin, ent. Ztg. 93:57-

115.

Buhr, H. 1941. Mecklenburgische Minen. IV. Nachtrag zu den Dipteren-Minen mit Einschluss

der in den Rostocker Botanischen Garten festgestellten. Arch. Ver. Freunde Naturg.

Mecklenb. 15:21-101.

Griffiths, G. C. D. 1972a. Studies on boreal Agromyzidae (Diptera). I. Phytomyza miners

on Saxifragaceae. Quaest. ent. 8:67-80.

Griffiths, G. C. D. 1972b. Studies on boreal Agromyzidae (Diptera). II. Phytomyza miners

on Senecio, Petasites and Tussilago (Compositae, Senecioneae). Quaest. ent. 8:377-405.

Hering, M. 1935-1937. Die Blattminen Mittel- und Nord-Europas einschliesslich Englands.

Verlag Gustav Feller, Neubrandenburg. xii + 631 pp.

Hering, E. M. 1957. Bestimmungstabellen der Blattminen von Europa einschliesslich des

Mittelmeerbeckens und der Kanarischen Inseln. Uitgeverij Dr. W. Junk, The Hague. 1185

+ 86 pp. (3 vols.).

Hulten, E. 1968. Flora of Alaska and neighbouring territories. Stanford University Press,

Stanford, California, xxii + 1008 pp.

Meijere, J. C. H. de. 1937. Die Larven der Agromyzinen. Dritter Nachtrag. Tijdschr. Ent.

80:167-243.

Sehgal, V. K. 1971. A taxonomic survey of the Agromyzidae (Diptera) of Alberta, Canada,

with observations on host-plant relationships. Quaest. ent. 7:291-405.

Spencer, K. A. 1969. The Agromyzidae of Canada and Alaska. Mem. ent. Soc. Can. no. 64.

311 pp.

Boreal Agromyzidae

9

Fig. 1-3. Phytomyza cnidii n. sp., holotype 6: 1, aedeagus and associated structures in lateral view (AEDAD aedeagal

apodeme, AEDH aedeagal hood, DPH distiphallus, Ml medial lobe, POG postgonite); 2, distal section and medial

lobe of aedeagus in anteroventral view; 3, ejaculatory apodeme. Fig. 4-6. Phytomyza conioselini n. sp., holotype <5:

4, aedeagus in lateral view; 5, distal section and medial lobe of aedeagus in ± ventral view; 6, ejaculatory apodeme.

Fig. 7-8. Phytomyza sitchensis n. sp., holotype 6: 7, aedeagus in lateral view; 8, ejaculatory apodeme.

10

Griffiths

Fig. 9. Phytomyza sitchensis n. sp., puparium in dorsal view. Fig. 10. Phytomyza conioselini n. sp., puparium in dorsal

view. Fig. 11. Phytomyza sp. (compare angelicae Kaltenbach), posterior spiracle of puparium in caudal view. Fig. 12.

Phytomyza sitchensis n. sp., posterior spiracle of puparium in caudal view. Fig. 13. Phytomyza conioselini n. sp., poste-

rior spiracle of puparium in caudal view.

Boreal Agromyzidae

11

1 cm.

14A

N

Fig. 14. Leaf of Conioselinum chinense (L.) with mines of Phytomyza sp. (compare angelicae Kaltenbach) (A) and

P. sitchensis n. sp. (B).

SOLITARY WASPS FROM SUBARCTIC NORTH AMERICA -

II. SPHECIDAE FROM THE YUKON AND NORTHWEST

TERRITORIES, CANADA: DISTRIBUTION AND ECOLOGY

ANDRE L. STEINER

Department of Zoology

University of Alberta Quaestiones entomologicae

Edmonton, Canada T6G 2E9 9 : 13-34 1973

Sphecid wasps of 35 species were collected in the Yukon and Northwest Territories

during the summers of 1967 and 1968. Literature about these species is reviewed. Latitude,

vegetation, soil type and slope of the study areas are analyzed as is their use by the wasps,

for feeding, preying, nesting and basking. Particularly well represented in these samples are

the subfamilies Crabroninae and Pemphredoninae, and the genera Ammophila, Podalonia

and Tachysphex. Included are circumpolar, Holarctic and Nearctic species. Some of the last-

named group range widely, with populations occurring as far south as Florida and Mexico.

Trente-cinq especes de sphegides ont ete recoltees au Yukon et dans les Territoires du

Nord-Ouest au cours des etes 1967 et 1968. Le texte comprend un compte-rendu de la

bibliographie relative a ces especes. On y trouvera une analyse de la latitude, de la vegeta-

tion, de la nature du sol, de la pente des regions etudiees ainsi que de Tutilisation qu’en

font les sphegides en fonction de leur comportement alimentaire et nidificateur, de la selec-

tion des proies et de leur exposition au soleil. Ce sont sur tout les sous- families Crabroninae

et Pemphredoninae et les genres Ammophila, Podalonia et Tachysphex qui sont bien repre-

sents parmi cette faune. On y trouve aussi des especes circumpolaires, holarctiques et

nearctiques. Pour le dernier groupe nomme, la distribution est tres large, avec des popula-

tions pouvant atteindre des regions aussi meridionales que la Floride et le Mexique.

Despite severe climatic conditions and a short season favorable for activity, many species

of the family Pompilidae are represented in the subarctic regions of North America (Steiner,

1970). This paper deals with another family of solitary wasps, the Sphecidae, having repre-

sentatives which were collected in the same study area.

This paper describes habitats and microhabitats, and provides data about the uses made

of these by different species of sphecids. These wasps are much more diversified in their

ecological and behavioral characteristics than are pompilids. Sphecids are well suited for

behavior-ecology studies, particularly for those concerning adaptations for avoiding com-

petition at the community level.

THE STUDY AREAS

General features and maps of the regions visited and of the localities sampled have been

previously presented (Steiner, 1970). Additional data are provided in Table 1.

Climax communities of the boreal forest and taiga do not appear to provide particularly

favorable habitats for solitary wasps but some natural or man-disturbed areas in the north

are potentially good habitats. These include pits, river banks, lake shores, flats, outwash

plains and forest edges. Most of the sphecids discussed were collected in such places. Figures

1 to 9 illustrate five locality types from which samples were collected.

Table 1 . Distribution per species and localities.

14

Steiner

('N apnqiBj puB) sgpiiBooq

c«

o

a>

a,

oo

a. -2

3

.a.

Q ^

03

3

3

.§

3

*• 2

=§ ^

^ §■

b a

Q ^

a.

o

•5

Co

a '£

8 §

■5.1

53 -a

?s 2

-3 sL

3 a*

3

3

•§

v.

-3

3.

^ K’ K’ Q ^ ^ ^ a, q,' ^

— rstro^fin\or^oooNO — rNiro-^t

Subarctic Sphecidae

15

* localities intensively sampled.

- localities sampled very incompletely and superficially.

16

Steiner

In the Northwest Territories study area, most localities are on plains and plateaus to the

west of the Canadian Shield. However, localities XII, XIII and XV are on the Canadian

Shield and locality I (Fort Smith) is near its western edge. In this area samples were col-

lected throughout the summers of 1967 and 1968. The Yukon study area was visited only

briefly after mid-August of 1968. Locality X (Fort Providence, in the Mackenzie River

valley, Fig. 1 and 2) was the most favorable area visited and was the most intensively

investigated.

Eleven habitats are recognized. They are characterized below in terms of slope, vegetation

and soil. Also described are the uses made of each habitat by its sphecid inhabitants. See

Table 2 for an explanation of letters in Figures 1 to 9.

Table 2. Explanation of letters in Figures 1 to 9. (Typical wasp habitat facies and micro-

habitats in some Northwest Territories sample localities)

a = Immature aspen interspersed with willow and shrub

b = Low spoilbank (small man-made ridge)

c = Flat area, covered with dense, low vegetation

d = Patches of Achillea millefolium (yarrow)

e = Flat area, sparsely vegetated

f = Patches of Epilobium species (fireweed)

g = Spruce/ Jackpine forest (interspersed with aspen)

h = Collapsed cutbanks (steep slope)

i = Patches of shrubby vegetation ( Rosa sp., Salix sp., Betula sp., Alnus sp., etc.)

j = Edge of mature forest (with shrub understory)

k = Decaying exposed tree roots and branches

1 = Decaying, partially buried logs

m = Spruce forest

n = Tall grasses

o = Vegetated spoilbank (gentle slope)

p = Vegetated ditchside

q = Non-vegetated man-made disturbed area

r = Edge of mature forest

s = Hummocky, richly vegetated area

t = Poorly drained flat bottom of shallow borrow pit

u = Elevated area left around base of large tree

v = Tension zone (ecotone); transition zone from bog to upland (dense low shrubby

vegetation)

w = Cutbank of borrow pits resulting from excavation

x = Spoilbank in the process of becoming vegetated

y = Flat vegetated flood plain of river

1). Recently disturbed, flat, unvegetated areas with friable (often sandy) soil ( q , Fig. 5

and 7). - These areas border the highway in dugouts, borrow pits, sand and gravel pits (Fig.

5), or border lake shores (Fig. 7) and river banks and were in localities I, VI, VII, XI, XII,

XIII and XIV.

Subarctic Sphecidae

17

Fig. 1-2. Flood plain of Mackenzie River with flat, low vegetated areas (Fig. 2) and juxtaposition of spoilbanks and

forest edge habitats, of sandy soil (Fig. 1). (Ft. Providence: X, 9 August 1967). Fig. 3. Shallow borrow pit along

highway, in the process of becoming vegetated; friable soil. (Frank Channel: XIV, 8 August 1967)

18

Steiner

Fig. 4-5. Man-made gravel/sand pit, along highway, with friable soil: general view (Fig. 5) and detail of gently sloping

area (Fig. 4) with various microhabitats. (Birch Lake: XI, 2 August 1967). Fig. 6. Deep borrow pit in compact, poorly

drained soil, with tension zone between bog and upland. (Rae: XV, 8 August 1967)

Subarctic Sphecidae

19

Fig. 7-9. Disturbed areas on sandy lake shore areas (Fig. 7-8) near forest edges, and extensive man-made bare and poorly

drained area, along the highway (Fig. 9). (Prelude Lake: XIII, 7 August 1967)

20

Steiner

If it is situated adjacent to a vegetated area this habitat is used occasionally by wasps for

basking early in the morning. Those most frequently observed were members of Astata

nubecula , members of the genera Ammophila, Podalonia and Tachysphex and a few pom-

pilids and crabronines. A few females of Tachysphex terminatus were seen digging in such

areas, but usually in sparsely vegetated areas.

2) . Recently disturbed, sloping, unvegetated areas of friable soil (h, Fig. 5 and 9). - These

often consist of steep ( h , Fig. 5) or gently sloping, man-made, collapsed cutbanks and spoil-

banks and were found in localities I, VII, XI (Fig. 5, h), XII, and XIII (Fig. 9, h).

Such areas resemble habitat type 1 but, when favorably exposed to the morning sun,

are used more consistently by wasps for basking. Males of Astata nubecula were at site h

(Fig. 5) perched on prominent features of the landscape such as boulders and stumps that

were also used as observation posts. Specimens representing the other taxa listed for habitat

1 were found in habitat 2, also.

3) . Man-made, poorly-drained, flat bottoms of borrow pits, usually in compact soil ( t ,

Fig. 6 and 9). - This habitat was extensive in locality XIII ( t , Fig. 9), less so in locality XV

( t , Fig. 6).

A relatively large amount of soil moisture rendered this type of habitat generally unsuit-

able for wasps, although a few females of Ammophila and Podalonia carried prey across

such areas. Drier sections of this habitat were used for nesting by a few females of some

species, such as Cerceris nigrescens (XV, Fig. 6), which usually nest in habitats with more

friable soil (XI, Fig. 4 and 5).

4) . Flat or gently sloping, sparsely-vegetated areas ( e in most Figures, y in Fig. 2). —

Usually of friable soil, these areas are: (a) man-made, occurring in pits (Fig. 4 and 5); or

(b) natural, on flood plains of rivers (y, Fig. 2) and lake shores. They were extensively

represented in most localities, particularly along flood plains of the Mackenzie (IX and X)

and of the Hay Rivers (VI), and shores of Great Slave Lake (XIV) and Prelude Lake (XIII).

These areas were used extensively as nesting grounds by females of Ammophila, Podalo-

nia, Tachysphex (particularly T. aethiops, the most common species of this genus), Nysson ,

Cerceris and some Crabroninae. A few nests of Oxybelus were seen here, but they were

more common on more sloping ground. Females of small species of Pemphredoninae and

Crabroninae, which prey on small insects, used these areas mainly for hunting and less

frequently for basking and feeding. Females of the larger wasp species hunted mainly in

more densely vegetated areas, which had a more abundant flora and insect fauna.

5) . Flat areas, densely covered with herbaceous vegetation (c in most Figures). — Most

localities included this type of habitat, but it was best developed along the Mackenzie River

(X, Fig. 1 and 2).

An abundant and diverse insect fauna was found in these areas, including acridoid grass-

hoppers, leafhoppers, small beetles, many flies, bugs, lepidopterous caterpillars, and repre-

sentatives of other groups. Most flowers on which the sphecid wasps fed occurred in these

areas.

Although many wasps hunted and fed in this type of habitat, nests were rare. Sphecids of

the genus Tachysphex and pompilids used this habitat most intensively as hunting ground.

Females of Tachysphex prey mainly on the immature acridids and tetrigids, abundant in

these places. Other hunters observed were females of Ammophila and Podalonia species.

Patches of fireweed (Epilobium, f, Fig. 1, 2, 4 and 8), yarrow ( Achillea millefolium , d ,

Fig. 1 and 2), and undetermined umbellifers, common in this habitat, were intensively ex-

ploited as a source of food by wasps of many species, particularly in the morning.

6) . Low spoilbanks and man-made ridges of friable soil in the process of becoming vege-

tated ( b , Fig. 1 and 2; o, Fig. 4; x, Fig. 5, 6 and 9). - This type of habitat provides nesting

Subarctic Sphecidae

21

grounds for many species. Along the Mackenzie River (X, Fig. 1 and 2) these areas were

used as nesting grounds by females of Oxybelus uniglumis quadrinotatus, Mimesa pauper,

Mimesa clypeata, Dryudella picta and Diploplectron peglowi. Dryudella males exhibited

territorial behavior, as described for Astata nubecula (Evans, 1970: 487-488). The only

specimens of Miscophus americanus from the study areas were found in a similar habitat

(locality XI).

These areas were used extensively by female insects which prey on sphecids: the inquili-

nous sphecids of the genus Nysson \ chrysidid wasps; and parasitoid Diptera. Other sphecids

did not extensively use these areas as hunting grounds.

Patches of yarrow {d, Fig. 1) and fireweed (/, Fig. 2) growing in these areas were ex-

ploited for food by many wasps. Basking was observed on the spoilbanks. On spoilbanks

situated adjacent to shrubby and sparsely-forested habitats (a, Fig. 1), interactions between

and within species were most intense, frequent and complex due to the addition of Pem-

phredoninae and Crabroninae, characteristic of shrubby habitats.

7). Shrubby, brushy and sparsely-forested areas, with understory {a, i, j, r, s, v, Fig.

1-9). — Such habitat includes stands of aspen interspersed with willow and shrub (a, Fig. 1),

edges of mature forest with shrub understory (/, Fig. 3, 5 and 9; r. Fig. 7) and ecotones

(tension zones) such as those between bog and upland, having dense, low, shrubby vege-

tation (v, Fig. 6).

These areas provide both nesting and hunting sites for those pemphredonines and crabro-

nines that are twig nesters, such as Mimesa sp. and Pemphredon bipartior for the former,

Ectemnius for the latter. But they provide only hunting sites for those crabronines that are

ground nesters, such as Crabro latipes. An occasional nest of Tachysphex aethiops and of

Podalonia robusta was found on the ground, but only in thinly-vegetated areas or at the

edges of this habitat type. Females of Crabro latipes and Ectemnius nigrifrons hunted in the

leaves of shrubs. Females of the philanthine Cerceris nigrescens nigrescens, which prey on

small weevils (Evans, 1970: 501; Muesebeck et al., 1951: 1009; Scullen and Wold, 1969:

212), used extensively as hunting grounds area v (Fig. 6).

Males of crabronines, especially C. latipes, were numerous and were probably attracted

by the hunting females. They attempted to mate with females and with other males particu-

larly at sites a (Fig. 1) and / (Fig. 9). Males of some undetermined pemphredonines behaved

similarly. A few males of the larrine Tachysphex aethiops were also found in such a habitat,

investigating the leaves of a small Alnus bush. They were perhaps feeding, as individuals of

several other species of Larrinae are known to lick exudations of sap from various plants or

shrubs.

Between bouts of hunting, the occasional crabronine basked in this habitat, flattening

itself on an exposed leaf or against a tree trunk. Early in the morning, however, they basked

mainly on boulders or on the ground in non-vegetated or sparsely-vegetated areas adjacent

to the brushy hunting grounds.

8.) Patches of shrubby vegetation (including populations of Rosa, Salix, Betula and

Alnus) ( i , Fig. 4 and 5) or hummocky, richly-vegetated areas ( s , Fig. 8). — These areas

provide suitable situations for the twig-nesting species. They were also visited by ground-

nesting crabronines and pemphredonines during their hunting trips. The few specimens seen

or captured of Lestiphorus cockerelli , Gorytes albosignatus and Alysson triangulifer were

found in this habitat. From circumstantial evidence, I think the latter were hunting, not

nesting, here. One individual of G. albosignatus was found digging the soil in a different

habitat (site b, Fig. 2), and insofar as is known Nearctic Alysson females nest in the soil

(Muesebeck et al., 1951: 980).

22

Steiner

9) . Decaying, often partially buried logs (/, Fig. 4) and decaying exposed tree roots and

branches ( k , Fig. 5 and 8). — These sites were visited by pemphredonines and crabronines

some of which inspected abandoned galleries of wood borers (hunting or nest-seeking be-

havior?). Female crabronines of Ectemnius arcuatus and E. dives are known to nest in logs,

timber and stems (Muesebeck et al., 1951: 1026-1027). A few wasps (pemphredonines,

crabronines, Ammophila azteca and others) basked in these places, and males of Astata

nubecula used them for observation posts.

10) . Vertical banks (sand cliffs; cutbanks; river cutbanks; cutbanks of borrow pits) (w,

Fig. 6). — In locality VI, females of the small pemphredonine Diodontus were found in-

specting, both in flight and by walking, such cutbanks and the burrows therein, possibly

for prey or for nesting sites. Other wasps found here consistently were: a small crabronine,

undetermined, possibly a member of the genus Crossocerus ; some vespids; and some chrysi-

dids. Occasional basking was also observed here.

11) . Dense, spruce-jackpine (g, Fig. 3, 5, 6, 7 and 8) and spruce stands ( m , Fig. 4 and 9). —

These areas were seldom visited by wasps.

Discussion. — When nesting, hunting, feeding and basking sites were close together, as in

locality X (Fig. 1), conditions appeared particularly favorable for many wasps. Indeed, the

speed with which prey capture and nest provisioning can be completed may be crucial,

particularly in northern climates where favorable weather is limited (Evans, 1970).

Considerable segregation in habitat and microhabitat was noticed among sphecid wasps,

much more so than among the spider wasps studied previously (Steiner, 1970). Competition

among species is thus considerably reduced, and is further reduced by behavioral specializa-

tion in prey selection and hunting techniques (Evans, 1970). Further segregation is achieved

by some stratification of the nest cells in the soil, at different depths according to species

(Evans, 1970). However, there was little species segregation at feeding and at basking sites.

ANNOTATED LIST OF SPECIES

Family Sphecidae

Subfamily Astatinae

Genus Diploplectron Fox

Known biology. - Females nest in open sandy places (Evans, 1957: 180). Prey consists

of adults or nymphs of Hemiptera. Williams (1946: 648) provides details.

Known distribution. — North America and South Africa (Evans, 1957: 180).

1 . Diploplectron peglowi Krombein.

Known distribution. — New York: Oswego Co. (Muesebeck et al., 1951: 939).

Author’s records. - NORTHWEST TERRITORIES. - X, 6, 2 99 31 July 1967. XIII,

2 66, 9 6 August 1967 (9 hunting); 6 7 August 1967.

Genus Astata Latreille

Known biology. - Females of this genus provision their nests with Hemiptera, especially

pentatomids and lygaeids (Muesebeck et al., 1951: 939).

2. Astata nubecula Cresson.

Known biology. — Females nest in bare, hard, stony soil, and provision their nests with

immature stink bugs ( Chlorochroa uhleri Stahl) carried in flight (Evans, 1970). Nymphs

Subarctic Sphecidae

23

of Thyanta have also been recorded as prey (Krombein et al., 1967: 387). Males exhibit

territorial behavior (Evans, 1970).

Known distribution. — Western United States, Oregon to New Mexico north to Idaho

(Muesebeck et al., 1951: 940). Wyoming: Jackson Hole (Evans, 1970). Alberta (Strick-

land, 1947: 129).

Author’s records. - NORTHWEST TERRITORIES. - VIII, d 10 August 1967 (returns

and lands on exactly the same spot); 9 2 August 1967 (exploring). XIII, 3 66 30 July

1968 (each lands on conspicuous, elevated objects such as stumps, fallen logs and rocks);

2 66, 9 7 August 1967. XIV, 6 8 August 1967 (returns and lands on same spot).

Genus Dryudella Spinola

3. Dryudella picta (Kohl).

Author’s records. - NORTHWEST TERRITORIES. - X, d, 9 17 July 1968 {in copula ,

the male carrying the female under him in flight in prey-like fashion, and landing from

time to time); <5 21 July 1967 (basking in the morning sun). XIII, 2 66 1 August 1967

(returning to and landing on same spots, repeatedly).

Subfamily Larrinae

Tribe Miscophini

Genus Miscophus Jurine

Known biology. — Females nest in sandy soil and provision their nest cells with tiny

spiders transported in a series of low flights (Kurczewski, 1 969).

4. Miscophus americanus (W. Fox).

Known biology. — Kurczewski (1969) presented a detailed study of the biology of this

species. Notes were published by Hartman (1905: 69-70), and Krombein (1952b: 328).

Adults are found on sand, on bare places and in woods. Females nest in well-packed sand

in slightly sloping areas. Prey consists of immature spiders of the families Epeiridae (Hart-

man, 1905) and Theridiidae (Kurczewski, 1969). The latter author provides data on prey

hunting behavior, nest provisioning, nest structure, and position of the egg on the prey.

Known distribution. — Eastern North America, from New York south to Florida and

west to Colorado (Muesebeck et al., 1951: 944) and Alberta (Strickland, 1947: 129).

Author’s records. - NORTHWEST TERRITORIES. - XI, 2 66 2 August 1967.

Tribe Tachytini

Genus Tachysphex Kohl

Known biology. — Prey species are orthopteroids, mostly immature acridids, but females

of some species also capture mantids and blattids (Muesebeck et al., 1951: 950).

5. Tachysphex aethiops (Cresson).

Known biology. - A nest was found on flat, friable sand 2 meters from a river bank.

The prey therein was an immature acridid grasshopper of the genus Trimerotropis (Evans,

1970: 489-490).

Known distribution. — Western United States in mountains (Muesebeck et al., 1951:

950). Wyoming: Jackson Hole (Evans, 1970).

Author’s records. - NORTHWEST TERRITORIES. - VI, d, 9 29 July 1967; d 12

24

Steiner

August 1967. X, 9 1 1 August 1968 (apparently hunting);? 25 August 1967. XII, 3 99 29

July 1968 (hunting and nest digging in sandy hillocks). XIII, 2 66, 2 99 (former investiga-

ting leaves of small Alnus bush) 30 July 1968; 9 6 August 1967. XIV, 2 66 8 August 1967.

6. Tachysphex quebecensis Provancher.

Known biology. — Prey: immature acridids of the genera Camnula and Melanoplus

(Krombein et al., 1967: 393).

Known distribution. — Transcontinental in northern United States and southern Cana-

da, Quebec to California, in north (Krombein et al., 1967: 393; Muesebeck et al., 1951:

952).

Author’s records. - NORTHWEST TERRITORIES. - X, 6 25 July 1967; 9 26 July

1967. XII, 9 29 July 1968. XIV, 6, 9 26 July 1968.

7. Tachysphex terminatus (F. Smith)

Known biology. - Adults are found in woods and in barrens (Krombein et al., 1952b:

330). Nests are in open places, in sand or in loose topsoil (Krombein et al., 1958: 188;

Muesebeck et al., 1951: 953). Prey includes tetrigids and immature acridids of many

genera (Muesebeck et al., 1951 : 953). The latter group includes Melanoplus and Tryxalus

(Krombein et al., 1958: 188; Kurczewski, 1966a), and Chorthippus curtipennis (Harris)

(Evans, 1970: 491). Female closes entrance while away from burrow (Evans, 1970). Male

behavior was studied by Kurczewski (1966b). Data about population ecology of this spe-

cies are recorded by Kurczewski and Harris (1968).

Known distribution. — Transcontinental, in the north from Quebec to British Colum-

bia; southward in the east to Georgia and westward to Arizona (Krombein et al., 1967:

393; Muesebeck et al., 1951: 953). Alberta (Strickland, 1947: 129).

Author’s records. - NORTHWEST TERRITORIES. - XIII, 9 30 July 1968 (starting

nest digging in the ground). XIV, 2 66 8 August 1967.

Subfamily Pemphredoninae

Tribe Psenini

Genus Diodontus Curtis

Known biology. — Nests are in cavities in logs, or stems of plants, such as canes of Rubus

(Muesebeck et al., 1951: 958; Spooner, 1948: 129-172). Females of Palaearctic species prey

on aphids and psyllids, which are carried ventrally by the middle legs (Spooner, 1948).

Prey records are not available for the Nearctic species.

8. Diodontus sp. (or spp.?).

Author’s records. - NORTHWEST TERRITORIES. - III, 9 14 August 1967. VI, 3 66,

2 99, 29 July 1967 (investigating vertical cutbanks); 6, 9 12 August 1967.

Genus Mimesa Shuckard

Subgenus Mimesa ( sensu stricto )

Known biology. - Females nest in soil. Prey consists of cicadellids, which the female

carries with her middle legs (Muesebeck et al., 1951: 958; Spooner, 1948).

9 . Mimesa pauper Packard.

Known biology. - Adults at edge of woods on high foliage (Kurczewski and Kurczew-

Subarctic Sphecidae

25

ski, 1963:146).

Known distribution. — Transition zone east of Rocky Mountains (Muesebeck et al.,

1951: 960).

Author’s records. - NORTHWEST TERRITORIES. - X, 9 1 1 July 1968 (investigating

insect burrows, holes, in the soil); 3 99 31 July 1967 (same remark). XII, one (sex?)

29 July 1968 (same remark). XIII, 4 66 (on leaves of small trees) 7 August 1967. XIV,

6 26 July 1968; 4 99 8 August 1967.

Subgenus Mimumesa Malloch

Known biology. — Females of Palaearctic species nest in cavities in logs and stems. Their

prey consists of delphacids and cicadellids, which the female wasp carries in her mandibles

(Spooner, 1948). Data about biology are not available for Nearctic species.

10. Mimesa clypeata (W. Fox).

Known distribution. — Western North America: California and Nevada north to

Alaska (Muesebeck et al., 1951: 961).

Author’s records. - NORTHWEST TERRITORIES. - IX, 9 1 August 1967 (investi-

gating the ground, holes, burrows). X, 9 26 July 1967.

1 1 . Mimesa species.

Author’s records. - NORTHWEST TERRITORIES. - X, 9 22 July 1967 (flying

around shrubs). XII, 6 4 August 1967. XIV, 299 8 August 1967 (flying around young

aspen and big boulders).

Tribe Pemphredonini

Genus Pemphredon Latreille

Known biology. — Females nest in twigs, deserted galls, abandoned beetle burrows, or in

rotten wood, and provision the cells with bugs of the family Aphidae (Muesebeck et al.,

1951: 965).

12. Pemphredon bipartior W. Fox.

Known biology. — Nests in twigs of sumac and elder. Prey: lEriosoma lanigerum

(Hausmann); Rhopalosiphum rhois Monell (Muesebeck et al., 1951: 966).

Known distribution. — Eastern United States, New York to Texas (Muesebeck et al.,

1951: 966).

Author’s records. - NORTHWEST TERRITORIES. - XIII, 2 66 7 August 1967.

13. Pemphredon montana Dahlbom.

Known distribution. — Holarctic. Nearctic Region: British Columbia. Palaearctic

Region: Europe (Muesebeck et al., 1951: 965).

Author’s records. - NORTHWEST TERRITORIES. - VII, 9 mid-July 1968.

Subfamily Sphecinae

Tribe Ammophilini

Genus Ammophila Kirby

14. Ammophila azteca Cameron.

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Steiner

Known biology. — Females prey on small larvae of sawflies and moths (recorded

families — Geometridae, Gelechiidae, Sphingidae) and carry the prey in flight (Evans,

1970: 485). Evans (1965) presented a detailed study of the biology of this species.

Known distribution. — Wyoming: Jackson Hole (Evans, 1970).

Author’s records. - NORTHWEST TERRITORIES. - III, d 14 August 1967. V,

9 14 August 1967. VI, 9 29 July 1967; 9 12 August 1967. VII, 9 11 August 1967.

IX, d 1 August 1967. X, 9 1 1 July 1968; 6, 9 17 July 1968; 3 dd, 9 22 July 1967

(feeding on Achillea sp.); 9 25 July 1967; d, 9 (in copula) 26 July 1967. XI, d, 9

2 August 1967; 9 8 August 1967. XII, 6 7 August 1967. XIII, 3 dd, 2 99 6 August

1967; 9 7 August 1967. XIV, d 26 July 1968; 6, 9 8 August 1967. XV, 9 8 August

1967. YUKON TERRITORY. - B, 9 15 August 1968. E, 3 99 10 August 1968.

G, 3 99 12 August 1968.

15. Ammophila mediata Cresson.

Known distribution. — Ontario to British Columbia, Michigan to Colorado (Muese-

beck et al., 1951: 976). Wyoming: Jackson Hole (Evans, 1970). Alberta (Strickland,

1947: 129).

Author’s records. - NORTHWEST TERRITORIES. - I, 2 99 15 August 1967.

IV, 9 28 July 1967. X, 6, 9 21 July 1967. XIII, 9 30 July 1968. XIV, d 26 July

1968; 9 8 August 1967. YUKON TERRITORY. - B, 9 15 August 1968. C, 9 9 August

1968.

16. Ammophila strenua Cresson.

Known distribution. - Western United States (Muesebeck et al., 1951: 977). Wyo-

ming: Jackson Hole (Evans, 1970).

Author’s records. - NORTHWEST TERRITORIES. - X, d 21 July 1967.

Genus Podalonia Spinola

17. Podalonia luctuosa (F. Smith).

Known distribution. - Canada, United States — western and northern tier of states

to Maine (Muesebeck et al., 1951: 977). Wyoming: Jackson Hole (Evans, 1970).

Alberta (Strickland, 1947: 128).

Author’s records. - NORTHWEST TERRITORIES. - II, d 14 August 1967. Ill,

d 14 August 1967. YUKON TERRITORY. - McGregor Creek, d 10 August 1968.

18. Podalonia robusta (Cr.)

Known biology. — Prey: noctuid larva (Krombein et al., 1967: 404 , see also Evans,

1963: 237).

Known distribution. — Canada; United States - western and northern tier of states

to Maine (Muesebeck et al., 1951: 978). Alberta (Strickland, 1947: 128).

Author’s records. - NORTHWEST TERRITORIES. - VI, 9 29 July 1967. IX, d

1 August 1967. X, 6 66, 9 11 July 1968; 9 17 July 1968; d, 9 22 July 1967; 3 99

25 July 1967; d, 9 (in copula ), 9 31 July 1967. XII, 6, 9 29 July 1968; d 7 August

1967. XIV, 9 8 August 1968. YUKON TERRITORY. - B, 2 99 (feeding on flowers

and hunting(?) on the ground, respectively) 15 August 1968.

Also seen (but not captured): an entirely black specimen of (?) Podalonia sp.

(XII).

Subarctic Sphecidae

27

Subfamily Nyssoninae

Tribe Alyssonini

Genus Alysson Panzer

19. Alysson triangulifer Provancher.

Author’s records. - NORTHWEST TERRITORIES. - IX, 9 1 August 1967 (investi-

gating leaves, in shrubs).

Tribe Nyssonini

Genus Nysson Latreille

Known biology. — Larvae of this genus are inquilinous in the nests of other sphecid wasps

and bees (Muesebeck et al., 1951).

20. Nysson lateralis Packard.

Known distribution. — Chiefly Transition zone east of Rockies (Muesebeck et al.,

1951: 983).

Author’s records. - NORTHWEST TERRITORIES. - VII, 9 1 1 August 1967. IX, 2

99 1 August 1967. X, 9 26 July 1967. XI, 2 99 8 August 1967. XIII, 9 6 August 1967.

21. Nysson subtilis W. Fox.

Known biology. — Adults found along trail in open areas (Krombein, 1952a: 181).

Known distribution. — Pennsylvania, Illinois (Muesebeck et al., 1951: 983).

Author’s records. - NORTHWEST TERRITORIES. - X, 9 22 July 1967; 9 25 July

1967; d, 9 31 July 1967. XI, 9 2 August 1967.

Tribe Gorytini

Genus Lestiphorus Lepeletier

22. Lestiphorus cocker elli (Rohwer).

Known distribution. — Eastern United States, northern tier west to Colorado (Mue-

sebeck et al., 1951 : 988).

Author’s records. - NORTHWEST TERRITORIES. - X, 9 31 July 1967.

Genus Gorytes Latreille

23. Gorytes albosignatus W. Fox.

Known distribution. — Western United States, North Dakota to Montana, south to

Nebraska (Muesebeck et al., 1951: 991). Wyoming: Jackson Hole (Evans, 1970: 494).

Alberta (Strickland, 1947: 126).

Author’s records. - NORTHWEST TERRITORIES. - X, 9 17 July 1968 (digging in

sand hill).

Subfamily Philanthinae

Tribe Cercerini

Genus Cerceris Latreille

24. Cerceris nigrescens nigrescens F. Smith.

Known biology. - Females nest in the ground, and prey on weevils of the following

28

Steiner

taxa: Hyperodes delumbis (Gyllenhal); Sitona hispidula (Fabricius); Gymnaetron sp.;

Gymnaetron antirrhini (Paykull) (Evans, 1970: 501; Muesebeck et al., 1951: 1009;

Scullen and Wold, 1969: 212). Scullen (1965: 494-495) lists the names of the plant

species visited by adults of C. nigrescens.

Known distribution. — This is the most widely distributed species of Cerceris in

North America, ranging from New England and adjacent southeastern Canada west-

ward to the Pacific coast, northward to Alaska and south to Nevada and North Caro-

lina (Muesebeck et al., 1951: 1009; Scullen, 1965: 492; Scullen and Wold, 1969: 212).

Alberta (Strickland, 1947: 130).

Author’s records. - NORTHWEST TERRITORIES. - I, 9 15 August 1967. VII,

2 99 1 1 August 1967. IX, <5, 9 1 August 1967. X, d 1 1 July 1968; 3 66, 2 99 22 July

1967; d, 9 25 July 1967; 9 31 July 1967. XI, 3 99 2 August 1967; 9 8 August 1967.

XII, d 4 August 1967. XIII, 9 30 July 1968; 9 6 August 1967. XIV, d 26 July 1968;

2 99 8 August 1967. YUKON TERRITORY. - E, d, 3 99 10 August 1968.

Subfamily Crabroninae

Tribe Crabronini

Genus Crabro Fabricius

Known biology. — Females nest principally in soil, though occasionally in rotten wood

and prey on flies (Muesebeck et al., 1951: 1015). Kurczewski and Acciavatti (1968) review

nesting behavior of the Nearctic species.

25. Crabro latipes F. Smith.

Known biology. - A detailed study of nesting behavior is provided by Kurczewski,

Burdick and Gaumer (1969). Nests are in open areas with sparse vegetation. Prey con-

sists of a wide variety of average-size flies of rather stocky build, such as individuals

of Musca domestica Linnaeus (Muesebeck et al., 1951: 1017) and Musca autumnalis

DeGeer (Kurczewski and Harris, 1968).

Known distribution. — Transcontinental in the north, in Canada, Alaska and the

Canadian and Transition zones of conterminous United States (Muesebeck et al., 1951:

1017). Alberta (Strickland, 1947: 127).

Author’s records. - NORTHWEST TERRITORIES. - VI, d, 4 99 29 July 1967.

VII, d 1 1 August 1967. X, d, 9 1 1 July 1968; d 17 July 1968 (on leaves of willows?

pouncing on other males of apparently the same species); 9 21 July 1967 (investigating

holes in the ground); 9 22 July 1967; 9 25 July 1967; d 26 July 1967; 2 66, 9 31 July

1967. XI, 3 66 2 August 1967. XII, 9 7 August 1967 (on leaves, shrub). XIII, d 6

August 1967 (on leaves, shrub); 4 66 1 August 1967. XIV, 2dd 8 August 1967 (on

leaves of small Alnus ?). YUKON TERRITORY. - G, d 12 August 1968.

26. Crabro sp.

Author’s records. - NORTHWEST TERRITORIES. - XIII, 9 30 July 1968.

Genus Crossocerus Lepeletier and Brulle

Known biology. - Females usually nest in soil, occasionally in cracks in walls or in aban-

doned beetle burrows in wood. Prey consists of small flies (Muesebeck et al., 1951: 1020).

27. Crossocerus species.

Subarctic Sphecidae

29

Author’s records. - NORTHWEST TERRITORIES. - IX, 31 July 1967. X, d,

3 99 31 July 1967. XIV, 9 8 August 1967.

Genus Ectemnius Dahlbom

28. Ectemnius arcuatus Say.

Known biology. — Females nest in logs. They prey on flies of the species Musca

domestica L. (Muesebeck et al., 1951: 1026; under the name Hypocrabro chrysargirus

(Lepeletier and Brulle)).

Known distribution. — Transcontinental in Transition and Austral zones (Muesebeck

et al., 1951: 1026; under the name Hypocrabro chrysargirus (Lepeletier and Brulle)).

Author’s records. - NORTHWEST TERRITORIES. - X, 9 17 July 1968 (feeding

on Epilobium sp. flowers).

29. Ectemnius dives (Lepeletier and Brulle).

Known biology. — Females nest in logs, timber and stems. Prey consists of muscoid

Diptera (Muesebeck et al., 1951: 1027). Kurczewski and Kurczewski (1963: 148)

observed males on flowers of Daucus carota and Achillea millefolium.

Known distribution. — Holarctic. Nearctic Region: transcontinental in Canadian

and Transition zones of Canada and United States (Muesebeck et al., 1951: 1027).

Alberta (Strickland, 1947: 127). Palaearctic Region: Germany, Austria, Switzerland

and Morocco (Leclercq, 1949: 11).

Author’s records. - NORTHWEST TERRITORIES. - X, 2 66 1 1 July 1968; d

25 July 1967; 2 66 26 July 1967; 3 99 31 July 1967.

30. Ectemnius lapidarius (Panzer).

Known biology. — Adults are in open woods and at the edge of woods, on flowers

of Solidago sp., and of Daucus carota (Kurczewski and Kurczewski, 1963: 148).

Known distribution. — Holarctic. Nearctic Region: Pennsylvania (Kurczewski and

Kurczewski, 1963: 148). Wyoming: Jackson Hole (Evans, 1970: 492). Alberta (Strick-

land, 1947: 127). Palaearctic Region: Finland, Germany and Austria (Leclercq, 1949).

Author’s records. - YUKON TERRITORY. — Lake Labarge, 9 6 August 1968. G,

9 12 August 1968.

31. Ectemnius nigrifrons (Cresson).

Known biology. — Recorded as prey is the fly species Syrphus ribesii (Linnaeus)

by Muesebeck et al., (1951: 1024; under the name Clytochrysus nigrifrons).

Known distribution. — Holarctic. Nearctic Region: transcontinental, chiefly in Tran-

sition zone (Muesebeck et al., 1951: 1024; under the name Clytochrysus nigrifrons).

Alberta (Strickland, 1947: 127). Palaearctic Region: Switzerland (Leclercq, 1949).

Author’s records. - NORTHWEST TERRITORIES. - II, 9 14 August 1967. V,

9 14 August 1967. IX, 6, 2 99 1 August 1967. X, 9 1 1 July 1968; 2 dd 17 July 1968.

XIII, 9 7 August 1967. YUKON TERRITORY. - C, 2 99 9 August 1968 (stalking

behavior: react to slightest movements by orienting responses; also intense visual scan-

ning; hunting behavior?); 2 99 10 August 1968 (same remark, concerning hunting

behavior?). E, 9 10 August 1968. G, 2 99 12 August 1968.

32. Ectemnius trifasciatus (Say).

Known distribution. — Transition zone of Canada and United States, east of the

30

Steiner

Cascade and Sierra Nevada ranges (Leclercq, 1949: 11; Muesebeck et al., 1951: 1027,

under the name Hypocrabro trifasciatus ). Alberta (Strickland, 1947: 127).

Author’s records. - NORTHWEST TERRITORIES. - III, 9 14 August 1967. X,

2 99 (investigating the ground) 17 July 1968. XIV, 9 26 July 1968.

33. Ectemnius species.

Author’s records. - NORTHWEST TERRITORIES. - X, 9 31 July 1967. XIII, 9

7 August 1967.

Genus Lestica Billberg

34. Lestica producticollis (Packard).

Known biology. — One 9 found in woods (Krombein, 1952b: 338).

Known distribution. — Transcontinental; in Canada and United States in Transition

and Upper Austral zones (Muesebeck et al., 1951: 1028; under the name Solenius pro-

ducticollis (Packard)). Alberta (Strickland, 1947: 127).

Author’s records. - NORTHWEST TERRITORIES. - X, 6 9 August 1967.

Tribe Oxybelini

Genus Oxybelus Latreille

35. Oxybelus uniglumis quadrinotatus Say.

Known biology. — Females dig nests in light, friable sand (Evans, 1970: 493). Prey

consists of flies of the following taxa: Symphoromyia sp ., Musca domestica Linnaeus;

Ophyra leucostoma Wiedemann; Sarcophaga rapax Walker; Hylemya cilicrura (Ron-

dani), and other muscids, and anthomyiids (Evans, 1970: 493; Muesebeck et al., 1951:

1033, under the name Oxybelus quadrinotatus). Evans (1962: 477) presents a detailed

study of prey-carrying behavior.

Known distribution. — Generally distributed throughout the United States and south-

ern Canada (Muesebeck et al., 1951: 1033; under the name Oxybelus quadrinotatus).

Author’s records. - NORTHWEST TERRITORIES. - V, 9 14 August 1967. X, 9

1 1 July 1968 (digging in sand); 9 25 July 1967 (digging in sand); d, 9 26 July 1967;

2 99 31 July 1967. XI, 9 2 August 1967. XIV, d 26 July 1968.

DISTRIBUTION PATTERNS OF THE SUBARCTIC SPHECID

FAUNA AND FACTORS AFFECTING ITS DIVERSITY

The samples of sphecid wasps from the Yukon and Northwest Territories here discussed

indicate that the fauna is impoverished in terms of number of taxa compared with that of

more southern climates. These subarctic samples comprise 35 species in 21 genera or sub-

genera. In contrast, at Jackson Hole, Wyoming (44°N., 5750 feet above sea level at Moran)

there are 94 species in 42 genera or subgenera (Evans, 1970), and in Alberta 160 species in

53 genera or subgenera (Strickland, 1947). The latter area is much more extensive and was

sampled over a longer period of time than were the subarctic areas described here, so part

of the difference in diversity between the two must be the result of these factors.

That the number of species included in most genera is higher farther south than in sub-

arctic areas is illustrated by the following examples. Following each generic name is, first,

the number of species from Jackson Hole, Wyoming, and second, the number of species in

the subarctic samples: Podalonia, 6 vs. 2; Ammophila, 9 vs. 3; Tachysphex, 5 vs. 3; Crabro,

Subarctic Sphecidae

31

4 vs. 2; Ectemnius, 9 vs. 6.

Although these comparisons demonstrate faunal impoverishment over a wide range of

latitudes, this phenomenon is not so well marked within the limits of the study area, except

perhaps in the Yukon, where the study area extended almost 2° of latitude farther north

than in the Northwest Territories study area (see Table 1 ; localities are listed from left to

right, in order of increasing latitude for each study area). A more conclusive comparison

should, however, involve both standardization of the conditions of sampling in time, space,

and season; and rating of the localities in terms of vegetation, local climate, and soil

conditions.

Comparison between rows rather than columns is probably more reliable: it gives an

indication of how common and/or widespread each species is, assuming that sampling biases

are equally distributed over species or nearly so.

Some groups of sphecids, represented by an abundance of species southward have few

species in subarctic regions. For example, there are few Sphecinae in the sample. Of four

tribes in this subfamily, only one, the Ammophilini, is represented in the study area. One

ammophiline, Ammophila azteca, was one of the commonest and most widespread sphecids

encountered. Wasps of this genus are also at high latitudes and altitudes in Europe. Another

subfamily with many taxa farther south is the Nyssoninae. Of six tribes, only three are

represented in the subarctic study areas.

In contrast, the subfamily Crabroninae is well represented, both in numbers of species

and in numbers of individuals in these samples (10 species from a total of 35 sphecid wasps,

or almost a third; and 39 species from a total of 160 Albertan species, or a quarter). Some

subarctic crabronines are Holarctic.

Other wasp groups from the study areas with northern affinities are Dryudella (subfamily

Astatinae), some Gorytini (species 22 and 23), and some Pemphredoninae (species 10 and

13). The last-named subfamily is represented by six species in my samples. Like crabronines,

pemphredonines are very abundant and widely distributed over the study areas.

Some of the species represented in the samples are widely distributed on the North

American continent, particularly latitudinally. Some are transcontinental such as Astata

nubecula, (ranging southward to New Mexico and California) and Miscophus americanus

(ranging southward to Florida and Texas). Wasps of the tribe Tachytini are well represented

in temperate as well as tropical areas of the world. Many are wide-ranging on this continent,

for example, Tachysphex quebecensis and T. terminatus (species 6 and 7). Another species

of the genus, T. aethiops, is, however, restricted to relatively high latitudes or altitudes.

Cerceris nigrescens and Oxybelus uniglumis also range widely. Among pemphredonines,

Mimesa clypeata (species 10) and Pemphredon bipartior (species 12) are wide-ranging, as are

a number of crabronines (species 25, 28, 30, 31, 32 and 34).

Routes of dispersal are probably river valleys for southern-based species, and the north-

south trending mountain systems for northern-based species.

In conclusion, the subarctic sphecid fauna comprises elements derived from cold-adapted

groups as well as wide-ranging species probably derived from warm-adapted groups. Al-

though diversity is limited, the fauna is nonetheless quite varied for high latitudes. What

makes it possible for these insects to live so far north? We do not know, but factors can be

suggested, in general terms. Local conditions of climate, soil, vegetation and microclimates

are likely important (Corbet, 1969; Geiger, 1965; Uvarov, 1931). An important behavioral

adaptation is probably that of basking, which enables a flying insect to accumulate suffi-

cient solar energy even when the air temperature is quite low (Baker and Hurd, 1968;

Clench, 1966; Digby, 1965; Downes, 1964; Hocking and Sharplin, 1965; Kevan, 1970;

Kevan and Shorthouse, 1970; Monroe, 1956; Parry, 1951; Richards, 1970). Physiological

32

Steiner

adaptations to the cold which make possible survival through the winter are also probably

important (Aoki, 1956; Asahina, 1959, 1966, 1969; Dubach et al., 1959; Losina-Losinsky,

1962; Salt, 1961; Scholander et al., 1953; Smith, 1961; S^mme, 1964; Tanno, 1964;

Ushatinskaya, 1957).

The next phase of study of the subarctic sphecid fauna should aim at elucidating these

factors.

ACKNOWLEDGEMENTS

This study was supported in part by the Heart Lake Biological Station; the Department

of Zoology, University of Alberta; NRC grant A 3499; and Boreal Institute, University of

Alberta, grant GR-1, 1968-70.

I wish to express my gratitude to W. A. Fuller, Director of the Heart Lake Biological

Station; H. E. Evans, Museum of Comparative Zoology, Harvard University, Cambridge,

Massachusetts; K. V. Krombein, Smithsonian Institution, Washington, D. C.; B. Hocking,

G. E. Ball, W. G. Evans, B. S. Heming, P. Kevan, and R. MacArthur, University of Alberta,

for their help and suggestions at various stages of this project and during the preparation

of the manuscript. J. S. Scott assisted with preparation of the photographs for publication.

I am also indebted to the following persons for assistance with determination of my

specimens: R. M. Bohart, University of California, Davis, California, (Sphecidae); A. S.

Menke, Smithsonian Institution, and the United States Department of Agriculture, Belts-

ville, Maryland, ( Ammophila species); F. D. Parker, Utah State University, Logan, Utah,

( Diploplectron species); and H. A. Scullen, Oregon State University, Corvallis, Oregon,

(Cercerini).

REFERENCES

Aoki, K. 1956. The undercooling point and frost resistance in the prepupa of a ruby-tailed

wasp, Crysis (Pentacrysis) shanghaiensis [in Japanese]. Low Temp. Sci. Ser. B. 14:121-124.

Asahina, E. 1959. Cold hardiness in overwintering insects [in Japanese], p. 99-113. In

M. Fukaya, M. Harizuka and K. Takewaki [ed.] Recent advances in experimental mor-

phology. Yokendo: Tokyo.

Asahina, E. 1966. Freezing and frost resistance in insects, p. 451-486. In H. T. Meryman

[ed.] Cryobiology. Acad. Press, London.

Asahina, E. 1969. Frost resistance in insects, p. 1-49. In J. L. Beament, J. E. Treheme and

V. B. Wigglesworth [ed.] Advances in insect physiology. Acad. Press, London.

Baker, H. G. and P. D. Hurd. 1968. Interfloral ecology. A. Rev. Ent. 13:385-414.

Clench, H. K. 1966. Behavioral thermoregulation in butterflies. Ecology 47:1021-1034.

Corbet, P. S. 1969. Terrestrial microclimate: ameliorations at high latitudes. Science 166:

865-866.

Digby, P. S. B. 1955. Factors affecting temperature excess of insects in sunshine. J. exp.

Biol. 32:279-298.

Downes, J. A. 1964. Arctic insects and their environment. Can. Ent. 96:279-307.

Dubach, P., F. Smith, D. Pratt and C. M. Stewart. 1959. Possible role of glycerol in the

winter-hardiness of insects. Nature, Lond. 184:288-289.

Evans, H. E. 1957. Ethological studies on digger wasps of the genus Astata (Hymenoptera,

Sphecidae). J. N. Y. ent. Soc. 65:159-185.

Evans, H. E. 1962. The evolution of prey-carrying mechanisms in wasps. Evolution. Lan-

caster, Pa. 16(4):468-483.

Subarctic Sphecidae

33

Evans, H. E. 1963. Notes on the prey and nesting behavior of some solitary wasps of Jack-

son Hole, Wyoming. Ent. News 74(9): 233-239.

Evans, H. E. 1965. Simultaneous care of more than one nest by Ammophila azteca Cameron

(Hymenoptera, Sphecidae). Psyche, Camb. 72:8-23.

Evans, H. E. 1970. Ecological-behavioral studies of wasps of Jackson Hole, Wyoming. Bull.

Mus. Comp. Zool. 140(7):45 1-5 1 1 .

Geiger, R. 1965. The climate near the ground. Harvard Univ. Press, Cambridge, Mass, xiv +

611 p.

Hartman, F. 1905. Observations on the habits of some solitary wasps of Texas. Bull. Univ.

Texas, No. 65. 72 p., 4 plates.

Hocking, B. and C. D. Sharplin. 1965. Flower basking by Arctic insects. Nature, Lond.

206(4980):215.

Kevan, P. G. 1970. High Arctic insect flower relations: the inter-relationships of arthropods

and flowers at Lake Hazen, Ellesmere Island, N. W. T., Canada. Unpubl. Ph.D. thesis,

Univ. Alberta. 399 p.

Kevan, P. G. and J. D. Shorthouse. 1970. Behavioral thermoregulation by High Arctic

butterflies. Arctic 23 (4): 268-279.

Krombein, K. V. 1952a. Preliminary annotated list of the wasps of Lost River State Park,

West Virginia, with descriptions of new species and biological notes. Proc. ent. Soc. Wash.

54(4): 175-184.

Krombein, K. V. 1952b. Biological and taxonomic observations on the wasps on a coastal

area of North Carolina (Hymenoptera: Aculeata). Wasmann J. Biol. 10(3):257-341 .

Krombein, K. V. 1967. Trap-nesting wasps and bees: Life histories, nests, and associates.

Washington, D. C.: Smithsonian Press, 570 p.

Krombein, K. V. et al. 1958. First Supplement to “Hymenoptera of America ...” (see

Muesebeck, C. F. W., below).

Krombein, K. V., B. D. Burks et al. 1967. Second Supplement to “Hymenoptera of Amer-

ica .. . ” (see Muesebeck, C. V. W., below).

Kurczewski, F. E. 1966a. Tachysphex terminatus preying on Tettigoniidae — an unusual

record (Hymenoptera: Sphecidae: Larrinae). J. Kans. ent. Soc. 39:317-322.

Kurczewski, F. E. 1966b. Comparative behavior of male digger wasps of the genus Tachy-

sphex (Hymenoptera: Sphecidae: Larrinae). J. Kans. ent. Soc. 39(3):436-453.

Kurczewski, F. E. 1969. Comparative ethology of female digger wasps in the genera Misco-

phus and Nitelopterus (Hymenoptera: Sphecidae, Larrinae). J. Kans. ent. Soc. 24(4):

470-509.

Kurczewski, F. E. and R. E. Acciavatti. 1968. A review of the nesting behaviors of the

nearctic species of Crabro, including observations on C. advenus and C. latipes (Hyme-

noptera: Sphecidae). J. N. Y. ent. Soc. 76(3): 196-212.

Kurczewski, F. E. and B. J. Harris. 1968. The relative abundance of two digger wasps,

Oxybelus bipunctatus and Tachysphex terminatus , and their associates, in a sand pit in

central New York. J. N. Y. ent. Soc. 76(2):81-83.

Kurczewski, F. E. and E. J. Kurczewski. 1963. An annotated list of digger wasps from

Presque Isle State Park, Pennsylvania. Proc. ent. Soc. Wash. 65(2): 141-149.

Kurczewski, F. E., N. A. Burdick and G. C. Gaumer. 1969. Additional observations on the

nesting behaviors of Crabro advenus Smith and C. latipes Smith (Hymenoptera: Spheci-

dae). J. N. Y. ent. Soc. 77(3): 152-170.

Leclercq, J. 1949. Contribution a l’etude des Crabroninae (Hym. Sphecidae) de l’Hemi-

sphere Nord. Bull. Instit. Roy. Belg. 25(16): 1-18.

Losina-Losinsky, L. K. 1962. Survival of insect at super-low temperatures. Dokl. Akad.

34

Steiner

Nauk. SSSR 147:1247-1249.

Monroe, E. 1956. Canada as an environment for insect life. Can. Ent. 88:372-476.

Muesebeck, C. F. W., K. V. Krombein, H. K. Townes et al. 1951. Hymenoptera of America

north of Mexico — Synoptic Catalog. U. S. Dept. Agric. (Agriculture Monograph No. 2),

Washington, 1420 p.

Parry, D. A. 1951. Factors determining the temperature of terrestrial arthropods in sunlight.

J. exp. Biol. 28:445-462.

Richards, K. W. 1970. Biological studies of Arctic bumblebees. Unpubl. M.Sc. thesis, Univ.

Alberta. 165 p.

Salt, R. W. 1961. Principles of insect cold-hardiness. A. Rev. Ent. 6:55-74.

Scholander, P. F., W. Flagg, R. J. Hock and L. Irving. 1953. Studies on the physiology of

frozen plants and animals in the Arctic. J. cell. comp. Physiol. 42, Suppl. 1:1-56.

Scullen, H. A. 1965. Review of the genus Cerceris in America North of Mexico (Hymenop-

tera: Sphecidae). Proc. U. S. National Mus. 1 16:333-548.

Scullen, H. A. and J. L. Wold. 1969. Biology of wasps of the tribe Cercerini, with a list of

the Coleoptera used as prey. Ann. ent. Soc. Am. 62(1): 209-2 14.

Smith, A. V. 1961. Biological effects of freezing and super-cooling. Edward Arnold Ltd.,

London, 462 p.

S0mme, L. 1964. Effects of glycerol on cold-hardiness in insects. Can. J. Zool. 42:87-101.

Spooner, G. M. 1948. The British species of psenine wasps. Trans. R. ent. Soc. London,

99:129-172.

Steiner, A. L. 1970. Solitary wasps from subarctic North America — I. Pompilidae from

the Northwest Territories and Yukon, Canada. Quaest. ent. 6:223-244.

Strickland, E. H. 1947. An annotated list of the wasps of Alberta. Can. Ent. 79:121-130.

Tanno, K. 1964. High sugar levels in the solitary bee, Ceratina [in Japanese, English sum-

mary]. Low Temp. Sci. Ser. B. 22:51-57.

Ushatinskaya, R. S. 1957. Principles of cold resistance in insects [in Russian]. Acad. Sci.

USSR Press, Moscow. 314 p.

Uvarov, B. P. 1931. Insects and climate. Trans. R. ent. Soc. Lond. 79:1-247.

Williams, F. X. 1946. Two new species of Astatinae, with notes on the habits of the group.

Proc. Hawaiian ent. Soc. 1 2(3):64 1-650.

A TAXONOMIC REVIEW OF THE EASTERN NEARCTIC

SPECIES COMPLEX PTEROSTICHUS (HAPLOCOELUS) ADOXUS

(COLEOPTERA: CARABIDAE)

G. G. PERRAULT

138, rue Houdan

92330 Sceaux, France

Quaestiones entomologicae

9 : 35-40 1973

Study of selected material of Pterostichus (Haplocoelus) adoxus auctorum shows that

two species are included under this name: Pterostichus adoxus (Say), 1823 and Pterostichus

tristis (Dejean), 1828. Abbreviated synonymy is presented for each species and lectotypes

are selected.

L’etude des specimens de Pterostichus (Haplocoelus) adoxus auctorum montre que deux

especes sont confondues sous ce nom: Pterostichus adoxus (Say), 1823 et Pterostichus

tristis (Dejean), 1828. La synonymie est etablie pour chaque espece et des lectotypes choisis.

When studying the Pterostichini I collected in northeastern United States in 1967-68, I

had difficulty identifying two forms of Pterostichus subgenus Haplocoelus. Specimens of

both forms keyed to P. adoxus Say (Lindroth, 1966:449). The data presented in this paper

show that included in the current concept of P. adoxus are two species, named P. adoxus

(Say) and P. tristis (Dejean).

I studied a total of about 200 specimens which includes types and other material from

the collections listed below and material collected by me. Types for the following nominal

species are in the Museum of Comparative Zoology, Harvard University, Cambridge, Mas-

sachusetts (MCZ): Feronia adoxa Say, Pterostichus rejectus LeConte, P. subarcuatus Le-

Conte, and P. sustentus LeConte. Type material for the following nominal species described

by Casey is in the National Museum of Natural History, Washington, D. C. (USNM): P.

zephyrus, P. tetricula and P. sufflatus. Type material of Feronia tristis Dejean is in the

Oberthiir Collection, Museum National d’Histoire Naturelle, Paris (MHNP). I also studied

specimens from the collection of J. Negre, Versailles, France.

Specimens were compared with one another by examination of external characteristics,

by measurements and by examination of the male genitalia.

The following measurements were made: (1) overall length; specimen extended from apex

of elytra to apex of mandibles. Few female specimens were available and no differences

were discovered between sexes. Thus, all specimens studied, male and female, are considered

together. (2) distance of posterior lateral seta from adjacent hind angle: h; measured parallel

to longitudinal axis of pronotum.

Mensural data are presented in the form of a histogram (Fig. 1) and a scatter diagram

(Fig. 2). Form of pronotum and median lobe are illustrated by line drawings made with a

stereobinocular microscope and camera lucida.

MATERIAL

METHODS

36

Perrault

overall length in mm

Fig. 1. Histogram illustrating variation in overall length (mm) for selected material of P. adoxus and P. tristis.

□ neotype adoxus

0-4f A type rejectus

V type sufflatus

03

0-2

ot-

adoxus

^ 4 4

+ 4 4 E2

"' 4 4- 44

4 4+ 4 4a

4 4 ^7444 j. h*'

444

\ 4

\ 4

" ' " A © i 4 X\

■ 4+v 4 44 4A+T +- 4n;

4 +- 4444 44 4444 4

4 .#

tristis

• lectotype tristis

O type subarcuatus

■ type sustentus

▲ adoxus (Say) LeC.

specimen A/LC

T type tetricula

(} type zephyrus

2-5

30

35

40

LP in mm

Fig. 2. Scatter diagram illustrating differences in the relationship between length of pronotum (LP) and distance of

posterior lateral seta from the hind angle of the pronotum (h) for selected material of P. adoxus and P. tristis.

Nearctic Pterostichus

37

RESULTS AND DISCUSSION

The specimens available for study are arrayed in two groups, as indicated by differences

in measurements and ratios (Fig. 1 and 2), in pronotal form (Fig. 3 and 4) and in the form

and proportion of the median lobe (Fig. 5-8). Table 1 includes a summary of these data plus

additional, less definitive diagnostic characteristics, the most distinctive being No. 4, 5, and

9. The names used are the oldest available for each of these groups.

Table 1. Characteristics and character states for distinguishing between P. adoxus and

P. tristis.

Evidence that these groups are probably specifically distinct is provided by the geographi-

cal distribution of each. Specimens of the adoxus type occur in an area from south Quebec

and Maine to Pennsylvania, along the Appalachian Mountains. From the few available data,

no altitudinal limits can be set. However, all the specimens I have seen were collected below

800 m. Specimens of the tristis type are found in an area extending from southern Canada

to Georgia and from the east coast to Wisconsin, both in the mountains and in the lowlands.

The specimens I have seen were collected from sea level (Eatontown, Monmouth County,

38

Perrault

Fig. 3-4. Pronotum, dorsal aspect. 3, P. tristis ; 4, P. adoxus. Fig. 5-8. Median lobe; a - left lateral aspect; b - apex,

dorsal aspect. 5, P. tristis, Lectotype; 6, P. adoxus. Neotype; 7, P. rejectus, Lectotype; 8, P. tristis, specimen A/LC in

LeConte collection.

Nearctic Pterostichus

39

New Jersey) to 2000 m (Mt. Mitchell, Yancey County, North Carolina). A specimen labelled

“Texas”, ex Collection Sicard in MHNP, is probably mislabelled. The two groups are sym-

patric over a wide area and I have collected both in the same habitat at the following locali-

ties in the United States: NEW YORK: Ulster Co., Claryville, Catskill Mountains (800 m).

NEW HAMPSHIRE: Carrol Co., Jackson, White Mountains (300 m). MAINE: Piscataquis

Co., between Millinocket and Mount Kathadin (500 m).

H. Goulet ( personal communication ) found the two species together at Lac des Isles,

about 60 miles north of Montreal, Quebec, about 200 m elevation and noticed that in that

locality P. tristis was found only under the bark of fallen deciduous trees and P. adoxus on

soil under stones. Specimens with intermediate combinations of characteristics are not

known. Thus, although populations of these two groups live in close proximity they prob-

ably do not interbreed, and therefore they are probably specifically distinct.

SYNONYMY AND TYPE SELECTION

Pterostichus adoxus (Say)

Feronia adoxa Say, 1823:46. Neotype in MCZ. (For details see Lindroth and Freitag, 1969:

340).

Pterostichus rejectus LeConte, 1852:236. Lectotype, here selected, male, labelled “type;

5612”. No locality data. Two paralectotype females (MCZ).

Pterostichus sufflatus Casey, 1920:187. Lectotype, here selected, female, labelled “TYPE

NO. 47040”. No locality data. (USNM)

Pterostichus tristis (Dejean)

Feronia tristis Dejean, 1828:324. Lectotype, here selected, first specimen in front of adoxus

box label in the Oberthur Collection, labelled as follows: “d, LeConte; adoxa Say tristis

mihi exarata mihi alim, in Amer. Bor., D. LeConte” (labels on green-colored paper, hand-

written by Dejean). (MNHP)

Feronia interfector Newman, 1838:387. NEW SYNONYMY. Type material should be in

the British Museum (Natural History), but R. B. Madge {in litt .) could not find it there,

and Lindroth (1966:467) believed it to be lost. The evidence for this proposed synonymy

is, therefore, indirect. It is derived from label data associated with a male P. tristis (here

designated specimen A/LC; Fig. 2 and 8), in the LeConte collection. This label reads

“adoxus (Say) LeC, tristis (Dej.), interfector (Nw)”. It indicates to me that LeConte was

familiar with the Newman material and that he regarded it as conspecific with P. tristis.

It also suggests that at the time this specimen was labelled, LeConte did not regard the

three species described by him (P. rejectus, P. sustentus and P. subarcuatus ) as conspecific

with P. adoxus , although he did synonymize all of these names in 1873 (p. 304).

Pterostichus sustentus LeConte, 1852:236. Lectotype, here selected, female, labelled “type;

orange disc [southern states] ; 5611”. One paralectotype female, same data (MCZ). NEW

SYNONYMY.

Pterostichus subarcuatus LeConte, 1852:238. Lectotype, here selected, female, labelled

“type; pink disc [middle states]; 5618”. (MCZ). NEW SYNONYMY.

Pterostichus zephyrus Casey, 1884:2. Lectotype, here selected, male, labelled “TYPE NO.

47041”. No locality data. (USNM). NEW SYNONYMY.

Pterostichus tetricula Casey, 1913:130. Lectotype, here selected, female, labelled “Bayfield,

Wise.; TYPE NO. 47039”. (USNM). One paralectotype, female, same locality. NEW

SYNONYMY.

40

Perrault

ACKNOWLEDGEMENTS

I thank the following for loan of material: J. F. Lawrence and P. J. Darlington, Jr.,

Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts; T. L.

Erwin, National Museum of Natural History, Washington, D. C.; A. M. Villiers, Museum

National d’Histoire Naturelle, Paris; and J. Negre, Versailles, France. G. E. Ball, Department

of Entomology, University of Alberta, discussed the work with me during his stay in Paris

in 1972, and edited the manuscript.

REFERENCES

Casey, T. L. 1884. Contributions to the descriptive and systematic coleopterology of North

America. I. II. Phila. p. 1-198.

Casey, T. L. 1913. Studies in the Cicindelidae and Carabidae of America. Memoirs on the

Coleoptera. IV. New Era Printing Company, Lancaster, Pa., 400 p.

Casey, T. L. 1920. Random studies among the American Caraboidea. Memoirs on the

Coleoptera. Lancaster Press Inc., Lancaster, Pa. 9, 529 p.

Dejean, P. F. M. A. 1828. Species general des Coleopteres de la collection de M. le comte

Dejean, Paris 3, 556 p.

LeConte, J. L. 1852. Synopsis of the species of Pterostichus Bon and allied genera inhabit-

ing temperate North America. J. Acad. nat. Sci. Phil. 2:225-256.

LeConte, J. L. 1873. The Pterostichini of the United States. Proc. Acad. nat. Sci. Phil.

302-320.

Lindroth, C. H. 1966. The ground-beetles of Canada and Alaska. Part 4. Opusc. ent. Suppl.

29:409-648.

Lindroth, C. H. and R. Freitag. 1969. North American ground-beetles (Coleoptera, Cara-

bidae, excluding Cicindelinae) described by Thomas Say: designation of lecto types and

neotypes. Psyche, 76:326-361.

Newman, E. 1838. Entomological notes. Ent. Mag., 5:168-181, 372-402, 483-500.

Say, T. 1823. Description of insects of the families of Carabici and Hydrocanthari of

Latreille, inhabiting North America. Trans. Amer. phil. Soc., new series, 2:1-109.

Book Review

SWAN, L. A. and C. S. PAPP. 1972. The common insects of North America. Harper and

Row, Publishers, New York, Evanston, San Francisco, London, xiii + 750 pages, text-fig.

1-1422, 8 color plates, 2 appendices, glossary, bibliography, indices of subject and common

names and of scientific names. Price $15.00 U. S. A.

According to the authors, the purpose of this volume is to provide an easy way to iden-

tify the more common insects of North America north of Mexico, emphasis being placed

on comparison of specimens with illustrations. According to a statement by the publishers

on the dust jacket, the volume is of “special interest because of its thorough coverage of

Canada.”

The text consists of an introduction of 32 pages, a “pictured key” to the insect orders

and chapters 1 to 23, each dealing with the taxa of a single order.

The introduction, designed to enlighten those potential users of the book who have not

had the benefit of formal training in systematic biology or in entomology, explains classifi-

cation of the animal kingdom and locates the insects and other arthropod classes in the

general system. Binomial nomenclature is touched on, and the usual erroneous statement is

made that Linnaeus devised this system. Insect structure, function, and development are

discussed briefly but reasonably well. Many structural features are illustrated with fully

labelled line drawings. The introduction concludes with an excellent, 1.5 page discussion of

the value of insects to ecosystems in general, and to man: as pollinators, as agents of biolo-

gical control of plants and of other insects, and as items of diet.

On page 3, the authors chide “people” who err in that they “do not think of insects as

animals . . . ”. On page 10, the authors make a similar error when they refer (line 5) to “man

and animals.” The statement should be “man and other animals.”

The authors neglect to inform their readers that because of the small size of most insects,

optical equipment might be required to examine a specimen in sufficient detail to make a

meaningful comparison with the illustrations and data provided in the text.

In discussing classification, the authors use the terms “broken down” and “divided” to

refer to the process of classifying. In reality, classification consists of organizing discrete

entities into collective groups, and further grouping of the initial groups. The terms in quo-

tation marks are without meaning in classification.

The “pictured key” consists of 15 pages of illustrations and brief descriptive statements

numbered sequentially, about the characteristics of each order. There are no directions for

proceeding from one step to the next. A key, on the other hand, is a flow sheet, with

specific directions at each point. In this section, pictures there are; key there is not.

Twenty- three orders are treated in the text. It appears that the authors used as a model

for the sequence of orders some publication produced prior to 1950. To many it might seem

immaterial that an antiquated system is used, but to me it seems unreasonable to return to

an arrangement clearly discordant with those proposed in the more recent literature.

The book treats superficially and illustrates specimens of about 1,500 species, about 1.5

percent of the North American insect fauna. The species represent 276 families.

Each family is characterized structurally and biologically in about one half page of text.

Within each order, most species represented by figures are numbered. For each numbered

species, data given are common name (in boldface), scientific name, geographical range, a

brief description of adults (including size in inches to one or two decimal places) and larvae,

and some information on biology. Economically important species are noted. For some

groups, keys are given (for example, worker termites of eight genera).

The number of species treated per supraspecific taxon depends in part upon conspicuous-

ness of individuals and popularity of the group with collectors. For example, specimens of

42

296 species of Lepidoptera are figured, included in 44 families. On the other hand, only 1 1 1

species of Hymenoptera are figured, included in 35 families. Three of four genera of North

American tiger beetles are treated, including 12 species of Cicindela. In eight pages, 26 spe-

cies of coccinellid beetles are treated, and figures are provided of an additional 36 Hyper-

aspis, 1 1 Scymnus and six Hippodamia species. Butterflies fare well, also: 153 species, in 51

pages.

In general, the illustrations adequately represent the aspect of the specimen figured. Each

figure is numbered, and associated with each is a vernacular name in capital letters, the

scientific name in italics and an indication of the size of the specimen. The color plates are

technically satisfactory, but they add little of value. Many of the insects illustrated in color

have been so illustrated previously.

Appendix I is a four page synopsis of a portion of the geologic time table, beginning with

the Devonian Period and ending with the Quaternary, summarizing major geomorphic events

(excluding continental drift), and major biological ones, including appearance of the insect

orders.

Appendix II is a list of names of orders and families represented in the book. Both this

and the glossary are useful.

A volume of this sort might be expected to serve as an entry to point to the entomologi-

cal literature on identification. Normally, this is accomplished by references in the text

keyed to a bibliography. Although the latter is provided, there are no text-references. Thus,

a person wanting to know more about, say dragonflies, has to fumble through 1 7 pages of

references to discover the publications by Needham and Westfall, and Walker.

The bibliography comprises two portions: one, labelled “General,” the other, “Techni-

cal.” This distinction eludes me because the categories must overlap by definition (many

“general” works must also be “technical”) and because I was unable to discern criteria used

by the authors in assigning publications to one of the two groups. But this is a minor objec-

tion. More important, the basis for inclusion or exclusion of references is not apparent. It

seems that no systematic effort was made by the authors to list the recent taxonomic litera-

ture of major consequence to their work, or to be consistent about what was included. For

example, Swain’s “Insect Guide” was included, but the more recent “Field Guide to the

Insects” by Borror and White was excluded. “A Key to the Wyoming Grasshoppers” was

included, but the “Acridoidea of southern Alberta, Saskatchewan and Manitoba” was ex-

cluded. The revision of Meloe by Pinto and Selander was included, but the revision of the

meloid genus Epicauta by Werner was excluded. A list of this type could be extended.

Another criticism is that no consistent sequence is used for listing several publications by

the same author: for some, the earliest publication is listed first; for others, the latest; for

still others, no arrangement is perceivable.

The planning denied to preparation of the bibliography is further illustrated by the body

of information associated with the periodical cicadas. The authors devote a bit more than

two pages of text to these species (pages 133-135), including a map and a table, based on a

USDA Economic Insect Report (cited in the text, but not in the bibliography). In spite of

the importance accorded by the authors to this species complex, they do not cite the im-

portant taxonomic paper by Alexander and Walker entitled “Evolutionary relationships of

17-year and 13-year cicadas ... ”, in the Miscellaneous Publications, Museum of Zoology,

University of Michigan, No. 121, 1962.

The bibliography would be of greatest value if the publications were arranged by taxo-

nomic groups because the purpose of this book is to enable a person to identify insects; part

of this task is to locate relevant references. In spite of these shortcomings, I found this sec-

tion interesting because it contains references to many papers that had previously escaped

43

my notice.

How does one distinguish a “common insect” from an uncommon one? The authors

neglect to provide this information, and thus do not inform the readers of the basis for

inclusion or exclusion of taxa. For the beetles, I think I was able to deduce one of the

criteria, by comparison of illustrations with those in Jaques’ book, “How to know the

beetles.” There was a remarkable degree of overlap among the species illustrated (and re-

markable similarity of illustrations of the same species between the two books). Also, illu-

strations of ichneumonids are strikingly similar to those in “Ichneumon flies of America

north of Mexico” by Townes and Townes. I suggest that one of the criteria for inclusion of

a species was that it had been illustrated in a previous publication. The illustrations were no

doubt drafted by the authors, but I believe that many were based on previously published

illustrations rather than on insect specimens.

Comparison with illustrations in a book is a successful technique of identification only

when the taxa of most specimens that might be referred to it are represented therein by

figures. If a person who uses the book cannot know what a “common insect” is he cannot

know that a specimen in hand represents a species described in that book. Because criteria

for commonness are not established, specimens of any of the 90,000 or so species of North

American insects might be referred to this volume which deals with only about 1,500 spe-

cies. Because of the resulting low degree of probability of actually being able to identify

insects to species with this book, and because the authors place emphasis on species identi-

fication, the volume seems of limited value for its announced purpose — and so I believe it

is. It could play a useful role in identification of higher taxa, but for this purpose its cover-

age is limited.

A book of moderate size intended to guide one in identifying material drawn from a large

fauna should not pretend that its operational level is the species. The family level is realistic.

Borror and White’s “A Field Guide to the Insects of America north of Mexico” (Houghton

Mifflin Company, Boston) is a guide to the families. It is concisely written, superbly illu-

strated by drawings based on specimens, has a list of references arranged by taxa, and costs

only $5.95. This is the volume for anyone needing to make identifications, who does not

recognize the families at sight.

What is the value of “The Common Insects of North America” for Canada? I bring up this

point only because of the publishers’ statement about “thorough coverage” of this fauna.

The coverage is in no sense thorough. Some important forest and crop pests in Canada are

not mentioned, and for those that are, brief notice is given and little of it is specifically

relevant to Canada. For many species, there is no indication as to their Canadian distribu-

tion. I do not blame the authors for this misstatement. It is just one more example of “truth

in advertising” as this concept is currently understood by North American business interests.

It is a pity that authors need risk their reputations by association with firms whose adver-

tising personnel are unable or unwilling to distinguish between truth and falsehood.

Who needs this book? Entomological bibliophiles and libraries intent upon acquiring

complete holdings of entomological literature need it. The experienced entomologist, ama-

teur or professional, who understands its severe limitations, might like to have a copy be-

cause the many illustrations facilitate identification, or one might find an interesting refer-

ence by browsing through its bibliography. The book, I think, creates its own niche, rather

than filling one based on need of those interested in entomology.

G. E. Ball

Department of Entomology

University of Alberta

Book Review

URSPRUNG, H. and R. NOTHIGER (Editors). 1972. The Biology of Imaginal Disks. Vol-

ume 5 in: Results and Problems in Cell Differentiation. A series of Topical Volumes in

Developmental Biology. Springer- Verlag, New York, Heidelberg, Berlin, xvii + 172 pp., 56

figures, 12 tables. Cloth 8vo $14.60 (U. S.).

This monograph reviews recent research on insect imaginal discs — chiefly those of

Diptera-Cyclorrhapha and principally those of Drosophila species. The book contains six

review articles, each with its own bibliography: (1) R. Nothiger: The larval development of

imaginal discs, (2) W. Gehring: The stability of the determined state in cultures of imaginal

discs in Drosophila , (3) A. Garcia-Bellido: Pattern formation in imaginal discs, (4) H.

Ursprung: The fine structure of imaginal discs, (5) J. W. Fristrom: The biochemistry of

imaginal disc development and (6) H. Oberlander: The hormonal control of development

of imaginal discs. Although each article stands alone, there is considerable overlap, parti-

cularly in the first three contributions. All of the authors are productive contributors to

the subject reviewed and are former students or associates of Ernst Hadorn, the Swiss

embryologist who first realized the heuristic value of imaginal discs and to whom the book

is dedicated. D. Bodenstein, in a eulogy recognizing Hadorn ’s 70th birthday, summarizes his

contributions at the beginning of the book. A fuller account together with lists of his pub-

lications and theses done under his direction can be found in Chen, P. S., P. Tardent and

H. Burla. 1971. Ernst Hadorn zum siebzigsten Geburtstag. Revue Suisse de Zoologie 79:

5-28.

One of the principal lacunae in our understanding of development in insects and other

eucaryotes is that of cellular determination. How are different cells, all containing the same

genetic information in their chromosomes, programmed for a specific fate during develop-

ment? An answer to this question has awaited a fuller understanding of how genes work

at the molecular level and the discovery of appropriate, eucaryote, experimental systems.

Practitioners of the science of biochemical genetics have come a long way towards providing

the first, while the imaginal discs of holometabolous insects seem to constitute the second.

Experimental imaginal disc research began when Ephrussi and Beadle developed a tech-

nique for transplanting discs dissected from donor larvae of Drosophila into larval or adult

hosts by means of a micropipette. When a disc is transplanted into a larva of the same age as

the donor, it develops synchronously with the host and undergoes metamorphosis within it.

On emergence of the adult the implant can be removed and examined. In all experiments,

the discs were found to differentiate auto typically, i.e. into the structure for which the

disc was originally determined. Similar results were obtained with fragments of discs, in-

dicating that each disc of the third (last) larval instar contains a mosaic of different cell

groups, each determined to form a specific part of the adult structure.

One of the principal advantages of using Drosophila species in this work is the availability

of a large number of genetic marker mutations. These can be easily recognized by their

effect on imaginal surface structures exemplified by coloured, crooked, or multiple hairs,

microtrichiae, and bristles.

Discs may be dissociated enzymatically or mechanically into small groups of cells or into

single cells. If such cells from identical discs of different mutant donors are mixed together

and injected into a wild-type host larva it is found, after metamorphosis, that the cells from

different donors collaborate to form normal but mosaic adult structures. The contributions

of individual cells to the development of the whole structure can be recognized because they

differentiate into bristles and hairs having the colour and shape of the donor phenotype.

Evaluation of other experiments involving the mixture of mutant cells from dissociated

discs of different kinds (e.g. wing and leg; haltere and eye-antenna) showed that a cell from

45

a given disc will only associate with other ( isotypic ) cells from the same kind of disc and not

with those ( heterotypic cells) from other kinds of discs. The association of isotypic cells

and separation of heterotypic cells is considered by Gehring to be achieved by cell migration

and selective adhesion of cells.

If imaginal discs are transplanted into the abdomens of adult flies rather than into larval

hosts, they proliferate into blastemas. The host’s haemolymph serves as a culture medium

which allows proliferation but does not induce differentiation, probably because of the ab-

sence of ecdysone. Such blastemas can be cultured indefinitely by dissecting them from the

host fly every two to four weeks, cutting them into fragments and injecting the fragments

into fresh host flies. Other fragments are injected into host larvae where they undergo differ-

entiation on metamorphosis of the host. These larval “test implants” provide information

about the capacities of the cultured cells for differentiation. Using this technique, Hadom

and his students have shown that the cultured cells maintain their capacity for normal (auto-

typic) differentiation even after several years of culturing, i.e. they maintain their state of

determination.

However, in cultured blastemas, occasional changes in cell heredity affecting determina-

tion occur. Some of the cells, when tested in larvae, at metamorphosis differentiate allo-

typically into organs other than those for which the cells were originally determined. For

example, a fragment of genital disc blastema might differentiate into antennal or leg struc-

tures. This change in cell heredity is called transdetermination.

Using these culturing techniques Hadom’s group showed that for each state of determina-

tion in a particular disc, there exists a probability of transdetermination in a specific direc-

tion. Sometimes these changes in prospective fate are reversible, sometimes not. The only

factor so far detected which influences the frequency of transdetermination is proliferation.

This suggests that cell divisions are a necessary prerequisite for it.

Naturally-occurring developmental abnormalities leading to the same effect as transde-

termination can be induced by homoeotic mutations. A common example is aristapedia in

which the arista of the antenna is replaced by a tarsus. Gehring suggests that a single mutant

“switch” gene could bring into action all the genes necessary for the differentiation of a leg

disc in a blastema previously determined to form head structures. Some homoeotic muta-

tions are temperature-sensitive. Temperature-sensitive alleles of the mutation ssa, for exam-

ple, cause parts of the antennal disc to develop into leg structures at 16°C and into antennal

structures at 25°C with the temperature-sensitive period lying in the third-instar. Gehring

suggests that the main problem for future research is the identification of the carrier of

determination.

In cyclorrhaphous Diptera, the somatic cells exhibit pairing of homologous chromosomes

similar to that occurring at synapsis during prophase I of meiotic cells. By treating prophase

cells with X-rays it is possible to induce mitotic recombination in them. Strains of Droso-

phila are used which are heterozygous for a recessive marker gene. If a cell is irradiated just

before it divides, crossing-over may be induced such that one or both of the daughter cells

become homozygous with respect to the mutant gene. The clone of cells arising from this

initial daughter cell will, with subsequent development, appear in the adult as a patch of

mutant tissue surrounded by wild-type tissue.

Using this technique Schneiderman and Garcia-Bellido and their students have shown that

oriented cell divisions, differential mitotic rates, and local differences in cell size are all

involved in producing changes in form in the discs during their development. They have also

shown that determination is a gradual, progressively-narrowing phenomenon. By irradiating

individual cells at different stages of development and then following what happens in the

irradiated area, these workers have been able to determine the number of blastoderm cells

46

in the embryo that give rise to each imaginal disc and to prove that determination of adult

structures begins during blastoderm formation.

Experiments using gynandromorph tissue can yield the same kind of information. Gynan-

dromorph tissues are mosaic and contain both male and female cells. In Drosophila they

arise when one of the two X-chromosomes is lost during development of a female embryo,

resulting in female (xx) and male (xo) tissue patches. If the insect was originally hetero-

zygous for x-linked cell marker mutations affecting bristle colour or shape, for example, the

mosaic is recognizable on the body surface of the adult fly because the recessive mutations

are uncovered through the loss of wild-type alleles. Though gynandromorphs are rare in

nature, they can be induced artificially in various ways.

Studies of the ultrastructure of imaginal discs, as reviewed by Ursprung, have revealed no

differences between cells of different discs. They have yielded evidence suggesting that the

surface increase accompanying disc eversion in the pupal stage results largely from a change

in shape of the epithelial cells comprising the disc; in larvae they are columnar; in pupae

cuboidal. Some mutants of Drosophila lacking portions of or complete appendages in the

adult have imaginal discs in which many cells die during development. Others have smaller

than normal discs.

As emphasized by Fristrom and Oberlander, Drosophila imaginal discs are almost ideal

material for studying the biochemical effects of hormones on differentiating tissues. They

are easy to culture in vitro and their only disadvantage, their small size, has been overcome

by the perfection of mass isolation techniques (up to 220,000 discs per day). Since synthe-

tic juvenile hormone and ecdysones are commercially available, many breakthroughs in our

understanding of the biochemistry of development are in the offing. Fristrom and his col-

leagues have found that /3-ecdysone is much more active than a-ecdysone in inducing the

synthesis of RNA, principally ribosomal, in cultured discs. Ecdysone apparently enters the

disc cells where it directly affects transcription. No “second messenger” such as cyclic AMP

which mediates the action of several different hormones in vertebrate systems has been

found. Increased protein synthesis results from increased RNA synthesis and these proteins

probably participate in the orientation, assembly, or function of microfibers evoking the

change in cell shape causing appendage eversion during pupation. It was formerly thought

that blood pressure was responsible for eversion in vivo, but discs evert just as successfully

when removed and cultured in vitro with /3-ecdysone. Juvenile hormone acts directly and

antagonistically with ecdysone on both synthesis and eversion. Younger discs are less sensi-

tive to both juvenile hormone and ecdysone than mature discs. If discs are cultured with fat

body or in media conditioned with fat body, the effects of both juvenile hormone and

moulting hormone are more rapid, suggesting that the fat body may influence the acquisi-

tion of competence by the disc.

Disc research has become a meeting ground for geneticists, developmental biologists,

entomologists, biochemists and endocrinologists. Thus, this book deserves and will probably

have a wide circulation. Although it is well produced, the book has several errors in typog-

raphy and style (“prepupal” is usually spelled “prepual” and “for example” is always abbre-

viated whatever the context). Some contributors to the book (e.g. Garcia-Bellido) presup-

pose more background on the part of their readers than do others (Gehring, Fristrom) and

some (Fristrom) write more clearly. A detailed table of contents compensates in some mea-

sure for the lack of both author and subject indices.

Bruce Heming

Department of Entomology

University of Alberta

Book Review

D’ABRERA, B. 1971. Butterflies of the Australian Region. Lansdowne Press Pty Ltd.,

Melbourne. 416 pp., 2362 color photographs, two pages of maps, three groups of line

drawings in the text, glossary, 160 references, index, one page corrigenda, one loose in-

i sert errata. Size 13%” x 10/4”, hard covers. Printed in Hong Kong. Price: $39.95 in the

! u. s.

Since Fabricius (1775) first described species of Lepidoptera from Australia, several

accounts have appeared summarizing knowledge of this fauna. The most extensive of these

is volume IX of Seitz’s (1927) “The macrolepidoptera of the World, The Indoaustralian

Rhopalocera”. D’Abrera’s “Butterflies of the Australian Region” is the first work to figure

all known species of butterflies inhabiting Australia, New Guinea, the Moluccas, New Zea-

land, and the South Pacific islands. Included are Lepidoptera of the following families:

Papilionidae, Pieridae, Danaidae, Nymphalidae, Libytheidae, Satyridae, Amathusiidae, Ly-

■ caenidae, and Riodinidae. The family Hesperiidae is excluded.

The book is organized into two parts, introductory and systematic. The first part includes

! short notes on how the text is organized, butterfly life history, mimicry and protective

, coloration, variation, nomenclature and classification. The history of lepidopteran collecting

and study in the Australian Region is briefly summarized. This section including glossary is

j 39 pages long with 3V2” wide margins on most pages. These margins are partly occupied by

13 groups of photographs and diagrams.

The second part, entitled “A guide to the identification of the butterflies of the Austra-

lian Region”, provides data about and photographs of adults of more than 900 species. The

i figures are color photographs of specimens deposited in various museums. Most figures are

1 of dorsal aspects. In addition, illustrated are the ventral sides, morphs of polymorphic spe-

: cies, and some immatures. Keys are not included and identifications must be made by com-

| paring specimens with figures. Ninety-one specimens represent “types” (indicated by a red

i dot opposite the specimen), and six represent paratypes (indicated by a yellow dot). Four-

teen taxa are described as new: three species and 1 1 subspecies.

The following data are given for each species: binomen and author, reference to original

description, abbreviated synonymy, geographic range, status, races, depository for figured

specimens, life history stages, and short description of known ones, and food plant, if

known. On each page, the text is marginal. Each illustration is beside the text relevant

to it.

This volume is little more than an updated illustrative companion to Seitz’s “Indo-

australian Rhopalocera”. The sequence of arrangement of taxa, in general, is antiquated.

The illustrations are of good quality, but the scale at which the individual photographs are

reproduced is not indicated and the statement on the cover flap, qualifying these as “natural

size” is untrue. For example, P. demoleus sthenelus figured on page 41, is shown to have a

wingspan of 20 inches. Because illustrations are unnumbered and no list of illustrations is

provided, figures in the first part can be located only by searching through the text. In the

systematic section they can be found by locating the name of the relevant taxon in the

index. While many of the photographs portraying immatures or adults in their natural

habitats are first rate, some like that on page 48 should not have been included, for they

show nothing worth recording.

Poor layout reduces the value of many of the photographs. This is particularly true of

figures where the center fold cuts through the right half of the butterflies illustrated. The

marginal half of most pages is reserved for the text accompanying the figures in the systema-

tic part. The high price will hamper the sale of this book in a highly competitive market.

Three other books were published on the same subject within the last three years, though

48

each with different emphasis. In spite of its shortcomings, “The butterflies of the Australian

Region” is an important reference for lepidopterists.

Joseph Belicek

Department of Entomology

University of Alberta

Publication of Quaestiones Entomologicae was started in 1965 as part

of a memorial project for Professor E. H. Strickland, the founder of the

Department of Entomology at the University of Alberta in Edmonton

in 1922.

It is intended to provide prompt low-cost publication for accounts of

entomological research of greater than average length, with priority

given to work in Professor Strickland’s special fields of interest including

entomology in Alberta, systematic work, and other papers based on work

done at the University of Alberta.

Copy should conform to the Style Manual for Biological Journals

published by the American Institute of Biological Sciences, Second

Edition, 1964, except as regards the abbreviations of titles of periodicals

which should be those given in the World List of Scientific Periodicals,

1964 Edition. The appropriate abbreviation for this journal is Quaest. ent.

An abstract of not more than 500 words is required. All manuscripts will

be reviewed by referees.

Illustrations and tables must be suitable for reproduction on a page

size of 93/4x63A inches, text and tables not more than 73/4x43/4 inches,

plates and figures not more than SV2 X5 inches. Reprints must be ordered

when proofs are returned, and will be supplied at cost. Subscription rates

are the same for institutions, libraries, and individuals, $4.00 per

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issues $1.00. An abstract edition is available, printed on one or both

sides (according to length) of 3X5 inch index cards (at $1.00 per

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volume) .

Communications regarding subscriptions and exchanges should be

addressed to the Subscription Manager and regarding manuscripts to:

The Editor, Quaestiones Entomologicae,

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E’D-CpWif

MUS. COMP. 200 U.

LIBRARY

Quaestiones

entomologicae

JUL 1 3

Harvard

LfNlUftnsijy

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

VOLUME IX

NUMBER 2

APRIL 1973

QUAESTIONES ENTOMOL OGICAE

A periodical record of entomological investigation published at the Department of

Entomology, University of Alberta, Edmonton, Alberta.

Volume 9 Number 2 April 1973

CONTENTS

Editorial — For Love or Money? 51

I Shorthouse— The insect community associated with Rose Galls of

Diplolepis p*)lita (Cynipidae, Hymenoptera) 55

Larson — An annotated list of the Hydroadephaga (Coleoptera: Insecta)

of Manitoba and Minnesota 99

Richards — Biology of Bombus polaris Curtis and B. hyperboreus Schonherr

at Lake Hazen, Northwest Territories (Hymenoptera: Bombini) 115

51

Dr. Ruby I. Larson

Cytogeneticist, Canada Department of Agriculture

Lethbridge, Alberta

Editorial — For Love or Money?

One of the more unfortunate features of our life and times is the increasing difficulty of

finding anybody willing to do anything without being paid for it, preferably at the going

rate or better. This is perhaps not surprising in respect of daily toil or labour involving the

sweat of the brow or the mobilizing of the mind but it becomes somewhat absurd when ex-

tended to such supposedly enjoyable activities as the playing of games and even to being en-

tertained. This situation arose as a side effect from the efforts of organized labour to im-

prove the lot of the so-called working classes. They have been laudable efforts, towards an

objective with which I have no quarrel; but I have said so-called working classes because I

think this term needs re-definition for our present day and age. It was introduced at a time

when the population of many countries could be divided into two groups, one much larger

and less influential, those who worked for a living; and the idle rich. In our present day pop-

ulations we have plenty of idle and plenty of rich but these two qualities are less frequently

found in the same person than they used to be. The so-called working class of today in-

cludes a substantial segment, perhaps best referred to as the idle poor, who no longer work

52

but who apparently enjoy a modest existence on funds from welfare, unemployment insur-

ance, or some more oblique dispensation of the taxpayer’s money. It seems necessary to as-

sume that such people either do not like work of any kind or at least have been unable to se-

cure work of a kind which they might enjoy. We may mention in passing that management

involves work.

Another result of the efforts of organized labour to improve the lot of the working classes

has been to change the meaning of the terms professional and amateur so that they become

essentially antithetic. A person who gets paid for what he does is a professional, a person

who does not is an amateur. In their original meanings these words were far from antithetic.

Professional meant simply a person, who, by public declaration or otherwise through his

training or official qualifications, indicated his intention and presumably ability to fulfil a

certain role. An amateur was a person who filled a certain role, although perhaps not one

recognized by society, simply because he loved filling it. The assumption that a person who

does something without being paid for it loves doing it, may or may not be justified. A third

pair of meanings of these two terms, also antithetic and recently acquired, makes the pro-

fessional a person who does a good job and the amateur a person who does an indifferent

one. These last meanings are in direct conflict with the original ones since, in my experience,

one is more likely to get good work done by a person who loves doing it than by a person

who is merely doing it for the money. It is principally for this reason that it is unfortunate

that people willing to do things without being paid for it are becoming increasingly scarce.

While entomology has certainly not been immune to the reduction in its population of

amateurs, it is my impression that it has suffered less than most other branches of scientific

work. Certainly it appears likely that there will be plenty of opportunity for amateurs, in

the two best senses of the word, to work in the field of entomology for many years to

come. This has many advantages. In the first place, amateurs in a field help to keep it in

touch with the public. Perhaps more important is the increasing proportion of our time a-

vailable for leisure activities promised us by technological advance for some time now,

though many of us see little sign of fulfillment of this promise. One of the dangers of this in-

creased leisure is that it can lead people to accept, by way of regular employment, some-

thing which they are not really interested in doing, thus increasing the risk of them becom-

ing members of the idle poor. Since routine, humdrum, repetitive occupations are clearly

those most readily taken over by machines and computers, it would seem reasonable to ex-

pect technological advance to make it easier for people to find more enjoyable and interest-

ing occupations than in earlier days, but there is no clear evidence that this is so. Perhaps

this is because there are too many people and not enough things that need doing. Or could it

be that the possibility of survival without work has been selecting for survival those people

who can get no enjoyment out of work of any kind, the hard-core of the idle poor? If so,

what price a guaranteed minimum income? I would suggest the smaller the price, the better.

Amateurs, in the original sense, are enthusiastic people; enthusiasm is infectious and one of

the most important qualities to be sought in a teacher.

Dr. Ruby Larson has always been an enthusiastic person. Her first employment was as an

impoverished country school teacher in Saskatchewan. From that position, she took a sum-

mer school course in biology from Dr. J. G. Rempel, then Professor of Biology at the Uni-

versity of Saskatchewan, now fulfilling a similar role from retirement at the University of

Victoria. This convinced her that biological research was the most exciting occupation in the

world. While a student at the university, she found summer employment counting wheat

chromosomes at the Swift Current Experimental Station of the Canada Department of Agri-

culture in connection with the cereal breeding work being conducted there by A. W. Platt

and Chris Farstad. This eventually led to her appointment as a cytogeneticist and her work

53

in this field in relation to the resistance of plants to insect and other damage is well known.

Nobody however, who has been in contact with Dr. J. G. Rempel could escape some en-

thusiasm for entomology. These two enthusiasms still constituted only a part of the total

enthusiasm which Dr. Larson put into the formation and operation of the Junior Science

Club of Lethbridge. Characteristically, she attributes the success of this club to the young

people who joined it but, going back to first principles, the young people who joined it did

so because of her enthusiasm. This enthusiasm also drew collateral support for the Club

from her colleagues at the Canada Department of Agriculture Research Station in Leth-

bridge and elsewhere.

The authors of all three papers in this issue of Quaestiones entomologicae were members

of Dr. Ruby Larson’s Junior Science Club of Lethbridge. As she puts it, the remarkable

thing is not that they became entomologists, that was inevitable, but that all three of them

have followed their first main interest; David Larson with his beetles mainly because of

their beautiful stricture; Ken Richards with his bees partly because of his association with

Gordon Hobbs; and Joe Shorthouse with his insect galls. It is of special interest that the Lar-

son paper is doubly amateur, representing as it does, the work of J. B. Wallis, in his day one

of Canada’s leading amateur entomologists. The breadth of interest of the Club is reflected

in the fact that doctors, teachers, architects and engineers, in addition to entomologists,

have come from among its members. It is for this and other reasons that we are pleased and

proud to dedicate this issue of Quaestiones entomologicae to Dr. Ruby Larson, personality,

teacher, scientist, biologist, cytogeneticist, and entomologist; professional and amateur, in

the best senses of both words, in all of these fields.

The story of Ruby Larson is a story of what the enthusiasm of an amateur, in the original

sense of the word, can accomplish It is dlso a story of the influences of teachers on stu-

dents, Rempel, via Larson, on many others. Such influences, as H. T. Pledge has pointed out

in his book, Science since 1500, have played a tremendous role in the history of science. It

is also a story which demonstrates for the benefit of teachers at all levels, the vital impor-

tance of enthusiasm.

Education of today, especially at the university level, must be flexible to be fair to stu-

dents who may be degree-labelled for life; they must have an opportunity to pursue that

which they really wish to pursue. But to be fair to the society in which these students will

have to find employment as well as to the student, it must also be broad, for despite techno-

logical advances we have a long way to go before our societies can accommodate a life of ac-

tivity on a specific individual interest for each and every one of its members. The most im-

portant thing to ask of life is the opportunity to do that which one is most interested in do-

ing; preferably to get paid for doing it but, to do it anyway. All too often, life will say no;

but love will find out the way.

Brian Hocking

THE INSECT COMMUNITY ASSOCIATED WITH ROSE GALLS OF

DIPLOLEPIS POUT A (CYNIPIDAE, HYMENOPTERA)

J. D. SH ORTH OUSE

Department of Biology

University of Saskatchewan Quaestiones entomologicae

Saskatoon, Saskatchewan 9: 55-98 1973

The single chambered gall of Diplolepis polita (Ashmead) (Cynipidae) is initiated in the

early spring on immature leaves of Rosa acicularis Lindl. (Rosaceae). D. polita larvae and

succulent gall tissues attract five additional insect species which, by their inter-relationships

within the galls constitute a community. Each species appears over a different period so that

the community undergoes succession and climax. Life cycles and roles of all members of the

community are discussed. Most of the D. polita larvae are replaced early in the season by

larvae of the inquiline Periclistus pirata (Osten Sacken) (Cynipidae). P. pirata larvae cause

additional cell proliferation and in the process of becoming enclosed in layers of cells, struc-

turally modify their host galls. Galls inhabited by P. pirata are larger than galls inhabited by

D. polita. Larvae of P. pirata are the main food source for four entomophagous inhabitants:

Eurytoma longavena Bugbee (Eurytomidae), Glyphomerus stigma (Fabricius) (Torymidae),

Torvmus bedeguaris ( Linnaeus ) ( Torymidae ), and Habrocytus sp. (Pteromalidae),

La Communaute d’Insectes Associes avec des Galles de Rose de Diplolepis polita ( Cynipidae,

Hymenoptire)

Sommaire

La galle de Diplolepis polita (Ashmead) (Cynipidae) d’une seule chambre s’initie de bonne

heure le printemps sur des jeunes feuilles de Rosa acicularis L indl. (Rosaceae). Les larves de

D. polita et les tissus succulents des galles attirent cinq autres especes d’insectes, qui par

leurs relations constituent une communaute. Chaque espice se manifeste pendant une peri-

ode differente de Vannbe et comme resultat la communaute subit une succession d’habitants

et une periode d’apogee. Les cycles vitaux et les roles de tous les membres communautaires

sont analyses. La plupart des larves de D. polita s’est remplacee tot dans la saison par des

larves de Periclistus pirata (Osten Sacken) (Cynipidae). Ces larves de P. pirata evoquent une

nouvelle proliferation de cellules et pendant le processus de se faire entourer par des couches

cellulaires, elles font modifier la structure de la galle-hote. Les galles habitees par P. pirata

sont plus grandes que celles habitees par D. polita. Les larves de P. pirata servent de nourri-

ture principale des quatre habitants entomophages: Eurytoma longavena Bugbee (Eury-

tomidae), Glyphomerus stigma (Fabricius) (Torymidae), Torymus bedeguaris (Linnaeus)

(Torymidae), et Habrocystus sp. (Pteromalidae).

Cecidology, the study of plant galls, has long been of great interest to biologists. Al-

though galls have been mentioned in the literature since ancient times (Hippocrates, 406-

377 B. C., wrote on the medicinal properties of galls) it was not until the late eighteenth

century that any attempt was made to explain the connection between galls and the insects

found in them. Malpighi was probably the first to explain that the stimulus for gall forma-

tion was of animal origin (Plumb, 1953). Cosens (1915) reviewed the older gall literature in

a paper in which he discussed the founding of cecidology. Plumb (1953) also presented an

excellent review of early cecidological literature and explained the development of theories

about the source of gall forming stimuli, sites of action, and gall developmental morphology.

56

Shorthouse

Although a great deal of literature on insect galls has been amassed, much of it is only sys-

tematic. Most cecidological workers in North America have occupied themselves with the

classification of galls and gall insects and have disregarded fundamental problems such as gall

initiation, developmental morphology, and the inter-relationships of the inhabitants com-

posing the gall communities. Checklists are prominent in North American cecidological liter-

ature and the most popular is by Felt ( 1 940). The most comprehensive treatise of European

galls and gall formers is by Buhr (1965).

Insect galls can be defined as atypical growths produced by plants in response to a foreign

stimulus. This stimulus, either chemical or physical, or both, can be provided by the larvae

or the adult gall former. Gall formers are found in at least 8 insect orders, but the majority

are restricted to the families Cecidomyiidae and Cynipidae. Of the approximately 1,450 gall

formers in North America (Felt, 1940), about 38% belong to the order Hymenoptera. Of

the hymenopterous gall formers, 91% belong to the family Cynipidae, the galls of which are

recognized by all students of cecidology as the most remarkable in variety and complexity.

For gall development to occur, the life cycle of the gall former must be synchronized

with the optimum galling conditions of the plant. One prime requisite for gall formation is

the presence of meristematic tissue. The plant must be in such a condition that the foreign

stimulus can alter normal growth patterns. Malyshev (1968) suggested that gall wasps can

convert relatively differentiated tissue back into the meristematic state. Wells ( 1 920) sug-

gested that the gall former actually causes the dedifferentiation of host tissue, preventing

the normal expression of host characters. Once dedifferentiation has occurred, stimuli from

the insects cause the gall to grow into its specific shape. All galls, especially the more com-

plex, have characteristic shapes and structures. The structure of a gall depends upon the ge-

nus of insect producing it rather than upon the plant on which it is produced. Kinsey

(1920b) suggested that many gall-causing Hymenoptera may be more readily identifiable by

their galls than by their own morphological characters.

Although a few gall insects are found on more than one host species, nearly all are specif-

ic to a single host genus. Cynipids have found optimal conditions on the oaks since 86% of

the known species are associated with this genus. Malyshev (1968) suggested that this can be

attributed to the fact that oaks are slow growing and have shoots that stay fresh and suscep-

tible to galling for a long time. Most of the remaining cynipid species are associated with

members of Rosaceae and 7% of these are restricted to the genus Rosa. A possible explana-

tion for this might be Malyshev’s suggestion that primitive Cynipidae caused galls on com-

mon ancestors of Rosales and Fagales and the two orders subsequently diverged.

Galls are not evenly distributed on various parts of their host plants. Besides being host

specific, gall formers restrict themselves to specific plant organs. Mani (1964) reported that

5% of the known cynipid galls on Quercus form on the roots, 22% on branches, 2% on flow-

ers, 4% on acorns, and about 63% on leaves. He also reported that over 80% of the galls on

Rosaceae are formed on leaves.

Kuster (1911) distinguished two kinds of galls on the basis of structure. He termed the

more primitive galls the kataplasmas and those more complex, the prosoplasmas. Both terms

have been widely used. The kataplasmic galls (e.g. those caused by aphids) are characterized

by a lack of both definitive tissues and constant external shape. Kataplasmic galls are com-

posed of homogeneous parenchyma cells, show little differentiation, and are structurally

similar to the meristematic tissues from which they develop. Prosoplasmic galls are charac-

terized by a definitive size and form. Their tissues, differentiated into well defined zones,

are fundamentally different from the normal host tissue. Most cynipid galls are proso-

plasmic. Wells (1921) presented evidence that prosoplasmic galls were phylogenetically de-

rived from kataplasmic galls.

Diplolepis rose gall community

57

Gall structure depends on many factors, including time of oviposition and number of eggs

laid in one area. Galls developing with one larva present are termed monothalamous and

those containing several larvae are termed polythalamous. In polythalamous galls, each larva

is individually surrounded by plant tissue.

Insect galls are often inhabited by numerous species besides the gall former because of the

attractiveness of localized concentrations of nutritive plant tissues. The inter-relationships of

insect gall inhabitants constitute one of the most important aspects of cecidology. One

European gall is reported to have over 75 species of insects associated with it (Mani, 1964).

One of the first tasks in studying inter-relationships of gall insects is to determine the

feeding habits of each species. Mani (1964) listed 33 different roles into which gall inhabit-

ants can be classified. I found five insect species associated with the Diplolepis polita galls in

my study area; they exhibit a variety of feeding habits. Both phytophagous and entomo-

phagous species are present. The gall-forming cynipids are phytophagous for their entire lar-

val stage. One of the inhabitants is also a phytophagous cynipid and although it is unable to

initiate galls of its own, it is able to cause further proliferation of gall tissues. Galls inhabited

by these insects are not only structurally modified, but also grow much larger. The four re-

maining species are entomophagous and feed on their hosts either as parasitoids or ectopara-

sites.

It must be stressed that associating entomophagous species with a particular gall does not

give information on host-prey relationships. Great care must be taken in rearing experiments

to determine these associations and the present study is one of few where relationships of

the entomophagous species in a gall community have been determined.

Little work has been done on the biology of insect galls in Alberta. There has yet to be a

checklist compiled for the galls of Alberta and Western Canada. Only the aphid galls have re-

ceived concentrated attention (Harper, 1959a, 1959b, and 1966; Cumming, 1968). A.C. Kin-

sey, in several of his works, mentioned receiving galls from Calgary, Alberta. Weld (1926) re-

corded that a worker in Toronto received galls of Diplolepis bicolor (Harris) and D. multi-

spinosus (Gillette) from Calgary.

The purpose of this paper is to examine the biology and inter-relationships of each species

found in the D. polita gall. It is also my objective to show that the associations of species

within the gall constitute a community. Each species has its role in the community and the

sequence of appearance of each species initiates changes in the community’s structure. Be-

cause various community attributes such as succession and climax can be examined with rel-

ative ease, studying galls may in many ways add to our general knowledge of community e-

cology.

All species studied in this work are new locality records for Alberta and greatly extend

known distributions. Long series of all species discussed have been deposited in the Strick-

land Memorial Museum, University of Alberta, the University of Saskatchewan Insect Col-

lection, and the Canadian National Collection of Insects in Ottawa.

STUDY AREA

All field work was conducted at the George Lake Field Station of the Department of En-

tomology, University of Alberta, 40 miles N. W. of Edmonton, Alberta (53° 57' N, 114° 06'

W). All galls used in the community inter-relationship studies were found within the one

square mile field station situated on the southern margin of the boreal mixed forest subzone

(LaRoi, 1 968). Fire is an important feature of the boreal mixed forest subzone and has in-

fluenced the ecology of George Lake. The forest is otherwise essentially untouched, with

58

Shorthouse

only some isolated logging prior to 1930. Ledum groenlandicum Oeder bogs and Car ex

species meadows are found in several places. There are a few open areas which allow bush

stratum species to grow densely. Principal trees of the upper stratum are Populus balsam-

ifera L. and P. tremuloides Michx. Other trees present, but less common, are Betula papyri-

fera March., Picea glauca Moench., Alnus tenuifolia Nutt., and several species of Salix. The

bush stratum is more diverse and the dominant species are Rosa acicularis Lindl., Rosa-

woodsii Lindl., Amelanchier alnifolia Nutt., Cornus stolonifera Michx., Ribes lacustre Pers.,

and Viburnum edule Michx. Common herbs are Epilobium angustifolium L., Heracleum

lanatum Michx., and several species of Solidago.

GENUS DIPLOLEPIS GEOFFROY

Dalla Torre and Kieffer (1910) and Weld (1952b, 1957, and 1959) gave excellent descrip-

tions of the family Cynipidae along with keys to the subfamilies and genera. A brief descrip-

tion of the genus Diplolepis was included by Dalla Torre and Kieffer. The main character

used to distinguish the genus is the plowshare-shaped hypopygium. Kinsey (1920b) gave da-

ta on the phylogeny of the cynipid genera and presented biological characters of each. So

far as known, Diplolepis species form galls only on Rosa.

There is considerable confusion in the literature about which generic name should be ap-

plied to cynipids forming galls on Rosa. Rhodites Hartig has been used extensively in cyn-

ipid literature, but Rohwer and Fagan (1917) established that Diplolepis Geoffroy had pri-

ority. Because Rhodites and Diplolepis are isogenotypic, Rhodites disappears in synonymy.

Some Europeans still use Rhodites as there is sentiment for having it placed on the conserv-

anda list, but Eady and Quinlan (1963) used Diplolepis in their key to the British species.

Kinsey and Ayres (1922) were the first North Americans to use Diplolepis. When Felt

(1940) republished his North American checklist of galls, he also changed to Diplolepis.

There has also been confusion as to whether Geoffroy (1762) or Fourcroy (1785) is the au-

thor of Diplolepis. Weld (1952a) reviewed the nomenclature problem and recognized Geoff-

roy.

Diplolepis is Holarctic in distribution. Dalla Torre and Kieffer (1910) and Eady and-

Quinlan (1963) gave keys to the European species. Dalla Torre and Kieffer also included a

number of North American species and provided brief descriptions of each. No inclusive key

to North American species has been published. All species descriptions are brief and require

extensive elaborations. Males are seldom mentioned in the literature. Undoubtedly new spe-

cies remain to be described and a complete revision of the genus may show some of the ex-

isting names to be synonyms. Felt (1940) recorded 25 species of Diplolepis, as well as many

varieties, as occurring in North America. There are now about 30 known species and two of

these (D. mayri Schl. and D. rosae L.) have been introduced. Weld (1957 and 1959) listed

the species found in various areas of the United States and gave brief descriptions of their

galls.

Diplolepis polita Ashmead and its Gall

Diplolepis polita was described by Ashmead ( 1 890) as forming galls on the leaves of Rosa

californica Cham, and Schlecht. As with other Nearctic Diplolepis, the recorded description

of D. polita is brief and inadequate. One of the key characters used to distinguish the species

is its smooth and shiny mesopleura. The mesoscutum, particularly the posterior region, is

not as rugose as in other species. Both Ashmead (1890) and Dalla Torre and Kieffer (1 910)

stated that males and females are entirely black. All females from George Lake have a red-

dish-brown abdomen as well as reddish-brown legs.

Diplolepis rose gall community

59

D. polita has been found only in North America. Ashmead (1890) examined specimens

from California, Dakota, and Colorado. Weld (1957) recorded polita as being found on the

Pacific coast but did not mention it (1959) as occurring in Eastern United States. D. polita

is not mentioned in Eastern North America checklists. Galls of D. polita were the most com-

mon of the Diplolepis galls found at George Lake in 1968 and 1969. I have collected speci-

mens throughout Alberta, but it appears to be most common in central and northern re-

gions. The previous most northern locality recorded was Ashland, Oregon (Bugbee, 1951).

Ashmead (1890) mentioned receiving galls of D. polita from Cockerell who had used the

manuscript name spinosellus. Cockerell (1890) stated thatD. spinosellus was a new species,

but gave no description of the gall former or the gall. Muesebeck et al. (1951) declared spin-

osellus Cockerell invalid. Krombein and Burks (1967) again used the name but gave no ref-

erence to descriptions of the gall former or the gall. Fullaway (1911) also made brief men-

tion of D. polita and its gall. According to Weld (1952a) Fullaway misidentified the polita a-

dults and instead considered them D. bicolor. Weld examined Fullaway’s specimens and

found those labelled D. bicolor were actually D. polita. Beutenmuller (1922) also obtained

some of Fullaway’s specimens described as D. bicolor and realizing they were not D. bicolor,

proposed the name D. occidentalis. Weld (1952a) examined Beutenmuller’s occidentalis and

confirmed its synonymy with polita.

The gall of D. polita is small (average diameter 4.0 mm), spherical, monothalamous, and is

spinulose and sometimes tuberculose. All galls collected were found on the adaxial surface

of the leaflets (Figs. 1 and 2), although McCracken and Egbert (1922) stated that they can

also be formed on stems. McCracken and Egbert also stated that the gall varies in size from 5

to 10 mm in diameter and often harbours inquilines. Their measurements were probably

from inquiline modified galls rather than unmodified D. polita galls. D. polita galls are usu-

ally found in clusters, several galls per leaflet (Fig. 2), although individual galls on a leaf have

been found. The largest number of galls found on one leaf was 39. Of all the galled leaves

collected, 61% were host to 5 galls or fewer. Galls growing close to one another often coa-

lesce.

Immature galls (Fig. 1) are often smooth or with weakly developed spines. They are soft

and composed of large succulent cells, many of which are visible to the naked eye. The po-

lita larva is tightly nestled in the interior of the gall where it feeds on the rapidly growing

cells. Cells next to the larva often appear much larger than other cells of the gall wall and it

is presumed that they play an important role in the larva’s nutrition. As the gall matures the

walls become brittle and the spines more conspicuous. The spines are easily broken off and

as a result galls handled in the laboratory for some time may appear spineless (Figs. 3-6).

The mature gall is hollow and smooth on the interior and the last instar larva has an in-

creased amount of space inside the gall (Fig. 3).

Nothing has previously been recorded on the anatomy of the gall other than brief com-

ments. Beutenmuller (1907) mentioned that the gall is thin walled and hollow. McCracken

and Egbert (1922) were the first to establish that the gall is monothalamous. My histological

studies (Ms. in preparation) show the gall to be prosoplasmic for the wall tissue is composed

of four well defined zones.

Two temporally separated groups of D. polita galls appeared at George Lake in both 1968

and 1969 seasons. Most galls appeared in the early spring on mature rose plants and in this

study are referred to as spring initiated galls. The second group of galls appeared later in the

season on new sucker shoots and are referred to here as sucker shoot galls. Sucker shoot

galls are somewhat different in appearance from spring initiated galls, often more densely

covered with long and hair-like spines. Although sucker shoots probably begin growth in the

60

Shorthouse

Fig. 1. Immature gall of Diplolepis polita on leaflet of Rosa acicularis Lindl. George Lake, Alberta. May 20, 1969. Fig. 2.

Mature galls of Diplolepis polita. George Lake, Alberta. August 10, 1969.

Diplolepis rose gall community

61

spring, they were first observed near the end of June in both seasons. Sucker shoots are ster-

ile and have larger and more succulent leaves than do older plants. They grow rapidly and

most attain a height of 0.9m by the season’s end. Their stems are densely spined and the tall

thin plants produce few side branches. New sucker shoots were more common around open

areas such as Ledum bogs and artificial clearings than in the forest.

It has been recorded by several authors that galls growing under various physiological con-

ditions differ in their colorations. Both greenish-yellow and red galls of D. polita were found

and the amount of sunlight received by the host plant appears to regulate colour. Cosens

(1912) stated that galls of Pontania pomum Walsh (Family Tenthredinidae) are poorly col-

ored if they grow in deeply shaded areas. D. polita galls growing in the shade of Populus

species are generally a light greenish-yellow. Galls on plants growing in open spaces such as

meadows, roadsides, and burned over areas are often bright red, especially when immature.

Galls appearing on sucker shoots and those growing in darkly shaded areas may be creamy

to pure white in color. Niblett (1943) noted this for the galls of D. eglanteriae Htg. and D.

rosarum Gir. As the D. polita galls mature they become brown.

Schroder (1967) found that galls of D. rosae were more numerous on roses growing under

stress. He reported that plants suffering from a lack of water supported more galls. These

plants were small, their yearly growth poor, foliage thin, leaves smaller than normal, and

often pale in color. They are unlikely to be the sucker shoot plants described in this paper.

Schroder found these plants growing in areas subjected to extreme insolation and although

they were common, their growth was poor. He observed ovipositions in both vigorous and

weak plants and found that no galls formed on the vigorous plants. He suggested that the os-

motic pressure of vigorously growing plants may be responsible for fewer galls. The rarity of

D. rosae galls on domestic roses also indicates that healthy plants are able to suppress gall

formation. D. polita galls appeared no more common on plants growing in open spaces such as

roadsides than they did on plants growing in shaded areas. Galls occurring in such areas

were, however, found to be much more brittle than galls growing in shaded areas.

In any study concerned with insect galls, it is vital that careful attention be paid to the

accurate identification of host plants. Although Diplolepis species are restricted to Rosa,

several species can form galls on more than one host species. Niblett (1943) recorded/), eg-

lanteria on 7 species of rose. Harrison (1922) exposed 16 species of rose to D. rosae and

found that oviposition took place only on members of one section. The 3 species of rose

found in Alberta are R. acicularis Lindl., R. woodsii Lindl., and R. arkansana Porter, all be-

longing to section Cinnamomeae. Lewis (1957) emphasized that the genus Rosa is one of

the most difficult groups to separate into distinct species. Species hybridize with ease giving

fertile offspring and the wide variation contributes to identification problems. Of the two

species found at George Lake R. acicularis is more common than R. woodsii and is generally

taller, less bushy, and has larger leaflets. R. woodsii flowers later in the season and usually

has more densely spined stems (both species determined by W. H. Lewis). Galls of D. polita

were found only on R. acicularis at George Lake. Lewis (1959) illustrated the Holarctic and

Nearctic distribution of R. acicularis and stated that it has the most extended native range

of any species in the genus. R. acicularis is native to Northern Europe, Asia, and North

America.

The Diplolepis polita Gall Community

I define an insect gall community as the assemblage of insect populations associated with

a collection of galls initiated by the same species of gall former. For the purpose of this pa-

per, the assemblage of insects obtained by making large random collections of galls within

62

Shorthouse

the study area, was considered to constitute the Diplolepis polita gall community. Mani (19

64) defined climax of gall community succession as being marked by a dominance of entom-

ophagous species. Climax in the D. polita gall community can also be defined as being re-

ached when the galls mature and fall to the ground, for once this stage is reached, oviposi-

tion and larval feeding activities cease. The periods of emergence and oviposition for all spe-

cies in the community occur in sequence. Emergence early or late in the season would re-

duce the reproductive success of that species.

The present study investigates five species found associated with the larvae and galls of D.

polita. These are Periclistus pirata (O.S.) (Cynipidae), Eurytoma longavena Bugbee (Eury-

tomidae), Glyphomerus stigma (Fabricius) and lorymus bedeguaris (Linnaeus) (Torymi-

dae), and Habrocytus sp. (Pteromalidae). Incidentals found associated with a very small per-

centage of galls were Eupelmella vesicularis Retz (Eupelmidae), Ormyrus sp. (Ormyridae),

and Tetrastichus rosae Ashmead (Eulophidae).

Gall inhabitants that feed on gall tissues and do not directly attack the gall former have

been termed inquilines. The term inquiline is derived from the Latin ‘inquilinus’ meaning

tenant or guest. According to Askew (1971), inquilinism is a form of commensalism, some-

where between parasitism and symbiosis. An inquiline lives in close spatial relationship with

its host, not feeding upon the host, but nevertheless frequently destroying it. In a commens-

al association all the advantages are to one of the partners and it is common that some com-

mensals do more harm to their hosts than depriving them of some larval food. Though

some workers have used inquiline in a somewhat different or broader sense (Triggerson

(1914), Malyshev (1968), Lyon (1969)), Askew’s definition seems most appropriate.

A close taxonomic relationship between the commensal and its host often exists in inqui-

linism. Muesebeck et al. (1951) listed the inquiline species of Periclistus, Ceroptres, Syner-

gus, and Euceroptres as being guests in insect galls, all being of the family Cynipidae. All

species of fbriclistus are restricted as inquilines of rose gall wasps as are Synergus of oak gall

wasps. It is probably safe to assume that inquiline cynipids had an ancestor capable of in-

ducing galls. This is indicated by the retention of the ability to induce cell proliferation in

many species such as P. pirata. Askew (1971) has suggested that inquilinism is on the road

to parasitism, demanding only that the inquiline become entomophagous rather than phy-

tophagous.

Some oak galls are inhabited by inquilines that do not interfere with the gall former. In-

stead they form irregular chambers inside the thick walls of their host gall (Stemlicht, 1968)

and do not come in contact with the gall forming larvae. Other oak gall inquilines occupy

the central chamber of the gall former (Askew, 1971). Inquilines such as Periclistus and Syn-

ergus species initiate chambers of their own inside galls and in the process obliterate the

chamber of the gall former. These species are incapable of initiating gall formation and are

completely dependent upon the gall formers for the provision of their larval food.

Osten Sacken (1865) was one of the First to question the inquiline behavior of Periclistus

species. Periclistus pirata was found in nearly all D. polita galls and all such galls were modi-

fied by the P. pirata larvae forming individual chambers. Because P. pirata does not feed up-

on tissues of D. polita, it cannot be described as a parasite or predator but instead fits

Askew’s definition. No galls inhabited by P. pirata contained the larva of D. polita. It is sug-

gested that the D. polita larva is killed when the P. pirata oviposits in the immature galls.

There is little uniformity in the literature as to usage of terms associated with entomo-

phagous species inhabiting galls. Common terms such as parasite and predator are often mis-

used. Smith (1916) distinguished parasites and predators on the number of hosts required to

complete development. He defined parasitic insects as those which pass their entire larval

Diplolepis rose gall community

63

stage within or upon a single host and predacious insects as those which require more than

one host to complete development. He also noted that a distinction between parasitism and

predation is of limited importance and it is wise to keep in mind that many species are call-

ed parasites only because they belong to parasitic groups and not by reason of their behav-

iour. Many students of galls use the term predator only when referring to birds or mammals

which break into galls. There is also a common feeding habit of entomophagous insects in-

termediate between parasites and predators for which the term parasitoid has been intro-

duced. Parasitoid has been suggested for those insects which destroy their hosts (which are

usually insects), are of a relatively large size compared with their hosts, and are parasitic as

larvae only (Doutt, 1959). It has not been widely accepted although has received some usage

in major works such as Askew, (1971). Its use has merit in gall studies because many of the in-

habitants are not true parasites nor can they be considered typical predators. Doutt (1964)

discussed the system for classifying entomophagous insects based on host relationships. Par-

asites attacking a phytophagous host are termed primary parasites. If the primary parasite is

attacked, then its enemies are called secondary parasites.

Eurytoma longavena, a common inhabitant of the D. polita gall, chews its way from

chamber to chamber often consuming two or more P. pirata during its development. Gly-

phomerus stigma and Torymus bedeguaris may feed ectoparasitically or as parasitoids. Blair

(1944) described G. stigma and a Torymus as ectoparasites. Habrocytus sp. was found feed-

ing only on individual larvae of P. pirata, that is. as an ectooarasite. Before examining the

community inter-relationships of the D. polita gall, it is first necessary to examine the biolo-

gy of each inhabitant species.

METHODS

Galls collected in both 1968 and 1969 were used for studying life cycles and the roles

played by each species in the community. Most of the galls collected in 1968 were used for

associating larvae of the inhabitants with their adults and studying life cycles of each. Galls

collected throughout the season were dissected to obtain larvae and determine their feeding

behavior. Most larvae were readily distinguished morphologically (Figs. 7-12) and for three

species, P. pirata, E. longavena, and G. stigma, the larvae were also identifiable by examining

the characteristic damage done to the gall tissues (Figs. 3-6). Eggs of only/’ pirata, E. long-

avena, and G. stigma were easily identified. When mature larvae were found, they were

placed in small pin-mounted gelatin capsules as described by Shorthouse (1972). Large num-

bers of larvae were stored in standard insect specimen boxes where they could be checked

for developmental changes. These specimen boxes were stored in a field laboratory under

nearly ambient conditions of temperature and humidity to obtain fall emergents. The re-

maining larvae were then returned to the university, stored at 4°C for 3 months, then incu-

bated at room temperature (approximately 22°C) when a 65% emergence was obtained.

Once the larvae pupated, the mandibles were removed from the larval cast skins and mount-

ed on slides. Mandibles of the larvae are structurally dissimilar (Figs. 13-18) and useful for i-

dentifications. A correlation among adult, larva, larval mandibles, and feeding behavior was

obtained in this manner for each species. Other collections of mature galls made in the fall

of 1968 were stored undissected in plastic vials. Fall emergents were removed as they ap-

peared and the galls then subjected to 4°C to break diapause.

Galls collected in the 1969 season were used for observing seasonal changes in the com-

munity composition. A total of 27 collections were made. Each collection was made by ran-

domly walking through areas of rose and collecting every gall observed. These walks were

2mm

Figs. 3-6. Mature Diplolepis polita galls. 3. Gall inhabited by Diplolepis polita. 4. Gall modified by Periclistus pirata. 5. Gall

modified by Eurytoma longavena. 6. Gall modified by Glyphomerus stigma, (a) Diplolepis polita larva; (b) Periclistus pirata

larva; (c) Habrocytus sp. larva; (d) Eurytoma longavena larva; (e) Glyphomerus stigma larva.

Diplolepis rose gall community

65

often more than 1,000 metres in length with the result that each collection was composed of

samples from numerous rose patches throughout the study area. When a gall or gall-cluster

was found, the entire leaf was picked and placed into an 1 8 ounce ‘Whirl-Pak’ bag. Size of

each collection was roughly governed in the field by collecting two bagfuls of galled leaves.

Only 1 1 of the 27 collections were used for the community study. The first 3 collections

were small because of the scarcity of galls in the early spring, but from June 6 until the end

of August, galls were sufficiently common that the two bags could be filled within two

hours. Because nearly all galls had matured by August 8 (Fig. 28), only one large collection

was used for August. Approximately 4,500 galls were collected in this manner. All galls were

returned to the laboratory, measured, dissected, and the contents examined, or the galls

were fixed in FAA solutions for later examination.

In this study, an empty gall is defined as one which does not contain a live inhabitant and

therefore cannot contribute to the community. Galls from which E. longavena or T. bede-

guaris emerged late in the season each had a tiny emergence hole and were considered sepa-

rately.

LIFE CYCLES OF GALL INHABITANTS

Diplolepis polita ( Ashmead)

Few data have been published on the biology of North American Diplolepis species and

nothing has previously been published on the biology of Diplolepis polita. Most North

American publications such as Bassett (1890), Beutenmuller (1907), Kinsey and Ayres

(1922), McCracken and Egbert (1922), Osten Sacken (1863, 1865) and Weld (1926, 1952a,

1952b, 1957, and 1959) are concerned mainly with species descriptions. Europeans and A-

sians have contributed much more to our knowledge of Diplolepis biology; notable exam-

ples being Blair (1944, 1945a), Callan (1940), Kuznetzov-Ugamskij (1930), Niblett (1943,

1947), Schroder (1967), and Yasumatsu and Taketani (1967).

Only 21 adults of D. polita were obtained throughout the study. All were reared from

galls stored in the laboratory. No adults were collected by sweeping or trapping in 1968 or

1969, nor were any adults observed ovipositing. No adults of other Diplolepis species were

collected in the study area either and it is assumed that field work began too late in the sea-

son. The earliest search for adults began May 7, 1969. Schroder (1967), in his study of D.

rosae (L.), found that emergence occurred over a period of from 2 to 6 weeks, extending to

8 weeks if the weather was cool. He found that more individuals emerged on warm sunny

days than on cool rainy days. He also observed that some females were able to pass a num-

ber of days at temperatures below the freezing point without harm. However, Kinsey

( 1 920a) found that most adult cynipids are killed by sudden changes of temperature or hu-

midity and that adults emerging during inclement weather would not oviposit. Niblett

(1947) suggested that late frosts are responsible for many casualties and in years when these

frosts occur, few galls are to be found. There may be inter-specific as well as generic differ-

ences in tolerance of inclement weather conditions.

Yasumatsu and Taketani (1967) observed and described the oviposition of D. japonica

(Walker) and estimated the time required for initial gall growth to occur after oviposition. It

is well established (Mani, 1964) that gall form: tion is due to larval feeding and if prolifera-

tion begins soon after the larva commences feeding, the period between oviposition and

hatching can be estimated. Yasumatsu and Taketani estimated that the egg stage of D. fa-

ponica lasts from 7 to 10 days. Callan (1940) experimented with D. rosae and found that

the first sign of gall formation was from 12 to 36 days after oviposition. Schroder (1967),

66

Shorthouse

studying the galls of the same species found hatching about 7 days after oviposition and that

the gall begins to grow 4 to 5 days later. I found the first D. polita galls May 20, 1969 so

probably oviposition occurred before May 10 in 1969 and as a result the first visible growth

occurred between May 20 and 25. Since no adults were collected when the field search be-

gan on May 7, D. polita emergence and oviposition probably takes place in late April or ear-

ly May. Alder and Straton (1894) suggested that adult life is shorter in species of gall wasps

which deposited eggs over a short period. D. polita adults lived for an average of 4 days in

the laboratory which suggests that their eggs are deposited over a short period of time. Kin-

sey (1920a) found that Diplolepis adults live for only a few days and must oviposit soon af-

ter emergence.

The eggs of D. polita are probably laid in or on the leaf primordia of slightly forced R. a-

cicularis buds. Schroder (1967) found that the eggs of D. rosae were deposited on the medi-

an vein of the pinnules as well as on the developing petioles. He found that the anterior ends

of the eggs are inserted into the epidermis of the developing leaflets, leaving the greater part

of the egg free between the folded leaves. A similar situation probably occurs with D. polita.

The eggs of D. polita are similar to the stalked eggs of other cynipids described by Berland

(1951). D. polita females must contain a large number of eggs for although their population

is low in the spring (Fig. 21), their galls were one of the most common in the study area. Ya-

sumatsu and Taketani (1967) found that D. japonica females contained an average of 331

eggs whereas Schroder (1967) found that 5 to 7 day old D. rosae females contained an aver-

age of about 780 eggs.

As with all Diplolepis species the larvae feed on host tissues and initiate formation of the

gall. Little data could be obtained on the time required to complete larval development

since this and the time of oviposition depend on factors such as condition of the host plant,

which undoubtedly differs from area to area. Hence periodic collections do not clearly in-

dicate the succession of larval instars. D. polita larvae have 12 body segments (Fig. 7), lack

setae, and undergo an estimated 5 larval instars. Mandibles of the last instar larva are triden-

tate (Fig. 13). The larvae grow rapidly and continue feeding on succulent gall cells until the

gall matures and hardens. Cosens (1912) stated that cynipid larvae feed only on cell con-

tents resulting in the occurrence of collapsed cells around the larva. No fecal material is

found inside the gall for the larval gut is blind. When the leaf tissue surrounding the gall ma-

tures, the galls fall to the ground where they are protected by snow through the winter. All

Diplolepis species overwinter as mature larvae. Laboratory reared specimens had a short pu-

pal stage lasting on the average about 10 days. Adults emerge inside the gall and must chew

their way through the wall to escape.

When the D. polita adults emerge in the early spring, it is assumed that they immediately

begin searching for oviposition sites. Callan (1940) found that most males of D. rosae ap-

peared before the peak appearance of females. It is well established that parthenogenesis oc-

curs throughout the genus and that males are rare, if found at all. Callan also suggested that

some species may exhibit geographic parthenogenesis, that is, males may be more numerous

in northern populations. Sex ratio of the 21 specimens I obtained was 0.714 (sex ratio=

number of females/ total number of individuals). Kuznetzov-Ugamskij (1930) recorded a

Diplolepis species from Asia with a sex ratio near 0.500 and in this species parthenogenesis

probably does not occur. Although most populations of D. polita may be parthenogenetic,

the occurrence of a comparatively high number of males indicates that some sexual repro-

duction occurs. Kinsey (1920a) suggested that in some primitive Diplolepis species, normal

sexual reproduction may take place, but in the genus as a whole, the male is gradually disap-

pearing and parthenogenesis is becoming the sole means of reproduction. The presence of

Diplolepis rose gall community

67

males in all collections of Diplolepis species found in central Alberta is consistent with the

geographic distribution of parthenogenesis as found by Callan (1940) and Schroder (1967).

Periclistus pirata (Osten Sacken)

The genus Periclistus Foerster consists of 7 North American species considered by most

authors (Muesebeck et ah, 1951) to be restricted to an inquiline habit in the galls of Diplo-

lepis species. Although the exact relationship between Periclistus species and the gall form-

ers are not known, it is accepted that the livelihood of Periclistus species depends on the

presence of Diplolepis galls. Periclistus larvae are phytophagous and feed on the same gall

tissue as do the gall formers; they cannot initiate galls. There have been no studies concern-

ed with specificity of Periclistus species for Diplolepis galls. Fullaway (1911) recorded P.

piceus Fullaway and P. calif ornicus Ashmead from galls of D. polita. Osten Sacken (1863)

described P. pirata and obtained the specimens from galls of D. ignota (O.S.). This present

study is the first record of P. pirata from the gall of D. polita.

P. pirata was an important occupant of the D. polita galls of George Lake in 1968 and 19-

69. Of the spring initiated galls 88.5% collected June 6, 1969 contained either eggs or larvae

of P. pirata (Fig. 21). Blair (1944) found that Periclistus sp. were present in nearly all the D.

rosae galls he examined. Although P. pirata larvae are phytophagous, in all galls examined in

which they were present, the D. polita larvae had been destroyed. Fig. 21 shows that as the

eggs of P. pirata become more abundant in gall collections, the number of galls containing a

live D. polita larva was reduced. Although the exact mechanism of this replacement is not

known, it has been recorded in other Diplolepis galls (Blair, 1945a). Because the D. polita

larva was always found dead in galls containing P. pirata eggs, oviposition by P. pirata fe-

males must kill the immature D. polita. Once the D. polita larva has been killed, it shrivels

and becomes difficult to detect.

P. pirata adults emerged early in the spring, probably two or three weeks after the D. po-

lita adults had emerged and oviposited (Fig. 19). The emergence of P. pirata is synchronized

with the appearance of the immature D. polita galls. Male P. pirata emerged before the fe-

males (Fig. 20). The first males were collected in the field May 16, 1969 the first females

May 20, 1969, and the first oviposition was observed May 23, 1969. By June 1, 1969, adults

of P. pirata were common and 263 were obtained by hand collecting. In the evening the a-

dults rested under the upper leaves of Rosa and could be easily dropped into collecting vials.

Sex ratio of the adults collected in this manner and from spring rearing experiments was

0.557. It is interesting that an inquiline cynipid species should have a population composed

equally of the sexes whereas the gall former populations are dominated by females. Copula-

tion was observed on many occasions both in the field and in the laboratory and it is there-

fore doubtful that parthenogenesis occurs. Over 300 observations of P. pirata ovipositing in

immature D. polita galls were made (see feature photograph, Ent. Soc. Can. Bull., 1970.2

(4): 102). If both immature and mature galls were present in a cluster, the immature galls

would always be chosen for oviposition first. P. pirata must have a lengthy emergence period

for although most of the population emerges in the spring, adults were found ovipositing in

galls up to August 7, 1969. The reappearance of immature galls in July of 1969 (Fig. 25) al-

lowed late emerging females the opportunity for oviposition. The July increase in the numb-

er of galls containing P. pirata eggs (Fig. 21) is due to the appearance of sucker shoot galls.

The mean number of eggs per gall for all collections is given in Table 1 . P. pirata females ovi-

posited readily in galls that contained eggs from other females. The largest number of eggs

found in a spring initiated gall was 23, largest number in a sucker shoot gall 16. This may be

68

Shorthouse

because more P. pirata females were present in the spring. The ease with which oviposition

could be induced in the laboratory and the observations of oviposition during rainy cool

weather, indicated that this species is much more hardy than Diplolepis species.

The egg of P. pirata is white, of the hymenopteriform type (Clausen, 1940), banana-

shaped, and stalked. The stalk, which is as long as the egg, is elastic and has a slight bulge at

the distal end. Upon hatching, the larvae distribute themselves around the inner walls of the

gall and commence feeding on gall tissues. P. pirata larvae initially feed on the same cells as

D. polita larvae and the area in which they feed is always marked by a layer of empty cells.

As the larvae continue feeding, gall tissue surrounds each individual to form an inner cham-

ber (Fig. 4). Blair (1945b) found that Synergus reinhardi Mayr (Cynipidae) modified the

galls of Cynips kollari Hartig (Cynipidae) in a similar manner. Galls containing P. pirata

chambers appear polythalamous and the original cavity, once containing a single D. polita

larva, is nearly obliterated (Fig. 4). P. pirata larvae (Fig. 8) are easily distinguished from

those of D. polita by the mandibles (Fig. 14). P. pirata larvae are not as active as D. polita

larvae and do not thrash as violently when disturbed. P. pirata inhabited galls fall to the

ground and receive the same winter protection under the snow. The seasonal change in the

percentage of galls with larvae and the mean number of larvae per gall is shown in Table 1.

The pupal stage of laboratory reared specimens lasted about 9 days. Many of the galls col-

lected in the fall of 1969 which overwintered in the laboratory at 4°C and were then moved

to 25°C were dissected 4 months after emergence had ceased. About 3% of the P. pirata in-

ner chambers contained live larvae. Under normal conditions these larvae may have been

destined for emergence later in the season or their presence may indicate that a small per-

centage of the P. pirata population remains in the larval stage throughout the season and e-

merges the following year.

Eurytoma longavena Bugbee

Eurytomids are one of the most common entomophagous groups associated with Diplol-

epis galls. Bugbee (1967) listed 82 species of North American Eurytoma and stated that 33

species attack Hymenoptera. He stated that at least 1 2 species are known to be phytopha-

gous and listed one species, E. pachyneuron Girault, suspected of being both parasitic and

phytophagous. Peck (1963) presented a comprehensive bibliography for 72 North American

species. Although most gall inhabiting eurytomids are considered parasitic, the lack of de-

tailed life cycle studies hinders such generalizations. Bugbee (1951) discussed 12 species

known from Diplolepis galls and warned that knowing the associated gall gives little data on

actual host relationships. Also in this paper he discussed the phylogeny of the Eurytoma

species associated with Diplolepis galls. He suggested that most Eurytoma are restricted to a

single species of gall former, but also listed several species known from more than one gall

and suggested that further studies will reveal more complex relationships. He pointed out

that some species may attack inquilines and other gall inhabitants besides the gall former.

Bugbee (1951) also stated that no complete life-histories have been worked out for any of

the Nearctic species associated with Diplolepis galls.

E. longavena was the most common and influential entomophagous occupant in the 1968

and 1969 D. polita gall community. This species was described by Bugbee in 1951 and was

found inhabiting D. bicolor galls growing on an undetermined species of Rosa. E. longavena

is known only from its type locality of Terrace, British Columbia and according to Bugbee

(pers. comm.) nothing is known of its biology. Three species of Eurytoma have previously

been associated with D. polita galls (Bugbee, 1951); these are E. flavicrurensa Bugbee, E. in-

certa Fullaway, and E. terrea Bugbee. E. longavena has not been previously recorded from

the D. polita gall.

Diplolepis rose gall community

69

Table 1. Incidence of Periclistus pirata eggs and larvae in the galls of Diplolepis polita.

George Lake, Alberta, 1969.

*means are calculated exclusive of galls without eggs or larvae

spring = galls initiated in the spring only

sucker = galls initiated on sucker shoots only

Larvae of E. longavena were found in 17% of the mature galls collected August 17-23,

1968 and in 32% of the galls collected August 22, 1969 (Fig. 21). Most of the empty galls

found in 1968 and 1969 were also a result of E. longavena activities. Gall collections de-

scribed under methods were used in tabulating incidence of E. longavena eggs, larvae, and

pupae in the galls of the 1 969 season (Table 2). E. longavena has two generations per year in

the study area, although only a small percentage of the total population is derived from the

second generation. From a large collection of 1969 spring initiated galls, 12.3% of the E.

longavena population emerged the same season and the remainder emerged the following

spring. Adults that emerged in the fall of the same season were able to oviposit in sucker

shoot galls. The E. longavena population found in sucker shoot galls emerges the following

spring. Clausen (1940) stated that the number of generations of Eurytoma per year is de-

pendent upon the hosts attacked and mentioned that E. monemae Ruschka may have three

generations per year.

Bugbee (1951) stated that the sex ratios for several species he studied were approximately

equal although females are usually more numerous. A total of 423 E. longavena adults were

70

Shorthouse

obtained in this study and the sex ratio was 0.912. The fall populations of E. longavena con-

sisted of females only and the sex ratio of all individuals collected, exclusive of fall emer-

gents, was 0.892. Niblett (1947) found that a small percentage of the E. rosae Nees popula-

tions emerged in the first year and that the sexes were in equal numbers. E. longavena fe-

males emerged late in the season from 5% of the mature galls collected in 1968 and from 7%

of the mature galls in 1969.

Fig. 20 indicates that E. longavena adults begin to emerge about the same time as P. pira-

ta and are probably present when the first D. polita galls appear. The first eggs were found

in galls collected May 25, 1969, although the earliest of 53 ovipositions observed in 1969

was June 5, 1969 (Fig. 19). Spring adults emerge over a long period of time as indicated in

Fig. 20, and may overlap the fall population. The last ovipositing female observed, August

9, 1969, thus may have been from either the spring or fall population. This extended activi-

ty period is also shown by the presence of eggs in sucker shoot galls (Table 2). These eggs

could have been deposited by either late spring emergents or fall emergents. The mean num-

ber of eggs laid per gall is also shown in Table 2. Dates without data were due to difficulties

in locating eggs amongst P. pirata chambers and uneaten gall tissues. As with P. pirata, E.

longavena would readily oviposit in galls containing eggs from previous females.

E. longavena eggs are similar in appearance to other Eurytoma eggs described by Clausen

(1940) and Phillips (1927). They are brown and have a small curled stalk at one end. The

stalk is about one-third as long as the egg. The eggs are white immediately after oviposition

but turn brown within 24 hours. Several E. longavena females were dissected and all con-

tained 6 eggs or less. The largest number of eggs found in a spring initiated gall was 1 1 and

the largest number in a sucker shoot gall was 4. E. longavena eggs were found only in im-

mature galls containing eggs or larvae of P. pirata. No eggs were found in galls containing a

D. polita larva, which indicates that the females can distinguish gall contents. Eggs were de-

posited along the inner walls of the gall and upon hatching the larvae immediately began

feeding on the eggs or larvae of P. pirata and later fed on any gall inhabitants they encoun-

tered. Because P. pirata was the most abundant species in the gall community, it was the

chief host of E. longavena. I observed several E. longavena larvae feeding even before they

were completely free of their egg shells. Later in the season E. longavena larvae were also

found consuming immature larvae of Glyphomerus stigma, Torymus bedeguaris, and habro-

cytus sp. E. longavena are also cannibalistic and though most galls attacked by this species

contained several eggs (Table 2), only one larva usually survived. Of the galls containing E.

longavena 94% produced a single adult. Galls producing two adults were large. Caltagirone

(1964) stated that when more than two Eurytoma sp. eggs were present in the Pontania gall

he studied, the larva that hatched first killed the remaining eggs.

E. longavena larvae are both entomophagousand phytophagous but they must feed on in-

sect tissue before they feed on plant tissue. This is shown by the fact that larvae often do

not survive if hatched in galls with completed P. pirata chambers. E. longavena larvae must

consume free host material before they are capable of chewing through chamber walls. Once

the larvae have reached a certain stage in their development, they are capable of feeding on

either plant tissues or insect tissues found inside the P. pirata chambers. Once the P. pirata

larvae are enclosed by gall tissues, the E. longavena larvae must chew through chamber walls

if more insect tissue is required. If the combined feeding activities of several E. longavena

larvae consume all insect host material before they are capable of phytophagous feeding, all

will perish and an empty gall results. Most galls contained sufficient immature P. pirata to

supply the E. longavena with food and also allow some P. pirata to escape and form cham-

bers. It is pertinent that the hosts are not killed at the time of oviposition, for the presence

Diplolepis rose gall community

71

of P. pirata larvae provide the E. longavena larvae with access to succulent parenchyma cells

and to other entomophagous species attracted to the chambers. Moser (1965), Caltagirone

(1964), and Malyshev (1968) reported that the gall inhabiting Eurytoma they studied stung

and paralysed the hosts at the time of oviposition. Blair (1944 and 1945a) suggested that E.

rosae found in galls of D. rosae were predators, but Niblett (1947) and Askew (1961) dis-

agreed.

When E. longavena larvae chew from chamber to chamber, uneaten plant tissues accumu-

late inside the gall and this condition is characteristic of Eurytoma damage (Fig. 5). Al-

though the presence of this tissue suggests that the larvae tear through chamber walls, I ob-

served many individuals ingesting plant cells, thus confirming that the species is phytopha-

gous. Varley (1937) recorded the same behavior for the larvae of E. robusta Mayr. Blair

(1954a) stated that the larvae of E. rosae feed on Periclistus larvae and chew into gall tissues

but did not record a phytophagous habit. Malyshev (1968) listed several other eurytomids

that initially feed on eggs and host larvae and then feed on gall tissues. It appears that once a

E. longavena larva consumes a certain amount of entomophagous material, it is capable of

continued development on gall tissues. Phillips (1927) in his study of E. parva (Girault), sug-

gested that this species, similar to E. longavena, is gradually breaking away from the ento-

mophagous habit and is becoming phytophagous. Blair (1945a) suggested the same for E.

rosae.

Mature E. longavena larvae are recognized by their distinct segmentation, abdominal pro-

trusions, and anterior sensory setae (Fig. 9). The larval mandibles (Fig. 15) are heavily scle-

rotized, triangular in outline, and each has one large denticle on the inner margin. E. long-

avena larvae, except for the fall emergents, overwinter inside the gall and pupate the follow-

ing spring. The pupal stage of laboratory reared specimens lasted about 1 2 days.

Glyphomerus stigma (Fabricius)

The genus Glyphomerus is monobasic containing the single species G. stigma. This spe-

cies, described in 1793 by Fabricius, has a Holarctic distribution and is known mainly as an

ectoparasite of gall inhabitants. Viereck (1916) recorded it from the gall of D. rosae and

Hoffmeyer (1930) from the gall of D. polita. Peck (1963) presented a bibliography of North

American records and Fulmek (1968) listed it as associated with 6 of the 7 European species

of Diplolepis. G. stigma is therefore not host specific for it has been recorded from 3 Near -

ctic galls (Peck, 1963) and 9 Palearctic galls (Fulmek, 1968). Blair (1945a) found the species

attacking Periclistus brandtii Er. in the galls of D. rosae. Other than these host records, very

little is known of its biology.

The same gall collections mentioned previously were used in tabulating the incidence of

G. stigma eggs and larvae in the D. polita galls collected in 1969 (Table 3). Larvae of G. stig-

ma were found in 21% of the mature galls collected August 17-23, 1968 and in 12.5% of the

mature galls collected August 22, 1969 (Fig. 21). G. stigma at George Lake is univoltine, al-

though Niblett (1947) found that a few adults associated with British galls emerged in the

fall of the first season. Fig. 20 indicates that the George Lake males emerge before females

and this was confirmed by field collections. Females are rapid fliers and unless they are

found with their ovipositors inserted deep into a gall, they are exceedingly difficult to cap-

ture. From gall emergence studies 139 adults were obtained and the sex ratio was 0.561. G

stigma has a lengthy period of emergence as indicated by the 42 oviposition observations in

Fig. 1 9 and the presence of eggs both in galls collected in spring and in sucker shoot galls

(Table 3). Niblett (1947) found that July and August was the normal emergence period for

British populations associated with D. rosae galls. I recorded the first oviposition May 23,

72

Shorthouse

Table 2. Incidence of Eurytoma longavena eggs, larvae, and pupae in the galls of Diplolepis

polita. George Lake, Alberta, 1969.

*means are calculated exclusive of galls without eggs

** estimate

spring = galls initiated in the spring only

sucker = galls initiated on sucker shoots only

1969 although the first eggs were not found until June 6, 1969 (Table 3). Laboratory emer-

gence studies (Fig. 20) indicated that the peak emergence occurs after that of P. pirata and

E. longavena.

G. stigma eggs are white to transparent and are banana-shaped with one end slightly thick-

er than the other. The thicker end also has a small knob at the tip. The long ovipositor of

the adult enables it to deposit eggs in large, thicker walled galls unavailable to E. longavena.

The eggs are laid on the inside surface of the gall cavity or directly on the host larvae. Once

the eggs hatch, the shells are almost impossible to detect and therefore Table 3 includes only

unhatched eggs. Some of the unhatched eggs may have been missed and this could be one

explanation for the sudden increase of galls containing larvae in the August 22, 1969 collect-

ion (Table 3). Of the 1969 attacked galls 80% contained a single egg and the remainder con-

tained two eggs. The larvae are cannibalistic and the first hatched consumes other eggs pre-

Diplolepis rose gall community

73

Table 3. Incidence of Glyphomerus stigma eggs and larvae in the galls of Diplolepis polita.

George Lake, Alberta, 1969.

spring = galls initiated in the spring only

sucker = galls initiated on sucker shoots only

sent, With the result that no more than one G. stigma larva was ever found per gall. Because

this species consumes all inhabitants of the D. polita galls in which they occur, it plays an

important role in the gall community. G. stigma larvae were found preying upon the larvae

of D. polita, P. pirata, E. longavena, T. bedeguaris, and Habrocytus sp., with P. pirata the

most important prey. Chamber wall tissue was consumed when the attacked gall contained

P. pirata, indicating that the species is phytophagous as well as entomophagous. Larvae of E.

longavena and Habrocytus sp. were consumed if they were found inside P. pirata chambers.

Galls attacked by G. stigma larvae have their interiors hollowed and other than being larger

(Fig. 6), are similar in appearance to normal galls containing only a D. polita larva. The lar-

vae consume most of the inner, more succulent tissues of the gall and as a result the dam-

aged gall has an interior lined with several layers of cell particles.

Blair (1945a) gave a brief description of the G. stigma larva. The mature larva is white,

tapers towards the anterior end and is clothed with long soft hairs (Fig. 10). The head is

cordiform and has two deep, elongated fossae that turn dark brown as the larva matures.

Mandibles of the mature larva are slender and curved with a denticle on the inner side some

distance before the apex (Fig. 16). Larvae overwinter inside the gall and pupate the follow-

ing season. The pupal stage of laboratory reared specimens lasted about 20 days.

74

Sh orthouse

Torymus bedeguaris (Linnaeus)

The genus Torymus includes both phytophagous and entomophagous species, the latter

mainly attacking gall makers and gall inhabitants. According to Huber (1927), genus Tory-

mus in North America is known to include 40 species that attack immature stages of Cyni-

poidea. Of the 106 Nearctic torymids listed by Peck (1963), 7 are recorded from Diplo-

lepis galls. Fulmek (1968) listed 103 European species associated with insect galls and re-

corded 6 of the 7 European Diplolepis as hosts. He recorded one Diplolepis species known

as the host for 10 Torymus species. T. bedeguaris, a Holarctic species, was found associated

with George Lake D. polita galls but it was not a common species in the community. Peck

(1963) listed three species of Diplolepis known as hosts of T. bedeguaris and Fulmek (1968)

listed 9 European gall formers as hosts. T. bedeguaris has not been previously recorded from

D. polita galls.

Only 132 T. bedeguaris adults were obtained in the two years and the sex ratio for the

series was 0.404. Clausen (1940) stated that there is a preponderance of females for all spe-

cies in which sex ratios are known. In the fall of 1968 39 adults emerged from galls collected

that spring and the sex ratio of these specimens was 0.435. No adults emerged in the fall of

1969 from galls collected that season, although 44 were obtained once diapause was broken.

Sex ratio of these specimens was 0.388.

All T. bedeguaris that emerged in the fall of 1968 did so in late August or early Septem-

ber. Emerging at this time, the adults would find sucker shoot galls available for oviposition.

Three T. bedeguaris were observed ovipositing in sucker shoot galls in 1968 and one in

1969. Eight other females were collected in the field in July and August of 1969, which in-

dicates that the species is active later in the season than other members of the community

(Fig. 19). Only 4 adults emerged from the 300 galls used in the spring emergence study (Fig.

20) and their emergence period was after that of most other species. Varley (1941) stated

that temperature influenced the fall emergence of T. cyanimus Boh., an ectoparasite of Uro-

phora jaceana Hering (Tephritidae). Varley (1937) stated that the larvae of T. cyanimus at-

tacked the full grown host larvae in August. He found some adults emerging in the fall, but

the majority passed the winter in the larval stage and emerged the following spring. In 1947

he reported that although most T. cyanimus adults emerged in May, no eggs or larvae were

found until August. As an explanation, he suggested that T. cyanimus may have an alterna-

tive host or the adults may wait from May until August before the eggs are matured and

laid. Moser (1965) reported that T. vesiculus Moser, an ectoparasite of Pachypsylla celtidi-

vesicula Riley (Psyllidae, Homoptera), has two generations per year with some of the first

and all of the second generation overwintering as mature larvae inside the gall. Although it is

strange that some T. bedeguaris emerged in the fall of 1968 and none emerged in the fall of

1969, the fact that so few specimens were obtained makes it difficult to discuss population

trends with any degree of confidence. T. bedeguaris may have two generations per year un-

der certain conditions. Their populations may have been high in the spring of 1968, and en-

suing conditions may have allowed many of them to emerge in the fall.

No T. bedeguaris eggs were identified in any of the 1968 or 1969 gall collections, al-

though they may have been confused with G. stigma eggs. Varley (1947) reported that T.

cyanimus often laid eggs in groups and although several larvae may be found feeding on the

same host, only one larva matured. Only four T. bedeguaris larvae were found in the August

22, 1969 collection (Fig. 21) and two of these were reared to adults. Ten other adults were

obtained from rearing larvae in gelatin capsules. Eight of these were found as first or second

instar larvae attached to paralysed larvae of either D. polita, P. pirata, G. stigma, or E. longa-

vena. The occurrence of first and second instar larvae late in August indicated that the spe-

cies may be capable of overwintering as an immature larva and continuing its development

Diplolepis rose gall community

75

the following season. Hosts of T. bedeguaris are completely consumed leaving only an emp-

ty cast skin. T. bedeguaris matures on a single host.

The mature larva of T. bedeguaris (Fig. 22) is white and clothed with more long hairs

than the larva of G. stigma. It bears both heavy and long sensory hairs and several rows of

long integumentary hairs in a band encircling each segment, giving it a distinctly hairy ap-

pearance. The posterior end does not taper abruptly as does the larva of G. stigma, nor does

it have the sunken fossae. The mandibles (Fig. 17) are narrow and acute without denticles

and are difficult to locate in the larval cast skins.

Habrocytus sp. (indet.)

The Pteromalidae contains some of the most common of the Chalcidoidea and many of

the species are known to attack larvae of Hymenoptera. The biology of most species re-

mains unknown and Peck (pers. comm.) stated that the entire genus Habrocytus requires re-

vision. Specific characters have yet to be worked out. Peck (1963) listed 31 species of Hab-

rocytus and recorded 3 associated with Diplolepis galls. Fulmek (1968) recorded 1 5 Europe-

an species and listed 4 associated with Diplolepis galls. The present study is the first record

of a. Habrocytus species from a gall of D. polita.

Habrocytus sp. larvae were found attacking only larvae of P. pirata, although I suspect

they attack D. polita larvae along with other inhabitants. P. pirata chambers are probably

completed and the larvae matured before Habrocytus sp. oviposits. Varley (1937) suggested

that H. trypetae Thoms, was not specific in its choice of hosts and would attack other para-

sites encountered. Blair (1944) found that H. bedeguaris Thoms, attacked full grown larvae

and pupae of Diplolepis and Periclistus and that cannibalism often occurred. Callan ( 1 944)

suggested that Habrocytus periclisti Callan was restricted to Periclistus brandti Ratzb., an in-

quiline in the galls of D. rosae. No eggs of Habrocytus sp. were found and the first larvae

were observed July 5, 1969 (Table 4) crawling over paralyzed P. pirata larvae. Only a single

larva was found per host and the number of larvae per gall is dependent upon the number of

P. pirata larva present (Table 4), the maximum recorded was 11. Superparasitism, as Varley

( 1 947) recorded for H. trypetae, was not observed, although if several eggs had been laid per

host, the first hatched could easily have consumed other eggs or larvae present. Urbahns

(1916) in his study of H. medicaginis Gahan also found only a single larva was able to devel-

op per host. They destroy their hosts quickly and only a round, black, pellet remains. This

pellet is readily visible in dissected galls (Fig. 4) and was used as the species indicator. Ur-

bahns (1916) stated that the larva of H. medicaginis can become fully developed in 6 days

after its first meal. Habrocytus sp. overwintered in the larval stage inside the gall.

Approximately 300 adults were obtained in this study and the sex ratio was 0.491. Callan

( 1 944) reported examining a series of H. bedeguaris reared from galls of D. rosae, in which

the sex ratio was 0.360 and a series of H. periclisti in which the sex ratio was 0.490. Habro-

cytus sp. adults emerged late in the season and were the last species in the D. polita gall

community to emerge under laboratory conditions (Fig. 20). Forty-two observations of ovi-

position were made, the earliest was June 22, 1969 and the last was August 20, 1969 (Fig.

19). Only two adults were observed ovipositing in sucker shoot galls. Sucker shoot galls may

not be readily attacked by Habrocytus sp. because few/* pirata larvae are able to form inner

chambers.

76

Shorthouse

Table 4. Incidence of Habrocytus sp. larvae in the galls of Diplolepis polita. George Lake,

Alberta, 1969

* means are calculated exclusive of galls without larvae

spring = galls initiated in the spring only

sucker = galls initiated on sucker shoots only

Habrocytus sp. is almost entirely univoltine and the adults emerge the following season.

Only six fall emergents were obtained in the two years and it is not known whether they

oviposited. Niblett (1947) found that a few adults of H. bedeguaris emerged in the fall of

the first year, but the majority emerged in July and August of the second year. Several ma-

ture galls incubated after exposure to 4°C for 3 months were dissected 4 months after in-

habitants had stopped emerging. Live Habrocytus sp. larvae were found in a few of the P.

pirata chambers indicating that the species may be capable of an extended larval stage and e-

mergence two seasons later. Many Habrocytus sp. larvae are probably consumed by larvae of

E. longavena, G. stigma, and T. bedeguaris. The decrease in percentage of galls containing

Habrocytus sp. larvae on August 22, 1969 (Table 4) is partly due to the feeding of these in-

sects. The inclusion of sucker shoot galls, which contain few if any Habrocytus sp. larvae, in

the August 22, 1969 collection, also decreased the percentage of galls with larvae.

The mature grub-like Habrocytus sp. larva lacks distinguishing features and has weak seg-

mentation (Fig. 12). The integument is smooth and sensory setae are reduced. The tiny

mandibles are simple and lack denticles (Fig. 18).

Diplolepis rose gall community

77

Figs. 7-12. Mature larvae of Diplolepis polita gall inhabitants. 7. Diplolepis polita. 8. Periclistus pirata. 9. Eurytorna long-

avena. 10. Glyphomerus stigma. 11. Torymus bedeguaris. 12. Habrocytus sp. Scale lines all 1 .0 mm.

78

Shorthouse

Figs. 13-18. Mandibles of mature larvae found in galls of Diplolepis polita. 13 . Diplolepis polita. 14. Periclistus pirata. 15.

Eurytoma longavena. 16. Glyphomerus stigma. 17 . Torymus bedeguaris. 18 . Habrocytus sp.

Diplolepis rose gall community

79

Diplolepis polita (estimated)

Periclistus pirata

Glyphomerus stigma

Eurytoma longavena

Habrocytus sp.

Torymus bedeguaris (estimated )

1 1 1 r

MAY JUNE JULY AUGUST

Fig. 19. Oviposition periods recorded for species associated with galls of Diplolepis polita. George Lake, Alberta, 1969.

SUCCESSION AND CLIMAX IN THE DIPLOLEPIS POLITA GALL COMMUNITY:

FATE OF MEMBER SPECIES

Mani (1964) briefly introduced the study of plant gall communities. He emphasized that

the predator-parasite complex of galls is often considerably larger than that of inquilines and

while some galls lack inquilines few, if any, are free from entomophagous inhabitants. The

classic paper on this subject is by Varley (1947) in which he discussed factors controlling

population density of the knapweed gall-fly. Although there are many factors regulating the

gall former population, such as weather and availability of oviposition sites, it is the objec-

tive of this section to examine the roles played by each member species in the gall communi-

ty.

D. polita is the central species in the gall community for it causes gall formation and with-

out the gall none of the subsequent species could exist. When the galls first appeared in the

spring, D. polita predominated. The first three collections in 1969 contained only larvae of

D. polita (Figs. 21 and 23). The D. polita larval population decreased once the eggs of P. pi-

rata appeared. By May 28, 1969, only 18.5% of the galls contained a live D. polita larva, al-

though only the eggs of P. pirata and E. longavena were present in the remaining galls (Fig.

21). If E. longavena occurred in galls without P. pirata, it would undoubtedly devour the D.

polita larva resulting in a further decrease in the D. polita population. The largest population

of P. pirata larvae was found early in the season (Fig. 23) and because they formed the main

food source of the entomophagous species, their dominance soon began to decline. E. longa-

vena had the greatest influence on the P. pirata population and the number of galls contain-

ing the former species had risen substantially by June 20 (Fig. 21). Varley (1947) found

that E. curta Walker was chiefly responsible for controlling the population density of the

knapweed gall-fly. Cannibalistic activities of E. longavena probably prevented a major

NUMBER OF INDIVIDUALS

80

Shorthouse

[

20r Torymus bedeguaris

10

0

20

10

0

20

10

0

20

10

0

20

10

0

20

10

0

Habrocytus sp.

n

n_Jl

s

n

4

Glyphomerus stigma

n

i 1 r

Eurytoma longavena

. „

n rpllip I ■ T f ■ , ■

_ Periclistus pirata

R

ai

Diplolepis polita

□ MALE

■ FEMALE

□ UNSEXED

T T I I I 1 1 1

0 2 4 6 8 10 12 14 16 18

TIME IN DAYS

Fig. 20. Spring emergence from 300 galls of Diplolepis polita stored at 3°C for 3 months then transferred to 22°. The day

of first emergence is day 1. Galls collected at George Lake, Alberta, 1969.

Diplolepis rose gall community

81

change in the proportion of E. longavena to P. pirata between June 12 and June 27 (Fig.

23). As the season advanced, feeding activities of E. longavena continued to reduce both the

P. pirata population (Fig. 23) and the number of galls containing P. pirata (Fig. 21). Once

all P. pirata in a gall were consumed, there was an increase in the percentage of galls contain-

ing only E. longavena (Fig. 21, July 5). E. longavena larvae perished if they did not obtain

sufficient food before the supply of P. pirata was depleted. This was indicated by a decrease

in percentage of galls containing £. longavena between July 5 and July 14 (Fig. 21). The

first larvae of G. stigma appeared June 27 although this species was not abundant until later

in the season. Larvae of Habrocytus sp. had only a slight influence on the P. pirata popula-

tion in the first few collections in which they appeared (Fig. 23).

The occurrence of sucker shoot galls is shown by the reappearance of P. pirata and E. lon-

gavena eggs in the July 14 collection (Fig. 21). P. pirata was as detrimental to D. polita in

sucker shoot galls as in spring initiated galls. The presence of sucker shoot galls did little to

increase the D. polita population, although it was beneficial to both P. pirata and E. longave-

na (Fig. 21). Only 6 galls containing a D. polita larva were found in the 107 sucker shoot

galls collected July 23. Fall emergence of E. longavena enabled this species to oviposit in a

large percentage of the sucker shoot galls and as a result the sucker shoot population of P.

pirata was reduced. Because G. stigma larvae consumed all occupants of the galls they inhab-

ited, their presence in 5% of the July 14 galls represented a substantial decrease in the num-

bers of other larvae. No G. stigma larvae were found in sucker shoot galls and therefore the

inclusion of sucker shoot galls in the July 23 collection lowers the percentage of galls con-

taining this species (Fig. 21). The presence of E. longavena and P. pirata in sucker shoot galls

increased the relative abundance of these species in the July 23 collection. Had these addi-

tional larvae not been present in sucker shoot galls, the proportion of G. stigma and Habro-

cytus sp. would have been higher. Most of the P. pirata larvae in sucker shoot galls were con-

sumed by mid- August and this also explains the further decrease in the numbers of this spe-

cies (Fig. 23). Because each Habrocytus sp. requires oneF. pirata larva, their presence help-

ed decrease the number of P. pirata in the July 23 and August 22 collections (Figs. 21 and

23).

Fig. 24 shows the inter-relationships of all species composing the D. polita gall communi-

ty. Although D. polita was the key species in the community, later in the season P. pirata

took over the central position of the food web. P. pirata had the greatest influence on the D.

polita population and the presence of this species greatly increased biomass in the communi-

ty. The remaining entomophagous species depended upon P. pirata as their chief source of

food. Habrocytus sp. larvae were second to those of E. longavena as the chief destroyers of

P. pirata. G. stigma and T. bedeguaris did not restrict their attack to any one species. Their

importance in the community is therefore dependent upon the number of inhabitants in

each gall attacked.

The most common cause of empty galls in 1968 and 1969 was food shortage. E. longa-

vena and G. stigma perished if the galls they inhabited did not contain adequate food for

their development. This was less likely for T. bedeguaris because they oviposited later in the

season when competition between entomophagous species was nearing its climax. The interi-

or of empty galls often contained remains and particles of gall tissue. From the two seasons

8% of the empty galls contained only the remains of first instar larvae of P. pirata and had

no internal chamber development. Abnormal environmental conditions in the gall cavity or

some unusual physiological condition of the plant tissue may have had a toxic effect which

killed the gall inhabitants. Sometimes P. pirata females may have killed the immature D. po-

lita larva and then failed to oviposit. About 15% of the empty galls in the August 22 collec-

tion contained fungus. This fungus may have initially attacked plant tissues resulting in the

Percent Percent Percent

82

Shorthouse

100-

90-

80-

70-

60-

50-

40-

30-

20-

10-

30-

20-

10-

II

mi

iv

IV,

Diplolepis polita

Eggs, 1969 Periclistus pirata

SSS Eu rytoma longavena

Glyphomerus stigma

Torymus bedeguaris

Habrocytus sp.

x^-r

B

Larvae, 1968

Larvae, 1969

o in oo •

(N CN <N <3

Fig. 21. Percentages of galls containing members of the Diplolepis polita gall community. Number of galls in each collec-

tion is indicated. George Lake, Alberta, 1968 and 1969. A. Eggs found in galls collected May to August, 1969. B. Larvae

found in mature galls collected August 17-23, 1968. C. Larvae found in galls collected May to August, 1969.

Diplolepis rose gall community

83

death of the inhabitants or it may have developed on dead inhabitants and subsequently

spread throughout the gall interior. Disease undoubtedly killed some inhabitants.

40-

3 30

<

b20

O

10

0

GALLS WITHOUT

n EMERGENCE HOLES

GALLS WITH

EMERGENCE HOLES

n n n n fl

.1

17-23 20 25 28 6 12 20 27 5 14 23 22

AUG. MAY JUNE JULY AUG.

1968 1969

SAMPLE DATES

Fig. 22. Incidence of empty Diplolepis polita galls. George Lake, Alberta, 1968 and 1969.

Once the galls matured and fell to the ground, no further population additions took place

and by analysing collections of these galls, climax of the succession was fixed. Also, after an-

alysing the percentage of galls containing each species (Fig. 21) and the relative abundance

of each species (Fig. 23), one can predict population trends for the following season. Pre-

dictions of this nature are dependent upon many factors, such as weather, which may have

varying effects on the emergence of each species. It was obvious that the occurrence of D.

polita in 9% of the 1968 mature galls was sufficient to allow for an abundance of galls in

1969. This indicates that the D. polita gall community is constructed to tolerate low num-

bers of the gall former. Several authors have found the same for other galls (Askew, 1961,

84

Shorthouse

23-VII

5* VII

22-VIII

14-VII

Diplolepis polita

Periclistus pirata

Eurytoma longavena

Glyphomerus stigma

Torymus bedeguaris

Habrocytus sp.

( larvae only)

Fig. 23. Incidence of species in the Diplolepis polita gall community expressed as percentage of the total populations of gall

inhabitants. George Lake, Alberta. 1969.

Diplolepis rose gall community

85

Fig. 24. Food web of species of the Diplolepis polita gall community. George Lake, Alberta, 1968 and 1969. Heavy lines

represent phytophagous habit, narrow lines represent entomophagous habit.

Evans, 1967; and Gordinier, pers. comm.). By the end of 1969, the D. polita population was

reduced and initial field observations in 1970 indicated that the galls were less common than

in 1969. The occurrence of P. pirata larvae in 27% of the 1968 mature galls was chiefly re-

sponsible for the decrease in the D. polita population by the end of 1969. The availability of

P. pirata larvae allowed an increase in the E. longavena population which in turn was partial-

ly responsible for the increase in empty galls. Few Habrocytus sp. larvae were found in 1968

mature galls and their abundance in 1969 was probably due to the increase in the P. pirata

population. The decrease in numbers of G. stigma from 1968 to 1969 may be due to some

unknown factor affecting only the biology of this species. For the 1970 community struc-

ture, I predict a large increase in D. polita, because of a decrease in the abundance of P. pi-

rata. A decrease in P. pirata larvae would also cause a decrease in all entomophagous species

and this again would reflect the importance of P. pirata in the gall community. This reduc-

tion of entomophagous species would allow for a 1971 increase in the P. pirata population

86

Shorthouse

which in turn would decrease the D. polita population again.

GROWTH OF DIPLOLEPIS POLITA GALLS

The objective of this section is to correlate seasonal changes in gall dimensions with gall

contents. In this section the growth rate of galls inhabited by a single D. polita larva is com-

pared with the growth rate of galls inhabited by P. pirata eggs and larvae.

All studies on gall growth were made in 1969 and the field search for both galls and a-

dults began May 7, 1969. No leaves of R. acicularis were out at this date, although those of

Populus and Salix were just appearing. The first R. acicularis leaves were found May 8,

1969, in areas of greatest insolation; by May 11, 1969, immature leaves were present on

nearly all rose plants. The first D. polita galls, 2 leaves with 2 galls on each, were collected

May 20, 1969. From May 25, 1969 on, galls were much more common. The diameters of all

galls in the 1 1 random collections described previously were recorded and correlated with

gall contents. The mean sizes of galls containing either a D. polita larva or P. pirata eggs or

larvae, for all collection dates, are presented in Tables 5 and 6. In another study, 80 one

square metre quadrats were randomly marked off in 4 different rose patches. A total of 1 34

galls on 30 leaves were found within these quadrats. Each gall was examined and measured

approximately every 7 days and data were obtained on their growth rate, shrinkage, senes-

cence, and leaf abscission. Once the galls had fallen, they were returned to the laboratory

for dissection. Unfortunately none of these galls contained a larva of D. polita.

Growth of Galls Inhabited by larvae of Diplolepis polita only

The mean diameter of 44 mature galls (Fig. 26) collected in 1968 and containing only a

single D. polita larva was 3.8 mm (S.D. 0.47). An estimate of the growth curve for normal

galls collected in 1969 containing a D. polita larva is shown in Fig. 26. By studying growth

curves of individual plot galls, it was estimated that the maximum size of normal D. polita

galls occurred around the middle of July. After this date there was shrinkage and the final

size due to gall maturation was reached by the middle of August. The average amount of

shrinkage in gall diameter for the 134 plot galls was 0.82 mm (S.D. 0.57) indicating that

some of the mature D. polita galls collected August 22, 1969 (Table 5) could have been as

large as 5.1 mm in diameter. Undoubtedly growth rate and condition of the host plant af-

fects growth rates of attached galls. Factors such as soil condition and availability of light

and water affects plant growth rates and must also influence growth rates of galls. Oviposit-

ing in buds not in an optimum condition for galling could affect gall size. Positioning of the

gall on the leaflets, the number of leaflets per leaf, and the number of galls per leaflet and

leaf, could also influence gall size. Galls growing on older plants may have a different growth

rate and final size compared to galls growing on younger plants and sucker shoots.

The first immature sucker shoot galls, all less than 2.9 mm in diameter, were found July

14, 1969. As the season advanced, immature galls on these shoots became more abundant

and several collections were made up to August 13. Fig. 25 shows an increase in the number

of immature galls less than 4.0 mm in diameter on July 14. Each successive collection con-

tained a decreasing number of immature galls less than 4.0 mm in diameter as maturation

processes began (Fig. 25), but the July 14 collection, and the three that followed, showed

an increase in the number of immature galls. This increase illustrates the appearance of suck-

er shoot galls. In the July 23 collection, 106 of the 296 galls were from sucker shoots and

6 of these contained a D. polita larva (Table 5). Sucker shoot galls less than 4.0 mm in di-

ameter from two further collections (July 28, 1969 and August 13, 1969) are included in

Fig. 25, but their contents are unknown. All galls in the August 22, 1 969 collection were

Diplolepis rose gall community

87

mature and the sucker shoot galls in this collection were combined with spring initiated galls

(Fig. 21).

Table 5. Mean diameters of galls containing one larva of Diplolepis polita. George Lake,

Alberta, 1969.

spring = galls initiated in the spring only

sucker = galls initiated on sucker shoots only

Immature galls appearing on sucker shoots in July could be due to a delayed hatching

mechanism, as mentioned by Yasumatsu and Taketani (1967). They found that/), japonica

has two periods of gall formation, each appearance of the galls being dependent on the

length of time before hatching. They reported that the first group of galls began developing

7 to 10 days after oviposition and the second group began developing 40 days after oviposi-

tion. It is possible that the increase in number of immature D. polita galls near the middle of

July is a result of such a delay. If all D. polita eggs were laid around May 1, this second

group of immature galls would be developing after approximately 70 days’ hatching delay.

Although all 13 immature galls collected July 14 contained P. pirata eggs, the remains of a

D. polita larva were found in 5 of them. It appears that hatching of the eggs laid in the buds

of sucker shoots is more delayed than in buds of older plants, perhaps because of some

physiological condition within the host plant.

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Shorthouse

COLLECTION DATES

Fig. 25. Seasonal change in the numbers of immature Diplolepis polita galls less than 4.0 mm in diameter, expressed as a

percentage of each gall collection. George Lake, Alberta, 1969.

Growth of Galls Inhabited by Periclistus pirata

Although many authors have discussed the position of ‘inquilines’ in gall communities,

few have mentioned their ability to increase gall size. Niblett (1947) was one of the few to

show this and stated that the Diplolepis gall he was studying showed a great variation in size

when inhabited by Periclistus larvae. Blair (1945) maintained the opposite for the inquiline

Synergus reinhardi Mayr in the galls of Cynips kollari Hartig, suggesting that inquiline larvae

may inhibit gall growth. Evans (1967) stated that if the Besbicus mirabilis (Kinsey) gall is in-

habited by the inquiline Ceroptres species, the immature gall ceases to grow and becomes

hard and brittle. Yasumatsu and Taketani (1967) found that galls of D. japonica attacked by

Periclistus sp. were irregular in shape, but they made no mention of size changes.

Eighteen of the 55 D. polita galls collected May 25, 1969, contained eggs of P. pirata and

Diplolepis rose gall community

89

in all 1 8 the larva of D. polita had been killed. In the May 28 collection, 44 of 54 galls con-

tained P. pirata eggs and their mean size was greater than that of the 10 remaining (Fig. 26).

Initiation of inner chamber development by the P. pirata larvae was first observed June 20,

1969 and by July 14, chambers were completed in 82% of the galls containing/*, pirata lar-

vae. The largest spring initiated gall containing/*, pirata was found July 5, 1969 and was

12.4 mm in diameter.

The increased size of galls containing /*. pirata eggs (Table 6) results from additional cell

proliferation. Substances that cause the proliferation could be injected into gall tissue at the

time of ovipositon or the eggs may secrete activating substances. Once hatched, larval feed-

ing activities also contribute to the increase in gall size. Even though predators may destroy

all gall inhabitants, gall size has usually already been influenced by the P. pirata larvae. Pre-

dation after P. pirata had influenced gall size, results in many large galls without P. pirata

chambers. Similar findings were also recorded by Niblett (1947).

Table 6. Mean diameters of Diplolepis polita galls containing eggs or larvae of Periclistus

pirata. George Lake, Alberta, 1969

spring = galls initiated in the spring only

sucker = galls initiated on sucker shoots only

90

Shorthouse

Galls formed on sucker shoots later in the season are also attacked by P. pirata. Another

characteristic of sucker shoot galls is that they do not attain the size of P. pirata enlarged

galls initiated in the spring. The largest sucker shoot gall found was 6.4 mm in diameter. Of

the 106 sucker shoot galls found July 23, 1969, 50% contained larvae of P. pirata, but 94%

of these galls had no inner chamber development. Mean size of these 53 galls was 4.6 mm

(S.D. 1.0), significantly smaller than the mean of 8.1 mm (S.D. 1.6) for spring initiated

galls (Fig. 26).

I 1 1 1

MAY JUNE JULY AUGUST

1969

1968

Fig. 26. Seasonal change in the mean size of galls containing a singl e Diplolepis polita larva (solid lines) compared to galls

containing Periclistus pirata eggs or larvae only (broken lines). Vertical lines indicate one standard deviation each side of

the mean. George Lake, Alberta, 1968 and 1969.

Gall Senescence and Abscission

Maturation of gall tissue affects nearly all the insects associated with a gall community.

Most of the phytophagous larvae are only capable of feeding as long as plant cells remain

soft and succulent. All feeding activities of D. polita and P. pirata larvae are terminated once

gall tissues mature. Gall maturation also offers gall inhabitants some protection from pred-

ators and parasites. Oviposition activities of predators and parasites are influenced by the de-

gree of tissue maturation. Askew (1961) found that as a gall matured, there was an increase

Diplolepis rose gall community

91

$5 to

<s

Fig. 27. Percentage of galled leaves with necrotic tissue in collections of spring initiated Diplolepis polita galls. George

Lake, Alberta. 1969. Fig. 28. Rate of gall maturation of spring initiated Diplolepis polita galls found in 80 random plots.

George Lake, Alberta. 1969. Fig. 29. Rate of gall abscission of spring initiated Diplolepis polita galls in 80 random plots.

George Lake, Alberta. 1969.

92

Shorthouse

in the time taken for a parasite to pierce the gall wall. He also measured gall hardness and

found that the walls of mature galls of Cynips divisa Htg. were 200 times more resistant to

crushing than were the walls of immature galls.

Ignoffo and Granovsky (1961) defined gall senescence as the process of turning brown

due to tissue necrosis. They considered a gall necrotic when seven-eighths of the surface was

brown. Mani (1964) stated that the nutritional deficiency of a leaf beyond the gall is first

observed when the gall begins to mature. Early maturation of a galled organ is one of the af-

fects of gall formation on the host and was observed in galled R. acicularis leaflets in the

present study. The percentage of galled leaves with necrotic tissue, found in the 1 1 major

collections of 1969, is shown in Fig. 27. Only the spring initiated galls are represented in

this graph. All galled leaves had necrotic tissues by July 28, 1969, whereas the first discolor-

ation of normal leaves was seen in the last week of August.

D. polita galls were considered mature when at least 75% of the gall tissue was dark

brown. The rate of gall maturation was determined by examining the 134 galls found in the

random plots described previously (Fig. 28). The first mature gall was observed June 28,

1969 and all were mature by September 6, 1969.

Once a mature gall has fallen to the ground, it can be considered immune to attack by

most predators and parasites though rodents undoubtedly consume some fallen galls. The

rate of gall abscission was also determined by examining the 134 plot galls. The first plot

gall had fallen by July 14, 1969 and all had fallen by September 28, 1969 (Fig. 29). Galls

growing in large clusters probably fall before galls growing singly because of their combined

weight. Large gall clusters often cause the entire leaf to hang vertically (Fig. 2). Galls with

their weight increased by P. pirata larvae probably fall before galls containing a single D. po-

lita larva. Ignoffo and Granovsky (1961) found that the gall of Mordwilkoja vagabuhda

Walsh (Aphididae) prevented the formation of an abscission layer and the gall may remain

on the host for 3 years. Yasumatsu and Taketani (1967) found the first galls of D. japonica

began falling 39 days after initiation. If the average initiation date for the D. polita galls can

be considered about the middle of May, then the first galls fell approximately 60-70 days af-

ter initiation. D. polita galls overwinter on the ground and because they fall before normal

leaf abscission occurs, their subsequent covering by the autumn complement of leaves helps

to protect the gall inhabitants against winter.

DISCUSSION

This investigation of the Diplolepis polita gall and its inhabitants has revealed many basic

features of cynipid gall ecology. It is apparent that by studying cynipid galls, one has the op-

portunity of gaining new information on such basic concepts of biology as community ecol-

ogy, insect-plant specificity, plant developmental morphology, and the evolution of special-

ized insect groups.

D. polita is the central character in the gall community although as the season advances

its dominance in terms of biomass is soon lost to other gall inhabitants. Periclistus pirata is

mainly responsible for the rapid decline in the D. polita population. The entomophagous in-

habitants that subsequently invade the galls depend more on the larvae of P. pirata as their

source of food than they do on D. polita larvae. It therefore can be shown that the gall for-

mer prepares requisite conditions for the inquiline, which in turn provides requisite condi-

tions for the entomophagous species. By the end of the season, the community consists of

much larger proportions of inhabitants other than the gall former and it is the relative pro-

portions of these inhabitants that determines community structure the following season.

Diplolepis rose gall community

93

The Diplolepis species complex has received little attention and the entire genus is in need

of taxonomic revision. Several of the names in use are incorrect. Species have been distin-

guished mainly by their external morphology and because the species exhibit limited varia-

tion, use of many of these characters may have led to the taxonomic problems.

A great deal has yet to be learned about the biology of D. polita and for that matter, all

Diplolepis species. Adult D. polita can undoubtedly be observed in the field if the researcher

is at the right place at the right time. Condition of the host plant at the time of oviposition

and gall initiation should be easily delineated. Surprisingly little is known about the induc-

tion of cynipid galls. Plant biochemists and morphologists would undoubtedly be interested

in learning of the chemical stimuli these insects have evolved to cause cell hypertrophy and

hyperplasy. The various stages of gall development from the time of oviposition to gall ma-

turity may reveal structural features that influence activities of the associated insects. The

presence of males in the D. polita population indicates that normal sexual reproduction oc-

curs and in light of theories of north-south gradations in parthenogenesis, larger populations

of all northern species should be examined. The appearance of sucker shoot galls also re-

quires further investigation. If ovipositing females can be handled in the field, controlled

ovipositions in both spring and sucker shoot host plants and subsequent observations of gall

development should reveal whether there is a delay mechanism in initiation of sucker shoot

galls.

From the data obtained on the biology of Periclistus pirata, it is apparent that inquilines

have an important role in cynipid gall ecology. They take an important position in the com-

munity and grossly modify the normal gall structure. Exactly how Periclistus disposes of the

Diplolepis larvae requires further investigation. Because D. polita galls were observed to en-

large even before the Periclistus eggs hatched, it appears that substances inducing further cell

hypertrophy come from either the ovipositing females or the unhatched eggs. Periclistus lar-

vae may exhibit cannibalism, but this has yet to be determined. It would be interesting to

compare the manners in which Diplolepis and Periclistus stimulate and modify the plant tis-

sues with which they are in contact. From observations of immature D. polita galls it ap-

pears that the immature Diplolepis larva is surrounded by plant tissues when it hatches and

subsequent feeding causes cellular hypertrophy and hyperplasy in all directions from the

larva. When Periclistus larvae feed, the plant cells are stimulated in such a manner that the

tissues grow up and around the individual larva. It will be interesting to determine whether

other Diplolepis galls are modified by Periclistus in a similar manner. If the structure of

other galls can be modified as extensively as those of D. polita, then many of the gall de-

scriptions and illustrations now in the literature for Diplolepis may be inaccurate.

Plant morphologists may benefit from studies of cynipid gall developmental morphology.

How these insects gain control of morphogenetic potentialities of host organs remains un-

known. Galls cannot be regarded as organs, but they are more than tissue abnormalities for

they have constant size and structure. An interesting problem for the morphologist would

be to determine whether galls and the tissues composing them can be regarded as ‘new’

structures, morphologically different from familiar structures. Galls of Diplolepis have cells

and tissues unlike those normally found in the host plant. We require more information on

the mechanisms that induce gall cells to divide without reference to the morphogenetic char-

acter of the host organ. It may even be easier for morphologists to study processes of form

determination in galls than in normal developmental processes in plants. In galls the induc-

ing agent is not part of the physiological mechanism of the plant, but rather is introduced

into the plant.

Insect galls afford numerous opportunities for studying insect-plant host specificity and

specificity of the inhabitants. It is now known how certain insects are attracted and restrict-

94

Shorthouse

ed to either one or several closely related hosts, but little, if any, attention has been given to

why a species is restricted to one organ of the host plant. Why, for example, is D. polita re-

stricted to leaves? Would gall tissue develop if immature larvae were transplanted into meri-

stematic tissues of stems or roots? D. polita was found only on R. acicularis, although R.

woodsii often grows alongside. Is D. polita restricted to R. acicularis or does only this host

have tissues susceptible to galling when D. polita oviposits? Transplanting larvae of Diplo-

lepis species into meristematic tissues of other hosts may yield interesting results.

Many theories on how the habits of several closely related hymenopterans may have e-

volved can be further studied by examining cynipid galls. For example, have the inquilines

that induce cell hypertrophy and hyperplasy lost the ability to cause galls, or are they in the

process of developing the ability to initiate their own galls? Transplant experiments may

provide clues. Two chalcidoid species, Eurytoma longavena and Glyphomerus stigma were

found to be both entomophagous and phytophagous in their larval stages. It would be inter-

esting to know if these species could complete their development on only one food. Has the

phytophagous habit of these species which are normally considered entomophagous, devel-

oped because of the reduced biomass in monothalamous galls, or do they exhibit the same

habit in other Diplolepis galls?

A useful contribution to our knowledge of community structure can hopefully be made

by continued studies of population assemblages in insect galls. Insect gall communities are

simple communities and their attributes can be determined with relative ease. Granted, my

definitions and use of community attributes differ somewhat from the classical usage devel-

oped by plant ecologists such as Whittaker (1970). But nearly all aspects of community e-

cology such as diversity, organization, succession, climax, productivity and biomass, and nu-

trient cycling can be articulated by examining gall communities. Further research will sup-

ply data on these various aspects of gall community ecology and it will be of great interest

to compare results and conclusions with those of other community ecologists. Zoogeo-

graphical studies of Diplolepis communities would be valuable. From distribution maps

(Lewis, 1959) it appears that R. acicularis spread into North America through Beringia.

They probably brought Diplolepis species with them and comparisons of communities from

Alaska to California should provide information useful to those postulating theories of how

natural communities evolved (Whittaker and Woodwell, 1972). Studying series of all known

Diplolepis species, their galls and communities, should reveal a great deal of information a-

bout the evolution of the rose gall complex.

In the past, most workers have taken the descriptive approach to studies of plant galls. It

has been my aim to consider galls in terms of ecology rather than simply gall morphology or

taxonomy of inhabitants. Before botanists can effectively study developmental processes as-

sociated with insect galls, information on the ecology of associated inhabitants is required.

Botanical information on galls will be found useful in studies of gall formers and associated

species. By combining knowledge from various disciplines of biology, we will be in a better

position to understand cynipid galls and the intricacies of these fascinating insect-plant rela-

tionships.

ACKNOWLEDGEMENTS

I wish to thank W. G. Evans of the University of Alberta for his guidance and encourage-

ment throughout this study. G. E. Ball, B. Hocking and D. D. Cass of the University of Al-

berta and A. M. Harper of the Canada Agriculture Research Station, Lethbridge, offered

their advice on many occasions. N. S. Church of the Canada Agriculture Research Station,

Saskatoon, and D. M. Lehmkuhl of the University of Saskatchewan read and criticized this

Diplolepis rose gall community

95

manuscript and provided numerous useful suggestions.

I also thank J. S. Scott of the University of Alberta and J. Waddington of the University

of Saskatchewan for their help with the graphs and photographs.

I also thank my companions at the George Lake Field Station, A. C. Carter, H. Goulet,

A. W. Thomas, and especially summer assistant, N. R. Chymko. I am also indebted to Mr. E.

Donald and family, neighbours of the field station, for the numerous courtesies and services

they provided.

I am also grateful to the following for identifying specimens: O. Peck and M. Ivanochko

(Cynipidae and Torymidae), E. E. Grissell (Torymidae), R. E. Bugbee (Eurytomidae), C. M.

Yoshimoto (Eulophidae and Ormyridae), R. D. Eady (Pteromalidae), and W. H. Lewis (Ro-

saceae). R. J. Lyon provided a great deal of information on Diplolepis (Cynipidae) and spent

much time examining specimens.

Finally I wish to thank the Boreal Institute of the University of Alberta for providing a

grant-in-aid for the summer of 1969 and the Institute for Northern Studies of the University

of Saskatchewan for providing publication funds.

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Triggerson, C. J. 1914. A study of Dryophanta erinucei (Mayr) and its gall. Ann. ent. Soc.

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AN ANNOTATED LIST OF THE HYDROADEPHAGA (COLEOPTERA: INSECTA) OF

MANITOBA AND MINNESOTA

BY J. B. WALLIS1 2

This paper is based on a manuscript written by the late J. B. Wallis during the 1920’s and

1930’s. One hundred and sixty-seven species of beetles belonging to the families Haliplidae,

Dytiscidae, and Gyrinidae are recorded from Manitoba and Minnesota. One hundred and

forty-one species are recorded from Manitoba and one hundred and six from Minnesota with

eighty species common to both areas. Notes on collection records and descriptions of the

habitat in which certain species are found are given.

Introduction by D. J. Larson

2

Quaestiones entomologicae

9: 99-114 1973

This list of species of Hydroadephaga of Manitoba and Minnesota is a brief condensation

of a large manuscript written by the late J. B. Wallis. The original manuscript contains de-

scriptions of all of the following taxa, keys to aid in their identification, and collecting and

distribution notes. To a large extent, Wallis’ keys and descriptions are summaries of previ-

ously published works such as H. C. Fall’s revisions of Coelambus (= Hygrotus ) (1919 \Aga-

bus (1922a), Gyrinus (1922b), and Hydroporus (1923). Almost all of Wallis’ original re-

search on water beetles has been published elsewhere, for example in his papers on Haliplus

(1933a), Hydaticus (1939a), Graphoderus (1939b), Ilybius (1939c) and in other papers

which contain descriptions of new taxa (1924, 1926a, 1926b, 1933a, 1933b, 1933c). Be-

cause of this, I consider the most important material in the manuscript to be the extensive

list of reliably identified species along with the collecting and distribution records. Fall’s pa-

pers contain many references to specimens which were sent to him by Wallis. Also Wallis’

collection contairs specimens identified by Fall. The result of this appears to be very close

agreement between Fall and Wallis in their species concepts.

The history of this manuscript has not been fully traced. R. D. Bird (1958; pers: com.

1971) stated that Wallis developed an interest in insects through contact with the Criddle

family in Manitoba, with Norman Criddle especially encouraging him to specialize in aquatic

Coleoptera. H. B. Leech visited Wallis in 1946 or early 1947. At this time Wallis spoke about

his manuscript on the water beetles of Minnesota and Manitoba. Leech (in litt., 1971)

states: “he wrote it in the late twenties at the request of the Minnesota people who supplied

a large collection to be identified and who were to publish it. Then came the depression and

with no likelihood of getting it into print he lost heart.” Wallis’ interest in this work con-

tinued at least to 1933, for citations of the literature to that date have been included. How-

ever, papers published by W. J. Brown (1937) and Leech (1938, 1939), although directly

relevant to the Manitoba fauna, were not cited by Wallis. The original copy of Wallis’ manu-

script has been deposited in the Department of Entomology, University of Alberta. Copies

of it are located in the California Academy of Sciences (H. B. Leech); Saint Cloud State Col-

lege, Minnesota (R. Gunderson) and the University of Calgary (D. Larson). Most of Wallis’

water beetle collection has been deposited in the Canadian National Collection, Ottawa;

however some specimens are housed in the insect collection of the Riveredge Foundation,

Calgary, Alberta and in the Strickland Museum, University of Alberta.

1 Deceased

2 Department of Biology

University of Calgary, Calgary, Alberta

100

Larson

In compiling the following list, I have followed Wallis’ manuscript as closely as possible.

Wallis’ names are used consistently, although changes in status are indicated in parentheses.

Also, all editorial comments that I have inserted are placed between square brackets. For

each included species, the following information is given: name, Leng catalogue number, list

of localities arranged alphabetically, and collecting notes. A few taxonomic notes are also in-

cluded. For the sake of completeness, addenda have been included of names of water beetle

species recorded from Manitoba in the literature since 1937 and therefore not given in

Wallis’ list.

FAMILY HALIPLIDAE

Genus Haliplus Latreille

Haliplus (s. str.) strigatus Roberts. (2321)

Localities: MANITOBA - common west of the Laurentian Highland, and as far north as

Le Pas.

Haliplus (s. str.) longulus LeConte. (2322)

Localities: MANITOBA - throughout the southern part, moderately abundant. MINNE-

SOTA - St. Anthony’s Park, rare.

Haliplus (s. str.) immaculicollis Harris.

Localities: MANITOBA - abundant throughout. MINNESOTA - abundant throughout.

Haliplus (s. str.) blanchardi Roberts (2319)

Localities: MINNESOTA - Itasca State Park, Ramsey Co., St. Anthony, St. Paul.

Haliplus (Paraliaphlus) borealis LeConte. (2317)

Localities: MANITOBA - Selkirk, Winnipeg. MINNESOTA - Pine City, St. Peter, not a-

bundant.

Haliplus (Paraliaphlus) triopsis Say. (2301)

Localities: MINNESOTA - Le Sueur Co., Ramsey Co., Red Wing.

Haliplus (Paraliaphlus) pantherinus Aube.

Localities: MINNESOTA - Ramsey Co., St. Paul, St. Peter.

Haliplus (Liaphlus) connexus Matheson. (2300)

Localities: MINNESOTA - St. Paul, one specimen only.

Haliplus (Liaphlus) apo st olicus Wallis.

Localities: MINNESOTA - Bussey’s Pond (University of Minnesota Campus), Green Lake,

Itasca State Park, St. Paul.

Haliplus (Liaphlus) subguttatus Roberts. (2306)

Localities: MANITOBA - generally distributed in southern half. MINNESOTA - Cramer,

Hubbard Co., Ramsey Co., St. Paul; abundant.

Haliplus (Liaphlus) canadensis Wallis

Localities: MANITOBA - Victoria Beach, Winnipeg, Winnipeg Beach.

Haliplus (Liaphlus) cribrarius LeConte. (2305)

Localities: MANITOBA - generally distributed as far north as Mile 256, Hudson’s Bay

Railway. MINNESOTA - Cook Co., Grand Rapids, Hibbing, Itasca State Park.

Genus Peltodytes Regimbart

Pel tody tes edentulus LeConte. (2337)

Localities: MANITOBA - generally distributed in southern half; abundant. MINNESOTA-

generally distributed; abundant.

Hydroadephaga of Manitoba and Minnesota

101

Peltodytes tortulosus Roberts. (2324)

Localities: MANITOBA - Winnipeg Beach (type locality); generally distributed in south-

ern third, though not abundant. MINNESOTA - Ely, Minneapolis, Pelican Rapids, Ramsey

Co., St. Paul (Track Pond).

It is interesting to note that this species seems to be extending its range for one can

scarcely suppose that it could have been overlooked in such a well hunted place as Toronto,

Canada, from which locality I recently received a number of specimens, and in 1931 it was

taken at Quebec City.

FAMILY DYTISCIDAE

SUBFAMILY LACCOPHILINAE

Genus Laccophilus Leach

Laccophilus maculosus (Germar ). (2351)

Localities: MANITOBA - moderately common in the southern part of the province; not

as yet taken north of about 100 miles from the International Boundary. MINNESOTA-

generally distributed and common.

Collecting notes: Overwintering is apparently in rivers as specimens were taken by Mr. E.

Criddle in the Assiniboine River at Aweme, Manitoba, on January 9, 1928.

Laccophilus inconspicuus Fall (2354) (= biguttatus Kirby)

Localities: MANITOBA - common throughout as far north as Le Pas; probably extends

much farther north. MINNESOTA - two records only - a single specimen from Benson (23.

viii.22) and one from Ramsey Co. (1 l.iv.22) - both taken by W. E. Hoffman; doubtless

occurs all through the northwestern part of the state.

SUBFAMILY HYDROPORINAE

Genus Hydrovatus Motschoulsky

Hydrovatus pustulatus Melsheimer.

Localities: MINNESOTA - Hennepin, Le Sueur Co., Mora, St. Paul.

Genus Desmopachria Babington

Desmopachria convexa Aube <23 74)

Localities: MANITOBA - Selkirk, Victoria Beach, Winnipeg. MINNESOTA - Hennepin

Co., Benson, Ramsey Co., St. Paul.

Collecting notes: Occurs in very shallow water among debris and roots.

Genus Bidessus Sharp

Bidessus flavicollis LeConte. (2385). [placed in genus Liodessus Guignot by Young, 1969.]

Localities: MINNESOTA - Lake Emily, Lake Jefferson, St. Peter; rare.

Bidessus af finis Say. (2390). [Placed in genus Liodessus Guignot by Young 1969.]

Localities: MANITOBA - common everywhere, as far north as Mile 214, Hudson’s Bay

Railway. MINNESOTA - Hennepin Co., Le Sueur Co., Minneapolis, St. Paul.

Collecting, notes: Found in shallow waters with muddy bottoms.

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Larson

Bidessus granarius Aube. (2398). [Placed in genus Urarus Guignot by Young, 1969.]

Localities: MINNESOTA - Bussey’sPond, “Minn.”, St. Paul.

Genus Hygrotus Stephens

Hygrotus acaroides LeConte. (2407)

Localities: MANITOBA - Winnipeg (type locality of race Winnipeg Wallis), Rosebank,

Thornhill. MINNESOTA - Benson, St. Paul.

Hygrotus fare tus LeConte. (2405)

Localities: MANITOBA - Winnipeg.

Collecting notes: The single Winnipeg specimen was found in a little pool in the bed of

a partially dried up rivulet in the woods (24.v. 1 922).

Hygrotus punctatus Say ( =sayi Balfour-Browne, 1 944)

Localities: MANITOBA - abundant everywhere. MINNESOTA - abundant everywhere

Hygrotus turbidus LeConte. (2408)

Localities: MANITOBA - not uncommon southward. MINNESOTA - Booker Co., St.

Paul. St. Peter; apparently rare.

Hygrotus dispar LeConte. (2409)

Localities: MANITOBA - Le Pas, Winnipeg; uncommon. MINNESOTA - Hennepin Co.,

Owatonna, Rochester; uncommon.

Hygrotus compar Fall. (191 74)

Localities: MANITOBA - Aweme, Winnipeg; rare.

Hygrotus suturalis LeConte. (2413)

Localities: MANITOBA - Generally distributed, not common in the south, rather plen-

tiful northward. MINNESOTA - Warroad.

Collecting notes: Found usually in clear water.

Hygrotus sellatus LeConte. (2414)

Localities: MANITOBA - in southern third; not common. MINNESOTA - Nicollet Co.

Collecting notes: In weedy ponds.

Hygrotus canadensis Fall. (19178)

Localities: MANITOBA - Winnipeg (type locality); common in southern half of province.

MINNESOTA - Hennepin Co., Hibbing, Ramsey Co., St. Anthony’s Park; probably all

through the northern half of the state.

Collecting notes: Prefers clear water.

Hygrotus patruelis LeConte. (2412)

Localities: MANITOBA - common in southern third, not yet taken in the northern two

thirds. MINNESOTA - Grand Rapids, Hennepin, Ottertail Co., St. Anthony’s Park.

Collecting notes: Prefers clear water.

Hygrotus nubilus LeConte. (2420)

Localities: MINNESOTA - St. Paul (one specimen).

Hygrotus punctilineatus Fall. (19183)

Localities: MANITOBA - occasional throughout southern part.

Collecting notes: This species is occasionally found in normal( fresh water) situations but

the only station where I have taken it commonly is in a small pood only a hundred yards

or so from the saline Cobb’s Lake near Baldur, Manitoba. I have not had the water of this

pond analysed but judging from the vegetation, its chemical contents, while clearly saline

or alkaline, differ greatly from the near-by Cobb’s Lake.

Hydroadephaga of Manitoba and Minnesota

103

Hygrotus tumidiventris Fall. (19182)

Localities: MANITOBA - distributed in waters of a certain type of alkalinity, occasionally

taken elsewhere.

Collecting notes: Specimens of tumidiventris were common in the locality described a-

bove under punctilineatus but this species is apt to be found in situations where the water is

more usual.

Hygrotus masculinus Crotch. (2419)

Localities: MANITOBA - abundant in Shoal Lake, 35 miles or so northwest of Winnipeg;

only accidental elsewhere.

Collecting notes: masculinus adults prefer water containing a high percentage of magnesi-

um salts. The only place in Manitoba where I have taken masculinus is in Shoal Lake, which

appears to be quite rapidly drying up. Much of its bed is now dry, and its water is strongly

reminiscent of epsom salts. While covering a number of square miles, it is everywhere very

shallow and contains no vegetation except algae, even the edges almost everywhere being

without rushes or grass. Naturally, being so shallow and extensive, it is subject to rapid fluc-

tuations of level according to the direction and strength of the wind, often fifteen or twenty

mintes making a difference between a dry area and six or eight inches of water. The bot-

tom is slimy mud plentifully sprinkled with stones. Specimens of masculinus appear to pre-

fer water five or six inches deep and to hide under and around stones.

Hygrotus salinarius Wallis. (19188)

Localities: MANITOBA - Baldur (Cobb’s Lake, type locality), Salt Lake near Strathclair,

southern end of Lake Winnipegosis.

Collecting notes: salinarius adults prefer water strongly impregnated with common salt.

Hygrotus unguicularis Crotch. (2421)

Localities: MANITOBA - probably throughout the province; rare southward but moder-

ately common at points on the Hudson’s Bay Railway, increasingly so northward to Mile

474.

Hygrotus dentiger Fall. (20764)

Localities: MANITOBA - Thornhill (one specimen).

Collecting notes: This species was taken by Mr. F. S. Carr in saline lakes in Alberta, and

I took it quite commonly in a pond of moderate salinity or alkalinity near Roche Percee,

Saskatchewan. In this pond were specimens of many species found also in fresh water so

that its saline content could not have been very great.

Hygrotus impressopunctatus Schaller. (2424)

Localities: MANITOBA - throughout the province except perhaps in the extreme north;

abundant. MINNESOTA - throughout the state,abundant.

Genus Hydroporus Clairville

Hydroporus (Heterosternus) undulatus Say. (2447)

Localities: MANITOBA - Aweme, Husavick, Mile 214 Hudson’s Bay Railway, Rosebank,

Winnipeg. MINNESOTA - apparently fairly generally distributed. [Note. - Wallis treated con-

similis LeConte as a valid species. However, he states that “extremes of these two species

(undulatus and consimilis) are easy to separate but I must confess after examining hundreds

of specimens from one locality ...that most of these could just as well be called one as the

other.” Here, the names consimilis and undulatus are treated as synonyms]

Hydroporus (Heterosternus) clypealis Sharp. (2452)

Localities: MINNESOTA - one female, Red Wing (30.ix. 1923, W. E. HoffmannL

Hydroporus (Heterosternus) vittatus LeConte. (2465)

Localities: MANITOBA - Fork Liver. Winnipeg (several stations). MINNESOTA - Grand

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Larson

Marais (one female doubtfully placed here).

Collecting notes: Only occasionally and locally being not rare. On one occasion I took

several dozen adults in little pools in the bed of a partially dried up creek in East Kildon-

an just north of Winnipeg.

Hydroporus (Heterosternus) sericeus LeConte (2466) (= superioris Balfour-Browne).

Localities: MANITOBA - generally distributed, except possibly in the extreme north.

MINNESOTA - generally distributed.

Collecting notes: Locally very abundant in clear but weedy water.

Hydroporus (Heterosternus) solitarius Sharp. (2467)

Localities: MANITOBA - Mile 214 Hudson’s Bay Railway.

Collecting notes: Specimens of this species were quite common in July 1917 in deep wa-

ter in the Piquetenay River, swimming close to the almost perpendicular surface of the rock

to which they frequently clung.

Hydroporus (Heterosternus) paugus Fall. (19220)

Localities: MANITOBA - 16 miles e. Aweme; Township 6, Range 9 East.

Collecting notes: Adults inhabit larch or spruce swamps, where the water is cold.

Hydroporus (Heterosternus) stagnalis Gemminger and Harold. (2521)

Localities: MANITOBA - Stonewall

Collecting notes: Taken only in an old quarry test hole in a limestone formation.

Hydroporus (Heterosternus) planiusculus Fall. (19224)

Localities: MANITOBA - 16 miles e. Aweme. MINNESOTA - Chester, Olmsted Co.,

St. Peter.

Collecting notes: Adults are moderately abundant in the water of a cold rivulet fed by

g spring issuing from the sand hills near the banks of the Assiniboine River about 16 miles

east of Aweme. There is a peculiar formation here known locally as the “Devil’s Punch-

bowl”, and at the bottom of this is the spring and rivulet mentioned above. Where the riv-

ulet expands and becomes more or less choked with specimens of Chara, adults of planiuscu-

lus may usually be found in some numbers.

Hydroporus (s. str.) dichrous Melsheimer. (2510)

Localities: MINNESOTA - Le Sueur Co., Rochester, St. Anthony Park, St. Paul, St. Peter.

Hydroporus (s. str. ) melsheimeri Fall. (251 1)

[Note. - The description of this species and the list of localities is missing from Wallis’

manuscript. The species almost certainly occurs in Manitoba.]

Hydroporus (s. str.) dentellus Fall. (2506)

[ Note. - This species is treated in Wallis’ key to the Minnesota-Manitoba species of Hy-

droporus, and is not in the text of the manuscript, probably because inserted page which

also contained the discussion of H. melsheimeri was lost. I have seen Manitoba specimens of

dentellus collected by Wallis (Aweme, Strickland Museum, University of Alberta.]

Hydroporus (s. str.) notabilis LeConte. (2518)

Localities: MANITOBA - fairly generally distributed throughout, at least as far north as

Mile 332 Hudson’s Bay Railway. MINNESOTA - Mendota

Hydroporus (s. str.) arcticus Thomson. (25 1 9)

Localities: MANITOBA - Churchill ( 2 specimens, 5 & 1 l.ix. 30, F. Neave). Fall (1923)

pointed out that this species may well prove to be but a race of notabilis LeConte.

Hydroporus (s. str.) niger Say. (2514)

Localities: MINNESOTA - Le Sueur Co., St. Paul, St. Peter.

Hydroporus (s. str.) columbianus Fall. (19215)

Localities: MANITOBA - not common, but widely distributed in the south.

Hydroadephaga of Manitoba and Minnesota

105

Hydroporus (s. str.) rectus Fall. (19209)

Localities: MANITOBA - Aweme, Mile 256 Hudson’s Bay Railway, Township 7 Range 1 IE.

Hydroporus (s. str.) despectus Sharp. (2495)

Localities: MANITOBA - Aweme, Winnipeg. MINNESOTA - one female from Itasca Park

appears to belong here.

Hydroporus (s. str.) tenebrosus LeConte. (2493)

Localities: MANITOBA - generally distributed well to the north, abundant. MINNESOTA

- Le Sueur Co. ( 1 specimen).

Hydroporus (s. str.) pervicinus Fall. (19207)

Localities: MANITOBA - Aweme, Onah, Township 7 Range 1 IE, Transcona. MINNESOTA

- Bengali, Hibbing, St. Paul.

Hydroporus (s. str.) tartaricus LeConte. (2491)

Localities: MANITOBA - Aweme, Hudson’s Bay Territory, Winnipeg.

Hydroporus (s. str.) signatus Mannerheim. (2508)

Localities: MANITOBA - Mile 332 Hudson’s Bay Railway. MINNESOTA - Hibbing, Ram-

sevCo., St. Anthony Park, St. Paul.

Hydroporus (s. str.) obscurus Sturm. (2492)

Localities: MANITOBA - Mile 214 and Mile 332 Hudson’s Bay Railway, Township 7 Rangs

HE.

Hydroporus (s.str.) badiellus Fall. (19206)

Localities: MANITOBA - Mile 214 and Mile 332 Hudson’s Bay Railway, Township 7

Range 1 IE.

[Note. - Wallis states that obscurus and badiellus at least as far as the species are understood

here, are very similar and aouear to always occur together. Perhaps, at least in Manitoba,

these should be treated as only one species.]

Hydroporus (s. str.) appalachius Sherman. (2498)

Localities: MANITOBA - Aweme, Mile 332 Hudson’s Bay Railway, Thornhill, Winnipeg.

MINNESOTA - Hennepin Co.

Collecting notes: This species is sometimes not uncommon in shallow pools in woodland

streams.

[Note. - Wallis lists occidentalis Sharp as occurring in Manitoba and Minnesota. This species

appears to be western in distribution and probably does not occur in this area. The records

for occidentalis probably refer to dark specimens of appalachius. ]

Hydroporus (s. str.) melanocephalus Gyllenhal. (19205)

Localities: MANITOBA - Churchill, Mile 256 and 332 Hudson’s Bay Railway, Township

7 Range 1 IE.

Collecting notes: A species occurring in the colder waters, commoner northward, quite

rare southward where it has been taken only in the cold sphagnum moss bogs east of Winni-

peg.

Hydroporus (s. str.) fuscipennis Kies. (2509)

Localities: MANITOBA - universally distributed so far as known; our commonest species.

MINNESOTA - St. Anthony’s Park.

Hydroporus (s. str.) striola Gyllenhal.

Localities: MANITOBA - generally distributed and abundant. MINNESOTA- Hennepin

Co., Hubbard Co., Mora Co.

Hydroporus (s. str.) glabriusculus Aube. (2500)

Localities: MANITOBA - Aweme, Mile 256 and 332 Hudson’s Bay Railway, Winnipeg.

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Larson

Hydroporus (s. str.) rufinasus Mannerheim. (2504)

Localities: MANITOBA - Mile 332 Hudson’s Bay Railway, Township 7 Range 11E.,

Victoria Beach, Winnipeg.

Hydroporus (s. str.) tristis Paykull. (2501)

Localities: MANITOBA - quite generally distributed.

Hydroporus (Deronectes) striatellus LeConte. (2431)

Localities: MANITOBA - second Cranberry Lake (near Cranberry Portage, Hudson’s Bay

Railway) - two specimens (27.viii. 1930), F. Neave.

Collecting notes: Second Cranberry Lake is of a different formation from most of our

northern lakes, being in limestone, whereas most of the others are in granite.

Hydroporus (Deronectes) griseostriatus DeGeer. (2430)

[Note. - In the manuscript, this species is included in the key to Minnesota-Manitoba species

of Hydroporus, however the text page dealing with it is missing. The species is no doubt

widely distributed in the area.]

Hydroporus (Deronectes) rotundatus LeConte [- elegans Panzer] .

Localities: MANITOBA - moderately common throughout. MINNESOTA - Brandon.

Collecting notes: in clear weedy streams.

Hydroporus ( Oreodytes) duodecimlineatus LeConte (probably a synonym of laevis Kirby.)

(2482)

Localities: Mile 474 Hudson’s Bay Railway.

Hydroporus ( Oreodytes) scitulus LeConte.

Localities: MANITOBA - Mile 332 Hudson’s Bay Railway.

Genus Laccomis desOozis

Laccornis conoideus LeConte. (2532)

Localities: MANITOBA - generally distributed in the southern part of the province. MIN-

NESOTA - Owatonna (one specimen).

Collecting notes: It is not uncommon in spring in ditches and in temporary ponds. Later

it is found in places where the water keeps fairly cold.

SUBFAMILY COLYMBETINAE

Genus Agabus Leach

Agabus seriatus Say. (2539)

Localities: MANITOBA - Aweme, Mile 474 Hudson’s Bay Railway, Thornhill. MINNESO-

TA - Hennepin Co., Le Sueur Co., St. Anthony’s Park, St. Paul, St. Peter.

Collecting notes: Specimens are found chiefly in waters flowing from cold springs,

though the species does not seem to be a denizen of the cold larch swamps.

Agabus triton Fall. (19232)

Localities: MANITOBA - Winnipeg.

Collecting notes: This species is not uncommon in ditches and ponds near Winnipeg in

early spring.

Agabus punctulatus Aube. (2551)

Localities: MANITOBA - everywhere, even being recorded from Nelson River. MINNE -

SOTA - “Minnesota”, Ottertail Co., St. Anthony’s Park.

[Note.- The specimens recorded from Nelson River Manitoba could represent colymbus

Leech 1938; I have not examined them.]

Hydroadephaga of Manitoba and Minnesota

107

Agabus semipunctatus Kirby. (2553)

Localities: MANITOBA - rather generally distributed as far north as Mile 24 [ typing

error; should perhaps read mile 214, a frequently mentioned locality]. Hudson’s Bay Rail-

way. MINNESOTA - “Minnesota”, Ramsey Co.

Agabus sharpi Fall (19234) (= falli Guignot)

Localities: MANITOBA - Winnipeg and vicinity.

Agabus disintegratus Crotch. (2557)

Localities: MINNESOTA - Rochester.

Agabus ambiguus Say.

Localities: MANITOBA- widely distributed in southern half and quite abundant. MIN-

NESOTA - Le Sueur Co.

Collecting notes: This species remains active throughout the winter, specimens having

been taken through a hole cut in the ice of the Assiniboine River near Aweme on January

9th, 1928, by Mr. E. Criddle.

Agabus congener Paykull. (2560)

Localities: MANITOBA - Churchill, Winnipeg.

Agabus discolor Harris. (2564)

Localities: MANITOBA - Aweme, Mile 332 Hudson’s Bay Railway, Onah, Winnipeg; rath-

er abundant. MINNESOTA - Duluth.

Agabus inscrip tus Crotch. (2559)

Localities: MANITOBA - Bird’s Hill, Mile 332 Hudson’s Bay Railway, Riding Mountain.

MINNESOTA - Lake Superior, White Fish Point.

Agabus canadensis Fall. (19237)

Localities: MANITOBA - abundant in southern half.

Collecting notes: In savannah and prairie associations.

Agabus subfuscatus Sharp.

Localities: MANITOBA - Aweme, Winnipeg. MINNESOTA - Ottertail Co. One female

should probably be referred here.

Agabus phaeopterus Kirby. (2566)

Localities: MANITOBA - quite generally distributed at least as far north as Mile 332, Hud-

son’s Bay Railway. MINNESOTA - Duluth.

Agabus bicolor Kirby (2567)

Localities: MANITOBA - Aweme, Mile 214 Hudson’s Bay Railway Township 1 Range

14E., Township 7 Range 1 IE.

Agabus confinis Gyllenhal. (2563)

Localities: MANITOBA - Bird’s Hill, Hudson’s Bay, Mile 214 Hudson’s Bay Railway, Rid-

ing Mts., Thornhill, Township 7 Range 1 IE., Township 14 Range 10E. MINNESOTA- Duluth.

Collecting notes: quite rare but widely distributed in cold water, usually in larch swamps.

Agabus infuscatus Aube. (2571)

Localities: MANITOBA - Churchill, Mile 332 Hudson’s Bay Railway.

Collecting notes: Just west of the railway bridge at Kettle Rapids in mid July an outcrop-

ping of rock was exposed in the bed of the Nelson River owing to the lowering of the water.

On the surface of this rock were several small puddles two or three feet wide and a foot or

so deep. Considerable amounts of slimy algae were floating on the surface of the water in

these pot holes, and formed several inches of sediment at the bottom. From three or four

of these unlikely looking puddles several dozen specimens of infuscatus were taken. Speci-

mens were not found elsewhere.

108

Larson

Agabus arcticus Paykull. (2576)

Localities: MANITOBA - Mile 214 Hudson’s Bay Railway and northward to Churchill.

Collecting notes: Specimens of this species were common in shallow grassy water in an

expansion of the Piquitenay River at Mile 214 Hudson’s Bay Railway.

Agabus ontarionis Fall. (19238)

Localities: MANITOBA - Aweme, Charleswood, Makinak.

Agabus ajax Fall. (19239)

Localities: MANITOBA - 16 miles east Aweme, Fort Churchill.

Agabus anthracinus Mannerheim. (2575)

Localities: MANITOBA - generally distributed probably into the far north, quite com-

mon.

Agabus nigroaeneus Erichson (2579) ( -erichsonii Gemminger and Harold).

Localities: MANITOBA - generally distributed; quite abundant. MINNESOTA - state lo-

cality only.

Agabus pseudo confertus Wallis. (20782)

Localities: MANITOBA - Bird’s Hill, Mile 17 Hudson’s Bay Railway, Township 7

Range 11E., Winnipeg.

Collecting notes: This species is an inhabitant of the true sphagnum bogs, being found in

the small holes in the swamps where a little clear water shows. It is an early spring species

and I have taken specimens by breaking an inch or so of ice from the surface and then

dredging among the moss which is itself largely imbedded in ice.

Agabus kenaiensis Fall. (20778)

Localities: MANITOBA - Bird’s Hill, Onah, Township 7 Range 1 IE.

Collecting notes: Like pseudoconfertus, this species is also an inhabitant of sphagnum

bogs.

Agabus minnesotensis Wallis.

Localities: MINNESOTA - Hennepin Co., (single type).

[Note. - Perhaps the type specimen was erroneously labeled as the species has since been

fnnnrl in the west onlv (Anderson. 1962L1

Agabus verus Brown [= clavicornis Sharp (J. Balfour-Browne, 1947)1(21729)

Localities: MANITOBA - Churchill.

Agabus clavatus LeConte (2577) (= antennatus Leech)

Localities: MANITOBA - Mile 214 Hudson’s Bay Railway, Stonewall, Thornhill, Win-

nipeg. MINNESOTA - Le Sueur Co., Ramsey Co., St. Paul, St. Peter.

Collecting notes: Adults are somewhat local and not usually at all common but I took

many specimens in company with arcticus specimens in shallow water among grass in a

widening of the river at mile 214 Hudson’s Bay Railway.

Apator ( = Agabus) bifarius Kirby. (2587)

Localities: MANITOBA - common throughout the province. MINNESOTA - Owatanna,

Ramsey Co., St. Anthony’s Park, St. Paul.

Genus Ilybius Erichson

Ilybius pleuriticus LeConte. (2590)

Localities: MANITOBA- generally distributed in southern half but rather uncommon;

more abundant at Mile 214 Hudson’s Bay Railway. MINNESOTA - Beaver Dam near Ely.

Ilybius angustior Gyllenhal. (2595)

Localities: MANITOBA - generally distributed and moderately abundant as far north as

the limit of trees. MINNESOTA - Olivia.

Hydroadephaga of Manitoba and Minnesota

109

Ilybius subaeneus Erichson. (2589)

Localities: MANITOBA - throughout the province at least as far north as within 90

miles of Hudson’s Bay. More abundant northward. MINNESOTA - Duluth, Grand Marais,

“Minnesota”.

Ilybius biguttulus Germar (2598)

Localities: MINNESOTA - Hibbing, Le Sueur, Two Harbors.

Ilybius fraterculus LeConte.

Localities: MANITOBA - abundant in southern half. MINNESOTA - throughout the

northern half of the state at least.

Ilybius discedens Sharp. (2597)

Localities: MANITOBA - on the Canadian Shield to Hudson’s Bay.

Collecting notes - In cold sphagnum bogs.

Genus Coptotomus Say

Coptotomus interrogatus Fabricius. (2610)

Localities: MANITOBA - everywhere as far north at least as Le Pas; very abundant. MIN-

NESOTA - apparently widely distributed and abundant.

Genus Scutopterus Crotch (= 'NeoscutopterusF. Balfour-Browne)

Neoscutopterus angustus LeConte. (2612)

Localities: MANITOBA - Thornhill, Township 7 Range 1 IE., Winnipeg

Collecting notes: Usually found in small mossy pools in larch swamps.

Neoscutopterus horni Crotch. (2613)

Localities: Aweme, Riding Mts., Township 7 Range 1 IE.

Collecting notes: Found in the same type of habitat as the preceding species.

Genus Rhantus Boisduval and Lacordaire

Rhantus sinuatus LeConte. (2620)

Localities: MINNESOTA - Le Sueur Co., St. Paul.

Rhantus plebeius Sharp (= binotatus Harris). (2616)

Localities: MANITOBA - generally distributed in southern half. MINNESOTA -Hibbing,

Grand Marais.

Rhantus notatus Fabricius. (2622)

Localities: MANITOBA - abundant throughout the southern portion; may be found

well towards Hudson’s Bay. MINNESOTA - Le Sueur Co., St. Anthony Park; probably

generally distributed.

Rhantus suturellus Harris [=wallisi Hatch cf. Hatch 1953]

Localities: MANITOBA - universally distributed at least as far north as Mile 214 Hud-

son’s Bay Railway. MINNESOTA - Beaver Dam, Hennepin Co., Lake Co., Olivia, Ramsey

Co., St. Peter.

Rhantus zimmermanni Wallis (= suturellus Harris cf. Hatch 1953)

[Locality list missing from manuscript.]

Rhantus tostus LeConte. (2624)

Localities: MANITOBA- abundant everywhere, at least south of Mile 214 Hudson’s Bay

Railway. MINNESOTA - probably generally distributed and abundant.

Genus Colymbetes Clairville

Colymbetes longulus LeConte. (2627)

Localities: MANITOBA - found sparingly on the western edge of the coniferous forests

and very rarely further west.

Collecting notes: Most specimens have been collected from ponds in larch swamps. A few

have been found elsewhere.

110

Larson

Colymbetes dahuricus Aube

[Note. - A single female collected at Mile 214 Hudson’s Bay Railway was assigned to this

species; it probably belongs to the species longulus LeConte.]

Colymbetes sculptilis Harris complex. (2632)

[Note. - Wallis separates the Manitoba specimens into three species: dolobratus Paykull

(Mile 474, Hudson’s Bay Railway); rugipennis Sharp (widely distributed), and sculptilis

Harris (widely distributed). According to Young and Severing 956), the names rugipennis

and sculptilis are synonyms. Also, Wallis’ specimens of dolobratus probably represent a

northern form of sculptilis. ]

Localities: MANITOBA - widely distributed. MINNESOTA - widely distributed.

Genus Dytiscus Linnaeus

Dytiscus fasciventris Say. (2636)

Localities: MANITOBA - throughout the southeast portion of the province. MINNESO-

TA - throughout the state.

Dytiscus hybridus Aube. (2637)

Localities: MANITOBA - Southern portion. MINNESOTA - throughout the state.

Dytiscus verticalis Say. (2638)

Localities: MINNESOTA - common southward and eastward.

Dytiscus sublimbatus LeConte. (2640) (= cordieri Aube )

Localities: MANITOBA - generally distributed in southern half, west of Canadian Shield.

MINNESOTA - Hubbard Co., Norman Co., Ottertail Co., Pine City; St. Peter.

Dytiscus anxius Mannerheim.

Localities: MANITOBA - fairly common in southern portion.

Dytiscus parvulus Mannerheim. (2642)

Localities: MANITOBA - southward only, rather rare. MINNESOTA - Hubbard Co., Ram-

sey Co., St. Peter.

Dytiscus dauricus Gebler. (2645)

Localities: MANITOBA - Husavick, and southeastward; Winnipeg.

Dytiscus harrisi Kirby. (2646)

Localities: MANITOBA - in the southeast portion of the province.

MINNESOTA - Isanti Co., Le Sueur Co., St. Anthony Park.

SUBFAMILY HYDATICINAE

Genus Hydaticus Leach

Hydaticus modest us Sharp.

Localities: MANITOBA - generally distributed. I have one specimen labelled H. B., hut

the species is not common north of Le Pas. MINNESOTA - generally distributed.

Hydaticus piceus LeConte. (2649)

Localities: MANITOBA- Victoria Beach and southeastward, Winnipeg. MINNESOTA-

Albert Lea, Le Sueur Co., Ramsey Co., St. Paul.

Genus Acilius Leach

Acilius semisulcatus Aube. (2651)

Localities: MANITOBA - abundant in southern half. MINNESOTA - abundant through-

out.

Acilius fra t emus Harris. (2652)

Localities: MINNESOTA - Anoka Co., Becker Co., Grand Marais, Hibbing.

Acilius mediatus Say. (2653)

Localities: MINNESOTA - Ramsey Co.

Hydroadephaga of Manitoba and Minnesota

111

Genus Thermonectes Crotch

Thermonectes ornaticollis Aube. (2654)

Localities: MINNESOTA - Le Sueur Co. (Fish Hatchery).

Genus Graphoderus Aube .

Graphoderus liberus Say. (2659)

Localities: MANITOBA - in the southern half; local, not common. MINNESOTA - prob-

ably universally distributed.

Graphoderus perplexus Sharp. (2661)

Localities: MANITOBA - throughout southern half. MINNESOTA - Grand Marais, Ram-

sey Co., St. Louis Co., St. Paul.

Graphoderus fasciatocollis Harris ( =fascicollis Harris)

Localities: MINNESOTA - Anoka Co., Beaver Dam, Hubbard Co., Le Sueur Co., Owa-

tonna, Ramsey Co., Stillwater, St. Paul.

Graphoderus manitobensis Wallis.

Localities: MANITOBA - Winnipeg (single male type).

Graphoderus occidentalis Horn. (2663)

Localities: MANITOBA - abundant in southern portion. MINNESOTA - Grand Marais,

Hubbard Co., Le Sueur Co., Ramsey Co., St. Anthony Park, St. Paul.

SUBFAMILY CYBISTERINAE

Genus Cybister Curtis

Cy bister fimbriolatus Say. (2667)

Localities: MINNESOTA - Le Sueur Co., St. Paul.

FAMILY GYRINIDAE

Genus Dineutus MacLeay

Dineutus discolor Aube. (2674)

Localities: MINNESOTA - Mora (one specimen).

Dineutus horni Roberts. (2681)

Localities: MINNESOTA - apparently throughout the state.

Dineutus nigrior Roberts. (2679)

Localities: MANITOBA - Victoria Beach ( one specimen). MINNESOTA - Ramsey Co., St.

Louis Co., St. Paul.

Dineutus assimilis Kirby.

Localities: MANITOBA - southern half. MINNESOTA - universally distributed.

Genus Gyrinus Geoffroy

Gyrinus minutus Fabricius. (2684)

Localities: MANITOBA - very abundant, north to Churchill. MINNESOTA - abundant

throughout the state.

Gyrinus ventralis Kirby. (2691)

Localities: MINNESOTA - Ramsey Co.

Gyrinus aeneolus LeConte. (2687)

Localities: MANITOBA - Township 7 Range 1 IE. MINNESOTA - Anoka Co., Hennepin

Co., Mora, Ramsey Co., Rochester.

Gyrinus dichrous LeConte. (2689)

Localities: MANITOBA - Berens River (east side of Lake Winnipeg). MINNESOTA -

Afton, Detroit, Itasca State Park.

112

Larson

Gyrinus latilimbus Fall. (19250)

Localities: MINNESOTA - Bengali, Cook Co., Two Harbors.

Gyrinus bifarius Fall. (19251)

Localities: MANITOBA - not taken in southern half of the province; moderately abund-

ant from Le Pas northward at least as far as the Kettle Rapids on the Nelson River. MINNE-

SOTA - Mora (one specimen).

Gyrinus confinis LeConte. (2685)

Localities: MANITOBA - abundant everywhere as far north as the Kettle Rapids. MINNE-

SOTA - Detroit, Itasca Co., Lake City, Lake Itasca.

Gyrinus aquiris LeConte. (2692)

Localities: MANITOBA - Husavick (one male). MINNESOTA - Detroit, Hennepin Co.,

Minneapolis, Ramsey Co., St. Paul.

Gyrinus maculiventris LeConte. (2695)

Localities: MANITOBA - extremely abundant as far north as Mile 214 Hudson’s Bay Rail-

way. MINNESOTA - apparently abundant, at least in northern half of state.

Collecting notes: This species remains active all winter, specimens having been taken at a

hole in the ice on the Assiniboine River near Treesbank by E. Criddle on January 9, 1928.

Gyrinus affinis Aube. (2696)

Localities: MANITOBA - widely distributed. MINNESOTA - Cook Co., Lake Co., Raw-

ichami River, St. Louis.

Gyrinus borealis Aube. (2707)

Localities: MANITOBA - recorded by Bell from the Nelson River 55° 50' N 99° 30'W,

but I have no means of checking the record.

Gyrinus pugionis Fall. (19255)

Localities: MINNESOTA - Babbitt, near Ely, Itasca State Park.

Gyrinus picipes Aube. (2704)

Localities: MANITOBA - Mile 214 Hudson’s Bay Railway.

Gyrinus lugens LeConte. (2707a)

Localities: MANITOBA - widely distributed. MINNESOTA - Kawishiwi River, Lake Co.

Gyrinus analis Say. (2700)

Localities: MANITOBA - Onah (a single female det. by H.C.Fall). MINNESOTA - Fort

Snelling, Minneapolis. Rochester.

Gyrinus opacus Sahlberg. (2702)

Localities: MANITOBA*- Mile 214 Hudson’s Bay Railway and northward.

Gyrinus wallisi Fall. (19256)

Localities: MANITOBA - Baldur, Stonewall, and northwards probably to the northern

limits of the province.

Gyrinus impressicollis Kirby. (2706)

Localities: MANITOBA - Mile 214 Hudson’s Bay Railway (Piquitenay River)

Collecting notes: In 1917 I took a few specimens on the Piquitenay River. These were

swimming on the rough surface of the deeper water in ones or twos and were seen nowhere

else. A few years later, a specimen or so turned up unexpectedly among some material taken

in the evening when crossing Long Pine Lake at Ingolf, Ontario. The experience on the

Piquitenay River was recalled and on my next visit to Ingolf search was made for impressi-

collis out in the deep waters some distance from shore, with such success that many were

captured. This information was passed on to Mr. W. J. Brown of Ottawa who also took it in

numbers on Lake Kazubazua, Quebec. Hence, it seems probable that if search be made in

the rougher waters some distance from the shores, impressicollis will be found to inhabit

most of the rocky lakes of the Laurentian Highland.

Hydroadephaga of Manitoba and Minnesota

113

ADDENDA

Hydroporus lapponum Gyllenhal. (19216)

Localities: MANITOBA - Churchill (Brown, 1937).

Agabus browni Leech

Localities: MANITOBA - Churchill (Leech, 1938).

Agabus colymbus Leech

Localities: MANITOBA - Churchill (Leech, 1938).

Agabus hudsonicus Leech

Localities: MANITOBA - Churchill (Leech, 1 938).

Agabus velox Leech

Localities: MANITOBA - Churchill (Leech, 1939).

REFERENCES

Anderson, R. D. 1962. The Dytiscidae (Coleoptera) of Utah: keys, original citation, types

and Utah distribution. Gt. Basin Nat. 22: 54-75.

Bird, R. D. 1958. John Braithwaite Wallis, 1877- .

Newsl. ent. Div. Dept. Agric. Can. 36(5): 2-3.

Brown, W. J. 1937. The Coleoptera of Canada’s Eastern Arctic. Can. Ent. 69: 106-1 1 1.

Fall, H. C. 1919. The North American species of Coelambus. Mt. Vernon, N.Y.: John D.

Sherman, Jr., pp. 1-36.

1922a. A revision of the North American species of Agabus together with a description of

a new genus and species of the tribe Agabini. Mt. Vernon, N.Y.: John D. Sherman, Jr.,

pp. 1-36.

1922b. The North American species of Gyrinus. Trans. Am. ent. Soc. 47: 269-306.

1923. A revision of the North American species of Hydroporus and Agaporus. Salem.

Mass., printed by S. E. Cassino Co., pp. 1-129.

Hatch, M. H. 1953. The beetles of the Pacific Northwest. Part 1: Introduction and Adeph-

aga. Univ. Wash. Pubis. Biol. 16: vii + 1 - 340.

Leech, H. B. 1938. Descriptions of three new species of Agabus from Hudson Bay (Cole-

optera: Dytiscidae). Can. Ent. 70: 123-127.

1939. On some Nearctic species of Agabus, with the description of a new species. (Coleop-

tera: Dytiscidae). Can. Ent. 71: 217-221.

Wallis, J. B. 1924. Two new species of Coelambus. Can. Ent. 56: 105-108.

1926a. The status of Gyrinus piceolus Blatchley (Coleoptera). Can. Ent. 58: 50.

1926b. Some new Coleoptera. Can. Ent. 58: 89-95.

1933a. Revision of the North American species (north of Mexico) of the genus Haliplus,

Latreille. Trans. R. Can. Inst. 19: 1-76.

1933b. Three new species of Hydroporus belonging to the vilis group. (Coleoptera). Can.

Ent. 65: 261-262.

1933c. Some new Dytiscidae (Coleoptera). Can. Ent. 65: 268-278.

1939a. Hydaticus modestus Sharp versus Hy da ticus stagnalis Fab. in North America

(Coleoptera, Dytiscidae). Can. Ent. 71: 126-127.

1939b. The genus Graphoderus Aube in North America (North of Mexico) (Coleoptera).

Can. Ent. 71: 128-130.

114

Larson

1939c. The genus Ilybius Er. in North America (Coloptera, Dytiscidae). Can. Ent. 71: 192

-199.

1950. A new species of Dytiscus Linn. (( oleoptera, Dytiscidae). Can. Ent. 82: 50-52.

Young, F. N. 1969. A checklist of the American Bidessini (Coleoptera: Dytiscidae- Hydro-

porinae). Smithson. Contr. Zool. 33: 1-5.

Young, F. N. and H. C. Severin. 1956. Evidence of intergradation between putative species

of Colymbetes in South Dakota (Coleoptera: Dytiscidae). J. Kans. ent. Soc. 29: 79-83.

BIOLOGY OF BOMBUS POLARIS CURTIS AND B.HYPERBOREUS SCHONHERR

AT LAKE HAZEN, NORTHWEST TERRITORIES

(HYMENOPTERA: BOMBINI)

K. W. RICHARDS 1

Department of Entomology

University of Alberta

Edmonton, Alberta, Canada

Quaestiones entomologicae

9: 115-157 1973

Adaptations for survival in a high arctic environment by Bombus polaris and B. hyper-

boreus are described. For B. polaris, adaptations related to low temperature are: structural

characteristics of adults— large size, long dense hair, and dark color; behavioral— nest con-

structed on surface sites, with entrance facing towards the sun at maximum elevation, and

flight by queens and workers close to ground surface. For the short season of growth, adap-

tations include: development of only a single brood of workers prior to production of sex-

ual forms; eggs of first brood all laid in a single cell, and larvae fed collectively ; extended

foraging activity by queens and workers in continuous daylight; and acceptance by for-

agers of a wide variety of flowers. B. hyperboreus is a nest parasite of B. polaris. Adults of

B. hyperboreus are similar to those of B. polaris in characteristics not associated with

nesting. Shortening of the life cycle of B. hyperboreus was achieved by elimination of the

worker caste.

This paper reports studies of various aspects of the biology of Bombus polaris Curtis,

1835 and B. hyperboreus Schonherr, 1809. The purpose of these studies was to determine

how these species have adapted to life in an arctic environment. Investigations were con-

ducted at Lake Hazen (81° 49' N, 71° 18' W), Ellesmere Island, Northwest Territories,

Canada, in the study area described by Savile (1964), from May 24 to August 20, 1967 and

May 30 to August 28, 1968.

NOMENCLATURE, SYSTEMATICS AND GEOGRAPHICAL DISTRIBUTION

The high degree of polymorphism exhibited by northern species of Bombus and lack of

communication between North American and European workers in describing new species

collected by early arctic explorers has resulted in lengthy synonymies. Not all such problems

are yet settled, and an explanation is required to justify use of one of the names in this

paper.

Because the name Bombus arcticus Kirby, 1824, used by Richards (1931) is a secondary

junior homonym of Apis arctica Quensel, 1 802, which in turn is a junior subjective synonym

of Bombus agrorum Fabricius, 1793, Kirby’s arcticus must be replaced by another name.

The name Bombus polaris Curtis, 1835, a junior subjective synonym of B. arcticus Kirby, is

available, and is used here as the valid name. O. W. Richards (pers. comm., 1969) accepts

this name change.

The species B. polaris, B. hyperboreus and five others belong to the subgenus Alpino-

bombus (Richards, 1931 and 1968). The group is in need of revision, a task rendered diffi-

cult by shortage of material and much variation in physical characteristics among the

species. Members of Alpinobombus are confined to arctic and alpine tundra in the holarctic-

^ Present address: Department of Entomology , University of Kansas,

Lawrence, Kansas 66044, U.S.A.

116

Richards

region. They are found in the Alps, Arctic Europe, Asia, Greenland, Arctic America, and in

the mountains of western North America as far south as Arizona. Bombus polaris (Fig. 1)

and B. hyperboreus (Fig. 2) are arctic and probably circumpolar forms. The apparent gaps in

their ranges in Siberia probably represent lack of collecting.

Me Alpine (1964, 1965a) indicated that members of only a small number of insect species

live in the northwest Queen Elizabeth Islands because of the environmental influences, and

that these are extremely tolerant of “harsh” arctic conditions. This harshness of the environ-

ment probably excludes members of Bombus from some areas. Generally the distributions

of B. polaris and B. hyperboreus are similar; B. polaris is recorded from areas where B.

hyperboreus is absent (i.e. Quebec and Labrador), whereas B. hyperboreus is recorded only

from areas where some other member ( B . balteatus or B. alpinus) of the subgenus is also re-

corded. Chernov (1966) and Brinck and Wingstrand (1949, 1951) illustrate this point. The

distribution of B. polaris and B. hyperboreus was determined from the literature and from

specimens in the Canadian National Collection and my collection. Records are published by

Strand (1905), Sladen (1919), Friese (1923a), Richards (1931), Hellen (1933), Braende-

garrd, Henriksen, and Sparc k (1935), Skorikov (1937), Henriksen (1937, 1939), Carpen-

ter and Holm (1939), Brinck and Wingstrand (1949, 1951), Yarrow (1955), Savile (1959),

Bruggeman (1958), Ander (1965), Chernov (1966), Swales (1966), and Mosquin and Martin

(1967).

NESTING

Artificial domiciles

To obtain enough bumblebees to study populations, flight activities, nest temperatures,

food preferences, and interspecific associations, attempts were made to attract queens to

artificial domiciles placed in their natural habitats. Colonies established in artificial domi-

ciles are easier to study than those in natural nests.

Materials and methods.- 1 used light weight domiciles of two designs (Fig. 3), each includ-

ing three parts: a masonite base, a body with entrance hole 19 mm or 25 mm diameter, and

a styrofoam top. Tops of one type were inverted blue flower pots, and bodies were pieces of

15 mm shellac-soaked cardboard tubing. Tops of the second type were rectangular white

boxes on bodies of XA in. plywood. Nesting material of upholsterer’s cotton was placed in-

side each box. Entrance directions varied. Black polyethylene tubing, one foot long, of

either 22 mm or 13 mm outside diameter, was connected to the entrance holes of some

domiciles to form entrance tunnels. The outer ends were cut obliquely to form landing plat-

forms. Domiciles were placed on the ground with small pebbles, soil, vascular plants and

moss over the tunnels. Permafrost barred the use of underground nests. In 1967 200 domi-

ciles in 10 localities and in 1968 180 domiciles in nine localities were used. The domiciles

were placed in the field when spring melt commenced. Because many domiciles were dam-

aged by arctic foxes, arctic hares, and musk oxen, fine-mesh chicken wire was secured over

the tops of the accepted domiciles to protect them.

Results. - Of the 380 domiciles used in two years, only five were occupied by B. polaris

queens. Each was of the blue styrofoam flower pot type with a large tunnel. Entrances to

three faced between 250° -280° and two faced between 100° -115° range, suggesting no

strong directional preference by the bumblebees which occupied them. The domiciles were

in four different habitats. At acceptance time, adjacent areas were free of snow, though

some melting snow drifts remained on N and NE-facing banks of Skeleton Creek and on

slopes in the higher fault zone.

Biology of Bombus polaris and B. hyperhoreus

117

Natural nests

The purpose of this investigation was to discover factors controlling nest initiation by

arctic bumblebees. Areas that queens rejected while searching and the habitats in which they

eventually established were also investigated.

In the arctic regions, data about nests of various Alpinobombus species have been re-

ported by: Jacobson (1898), Friese (1904, 1908, 1923a, and b), Johansen and Nielsen

(1910), Frison (1919), Friese and Wagner (1912), Sladen (1919), Brinck and Wingestrand

(1951), Freuchen and Salomonsen (1958), L0ken (1961), and Milliron and Oliver (1966).

Materials and methods. -Queens in search of nesting sites were observed, and these data

were recorded: amount of time spent searching, habitats and sites investigated, possible rea-

sons for rejection of particular sites, temperatures and moisture conditions of the soil, avail-

able nesting material, and directions of tunnel entrances. Each queen in natural and artificial

domicile nests was marked with nail polish on a particular part of the body as soon as the

nest was found so that she could be recognized again.

Date of nest establishment was calculated by subtracting one to six days from the date on

which the nest was discovered, depending upon kind and amount of progress at the time

(Hobbs, 1964b). Data from the accepted artificial domiciles are included to indicate peaks

of establishment.

Areas in which nests occur. -Of 94 natural nests located at Lake Hazen, 92 were on the

surface of the ground, one was on a caribou rug in the sleeping tent, and one was in an aban-

doned lemming burrow. Nests were in marsh and sedge meadows along streams (Skeleton

Creek and creek no. 51) and beside pools and tarns (i.e. those in T6, Q7, P6, M10 of Fig. 6).

Three nests described by Milliron and Oliver (1966) were also in marsh meadows.

General characteristics and vegetation of marsh and sedge meadows (Fig. 7) and marginal

areas around each of the pools and tarns have been described by Savile ( 1 964) and Oliver

and Corbet (1966). Plants found^ in the moister meadows are of Juncus albescens (Lge)

Fern., J. castaneus Sm., J. biglumis L., Eutrema edwardsii R. Br., Cardamine pratensis L.,

Saxifraga hirculus L, and Ranunculus trichophyllus Chaix. The principal mosses are Dre-

panocladus brevifolius (Lindb.) Warst and Bryum s pp. The drier meadow areas are charac-

terized by the dominant Carex aquatilis Wahlenb. var stans (Drei.) Boot, with varving

amounts of Eriophorum scheutzeri Hoppe, E. triste (Th. Fries. )Hadac and L0ve, J. biglumis,

Arctagrostis latifolia (R. Br.) Griseb., Polygonum viviparum L., Salix arctica Pall, and lesser

amounts of Equisetum arvense L., E. variegatum Schleich., Pedicularis arctica R. Br., P.

hirsuta L., Cerastium beeringianum Cham, and Schlect., Saxifraga nivalis L., S. rivularis L.,

and Ranunculus sulphureus Sd. The principal bryophytes are Drepanocladus revolvens (Sw.)

Wamst, Orothecium chryseum (Schultes) BSG., and Campylium arcticum (Williams) Broth.,

and Bryum spp. The vegetation forms a closed cover over the partly decaying organic mater-

ial (Day 1964).

In meadows and marginal pool areas natural nests were on small flat areas, in depressions,

and beside small hummocks of moss or other vegetation. Variations in structure were numer-

ous. The majority (P>0.005) of nests (Fig. 5a) examined had entrances which faced in the

180° to 270° quadrant; more (Fig. 5b) faced between 225° and 270° than in the 180°-

225° sector. A possible explanation is the sun orientation at the daily temperature peak.

The maximum diel soil surface temperature occurred between 1300 and 1600 hours (Corbet

1966, 1967a, b) when the bearing of the sun progresses from 195° to 240° . The highest

temperatures around a conical mound of moist moss were at the west, not at the south

where the sun’s altitude was greatest: surface temperature of the north slope remained al-

most steady through 24 hours. Also, moss mounds and Dryas hummocks in sedge meadows

Throughout the text, flower nomenclature follows that of Porsild (1964) and Savile

(1964) and brophyte nomenclature is that of Brassard (pers. comm. 1969).]

118

Richards

were the warmest areas and showed exceptionally marked diel periodicities of surface tem-

peratures (Corbet, 1967b). Moderate to light moisture conditions were not deterrent fac-

tors in nest establishment for though the nests often became waterlogged beneath they re-

mained comparatively dry on the surface. Nests in the sedge meadows and along the north

banks of the streams were drier; the moisture in and around these nests resulted from the

later permafrost melt.

In an attempt to discover why surface sites in meadows and marshes were preferred to

rodent burrows (the traditional sites of bumblebee nests in temperate areas), temperature

data were obtained for both types of sites. Data are presented in Table 1 for abandoned lem-

ming holes at least 30 cm deep which queens had investigated. In a marsh and sedge meadow

(M7, Fig. 6) 75 readings were recorded from a grid 25 meters long by eight meters wide

(Table 2). Temperatures at the soil surface fluctuate daily, especially during June at Lake

Hazen (Powell, 1961 ; Corbet, 1967b) which may influence the temperature of the marshand

sedge meadows. Soil temperatures measured one foot beneath the surface (Powell, 1961)

were lower than temperatures in lemming burrows. The latter were more exposed and more

influenced by surface temperatures and solar radiation. Nonetheless, temperatures in the

lemming burrows were lower than those in the marsh meadow.

Table 1. Temperatures in abandoned lemming holes when they were investigated by B. pol-

aris queens; June 3 to June 23, 1967, and June 15 to July 3, 1968, at Lake Hazen, N. W. T.

Table 2. Soil temperature ranges at 5 cm depth in a marsh and sedge meadow (M7, Fig. 6) at

1400 his. on three different days in 1968 at Lake Hazen, N.W.T.

Biology of Bombus polaris and B. hyperboreus

119

Moisture conditions of the burrows examined by queens were also investigated. About 75

per cent of the burrows contained ice or permafrost, or were extremely damp in some part.

In contrast, surface moss and liverworts of the marsh meadows were not so affected by ice

I and permafrost, although they were briefly inundated by the fluctuating water level during

snow melt. In summary, surface areas of marsh and sedge meadows were warmer and drier

than were most of the lemming burrows.

Emergence of queens and the search for nest sites. - Physiological factors associated with

I egg development probably stimulate a queen to seek a nest and start a colony (Medler

1962a). The ovaries of B. (Bombus) lucorum L. queens in Surrey, England did not develop

until after a hibernation period (Cumber 1949) followed by an active feeding period of al-

most three weeks resulting in a noticeable increase in weight and in a swelling of the ovar-

ioles (Cumber 1949, 1953). At Hazen the pre-feeding period is about a week. After their

ovaries mature the B. polaris queens search for nesting sites.

The first B. polaris queens were observed May 27, 1967 and June 14, 1968, and the first

B. hyperboreus queens were observed on June 9, 1967 and June 21, 1968. During the first

few days of the season, queens visited for nectar and pollen the flowers of Saxifraga oppo-

sitifolia, the only ones in bloom at the time.

Hunting begins when Salix arctica blooms, which is within a week after the first flowers of

Saxifraga oppositifolia appear. In 1967, searching by B. polaris queens extended from June

3 to June 23, and in 1968 from June 15 to July 3. Searching for nests of B. polaris by

queens of B. hyperboreus extended from June 15 to July 16 in 1967, and from June 23 to

July 10 in 1968. Searching by B. polaris queens reached a peak four to six days after beginn-

ing and then declined for the rest of the period. For example, on June 19, 1968, 24 B. pol-

aris queens were observed seeking nests, whereas on June 30, only two such queens were ob-

served. The activity of B. hyperboreus queens reached a peak seven to 1 1 days after they

became active.

Queens searched in cracks in the soil in clay banked areas, in rocky areas, in marsh and

sedge meadows, around Dryas hummocks, and in abandoned burrows of lemmings. They

even searched the walls and caribou rug floors of the tents. Queens flew throughout the 24

hour period if the weather was favorable, usually less than 25 cm and not more than 30 cm

above ground level. They alighted now and then to inspect promising sites more closely.

The lemming burrows investigated were usually those with south to northwest-facing

entrances (Fig. 4). Diameters of these tunnels were in the 4-8 cm range. Investigation of a

burrow usually occupied less than 30 seconds, but some queens spent up to three minutes in

this activity.

Suitable material for construction of bumblebee nests was found in 27 lemming burrows.

This material had probably been used in construction of lemming nests and included dried

or decaying grasses, sedges, mosses hair, and feathers. Marsh meadows also provided nest-

ing material of mosses, leaves, grasses and sedges. Suitable nesting material was absent from

sparsely vegetated areas of saline clay, sand, gravel, Dryas-Kobresia habitats, mud and gravel

deltas, and most Dryas hummock habitats.

Nest establishment for nine B. polaris queens observed began on June 7 and ended June

17, 1967. The peak occurred June 9 or June 10 when four of the queens established. In 19-

68, for eight queens, the period was from June 15 to 25 with no obvious peak. Three

queens, while in the process of establishing, abandoned their nests, apparently because of

disturbances during examination. Four queens failed to return to their nests after a

light snow storm on June 29, 1968 when the screen temperature was 1.0 C.

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Richards

Nest construction. -Upon accepting a suitable location to establish a natural nest (Fig. 7)

or occupy a domicile, a queen completely rearranges the nesting material to form a brood

chamber. To loosen and rearrange the moss she pulls it with her mandibles and fore-legs and

pushes it under her body to the desired position with her mid- and hind-legs. Queens (and

later, assisting workers), continue to rearrange the material so long as the colony is expand-

ing. Most nesting material consists of mosses and liverworts. Names of the principal bryo-

phytes from 92 nests are presented in Table 3. The principal vascular plant-nesting material

consisted of dried blades of Carex aquatilis and Eriophorum spp. with Equisetum spp.,

Dryas and Salix in some nests. For a complete list of mosses and liverworts from the nests

with relative abundance, see Richards (1970). In 1967, representatives of 46 mosses and

four liverworts with an average of 7.1 (range 2-14) species were collected from each of the

nests; in 1968, representatives of 48 mosses and eight liverworts with an average of 6.7

(range 3-13) species per nest were collected.

Table 3. The principal bryophytes from 92 natural nests of B. polaris at Lake Hazen, 1967

1968.

Number of nests

with moss

Nests changed as the season progressed. Earlier the external dimensions were small, about

5 cm diameter, whereas later some were as large as 1 5 cm. The external covering of the nest

cavity was convex-oval with the mosses and dried sedge leaves intermixed to form a thick

and tightly constructed surface. The covering of some nests reached a height of 5 cm. Nest

cavities were in shallow depressions covered with dried moss, leaves, roots and occasionally

peat. The queen and workers excavated parts of the moss mounds beside the nests where the

bees defecated.

Biology of Bombus polaris and B. hyperboreus

121

Discussion.- The factors controlling nest initiation are complex. Whatever physiological

factors are involved must take effect either promptly after emergence from hibernation or

become effective during the fall feeding period and then again before or during spring emer-

gence. Queens investigate all major habitats for a suitable site. That queens search such areas

as lemming holes suggests that in some localities B. polaris queens regularly establish under-

ground. This is also suggested by a few publications (Johansen and Nielsen, 1910; Frison,

1919). Another possibility is that underground searching is a trait inherited from the ances-

tral stock of B. polaris.

The habitat in which the queen establishes her nest has the following characteristics; it is

warmer than habitats of compact soil, has more suitable nesting material than is present in

Dryas-Kobresia habitats, or in sand and gravel knoll habitats; it is adjacent to food sources;

and it lacks the excessive moisture of the clay, sand, gravel, and Dry as hummock areas dur-

ing nest establishment. I believe that low temperatures of soil or lemming burrows inhibit

queens from establishing in such places, even if other factors are favorable. While founding

queens are not specific in choosing nesting material, they seem to prefer leaves and bracts of

mosses and liverworts.

Interestingly, queens investigated lemming burrows with entrances facing the same direc-

tion as sites on which most nests were located. The patterns of distribution and behavior of

bumblebees are correlated with the aspect of the slope they inhabit, as is true of many other

organisms living in the arctic environment (Corbet, 1967).

COLONY DEVELOPMENT

Colony composition

Among the potential aspects for adaptations of arctic and alpine bumblebees are the phys-

iological and morphological stages of a colony. Because the climatic environmental factors

(i.e. low temperature, small heat budget, and reduced growing season) are more severe than

in other regions the adaptive developmental changes have become vital in allowing the bum-

blebees to survive in the high arctic.

There has been little previous work on Alpinobombus brood-rearing behavior and all but

two of the authors who have investigated the various species, described only the contents of

nests at a particular stage when they were collected (Sladen (1919); Frison (1919 and

1927a); Brinck and Wingstrand (1951); Johansen and Nielsen (1910); Milliron and Oliver

(1966); Hasselrot (1960); Hobbs (1964a and b); L^ken (1961M

Materials and methods.- Observations and rough sketches were made and photographs

were taken of five artificial domicile nests and 24 natural nests at intervals of two to five

days throughout the seasons. I determined arrangement and number of eggs, larvae, pupae,

and pollen by dissecting the wax-pollen canopy covering the broods. Nests were periodical-

ly collected and the brood was examined in the laboratory.

Before I gained experience in removing the protective covering and accidentally touched

the brood, the workers removed the eggs or larvae from a recently exposed wax-pollen cell

with their mandibles and carried them outside the nest. These larvae were examined and

collected. I also observed workers eating the exposed eggs in egg cups an activity which led

Brian (1951) to suggest cannibalism. When I exercised care and did not touch the eggs or

larvae, the workers repaired the wax-pollen covering over them. During observations, expo-

sure of the nest resulted in temperature drops comparable with those resulting from absence

of a queen during foraging (Fig. 17); such additional cooling is not considered to be cn im-

portant stress factor on brood development. In one instance, a 1 5 minute observation result-

ed in a drop in nest temperature of 8.25 C (30.25 to 22 C), in another, a 17 minute observa-

122

Richards

tion resulted in a decline of 10.0 C (31 to 21 C).

First brood.- The following is a summary of observations: After a queen of B. po laris

completely rearranges the moss or upholsterer’s cotton nesting material, she visits flowers

of Saxifraga oppositifolia or Salix arctica and returns to the nest with two pollen pellets.

After placing these pellets side by side she builds the wax-pollen first brood cell (Fig. 8) on

top of them. The honey pot was built after the brood cell by the nine queens observed. The

honey pot is built separately from the brood and in line with the longitudinal axis of the in-

cubation groove. It is sufficiently near the groove that the queen while incubating drinks from

it without leaving the brood. Before she builds the honey pot the queen stores honey in the

moss and cotton nesting material, providing a small food reserve which also acts as an early

insulating layer. Eggs are usually deposited vertically in a single cup, at one time. In one in-

stance the egg cell was probably reopened and additional eggs were laid, for they were in two

separate groups within the cell. Egg cells have a definite area under them for fresh moist

pollen during larval development. Thus the queen at this stage is a “pocket-maker”. The

wax-pollen canopy covering the egg cells is a dark brown, rough mass colored by the yel-

lowish pollen of S. oppositifolia and S. arctica. The incubation groove, although poorly

formed on the egg cell, was present in all nests observed.

First brood larvae are fed moist, easily manipulated pollen by the queen. She pushes it

under the brood in several places, especially under the sides of the incubation groove. Larvae

in front of the incubation groove (toward the nest entrance) were larger and developed

more rapidly than those toward the back, because most pollen is pushed under larvae in the

former position and the honey pot is nearer to them. After one to three days growth, the

larvae lie side by side in a curled position (Fig. 9) allowing easy access to the pollen below.

When pollen is plentiful the larvae eat into the mass until they become completely enclosed.

Last instar larvae construct separate cells immediately before spinning cocoons and are then

fed individually with a mixture of honey and pollen. Often they are such a size that the wax-

pollen canopy does not completely cover them. In some nests the color of the wax-pollen

covering first the egg cells and then the larvae changes from a dark brown to a lighter tan,

owing to a change in the pollen supply.

Cocoons are separated from each other by flimsily spun silk and by the wax -pollen cover-

ing. When no fresh pollen is brought into the nest, that already present begins to harden and

dry. The incubation groove, evident from the initial egg cell construction, is more pronounc-

ed from mid-larval stage to worker emergence.

First brood larvae usually become workers. In one instance, however, three workers andl 1

males (imagines and late pupae) were present when a nest was collected. This nest was still

developing, for it had an egg cell with 1 2 eggs and eight two-or-three-day old larvae. The

queen had abandoned the nest, apparently because there were fewer workers to assist her in

food gathering and incubation, and the few emerged males and workers had depleted the

honey. The workers which emerge first are those in the front of the brood nearest the honey

pot on the base of the incubation groove. Succeeding emergences progress posteriorly

through the incubation groove to the brood furthest from the honey pot. Hence the larger

workers emerge first, the small ones later. Marked variation in size of workers within one

nest has been reported for other species by Sladen (1912), Cumber (1949), and Medler

(1962b, 1965).

Second and third broods (sexuals).- Second and third brood egg cells, also made of wax-

pollen material, are built on the outside tops of the cocoons that form the sides of the

incubation grooves (Fig. 10). The egg cells are constructed one to two days before or after

the first brood larvae begin to spin. Eggs at this time are laid in a horizontal orientation

(Figs. 11, 12), with most side by side or on top of each other. In 10 of 13 instances, the

Biology of Bombus polaris and B. hyperboreus

123

first eggs were placed in the egg cell which occupied the front half of the incubation groove.

Usually, one of the ridges of an incubation groove was completely covered with egg cells

before any were built on the other ridge. Additional egg cells were not built on the same

ridge, until after the first eggs had hatched. As a result, brood in different stages of develop-

ment can be on the same ridge. No pollen is placed in or under the egg cells to prime the

eggs of the second or third broods. The egg cells are small, thick, dark brown wax-pollen

cups 6-7 mm long and 3-4 mm wide according to the number of eggs laid in them.

When larvae on the fore part of the incubation groove emerge the wax-pollen canopy cov-

ering is not extended to any adjacent or posterior group. Therefore, the larvae beneath the

canopy on any ridge are separated and of different sizes on each side of the incubation

groove. Fresh moist pollen is pushed into pollen pockets(Fig. 13) beneath the larvae on the

edges of the brood when the first eggs hatch. The pollen pots for feeding male and queen

larvae are larger than the pollen receptacles under the worker larvae. Bees at this stage of

development become “pollen storers.” The curled position and pollen diet of second and

third brood larvae are identical to those of first brood larvae. Although they built separate

cells the last instar larvae were not always completely covered with wax as were the earlier

instars. All larvae are fed pollen; larvae of sexuals are also fed honey from the honey pots.

Last instar larvae are fed by workers through holes in the wax-pollen canopies.

While the first male cocoons were being spun (Fig. 14), egg cells were still being built;

eggs in these cells either did not develop or they were removed by workers. None of the sec-

ond or third brood in any observed nests were workers. The first egg batches of the sexual

brood produced males. Subsequent batches, laid on the remaining part of the incubation

groove or on the canopy covering the last instar larvae and early cocoons of the second

brood, produced queens (Fig. 15). Males remained in the nest up to two days whereas the

queens remained up to seven. The original established queen remained with her nest until

after the first males emerged.

Honey pots.- Queens built two or three additional honey pots up to 2-3 cm high and

1-1.5 cm diameter before the workers emerged. These were usually Vi to % full. During the

season the capacity of each honey pot varied, as first the queen and later the workers added

or removed wax. Two small honey pots, one taken from each of two nests while the first

brood was in the egg stage, had capacities of 1.12 cc and 1.73 cc. After the workers emerged,

as many as 14 honey pots were built (Fig. 16), each usually Vi to % full of honey, less full in

times of. unfavorable weather. Honey was also stored in some of the old worker cocoons and

the empty male cocoons, but these storage areas were at most % full.

Pop ulation structure.- The population of eggs, larvae, and pupae of the first, second, and

third broods for each year are indicated in Table 4. Reasons for the variation in egg numbers

are unknown. The ratio in spring of females to workers produced was 1:16, and in fall the

ratio was 1 : 3.03 (1967) and 1:1.76 (1968). The numbers of eggs per cell of male and female

broods is given in Table 5. Differences in numbers between years and in sex ratio were not

statistically significant at the 95% level. The ratio of the number of larvae of sexual forms to

the number of workers in 1967 was 1.57; in 1968 it was 1.53. Sex ratios within each nest

were in 1967, 2.0 males per female and in 1968, 1.6 males per female. Although queens

were successful in rearing almost all first brood workers to maturity, they were less success-

ful with second and third broods. Of those nests studied during the entire seasons of 1967

and 1968, mortality before emergence of adult males was 57.1% and 37.2% and of adult fe-

males was 62.5% and 41.0%.

124

Richards

Table 4. Numbers of first, second, and third broods of eggs, larvae, and pupae from nests of

B. polaris at Lake Hazen, N.W.T., for 1967 and 1968.

Table 5. The numbers of eggs per egg cell of B. polaris male and female broods per year in

the artificial domicile nests and natural nests at Lake Hazen, N.W.T.

Times required for development of each stage and each brood of B. polaris for each year,

are given in Table 6.

Discussion.- Species of Alpinobombus differ from one another and from members of

other subgenera in oviposition characteristics, colony size, brood composition, and length of

time required to complete stages of the life cycle. As an illustration of subgeneric differences,

data for B. balteatus (Hobbs, 1964a, b) are aligned with my data for B. polaris (Table

7).

Biology of Bombus polaris and B. hyperboreus

125

Table 6. Duration of development in days of eggs, larvae, and pupae of first, second, and

third broods of B. polaris at Lake Hazen, N.W.T., 1967 and 1968

Table 7. Differences between the composition and development of brood of B. polaris and

B. balteatus (data for B. balteatus after Hobbs 1964b).

B. polaris B. balteatus

First brood

- all eggs deposited at one time

- average no. of eggs in four broods 16.24

(range 1 5 - 17)

- average no. of larvae in 13 broods 15.9

(range 14 - 19)

-eggs laid on same ridge of incubation

groove were of different ages and castes

-wax-pollen canopy covering larvae of

adjacent cells not extended to cover all

larvae on the same ridge

-only one brood of workers produced

before males and queens

-not all eggs deposited at one time

-average no. of eggs in eight broods 1 1

(range 7-21)

-average no. of larvae in five broods 14

(range 12 - 15)

brood

-eggs laid on same ridge of incubation

groove were of different ages but same

caste

-wax-pollen canopy covering larvae of

adjacent cells extended to cover all lar-

vae on the same ridge

-sometimes more than one brood of

workers produced before males and

queens

126

Richards

Queens of Alpinobombus colonies are the onlv bumblebees known to place all eggs of the

first broods in single egg cells. The numbers of eggs laid per cell for second and third broods

of B. polaris (and probably for other species of Alpinobombus) were the same as those laid

in succeeding worker and sexual cells by queens of the subgenera Bombus and Cullumano-

bombus, and were greater than the numbers laid by queens of Subterraneobombus, Fer -

vidobombus, Pyrobombus and Bombias. The last-named is the only subgenus known of

which queens lay only a single egg per cell for the second and succeeding broods (Hobbs,

1964a, 1965a).

More eggs per brood are laid by queens of B. polaris in the first three broods than are laid

by queens of most other species of bumblebees. However, because of environmental factors

restricting total number of broods, seasonal egg production overall is less for arctic species

than for those inhabiting warmer areas, farther south.

Colonies of Alpinobombus species produced more workers in the first broods than did

colonies representing most other sub-genera studied by Hobbs (1964-1968). It is difficult to

estimate values for workers per queen in fall for species with colonies living under warm-

er conditions in lower latitudes because of the wide variation in numbers of workers among

different colonies. Generally, however, values for this ratio are about the same for both

temperate and arctic-alpine species.

Colonies of Alpinobombus species are small because usually only a single brood of work-

ers is produced before a queen begins to produce the sexual broods (Hobbs, 1964a). Queens

of other subgenera produce at least two worker broods— hence more workers— than Alpino-

bombus colonies produce. Among other subgenera, limited data suggest that number of work-

ers per colony varies with latitude: the Holarctic arctic-alpine B. (Pyrobombus) sylvicola

Kirby has as many as 139 worker cocoons (Hobbs, 1967b), wheras the tropical B. medius pro-

duces as many as 2183 workers (Michener and La Berge, 1954).

The paucity of workers led Friese (1902, 1908, 1923a and b) and Friese and Wagner

(1912) to suggest that arctic bumblebees including Alpinobombus species are tending

toward a solitary mode of life. I do not believe that this is so. Rather, I think that the re-

duced colony size is adaptive to life under arctic conditions. Certainly, queens of these

northern taxa behave toward their broods as social insects, just as do their southern counter-

parts.

Hasselrot (1960) reported that rates of development of the various life stages and larval

instars of a selection of Bombus species were similar to one another. Mean values for all

species studied were as follows: egg stage - 3.4 days; larval stage - 10.8 days; pupal stage-

1 1.3 days; total average time of development - 24.5 days. The rate of development is com-

parable to the above data for the life stages of B. polaris (cf. Table 6).

Brood reduction in the form of mortality of immatures of B. polaris at Lake Hazen was

the result of cannibalism and lack of food. The latter is related to periods of unfavorable

weather (about 4.0 C to 2.0 C, wind 8-12 mph, and complete cloud cover) during which

workers had difficulty in foraging and the food supply within the colonies became depleted

(i.e. honey pots less than 1/3 full and scarcity of fresh moist pollen). Under these condi-

tions, workers demolished egg cups and eggs and removed larvae from the nest, thus proba-

bly killing them. Food sources were also influenced by unfavorable weather as nectar secre-

tion was reduced and stamens became devoid of pollen.

Mortality data for the observed colonies of B. polaris are insufficient for detailed analysis

because not all second and third brood egg cells were opened to determine the maximum

number of eggs laid. However, the data available suggest that colonies in artificial domiciles

had fewer deaths than had colonies in natural nests, possibly because of the insulating effect

of the styrofoam tops. Cumber (1949) estimates at least 50% mortality before emergence of

Biology of Bombus polaris and B. hyperboreus

127

adults for bumblebees in general, and Brian (1951) estimates mortality of B. agrorum Fab-

ricius colonies based on all broods except the last, for two years, at 64 and 69 per cent.

At Lake Hazen, bumblebees are one of the few groups of insects to use most of the peri-

od of active growth. Adults are active soon after the first flowers bloom in June, continuing

until early in August when few flowers are left. Progression of brood development parallels

progression of weather and plants.

Flight activity

This is a sensitive indicator of foraging conditions in the field, and of the ability of colo-

nies to exploit available food sources. Rate of food acquisition strongly influences reproduc-

tive capacity, and tempo of most activities within colonies of honey bees (Gary 1967), and

the same principles may be applied to bumblebee flight activitiy. The purpose of this study

was to characterize weather conditions affecting the foraging of arctic bumblebees, adapta-

tions of the bees to the weather conditions, frequency of flight per 24 hours, and type of

food (pollen or nectar) exploited.

The effect of weather and general climatic conditions on bumblebee flight is important,

1 especially in arctic areas, where continuous daylight during the summer permits maximum

frequency and duration of this activity (Jacobson 1898 in Friese 1904, 1908, 1923a, b, and

Friese and Wagner 1912; and Johansen and Nielsen 1910;Sladen 1919;Frison 1919;L$ken

1949, 1954; Longstaff 1932; Bruggeman 1958; Freuchen and Salomonsen 1958; Savile

1959; Hasselrot 1960; Gavriliok 1961; Downes 1964; Hocking and Sharplin 1964; and Mill-

iron and Oliver 1966). Structural features of possible adaptive significance, such as large

size, hairiness, and melanism, and some physiological factors affecting foraging in cooler

weather are discussed.

Materials and methods.- In 1968, 24 hour observations were made every six days for 36

days at an artificial domicile. These six series of observations were made at the following six

stages of development: (1) first brood mid-larval; (2) first brood early-pupal and second

brood egg; (3) first brood emergence, second brood late-larval to early pupal, third brood

egg; (4) second brood late-pupal, third brood mid-larval; (5) second brood early emergence,

third brood pupal; and (6) third brood late-pupal to early emergence. Six supporting series

of observations from a natural nes t at unspecified intervals of hour and day, and occasional

flight activity observations at an artificial domicile in 1967 were also made. The brood com-

position and population of the nests were recorded the day before each observation. Thus,

the flight activity of bumblebees at major brood development periods was characterized.

Flight was observed from a seated position far enough from a nest to avoid disturbing the

bees but near enough to recognize the caste (queen or worker) and presence or absence of

pollen on legs. When reference is made to ‘pollen load’ or ‘pollen-gathering’ it is assumed

that a bumblebee was often also carrying nectar (Brian 1952;Free 1955b). The terms ‘nec-

tar load’ and ‘nectar-gathering’ are used only when the forager in question has not been

gathering pollen (Free 1955b). For each flight the times (Eastern Standard) of departure

and return, the caste of the bee and whether it carried pollen, were recorded. Air tempera-

ture was taken initially with a thermistor probe and later with a dial thermometer. Wind ve-

locity in mph and wind direction were estimated with a portable floating ball-type anemom-

eter. All were taken near the nest at a height above ground of 20-30 cm. Cloud cover was

estimated visually. Several readings of air temperature, wind velocity and direction, and sev-

eral estimates of cloud cover throughout each hour of observation were averaged to increase

reliability of the data. Solar altitude was taken from Corbet (1966).

For each bee observed away from the nest entrance the following notes were taken: spe-

cies, caste or sex, time, air temperature, flying height above ground, wind direction and ve-

128

Richards

locity at this height, and cloud cover.

Frequency of food collecting at the nest entrance.- The integral components affecting

flight activity were the responses of foraging bees to intra-nest stimuli and to meteorological

conditions. Various combinations of light, temperature, wind, and humidity affected bum-

blebee flight, and might be sufficient to bring about periodicities in flight. Thus, diel perio-

dicities of weather factors near the ground at Lake Hazen (Corbet 1966, 1967b) are consid-

ered (Table 8). Jackson (1959a, b) reported for Lake Hazen that for 76% of the observa-

tions the average wind velocity from June 1 to August 2 was 5 mph or less. The predomi-

nant wind direction was NE, along the Lake Hazen trough (Jackson 1959b, 1960; Corbet

1966, 1967b) followed by ENE, E, and NNE (Corbet 1966). Cloud cover did not exhibit

diel periodicity, but a tendency was noted for opacity to increase slightly between 1300 and

2200 hours (Corbet 1966)

Many of the diel fluctuations were obscured by weather trends persisting longer than a

day, such as barometric pressure, wind velocity, and cloud cover. The most regular are those

resulting directly from solar radiation at or near the soil surface (Corbet 1966).

Table 8. Diel ranges of times of maxima and minima of weather factors near the ground at

Lake Hazen, N.W.T. (after Corbet 1966).

The frequency of flight and the number of pollen and nectar loads collected on various

days by foragers of B. polaris from an artificial domicile are shown in Figs. 1 7-22 and those

for a natural nest are shown in Figs. 23-26.

Before workers emerged, the queen (Fig. 17) flew at all hours with approximately equal

frequency, collecting more pollen than nectar to feed the first brood larvae. The queen was

absent from the nest approximately 30 minutes on each of 20 foraging trips. After each for-

age she deposited pollen and nectar into pollen pocket(s) and/or honey pot(s) and warmed

the brood to a temperature comparable to that of the nest before her forage. When the first

brood was in the early pupal stage (Fig. 1 8) the queen collected pollen, but the frequency

of her flights was less than during larval development.

The climax of worker foraging activity occurred between July 6 and July 16 when nutri-

tional requirements of second and third brood larvae were maximal (Fig. 19 and Fig. 20).

These larvae were fed mixtures of pollen and nectar. The proportions of pollen loads to nec-

tar loads collected by foragers for these two days were 2.20:1 and 1.14:1 respectively.

Thus, there is perhaps a difference in the proportions of pollen to nectar fed to the second

and third brood larvae. Throughout July 6 the amount of pollen foraged remained nearly

the same, whereas nectar gathering reached a peak during 1200 to 1600 hours. On July 12,

Biology of Bombus polaris and B. hyperboreus

129

however, the pollen gathering peak was between 1000 and 1400 hours and nectar-gathering

was proportionately higher most other times. Collectively nectar- and pollen-gathering oc-

curred between 1200 and 1600 hours on July 6 and between 0900 and 1500 hours on July

12. The highest number of worker bumblebees (37) passing through the nest entrance in one

hour was counted at about 1400 hours on July 6. On July 12 workers flew until 2330 and

no foragers spent the night away from the nest.

The queen and workers foraged primarily for nectar when adults of the second brood

began to emerge and during the third brood pupal period (Fig. 21). This corresponds to in-

crease in number of honey pots, to feeding of second and third brood larvae quantities of

honey, and to feeding of newly emerged adult males. Males ate honey immediately after

emergence. The queen flew 24 hours a day, but at infrequent intervals. She remained away

from the nest for longer periods than during the period of sexual larval development and de-

parted permanently soon after the first males emerged.

Peak flight activity for workers was between 1000 and 1700 hours and was not, as yet, a

complete 24 hour activity. Flight activity was restricted by food shortage in keeping with

reduced nutritional requirements. The population of the nest was also reduced because

some foraging workers had died.

On July 24 (Fig. 22), newly emerged third brood queens and some workers were ob-

served in flight at the nest entrance. All collected only nectar. The new queens and workers

flew throughout the 24 hour period with no definite peak in activity. The nutritional re-

quirements of the nest were low as the development of the colony was completed, but nec-

tar-gathering was necessary for maintenance of the sexual and worker forms and for preser-

vation of a sufficiently high nest temperature for emergence of the remaining fall queens.

Once males had left the nest they did not return.