sun

Gzoo

harvard university

LIBRARY

OF THE

Museum of Comparative Zoology

Quaestiones

entomologicae

MUS. COMP. ZCCL.

LIBRARY

FEB 4 W2

harvard

UNIVERSITY

A periodical record of entomological investigations,

published at the Department of Entomology, Uni-

versity of Alberta, Edmonton, Canada

VOLUME VII

1971

CONTENTS

Guest Editorial — Northwestern Caddisflies 1

Nimmo — The adult Rhyacophilidae and Limnephilidae (Trichoptera) of Alberta

and Eastern British Columbia and their post-glacial origin 3

Editorial — Publish or Perish? 235

Frank — Carabidae (Coleoptera) of an arable field in central Alberta 237

Chance — Correction for drag of a flight mill, with an example for

Agrotis orthogonia Morr. (Lep. Noctuidae) 253

Sehgal — Biology and host-plant relationships of an oligophagous leaf miner

Phytomyza matricariae Hendel (Diptera :Agromyzidae) 255

Erwin — Notes and corrections to a reclassification of

bombardier beetles (Carabidae, Brachinida) 281

Book Review 282

Book Review 283

Book Review 284

Guest Editorial — Excellence has no Nationality 287

Sehgal - A Taxonomic Survey of the Agromyzidae (Diptera) of Alberta, Canada,

with Observations on Host-Plant Relationships 291

Nimmo - Corrigenda on the Adult Rhyacophilidae and Limnephilidae (Trichoptera)

of Alberta and Eastern British Columbia and their Post-Glacial Origin 406

Thomas - An Apparatus and Method for the Field Separation of Tabanid Larvae

(Diptera: Tabanidae) from Moss 407

Announcement 409

Editorial — Four Men and a Moth 411

Jacobson — The pale western cutworm, Agrotis orthogonia Morrison

(Lepidoptera: Noctuidae): a review of research 414

Cheung — Purification and properties of arginine phosphokinase

from honeybees Apis mellifera L. (Hymenoptera, Apidae) 437

INDEX

in

Achillea 270, 276

A. millefolium 256, 335, 369, 386

A. sihirica 256-258, 262, 266-274, 305,

335, 369, 386

Actaea rubra 306, 384

Agonum cupreum 239, 242, 246

A. cupripenne 239

A. placidum 239, 242, 246

A. quadripunctatum 239

A. retractum 239

Agromyza 292, 294, 296-303, 384

A. albertensis 296-297

A. albipennis 296-298, 302, 392

A. ambigua 297, 300

A. ambigua group 296, 384

A. aprilina 296-298

A. brevispinata 296-299, 392

A. canadensis 296, 299, 386

A. fragariae 296-297

A. hockingi 296-300, 392

A. kincaidi 296-300, 392

A. lucida 298

A. masculina 296-297 , 300-302

A. nearctica 296-291, 301, 392

A. nigripes 296-298, 302

A. nigripes group 296, 384

A. populoides 296-297 , 302, 385

A. pseudorep tans 296-297 , 302

A. pseudorufipes 299

A. rep tans 302

A. reptans group 296

A. rubi group 296

A. rufipes group 261, 296

A. spiraeae 296-297, 300-302, 383

A. spiraeae 296

A. sulfuriceps 296,. 303

A. ulmi 298

A. urticae 302

A. vockerothi 296-297, 301-303

Agromyzidae

Alberta 291-405

adult characteristics 293-294

host-plant relationships 255-280,

291-405

taxonomy 291-405

Agropyron 322-323

Agropyron repens 387

A. smithi 387

Agrotis delorata 416

A. orthogonia 253-254, 41 1-436

A. orthogonia duae 416

Ahmad, T. 256

Ainslie, C.N. 422

Algonquina 107, 116, 143-144

Allegophylax 145

Allen, P. 256, 265

Allium cepa 268, 272, 274, 278

Allomyia 57-60

Amara 238, 247

A. apricaria 240, 242, 247

A. apricaria group 248

A. aulica group 248

A. avida 240, 242, 247

A. convexa 240

A. cupreolata 240

A. ellipsis 240, 242, 248

A. far eta 240

A. lacustris 239

A. laevipennis 240

A. latior 240, 242, 247

A. littoralis 240, 242, 248

A. lunicollis group 248

A. obesa 240

A. pallipes 240

A. patruelis 240, 242, 248

A. quenseli 240

A. sinuosa 240

A. torrida 239, 242, 246-248

Amaranthus 344

Amauromyza 292, 295, 326-321, 384

Amelanchier canadensis 317, 384

Amphicosmoecus 5, 50-51, 55-56, 207, 216

A. canax 55-56, 159, 162, 184, 203, 207, 213

Amphioxus lanceolatus 437

A. abnormalis 321

A. riparia 326-327 , 396

A. shepherdiae 326-327 , 386, 396

A. subinfumata 326-327

Anabolia 5, 83, 101, 124-128, 138, 210, 217

A. bimaculata 125-128, 177, 192, 203, 210,

213

A. bimaculata group 210

A. consocia 125-126, 176, 191, 204, 210, 213

A. macula ta 128

A. medialis 126

A. osburni 125-127, 161, 177, 191, 204, 210,

213

A. sordida 2 1 0

IV

Anabolina 108, 1 12, 120-121

Anacyclus 256

Anderson, N. H. 20, 23, 28, 53, 61, 154,

201.

Andrewartha, H. G. 418, 429.

Andrews, P. 440

Anemone 384

A. canadensis 359, 374, 385

A. multifida 372, 385

A. riparia 359-360, 385

Angelica 353

A. arguta 382, 386

Anisogamus 96

Anolopsyche 108

Anthemis 256

Anthrax molitor 428

Antineura 344

Antirrhinum 266

Apan teles griffini 428

Apatania 5, 64-71, 156, 207, 217

A. aenicta 69

A. alberta 65, 70-71, 165, 185, 203, 207,

214

A. arctica 65-66

A. auricula 66

A. complexa group 70-71

A. crymophila 65, 69-70, 165, 185, 204,

207.213.221

A. dalecarlica 66

A. fimbriata group 65-66

A. frigida 67

A. groenlandica 66

A. inornata 66

A. pallida 61

A. palmeni 66

A. shoshone 65, 68, 165, 185, 204, 207,

213.221

A. shoshove 406

A. stigmatella 65-68, 165, 185, 204, 207,

213.221

A. stigmatella group 67-68

A. wallengreni group 69-70

A. zonella 65-66, 159, 164, 185, 204, 207,

214. 221

Apataniinae 5, 50, 64-71

Apataniini 5

Apatelia 65-69

Apatidea 65-67

Apidae 437-446

Apis mellifera 437-446

Apium 271

Apolopsyche 108, 116

Aptinoderus 281

A. cyaneus 281

Aptinomorphus 281

Aptinus 281

Aquilegia 266, 271, 352, 357, 362-363, 384-

385

A. brevistyla 355

A. formosa 355-356

Aralia 353

A. nudicaulis 357, 385

Arctocia 126

Arctoecia 125-126, 138

Arctopora 5, 83, 133-135, 211,217

A. pulchella 134-135, 178, 192,204,211,

213

A. salmon 21 1

A. trimaculata 21 1

arginine phosphokinase 437-446

Arnett, R. H., Jr. 283-284

Arnica 352

A. cordi folia 358, 386

Artemisia 258, 266-269, 272-277, 325

Asclepias syriaca 278

Aster 295,312,328,333

A. ciliolatus 267, 351, 360, 386

A. conspicuus 358, 386

Asterias rubens 437

Asynarchus 5, 83, 128-132, 143,210,217

A. aldinus 129, 131-132, 178, 192, 203, 210,

213

A. bat chew ana 131

A. conerus 131

A. contumax group 130

A. curtus 129-131, 177, 192, 203, 210, 213

A. lapponicus 132

A. lapponicus group 130

A. mutatus 129-130, 177, 192, 204, 210, 213

A. planifrons 131

Atecyclus septemdentatus 42>1

A ty lotus 407-408

Avena 323

Badister obtusus 240

Bajkov, A. 76

Balduf, W. V. 76, 95, 1 19, 124-125, 128, 146

Baldwin, E. 437-439, 444-445

Ball, G. E. 54, 62, 97, 102, 113, 116, 124,

137, 143, 197-198

Banks, N. 19, 23-27, 33, 52-54, 57, 61-62,

67-68, 76-79, 89-92, 95-1 13, 1 16-130,

134-146, 149-150, 153-156

Bannister, L. H. 437

Baptisa tinctoria 331

Barnes, W. 416

Bayrock, L. A. 197-198

Beirne, B. F. 429

Bembidion 244, 248, 250, 251

B. bimaculatum 239-242

B. canadianum 239, 242-244

B. grape i 239

B. mutatum 239, 242, 244, 251

B. nitidum 239-243

B. nudipenne 239

B. obscurellum 239, 242-243, 251

B. quadrimaculatum oppositum 239, 242-

244, 25 1

B. rapidum 239

B. rupicola 239, 242-243, 251

B. sordidum 241

B. timidum 243

B. versicolor 239, 242-243

beetles

bombardier 281

Colorado potato 276-278

Benjamin, F. H. 416

Berecyntus bakeri 428

Berry, E. W. 124-125

Betten, C. 19, 23-27, 33, 53-54, 57, 61-62,

67-68, 76, 79, 89-92, 95-105, 108-113,

116-125, 128-130, 134-146, 149-150,

153-156

Betten, C., et al. 196

Bidens cernua 307, 386

Birch, L.C. 418,429

Blakely, P. E. 418-424, 427, 431

Blethan, S. L. 437, 444

Blickle, R. L. (see Morse).

Bombyliidae 428

Bonnetia compta 428

Bowman, K. 416

Brachinida 28 1

Brachinus adustipennis 28 1

B. fulminatus 281

B. kavanaughi 281

B. micro americanus 28 1

B. senegalensis 28 1

B. vulcanoides 281

Braconidae 428

Brady cellus congener 240

B. lecontei 240

Brady tus 248

Branch, H. E. 95

Braschnikov, W. C. 3 10

Brassica 274

B. khaber 267, 272

B. oleracea 273-275

Brauer, F. 67,98, 102

Brimley, C. S. 126

Brooks, A. R. 427-428

Brosemer, R. W. 438

Brown, A. W. 431

Brues, C. T. 119, 140

Bryson, R. A. 199

Buhr, H. 277

Burks, B. D. 196

Butomomyza 317

Butt, B. A. 424

caddisflies, Alberta species 1-234, 406

origins and relationships 196-223

taxonomy 3-195

Calamagrostis 321

Calathus ingratus 239, 246

Callinectus sapidus 437

Calliphora erythrocephala 437

Calosoma calidum 239-241

Calycomyza 292, 295, 325-326, 384

C. althaeae 325

C. artemisiae 325

C. cynoglossi 325

C. menthae 325, 386

C. solidaginis 325, 387

C. sonchi 325-326, 387

Campanula 267

Cancer pagurus 437

Cannabis 277

Cantu, S. 424

Carabidae 237-252, 281

Alberta 237-252

of arable fields 237-252

Carabus complanatus 281

C. maeander 239

C. serratus 239

C. taedatus 239

Caragana arbor escens 267

Carcinus maemas 437

VI

Care x 319-32 1

Carlson, C. W. 438

Carpenter, G. D. H. 97

Centrostephanus rodgersii 437

Cerodontha 292, 295, 317-325, 384

C. angulata 317-319

C. calamagrostidis 319, 321-322

C. capitata 319, 323

C. chaixiana 318-320

C. dorsalis 318, 324

C. eucaricis 317-320

C. flavocingulata 320

C. frankensis 295, 317-320

C. gibbardi 317-320

C. incisa 3 1 9-322, 387

C. inconspicua 319, 322

C. lateralis 323

C. longipennis 319, 324

C. muscina 319, 322-323

C. occidentalis 318, 324-325, 395

C. sc/rp/ 317-321, 387

C. semipostica 3 1 9

C. superciliosa 319, 323, 387

C. ultima 318, 321

Chaetotaulius 118-119

Chalcididae 428

Chance, M. A. 253-254, 425

Chelonus 428

Chenopodium 267

C. 27 1

Cheung A. C. 437-446

Chilostigma 149-150, 154

Chilostigmini 5, 82, 148-156

Chilostigmodes 5, 148, 150-151, 212, 217

C. areolaris 150

C.areolata 150-151, 181, 194, 204,212-214

C. forcipata 212

Chlaenius alternatus 240

Chlamys opercularis 437

Chryanda 5, 50, 143-144, 211, 216

C. centralis 143-144, 180, 193, 204, 211,

214

C. cordon 144

C. pallida 144

C. parvula 144

C. signata 144

Chrysanthemum 256, 258, 266-269, 272,

276-277, 369, 386

Chrysothamnus 425

Chrysops 407-408

Cicindela aequinoctialis 28 1

C. limbalis 239

Cirsium arvense 425

Claassen, P. W. (see Needham).

Clematis 384

C. verticillaris 267, 361, 385

Clifford, H. F. 96, 1 12, 1 16, 120-124, 132-

133

Clistoronia 5,83,120-122,210-211,216

C. caroli 121

C. flavicollis 2 1 0

C. formosa 2 1 1

C. maculata 2 1 1

C. magnifica 121-122, 176, 191, 203, 210,

214

Clistoroniella 83

Clostoeca 143

Cohen, M. 256, 263

Coleoptera 237-252

collecting methods

caddisflies 11-12

ground beetles 238

rove beetles 238

spiders 238

tabanid larvae 407-408

Colpotaulius 110-112, 125-126, 133

control, cutworms 430-43 1

Cook, W.C. 41 1,41 5, 418, 42 1-431

Cooley, R. A. 424

Copidosoma bakeri 428

Coquillet, D. W. 344

Corbet, P. S. 65

Comus 353

C. canadensis 354, 385

C. stolonifera 354, 385

Co tula 256

Craig, D. A. 282

Crastemyza 3 1 7

Crepis 383-384

C. gracilis 381,386

C. tectorum 366, 386

Criddle, N. 108, 118, 154, 156

Crumb, S.E. 417, 429

Cucumis 267

Curtonotus 248

cutworms

pale western 41 1-436

predators 237

Vll

red-backed 237

Cymindis cribricollis 240

C. planipennis 240

Dahlia 272

Darlington, P. J., Jr. 281

Davich, T. B. 256

Davis, M. B. (see Betten et al. )

Davis, P.H. 265,383

Dejean, P. F. M. A. 281

Delphinium 267 , 363, 384-385

Dendromyza 3 1 6

Denning, D. G. 21-35, 53, 57, 61-62, 67, 73,

76, 79, 99, 103, 108, 1 10, 1 14, 1 19-122,

128, 131, 140-143, 149, 152-155, 196,

201,207-208

Depner, K. R. 419

DePew, L. J. 431

Deschampsia caespitosa 332, 387

Desmotaulius 127-128

Dethier, V. G. 276, 356, 427

Dicosmoecinae 5, 50-64

Dicosmoecus 5, 50-53, 206, 216

D. atripes 51-53, 162, 184, 203, 206

D. jucundus 51-53, 159, 162, 184, 203,

206,213

Di Jeso, F. 437

Diptera 255-280, 291-405, 407-408, 428

Disease, cutworms 427-429

Dizygomyza 292, 316-321

Dodds, G. S. 19-20, 23, 26-27, 61-62, 76,

90, 100, 124, 128, 138-140, 143

Dohler, W. 146

drag-flight 253-254

Dugesiella hentzi 437

Eames, A. J. 265, 383

Ecclisomyia 5, 50-51, 61-64, 207, 217

E. bilera 207

E. conspersa 61-63, 164, 185, 203, 207, 213

E. digitata 207

E. kamtshatica 207

E. maculosa 61-62, 159, 164, 185, 203, 207,

214

E. scylla 207

E. species one 61-64, 164, 185, 203, 207

Ecclisopteryx 149

Echinus esculentus 437

Edwards, D. K. 425

Ehrharta 323

Elkins, W. A. 119, 145

Elodi, P.437,444

Elson, J. A. 198-199

Elymus 323

Enderlein, G. 309,312,342

Ennor, A. H. 437-438, 444-445

Enoecyla 150

Enoicyla 145, 149-150

Erigeron 425

Ernestia radicum 428

Erwin, T. L. 281

Escherichia coli 437

Eshbaugh, F. P. 415

Essig, E. O. 23-27, 33, 53-54, 57, 61-62, 89,

97, 100-102, 105, 110, 121, 137-138, 141-

145, 149

Etnier, D. A. 92, 95-101, 104, 107-1 12, 1 19-

BO, 133, 145-146

Eupagurus bernhardus 445

Eupatorium 277-278, 333

Euxoa ochrogaster 237

Eyer, J. R. 415, 421, 426-427, 431

Fabricius, J. C. 149

Fallen, C. F. 296, 302, 350, 375

Farstad, C. W. 419-421

Faulkner, L. R. 431

Feeney, P. P. 256, 278

Fernald, M. L. 198

Fink, S. C. 438

Fischer, F. C. J. 1 1, 15, 19,22-23,52-156

flight mill 253-254

flight ranges 253-254

flight speeds 253-254

Flint, O. S., Jr. 23, 33, 52-54, 62, 65-67, 73,

76, 95, 99-101, 115, 120, 124-125, 128-

BO, 134, 140-142, 145-146, 149, 154,

196-199

forecasting, cutworms 430

Forsslund, K. H. 97, 102, 104

Fowler, K. S. 437

Fraenkel, G. S. 256, 276

Frank, J. H. 237-252

Frey, R. 327-328, 344

Frick, K. E. 265, 291-294, 298-299, 302,

309, 312-313, 316-328, 331-335, 343-347,

355, 364-365, 369, 375, 380, 383-384

Friend, W. G. 256

Friganea 1 1 9

Frison, T. H. 196

Fristrup, B. 97

Vlll

Frost, S. W. 298, 328, 331, 354, 366, 380

Galeopsis 277-278, 333

G. tetrahit 267, 272, 274, 278

Galium boreale 343-344, 386

Gammack, D. B. 444

Geum allepicum 296

Gibson, A. 415, 421

Giddings, J. L. 197

Giraud, P. 309

Glossosomatidae 16

Glyphopsyche 5, 148-156, 21 1-212, 216

G. bryanti 149

G. inter cissa 149

G. irrorata 149-150, 160, 181, 194, 204,

211-212

G. missouri 211-212

Glyphotaelius 124, 149

Gnaphosa sp. nr. muscorum 250

G. parvula 250

Gonia aldrichi 428

G. breviforceps 428

G. brevipulli 428

G. capitata 428

G. longi for ceps 428

G. longipulli 428

Goniotaulius 65, 91, 102, 1 19, 125, 127,

149

Goodrich, A. L. 53

Grammotaulius 5, 83, 122-123, 140-141,

210,217

G. betteni 210

G. interregationis 122-123, 176, 191, 204,

210,214, 221

G. lorettae 210

G. praecox 122-123

G. sibiricus 123

Grant, D. R. 197-199

Gravenor, C. P. 197-198

Gray, H. E. 425,427

Greene, C. G. 316

Greenslade, P. J. M. 250

Griffiths, D. E. 437-438, 444-445

Griffiths, G. C. D. 261, 277, 297-298, 361,

364, 370, 375,380-381,383

Grindelia squarrose 425

Groschke, F. 319

Guppy, R. 33-34, 62, 95, 99-103, 107, 110,

121, 137, 140, 143, 149, 155

Gupta, P. D. 278

Gupta, R. L. 256

Gurney, B. 277

Gutierrezia 425

Haematopota 407-408

Hagen, H. A. 53, 67, 91, 95, 99-102, 109,

112, 118-119, 122-129, 137, 145-146,

149-150

Halescus 121, 146

Handlirsch, A. 95

Hansen, F. E. 276

Haplomyza 292, 295, 344

H. t ogata 344

Hardwick, D. F. 419, 426-427

Hardy, J. 380

Harmston, F. C. (see Knowlton).

Harpalellus basilaris 240

Harpalus 238

H. amputatus 240, 242, 249

H. desertus 240

H. funerarius 240

H. pleuriticus 240, 242, 249

H. uteanus 240

Harris, P. 256

Harris, T. W. 101, 145

Harvey, T. L. 431

Hawn, E. J. 429

Helianthus 266, 274-277, 333, 425

H. annuus 258, 266-269, 272

Heliocidaris erythrogamma 437

Heliothis zea 276

Heming, B. S. 284-286

Hendel, F. 294, 300, 303, 309-328, 333, 343-

345,360, 364, 369, 375,379

Hennig, W. 383

Henrickson, K. L. 97

Heracleum 35 1 , 367

H. lanatum 379, 386

Hering, E. M. 256, 261, 276, 31 1, 335, 340,

346, 382-383

Hesperophylax 5, 139-142, 211,216

H. alaskensis 140

H. consimilis 140-141, 180, 193, 203, 211

H. designatus 140, 21 1

H. incisus 140-142, 180, 193, 203, 211, 214

H. magnus 21 1

H. minutus 21 1

H. occidentalis 140-141, 179, 193, 203, 211,

214

Hexomyza 292-295, 309, 384

IX

H. albicula 309

H. schineri 309, 385

Heywood, H. B. 196

Heywood, V. H. 265, 383

Heydemann, B. 250

Hisaw, F. L. (see Dodds).

Hocking, B. 235-236, 253, 41 1-412, 419-

420, 427

Hoerner, J. L. 431

Holothuria forskali 437, 445

Homarus americanus 437

H. vulgaris 437, 444

Homophylax 5, 50, 78-81, 207-208, 216,

406

H. acutus 78-80, 160, 167, 186, 203, 208,

214

H. adriana 208

H. andax 208

H. baldur 78-81, 160, 167, 186,203, 208,

214

H. crotchi 78-79, 160, 167, 186, 203, 208,

214

H. flavipennis 208

H. insulas 208

H. nevadensis 208

H. rentzi 208

honeybees 437-446

Hope, F.W. 281

Hopewell, W. 43 1

Hopkins, A. D. 276

Hopkins, D. M. 197

Hordeum 323

H. vulgare 268, 272, 274

host-plant relationships 255-280, 382-387

Hsiao, T. H. 256, 276

Hubenthal, W. 281

Hussey, N. W. 277

Hutchinson, J. 265, 383

Hybomitra 407-408

Hyland, K., Jr. 146

Hymenoptera 428, 437-446

Hypnotranus 57, 107

Ichneumonidae 428

Icteromyza 292, 317-318, 323-324

Ilex 378

Imania 5, 50-51, 56-61, 207, 217

I. acanthis 207

I. bifosa 51, 60-61, 159, 163, 184, 203,

207,214

I. bifosa group 60-61

I. cascadis 57-59, 159, 163, 184, 203, 207,

214

I. cidoibes 207

I. gnathos 207

I. hector 57-60, 159, 163, 184, 203, 207, 214

I. renoa 207

/. sty lata 57

/. tripunctata 57-58, 159, 163, 184, 201-203,

207,214, 220

I. tripunctata group 56-60

information storage and retrieval 283-284

Jacobson, G. C. 102, 122

Jacobson, L. A. 253, 41 1, 414-436

Jasus verreauxi 437, 444

Jeannel, R. 281

Jermy, T. 256, 276, 278

Johnson, C. W. 102, 104, 119, 125, 146

Juncus 323, 324

Kaltenbach, J. H. 302, 333, 370

Kaplan, N. O. 437, 444

Karl, O. 320

Kassab, R. 437, 444

Kasting, R. 422, 427, 429

Keller, J. C. 256

Kennedy, J. S. 256

Khan, M. R. 422

Kimmins, D. E. 16, 67

Kirby, W. 102

Kjellgren, B. L. (see Betten et al.).

Klotz, C. 444

Knowlton, G. F. 34, 53-54, 99, 140-143, 154

Kolbe, H. J. 66, 102, 112, 119, 122

Kolenati, F. 102

Kowarz, F. 380

Krafka, J. 126

Krafka, J., Jr. 95, 126, 128

Krivada, W. V. 150

Kruger, J. P. 252

Lactuca 218

L. scariola 366

Lamp sana 333

Landon, M. F. 444

Lathy rus 334

L. odoratus 267 , 271

L. ochroleucus 334, 380, 385

Layne, E. 438

leaf miners, biology 255-280

host-plant relationships 255-280, 382-387

X

taxonomy 291-405

Leech, R. L. 198

Leimnephila 1 1 9

Lemurimyza 292, 295, 341-342

L. dorsata 341

L. pacifica 341

L. pallida 341-342

Lenar chulus 134

Lenarchus 5, 83, 134-139, 211,217

L. brevipennis 135, 138-139, 179, 193, 203,

211,214

L. crassus 135-136, 178, 193,204,211,214

L. expansus 2 1 1

L.fautini 135-137, 179, 193,203,211,213

L. gravidus 2 1 1

L. intermedins 137

L. rho 211

L. rillus 2 1 1

L. vastus 135-138, 179, 193, 203, 211, 213

Leonard, F. A. (see J. W. Leonard).

Leonard, J. W. 23,95-101, 104, 109, 119,

124-127, 145-146, 149, 154

Lepidoptera 237, 253-254, 41 1-436

Lepneva, S. G. 154

Leptacinus batychrus 250

Leptinotarsa decimlineata 276

Leptocella 14

Leptoceridae 14

Lewis, S. K. 437

life history, cutworms 416-427

Likens, G. E. (see McConnochie).

Limnephila 1 1 9

Limnephilidae 3-234, 406

species one 157, 183, 195, 203

species two 157, 183, 195,203

species three 157-158, 161, 183, 195,203

species four 158, 161, 183, 195,203

species five 158, 183, 195,203

Limnephilinae 5, 50, 81-158

Limnephilini 5, 82-142

Limnephilus 5, 13, 67, 83-122, 125-141,

149-150, 208-210,217

L. acnestus 209

L. acrocurvus 2 1 0

L. acula 210

L. ademus 209

L. affinis 1 1 8

L. alberta 86-88, 1 14, 174, 190, 203, 210,

214

L. alberta group 1 14, 210, 217

L. americanus 89

L. argenteus 85,88, 113-114, 174, 190, 204,

210.213

L. argenteus group 1 13-1 14, 210, 217

L. arreto 2 1 0

L. asiaticus group 115, 209, 217

L. assimilis 2 1 0

L. assimilis group 107-109, 210, 217

L. canadensis 85-87, 120, 175, 191, 204, 209,

213

L. castor 209

L. cockerelli 86, 105-106, 171, 189, 203,

209. 214

L. coloradensis 209

L. congener 99

L. decepta 101

L. dispar 209

L. diversus 209

L. diversus group 120, 209-210, 217

L. elegans 97

L. elongatus 208

L. externus 84, 87, 99-100, 170, 188, 203,

208.213.221

L. externus group 99-100, 208-209, 217

L. extractus 85, 88, 91, 168, 186, 204, 208,

213

L. exulans 98

L. fagus 209

L.femoralis 85,87, 102-103, 171, 188,203,

209.213.221

L. fenestratus 209

L. fenestratus group 115-117, 209, 217

L. forcipatus 1 1 8

L. frijole 209

L. hageni 84, 88, 91-92, 168, 187, 204, 208,

214

L. hingstoni 104, 406

L. hyalinus 85, 88, 109-1 10, 173, 189, 203,

209. 213

L. incisus group 109-1 13, 209, 217

L. indivisus 84, 88, 95-96, 169, 187, 204,

208.213

L. infernalis 85, 87, 96-97, 169, 187, 203,

208,213

L. janus 85-86, 1 1 1-112, 173, 189, 204, 209,

213

L. kennicotti 85, 88, 1 16-117, 175, 190, 204,

209,213,221

XI

L. labus 84-86, 115, 174, 190, 204, 209,

213

L. lopho 86-88, 107, 172, 189, 203, 209,

214

L. lunonus 209

L. luridus 209

L. luridus group 101-103, 209, 217

L. luteolus 99-100

L. macgillivrayi 89

L. merinthus 112

L. minusculus 85-86, 111, 116, 174, 190,

203,213,209

L. miyadii 98

L. moestus 85, 88, 104-105, 171, 188, 204,

209,213,406

L. morrisoni 209

L. morrisoni group 209, 217

L. nebulosus 102

L. nigriceps 85,88, 117-118, 161, 175,

190, 204,210,214

L. nigriceps group 1 1 7- 1 1 8, 2 1 0, 2 1 7

L. nogus 86-87, 103-104, 171, 188, 203,

209, 213

L. nogus group 103-104, 209, 217

L. notatus 98

L. occidentalis 210

L. omatus 85-87, 97-98, 170, 187, 203,

208.213

L. omatus group 97-98, 208, 217

L. oslari 100

L. pallens 209

L. pallida 108

L. partitus 85-87, 92-93, 161, 168, 187,

204, 208,214

L. parvulus 86-88, 107-108, 172, 189, 204,

210.213

L. perforatus 102

L. perpusillus 84, 87, 112-113, 173, 190,

203.209.213

L. picturatus 85-87, 98-99, 170, 188,

204, 208,214, 221

L. picturatus group 98-99, 208, 217

L. pilosula 1 1 8

L. productus 210

L. pulchellus 406

L. rhaeus 1 1 2

L. rhombicus 85-87, 118-120, 175, 191,

203,208,214, 221

L. rhombicus group 1 18-120, 208, 217

L. roberti 108

L. sansoni 84, 87, 90-91, 168, 186, 203, 208,

214

L. secludens 85,88, 110-111, 173, 189,204,

209,213

L. sericeus 84, 86, 100-101, 170, 188, 204,

209,213

L. sericeus group 100-101, 209, 217

L. sitchensis group 104-107, 209, 217

L. spinatus 85-87, 108-109, 172, 189, 203,

210, 214

L. stigma group 95-97, 208, 217

L. stipatus 102

L. striola 1 1 8

L. subcentralis group 88-95, 208, 217

L. subguttatus 95

L. sublunatus 84, 87, 89-90, 95, 161, 167,

186, 203,208,214

L. subpunctatus 102

L. susana 84, 87, 93-94, 169, 187, 203, 209,

214

L. sylviae 209

L. taloga 210

L. tarsalis 209

L. tersus 100

L. thorns 208

L. valhalla 85, 88, 106, 172, 189, 203, 209,

214

L. sp. one 88, 94-95, 169, 187, 203

Limnophila 1 1 9

Limnophilus 89-92, 95-122, 134, 137, 141

Limulus polyphemus 437

Lindroth, C. H. 54, 62, 97, 102, 1 13, 1 16,

124, 137, 143, 198-200, 238, 241-251

Lindsay, I. S. 418,427,431

Ling, S. 30-31,34

Linnaeus, C. 1 18

Lioy, Da Paolo 3 1 6

Lipke, H. 256

Liriomyza 292-295, 328-341, 384

L. arcticola 339

L. balcanica 330

L. balcanicoides 328-333, 396

L. baptisiae 329-333

L. bifurcata 328-337, 397

L. cannabis 277

L. conspicua 329, 332

L. cordiUerana 329, 332, 387

L. eboni 329-333

Xll

L. edmontonensis 329, 333, 339

L. eupatorii 277 , 329, 333, 397

L. felti 294

L. fricki 330, 333-334, 384-385

L. kenti 330-331, 334-337

L. lathyri 328-330, 334-335, 340-341,

385, 397

L. lima 329, 335

L. melampyga 341

L. millefolii 329, 335-338, 384-387

L. montana 329-, 333, 336

L. munda 333

L. nordica 330-331, 334-337

L. pictella 333

L. pilosa 329, 336

L. pusilla 261

L. quadrisetosa 33 1

L. senecionivora 328-331, 334-337, 387, 398

L. septentrionalis 329, 337-338

L. singula 295, 329, 338

L. sinuata 328-329, 335, 338-339, 398

L. smilacinae 330, 339, 340, 387

L. socialis 329, 339

L. sylvatica 328-329, 339-340, 398

L. taraxaci 328-330, 334, 340-341

L. trifolii 333-334, 340-341

L. undulata 330, 340

L. veluta 328-330, 334, 340-341

L. viciae 329, 341,385

Lloyd, J. T. 95, 1 19, 124-126, 146

Loew,H. 319, 324

Lohman, K. 437

Lonicera 384

L. dioica 346-349, 386

L. involucrata 346-347, 365, 373, 386

L. tartarica 346, 386

Loricera pilicornis 239

Loveland, R. P. 282

Lundbeck, W. 344

Lupinus 267, 271, 273

L. sericeus 367, 385

Lycopersicon esculentum 267

Macquart, J. 389

Macrotaulius 83, 124

Maia squinado 437, 445

Maianthemum

Malloch, J. R. 298-300, 312, 322, 344-347,

368

Malva sy Ivestris 278

Manduca sexta 276

March, R. B. 429

Marcus, F. 437

Martin, J. E. H. 196

Martynov, A. V. 53, 57, 125-128, 149

Matricaria 256, 266, 276

M. chamomilla 263

M. matricarioides 256-258, 262, 266-269,

272, 369, 387

Matsuda, R. 284-286

Matsumoto, Y. 256

Mayr, E. 294

McAlpine, J. F. M. 196

McConnochie, K. 95-97, 104, 125, 146, 154

McDonald, H. 431

McDonald, S. 431

McDunnough, J. H. 416

McGinnis, A. J. 422, 427

McLachlan, R. 53, 67-69, 91-92, 95, 98-99,

102, 109, 112, 119, 125, 128, 145-146,

149-150

McMillan, E. 420, 427-430

Medicago 333

Meigen, J. W. 297, 302, 312-314, 321-322,

328,345-347,354,375

Meijere, J. C. H. de 263, 373

Meinert, F. 102

Melanagromyza 292-294, 303-309, 384

M. achilleana 303-306, 386, 393

M. actaeae 303-306, 384-385, 393-394

M. bidenticola 303-307, 386, 394

M. fastosa 304, 307-308

M. laetifica 304, 307-308

M. martini 303-305, 308, 385

M. matricarioides 304

M. miranda 305

M. occidentalis 304, 308

M. setifrons 303-304, 308

M. shewelli 304, 309

Melander, A. L. 308, 312, 324, 335, 343-344

Melanoplus bruneri 437

Melitotus 333

M. officinalis 21 1

Mentha arvensis 325, 386

Merkley, D. R. 89-92, 95-121, 125-138

Mertensia 353

M. paniculata 299, 370, 386

Me table tus americanus 240-242

Metaphyto genalis 428

xm

Meteorus leviventris 428

M, vulgaris 428

Metopomyza 292, 295, 342-343

M. flavonotata 342

M. griffithsi 342-343, 399

M. interfrontalis 342-343

Mez, C. 383

Micaria sp. nr. alberta 250

Michel, C. E. 95

Michelbacher, A. E. 256

Mik, J. 328

mill drag 253-254

Miller, L. A. 407

Milliron, H. E. 95

Milne, D. J. 96, 116-120, 127, 130, 146

Milne, L. J. 19, 22-28, 31-35, 53-54, 57, 61-

62, 67-68, 76, 79, 89-92, 95-113, 116-

146, 196

Milne, M. J. 31

Mitchell, B. 241

Monarda 325

Morales, M. F. 439

Moreland, B. 437, 444

Morrison, H. K. 416, 424

Morrison, J. F. 437-438, 444-445

Morse, W.J. 23,91,95-97, 101, 104, 107-

109, 124-127, 130, 137, 145-146, 149,

154-156

Mosely, M. E. 16, 89-92, 95-97, 102, 112,

119, 122-125, 128, 145-146, 149-150

Moyhuddin, A. I. 279

Munroe, E. G. 197-198

Musca 238

Muttkowski, R. A. 26, 95, 1 19, 126, 145-

146

Myxicola infundibulum 445

Nakahara, W. 97

Napomyza 292, 295, 349-350

N. immanis 349

N. lateralis 349

N. nugax 349-350

N. plumea 349-350

natural enemies, cutworms 427-429

Nayar, J. K. 256

Neave, F. 62, 76, 97-99, 108-109, 128,

140, 156

Needham, J. G. 95, 146, 196

Nemorimyza 292, 295, 327-328, 384

N. posticata 295, 328, 387

Nemotaulius 5, 83, 123-124, 210, 217

N. hostilis 124, 161, 176, 191, 203, 210, 213

Neophylacinae 5, 50, 71-77

Neothremma 5, 71-72, 75-77 , 207, 216

N. alicia 76-77, 160, 166, 186, 203, 207, 214

N. didactyla 207

N. galena 207

N. laloukesi 11, 183, 186, 203, 207, 214, 406

Nephrolepis 267, 270, 271

Nephrops norvegicus 437

Neuronia 145

Nicotiana tabacum 261, 271, 273

Nihei, T. 439

Nimmo, A. P. 3-234, 406

Noctuidae 237, 253-254, 41 1-436

Noda, L. 439

Nothofagus 200

Notiophilus aquaticus 239

N. semis triatus 239

Nowakowski, J. T. 256, 261. 276-277, 291,

302, 317-328, 343-346, 375, 382-383

Oatman, E. R. 256

Odum, E. P. 25 1

Oligophlebodes 5, 71-75, 207, 216

O. ardis 207

O. minuta 207

O. mostbento 207

O. ruthae 72-73, 160, 166, 185, 203, 207,

214

O. sierra 12-1 A, 160, 166, 186, 203, 207, 213

O. sigma 207

O. zelti 72-75, 166, 186, 203, 207, 214

Omophron americanum 25 1

Onocosmoecus 5, 51-55, 206-207 , 217

O. quadrinotatus 206

O. unicolor 54-55, 162, 184, 204, 206, 213,

221

Ophiomyia 292, 295, 310-316, 384, 385

O. asterivora 3 1 2

O. banffensis 3 1 0-3 1 2

O. decima 3 1 0-3 11,314

O. labiatarum 311-312, 395

O. madizina 3 1 2

O. maura 31 1-312, 316, 387, 395

O. monticola 3 1 0-3 1 2

O. nasuta 310-312

O. nona 311-313

O. pinguis 3 1 2

O. praecisa 310,313-315

XIV

O. prima 31 1-313, 316

O. pulicaria 310, 314, 395

O. pulicaria group 295

O. pulicarioides 310, 314

O. secunda 310, 314

O. septima 311,314

O. sexta 311, 314-315

0. stricklandi 310, 315, 395

O. undecima 311, 31 5-3 1 6

O. wabamunensis 311,316

O. youngi 312

Orcutt, A. W. (see Betten et al.).

Oriol, C. 444

Oxytropis camp estris gracilis 372-373

O. splendens 372, 385

Packard, A. S. 102

Pagurus bemhardus 437

Palaemon serratus 437

Paniscus 428

Papaver 267

Parachiona 107, 143-144

Parahidippus marginatus 250

Paralenarchus 83, 135-139,211,217

Parapatania 67

Parapheropsophus 281

Paraphytomyza 292, 295, 345-349, 384,

385

P. flavocingulata 346

P. lonicerae 346, 386

P. luteoscutellata 348

P. nitida 346-347

P. orbitalis 346-348, 386, 399

P plagiata 346-348, 386, 399

P. spenceri 346-349, 386, 399

P. tremulae 346

parasites, cutworm 428

Pardo sa groenlandica 250

P. sp. nr. metlakatla 250

P. moesta 250

P. sp. nr. saxatilis 250

Parker, J. R. 415, 418-431

Parks, T. H. 256

Patrobus lecontei 239

Pecten maximus 437 , 445

Peleteria anaxia 428

P. haemorrhoa 428

P. robusta 428

Pennak, R. W. 119, 124-127

Penstemon 352

P. confer tus 373, 386

P, procerus 373, 386

Periplaneta 238

Petasites sagittatus 373, 387

Peterson, L. K. 421

Pewe, T. L. 197

Pfadt, R.E.431

Phalaris 322

P. arundinacea 387

Phanocelia 5, 148, 155-156, 212, 216

P canadensis 156, 160, 181, 194, 204, 212,

214

Pheropsophidius 281

Pheropsophus 281

P. complanatus 28 1

Philarctus 5, 83, 132-133, 211, 217

P quaeris 133, 178, 192, 204,211,213

Philocasca 5, 143, 147-148,211,216

P. thor 147-148, 161, 181, 194, 203,211,

214

Philonthus concinnus 250

P. furvus 250

P. occidentals 250

Phleum 322

Pholcus phalangioides 437

photomicrography 282

Phryganea 65-67, 98, 101, ,1 17-1 19, 122, 145,

149

Phytagromyza 346-347

Phytobia 292, 295, 316-317

P. amelanchieris 316-317, 384

P. confessa 295,31 6-3 1 7

P. flavohumeralis 3 1 6-3 1 7

Phytoliriomyza 292, 344-345

P. arctica 344-345

P. formosensis 344

P. immaculata 344

Phytomyza 292, 295, 350-382, 384

P. affinalis 256, 359, 365

P. agromyzina 353, 354, 385

P. albipes 375

P. aquilegiae 355

P. aquilegiana 352-355, 358, 378, 385

P. aquilegioides 351-352, 355-356, 358, 378,

385,400

P. aquilegiophaga 352, 356, 359, 367, 385

P. aquilegivora 353, 356-357, 381, 385

P. aralivora 353, 357, 385

P. arnicivora 352, 357-358, 386, 400

XV

P. asterophaga 351, 355, 358, 360, 379,

386

P. atricomis 380, 383, 263, 277

P. atripalpis 379

P. banffensis 352, 355, 358

P. blairmorensis 352, 356-359, 365, 367,

400

P. canadensis 353, 359-360, 371, 385

P caprifoliae 354, 360, 364, 373, 375, 386

P. chrysanthemi 263, 380

P ciliolati 351, 358, 360, 386

P. cineracea 353, 360-361

P. clematiphaga 350, 361, 368, 375, 385

P. colemanensis 352, 361-362, 373-374,

401

P columbinae 351, 362-363, 385, 401

P. crassiseta 386

P. delphinivora 352, 363, 385

P. edmontonensis 353, 363-364, 401

P. erigerontophaga 360

P. evanescens 354, 364

P. flavicornis 350, 368

P. flavoscutellata 375

P. fuscula 352, 357, 364, 379

P. gelida 379

P gregaria 354, 364-365, 370, 386, 401

P. hordeola 277 , 381

P. ilicis 378

P. illustris 351, 356, 359, 365

P. intermedia 370

P. involucratae 353, 365, 370

P. jasperensis 353, 365-366, 371, 380, 402

P. lactuca 351, 366, 386-387, 402

P lanad 261, 351, 367

P. lupini 352, 359, 367, 385

P lupinivora 353, 367, 372, 385

P. luteiceps 350, 351, 368, 371, 385, 402

P. major 350, 361, 368-369, 403

P matricariae 255-280,. 35 1, 369, 373, 378-

379, 384-387

biology 259-262

immature states 263-265

host-plant relationships, adult females

265-269, 276-277

host-plant relationships, larvae 270-275,

277-278

P. mertensiae 353, 364, 369-370, 374, 386,

403

P. merula 354, 370

P milii 353, 370, 403

P. minuscula 356, 381

P. miranda 351, 370-371

P misella 352, 371, 404

P. modica 363

P. muldfidae 353, 371-372, 385, 404

P. nepetae 369

P. notopleuralis 354

P oxytropodis 354, 367, 372-373, 385, 404

P. pallipes 363

P. pedicularicaulis 365-366, 371

P. penstemonis 352, 361, 373-374, 386

P. periclymeni 354, 364-365, 373-375, 386

P. petasiti 351, 373-374, 387

P plantaginis 352, 361, 373-374, 386

P prava 351, 353, 364, 369, 374, 385

P. puccinelliae 364

P. queribunda 354, 364, 375

P. ranunculi 350, 361, 368, 375, 385

P. riparia 35 1 , 376, 404

P. rufipes 368

P. sehgali 353, 369, 376-377

P. senecionella 352, 377, 387, 405

P. solidaginivora 351, 378

P. solidaginophaga 352, 378-379, 387, 405

P spondylii 351, 367, 369, 373, 376, 379,

386

P. subalpina 352, 379-380, 405

P subtenella 352, 365, 371, 380

P subdlis 352, 380-381,385

P. syngenesiae 277, 352, 374, 377, 380-383,

386-387

P. syngenesiae group 263

P. thalic tricola 356

P. thalictrivora 353, 356, 381, 385

P timida 352, 363, 381, 405

P. urbana 352, 380-382

P. vibeana 375

Pieris brassicae 256

P. rapae 256

Pisum sativum 267 , 271-278

Plantago major 374, 386

Pladphilax 145

Platyphylax 140, 145, 149-150, 153

Plutella maculipennis 218

Poa 320

Poecilanthrax sackenii 428

P. willistoni 428

Poemyza 292, 317-318, 321-323

XVI

Polycelis cornuta 445

Populus 302, 309

P. tremuloides 346, 385

P. tremulus 346

Porosagrotis del ora ta 416

P. orthogonia 411,416

P. orthogonia duae 4 1 6

Porthetria dispar 437

Por tunas depur a tor 437

Potamobius astacus 437, 444

P. leptodactylus 437

Potentilla 267, 271, 296, 385

Pradel, L. A. 437, 444

Praspedomyza 292, 295, 343-344

P. galiivora 343-344, 386

Prest, V. K. 197-199

Proctor, W. 95-97, 104, 119, 125-126, 146

Protopheropsophus 281

Provancher, L. 89, 95, 102, 145, 149

Pruess, K. P. 415

Pseudomeriania nigrocornea 428

Pseudomonas aeruginosa 429

Pseudonapomyza 292, 295, 345

P. atra 345

P. lacteipennis 345

Pseudostenophylacinae 5, 50, 77-81

Psychoglypha 5, 148, 151-155, 212, 216

P alaskensis 152-155, 182, 195, 204, 212-

213

P. avigo 212

P bella 212

P. ormiae 212

P prita 152-153, 182, 194, 203, 212, 214,

406

P. rossi 2 1 2

P schmidi 152-153, 182, 195, 203, 212,

214

P. subboreale 154

P ulla 152, 155, 182, 195,203,212,214

Pteridomyza 294

Pterostichus adstrictus 239, 242, 245-247

P. corvus 239, 245

P. femoralis 239

P. lucublandus 239, 242, 245

publishers, Canadian 235-236

Putman, J. D. 53

Pycnopsyche 5, 143-147,211,216

Pguttifer 145-147, 181, 194, 203,211-

213

P. guttifer group 2 1 1

P. lepida group 21 1

P. similis 146

P subfasciata 145-146, 180, 193, 204, 211,

214

Quedius spelaeus 250

Radema 66-69

Rampton, U. N. 197-199

Ranunculus 360, 364, 384

R. abortivus 375, 385

R. acris 361

Rawson, D. S. 119, 146

rearing methods, cutworms 427

Regnouf, F. 437, 444

Rhyacophila 13, 16-48, 205-206, 215-216

R. acropedes 17, 19, 23-24, 38, 42, 47, 203,

205,214

R. acropedes group 23-24, 205, 216

R. alberta 17-20, 38, 41, 203, 205, 214

R. alberta group 19-22, 205, 216

R. amabilis 205

R. angelita 18-19, 33-34, 40, 44, 48, 204,

206,221

R. angelita group 33-34, 206, 216

R. anomala 26

R. belona 18, 25, 38, 42, 46, 203, 205, 213

R. betteni group 31-32, 206, 216

R. bifila 18, 26, 38, 42, 46, 203, 205, 213

R. bipartita 33

R. bruesi 3 1

R. chilsia 17, 32, 44, 47, 203, 206, 214

R. coloradensis 18, 26, 38, 42, 47, 203, 205,

213

R. complicata 3 1

R. doddsi 30

R. glaciera 16, 19-22, 38, 41, 46, 203, 205,

214

R. harmstoni 205

R. hyalinata 17-18, 27, 39, 42, 47, 203, 205,

214

R. hyalinata group 27, 205, 216

R. insularis 205

R. invaria group 24-26, 205, 216

R. iranda 206

R. kemada 205

R. kincaidi 205

R. milnei 17, 36-37, 40, 45, 48, 203, 206,

213

R. oregonensis 34

XVII

R. pellisa 17-19, 30-31, 39, 43, 47, 203,

206, 214

R. perplana 206

R. rickeri 17-19, 28, 39, 43, 47, 203, 206,

214

R. sibirica group 28-31, 205-206, 216

R. sonoma 205

R. stigmatica 26

R. tucula 17-21, 38, 41, *46, 203, 205, 213

R. vaccua 17-18, 31-32, 39, 44, 48, 203,

206.213

R. vagrita 17-19, 35-36, 40, 45, 48, 203,

206, 213

R. vagrita group 35-37, 206, 216

R. vao 205

R. vemna 17-19, 24, 40, 45, 48, 203, 205,

406

R. vepulsa 17-19, 28-29, 39, 43, 47, 203,

206.214

R. verrula 17-19, 34-35, 40, 45, 48, 203,

206.213

R. verrula group 34-35, 206, 216

R. vobara 17-19, 32-33, 40, 44, 47, 203,

206.214

R. vobara group 32-33, 206, 216

R. vocala 205

R. vofixa 17-18, 22, 38, 41, 46, 203, 205,

213

R. vofixa group 22, 205, 216

R. vuzana 206

R. sp. one 18, 37, 40, 45, 48, 203

R. sp. two 19, 37, 40, 45, 48, 203

Rhyacophilidae 3-234, 406

Rhyacophilinae 1 6

Ricker, W. E. 95, 128, 146, 196

Rivard, I. 244-247,250-251

Robert, A. 89, 95-97, 101, 104, 109, 1 19,

124, 127, 134, 146, 149

Robin, Y. 437, 444

Robineau-Desvoidy, J. -B. 346, 374, 379

Rock, P. J. G. 421

Rondani,C. 317, 324

Rosa acicularis 296

' Roselle, R. 415

Ross, H. H. 1-2, 4, 12-16, 19-36, 52-62, 66-

69, 72-73, 76, 79, 89-92, 95-146, 149-

156, 196-198, 201, 205-207, 220-222

Rotramel, G. L. 196

Rowe, J. S. 220-221

Rubiomyza 345

Sabella pavonina 444-445

Salix 309

Salsola pestifer 425

Salt, R. W. 421

Sasakawa, M. 296-298, 303, 312, 316, 375

Saville, D. B. O. 198

Say, T. 100-101, 145

Schaaf, A. C. 427

Schaupp, F. G. 245

Schi<^dte, J. C. 247

Schmid, F. 4, 12-15, 19-36, 49, 52-69, 72-73,

76-79, 83, 89-92, 95-146, 149-156, 206-

211

Schoonhoven, L. M. 256

Schrank, F. von P. 375

Scirpus 321, 387

Seamans, H.L. 411,415-431

Sehgal, V. K. 255-280, 291-405

Senecio 381, 383, 387

S. conge stus pallustris 377, 387

S. pauciflorus 377, 387

S. vulgaris 268, 272

Shepherdia 327

S. canadensis 327 , 386

Shewed, G. E. 299, 302, 324, 344

Shizukoa 316

Sibley, C. K. 95-97, 119, 124-126, 134, 146

Silene noctiflora 271

Siltala, A. J. 95

Simpson, C. B. 95

Simpson, G. G. 14

Sipunculus nudus 437, 444

Siridomyza 3 1 2

Smilacina stellata 268, 339, 387

Smith, G. M. 95, 119, 126, 146

Smith, J.B. 145-146,416

Smith, S. D. 20-36, 55, 67-68, 73, 137, 152,

201, 211

Smulyan, M. T. 256, 263, 380

Snodgras, R. E. 145

Solanum tuberosum 267, 273

Solidago 295, 312, 325, 328, 333, 378, 386,

268,273,278,425

S. lepida 378-379, 387

Sonchus 326, 384, 425

S. arvensis 273

S. asper 345

S. uliginosus 268, 272, 275, 366, 387

XV 111

Stfndemp, H. P. S. 252

Sorenson, C. J. 415, 421-422, 425-429

Spencer, G. J. 53, 57, 60-62, 72-73, 79, 95,

99-103, 107-110, 124, 127, 131-133,

136-137, 140-143, 149, 155

Spencer, K. A. 256, 278, 291-294, 297-

350,354

spiders, Alberta 237, 250

spiders, arable fields 237, 250

Spirographis spallanzanii 444

Sprules, W. M. 125, 146

staphylinid beetles, Alberta 231 , 250

staphylinid beetles, arable fields 237 , 250

Stegmaier, C. E. 333-334

Steinhaus, E. A. 429

Stenaptinus 273

Stenolophus comma 240

Stenophylacini 4, 82, 142-148

Stenophylax 96, 107, 116, 121, 125, 138,

145-146

Stem tor coeruleus 437

Strand, A. L. 415, 418-431

Strickland, E. H. 291, 41 1, 419, 427-430

Stride, G. O. 256

Strobl, G. 303

Styella mammiculata 437

Sugiyama, S. 256

Sympetrum rubicundulum 437

Symphoricarpos 360, 384, 386

S. albus 346, 347, 386

S. occidentalis 349, 386

Synuchus impunctatus 239

Szorenyi, E. 437, 444

Tabanidae 407-408

Tachinidae 428

Tacky porus 250

Takhtajan, A. 265, 383

Tanacetum 256, 266, 270, 274-477

T. vulgare 256-261 , 266-275, 335, 369,

387

Taraxacum 326, 384, 387

T. officinale 268, 272, 313, 340, 366

Tatchell, E. C. 437

Tauber, C. A. 256, 258,261

Tauber, J. M. 256, 258,261

Teskey, H. J. 407

Tetrahymena pyriformis 437

Thalictrum 352, 384-385

T. venulosum 356, 362-363, 381, 384, 385

Thienemann, A. 67

Thlaspi arvense 267, 27 1

Thoai, N. V. 437, 444

Thomas, A. W. 407-408

Thomson, C. G. 133

thorax, evolution 284-286

thorax, morphology 284-286

Thornley, H. F. 415, 421-422, 425-429

Thorsteinson, A. J. 256, 278

Thut, R.N. 28, 31,34, 36

Tilden, J. S. 261

Tilman, H. W. 200

Trechus quadristriatus 241

Trehan, K. N. 280

Trichocellus cognatus 240, 242, 249

Trichoptera 1-234,406

Tri folium repens 334, 385

Triticum 323

T. aestivum 387

Trochosa terricola 250

Tropaeolum 267

Tylomyza 3 1 2

Typha latifolia 268, 272, 274

Uhr, M. L. 437

Ulmer, G. 19, 26-27, 53-54, 57, 67-69, 91, 95-

109, 112, 119-129, 137, 143-146, 149-150,

154

Ulmus americana 296-298, 385

Unzicker, J. D. 26,67,76, 145

Urtica 296, 302, 305, 308, 368

U. gracilis 385

Veronica 386

Verschaffelt, E. 256

Viala, B. 437

Vida americana 267 , 334, 341, 385

Vigna 334

Villa alternata 428

V. willis toni 428

Virden, R. 437-439, 444-445

Vorhies, C. T. 119, 145

Wagneria rohweri 428

Walkden, H. H. 416, 419-421, 427-429

Walker, E. M. 196

Walker, F. 67, 91-92, 95, 102, 1 12, 1 19, 122,

125-127, 145-146, 149-150

Wall, R. E. 430

Watts, D. C. 437-439, 444-445

Watts, R. C. 444

Webster, F. M. 256

Wendland, W. M. 199

Wenner, B. J.431

Westfall, M. J. 196

Westgate, J. A. 198-200

Westwood, J. O. 349

Whitehead, D. R. 283-284

Whittaker, R. H. 256, 278

Wiggins, G. B. 89, 97-99, 102-104, 109,

116, 119, 124, 133, 143, 146, 201, 211

Wray, D. L. 125, 145

J Wyman, M. 287-289

Xyraeomyza 344

Xysticus calif ornicus 250

Zea 323

Zele 428

Zetterstedt, J. W. 65-67, 98, 1 17, 122, 132,

323, 364

Zinnia 266, 268, 273, 275

Zizania 322

.

Quaestiones

MU 3. COMP. ZOO

LIBRARY

MAR 29 1971

HARVARD

UNIVERSITY

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

VOLUME VII

NUMBER 1

JANUARY 1971

QUAESTIONES ENTOMOLOGICAE

A periodical record of entomological investigation published at the Department of

Entomology, University of Alberta, Edmonton, Alberta.

Volume 7

Number 1

19 February 1971

CONTENTS

Guest Editorial — Northwestern Caddisflies 1

Nimmo — The adult Rhyacophilidae and Limnephilidae (Trichoptera) of Alberta

and Eastern British Columbia and their post-glacial origin 3

Guest Editorial — Northwestern Caddisflies

It is a good half century since the world famous Russian trichopterist, Dr. A. B. Marty-

nov, declared that the Trichoptera were an ideal group from the standpoint of obtaining

meaningful bio-geographic inferences. Dr. Martynov himself never followed up his historic

statement on the Trichoptera, but instead gradually became engrossed in the study of fossil

insects. His prophetic remark has, nevertheless, been borne out as group after group of the

caddisflies have been studied on a phylogenetic and bio-geographic basis.

As controversies have emerged in recent years concerning the past history of the conti-

nental masses, Dr. Martynov’s special field of caddisfly study, the biota of the far north, has

gradually become of increased importance in contributing information of unusual interest

concerning inter-continental dispersals. The northern caddisflies of Europe have been well

known for over a century, thanks to the pioneer work of Zetterstedt in Scandinavia and

McLachlan in England. These investigators assembled and studied material and information

on the northern fauna of their native lands and regions. Early in his career, Martynov him-

self published many papers making known the characteristics and distribution of the Tri-

choptera of Russia and especially the fauna of Siberia.

During this time, little was discovered concerning the Trichoptera of northern North

America. Barnston and Kennicott made the first extensive northern collections, the former

in the area immediately south of Hudson Bay, the latter in north central areas of Canada,

including Great Slave Lake. In more recent years, limnological investigators added many

valuable collections and lately entomologists have collected many caddisflies in the Govern-

ment of Canada’s massive northern insect survey.

Although taxonomic problems concerning many northern species of Trichoptera have

been elucidated by a variety of authors, one facet necessary to achieving a synthesis of the

Holarctic caddisfly fauna has been sorely lacking. This is a thorough study of the group for

northwestern North America. For several years such a study appeared to be in the offing

and was actually prepared by J. Jared Davis. But because of publication difficulties and

other obstacles this useful manuscript never graced the printed page.

The study of the Alberta caddisflies presented in this publication is, therefore, a timely

and invaluable contribution to our knowledge of the Trichoptera of the far north. It gives

us, first, a basis of discriminating identification for all the species of the region for the

families treated, and second, geographic and ecological parameters that will be helpful in

integrating these species with their relatives in other parts of the northern Holarctic range.

The excellent illustrations of Alberta specimens will be of inestimable value in subsequent

studies of intra-specific variation and its implications concerning post-Pleistocene coloniza-

tion of deglaciated areas.

In this wise, Dr. Nimmo’s study will become a keystone in a synthesis of the evolutionary

history of the northern biota.

Herbert H. Ross

Department of Entomology

University of Georgia

November 27, 1970 Athens, Georgia

THE ADULT RHYACOPHILIDAE AND LIMNEPHILIDAE

(TRICHOPTERA) OF ALBERTA AND EASTERN

BRITISH COLUMBIA AND THEIR POST-GLACIAL ORIGIN

ANDREW PEEBLES NIMMO

Hancock Museum Quaestiones entomologicae

New Castle-upon-Tyne, England 7 : 3-234 1971

Of the Rhyacophilidae 22 species and of the Limnephilidae 91 species are recorded here

from the area, making a total of 113 species. Each species is described, and keys are

provided for identification of adult specimens to species.

Seven species of Limnephilidae are described as new: Imania hector; Apatania alberta;

Homophylax baldur; Oligophlebodes zelti; Limnephilus susana; Limnephilus valhalla; and

Philocasca thor.

The post-glacial origin of this fauna is examined, taking into consideration the possible

effects of past and present climatic patterns, extent of glacial ice masses and locations of

possible refugia, and locations and drainage patterns of major glacial and post-glacial lakes.

Also examined are the 12 range patterns exhibited by the species, and the distributions of

each species relative to the other species in its genus or species group. The 12 range patterns

form two main groups: one group of six is restricted wholly to the western Cordillera of

North America; and the remaining six are more widely distributed, being largely trans-

continental in extent. Altitudinal distributions are also briefly examined.

The conclusions reached are that only 5% of the present fauna is derived postglacially

from the Beringian refugium, while 95% is derived from North America south of the

southern limit of glacial ice. Dividing the 95% portion further, 61%) is derived from the

western Cordillera of the United States, 8% from eastern North America, 7%> from the

central Great Plains, 18% from all of North America south of the ice, i.e. from trans-

continental species, and l%o is of uncertain derivation. (Traduction franqaise a la page 234).

CONTENTS

Materials p. 5

Methods p. 11

The Family Rhyacophilidae Stephens p. 16

The Family Limnephilidae Kolenati p. 49

Origins and relationships of the fauna p. 196

References p. 224

Purpose of the study

The primary objective of this study is to determine the composition of the fauna of two

families of Trichoptera, the Rhyacophilidae and Limnephilidae, in Alberta and eastern

British Columbia. Secondly, by an examination of species distributions and relationships

it is hoped to elucidate the post-glacial origins of the fauna.

While these are the major objectives of the study, there are some subsidiary benefits to

be derived from the results. The first is the additional knowledge of North American Tri-

choptera which accrues, as to distributions, correlation of the sexes in species in which the

female was previously unknown, and the discovery of new species. A second advantage of

such a compilation as this, on a regional fauna, is the facilitation of ecological and other

4

Nimmo

studies of the adult Trichoptera of the area. Identifications should be possible without

recourse to a scattered and difficult literature.

The third benefit is facilitation of studies of the immature stages of the species of the

two families in the study area. The immature stages of most species are unknown at present,

and it is hoped that the identification facilities supplied in this study will permit the

immatures to be correctly correlated and identified. Once this is done ecological studies on

the immature stages can be carried out.

The taxa studied

Originally I had hoped to examine all families of Trichoptera in the study area, but

the large number of species, estimated at close to 200, and limitations of time precluded

this. Consequently two families were decided upon: the Rhyacophilidae and Limnephili-

dae. A minimum of 113 species of these two families is recorded here from the study

area.

Apart from my intrinsic interest, these two families were selected for the following

reasons. The Rhyacophilidae are a distinctly mountain group of Trichoptera and should

thus prove useful in tracing faunal changes in the study area. The Limnephilidae occur in

both mountains and plains, with distinctive large faunas in each area, and should prove

useful in elucidating faunal changes in both areas.

The Rhyacophilidae are represented in the study area by 22 species of one genus, Rhya-

cophila. These species represent 1 1 species groups which, in the text, are presented in the

sequence of Ross (1956). The Limnephilidae are represented in the study area by 91 species.

These variously represent a total of five subfamilies, four tribes, and 26 genera. In this study

Schmid’s (1955) order of presentation is used. Table 1 presents the names and organisation

of the higher taxa of the Limnephilidae of the area.

The study area

Geographically the investigation embraces the Province of Alberta and the Rocky Moun-

tains of eastern British Columbia. The western limits in British Columbia comprise the

line from Kimberly to Golden, thence to Revelstoke, to Avola, and finally to the Mount

Robson area. By these limits the northern portion of the Selkirk Mountains is also included.

Some information is also included which was derived from a collection from the Simpson

Islands of Great Slave Lake, Northwest Territories. The lake is just over 50 miles north of

the northern boundary of Alberta and the area was, until recently, difficult of access. It is

felt that any faunal information on Trichoptera from the lake would be applicable to

northern Alberta and I took the opportunity in 1964 of arranging with Mr. D. J. Larson,

Lethbridge, Alberta, to collect adult Trichoptera for me while in the area.

Figure 1 illustrates the positions of the localities from which insects recorded here were

taken. Figures la and lb are enlargements of certain portions of Fig. 1 in which too many

localities are recorded for inclusion in that figure. In Fig. 1,1a, and lb, many localities are

listed as such-and-such a lake or river. This refers to the point at which the nearest road

touches on, or crosses these bodies of water.

The study is limited to the area outlined above for two main reasons. It is a convenient

delimited area located immediately across the mountain and plains routes between Alaska

and the remainder of North America, and can be expected to yield evidence of faunal

changes or dispersals due to glaciations or climatic changes. Also, embracing as it does both

mountain and plains regions (Fig. 2), and ranging from boreal forest in the north, through

aspen parkland, to near desert grassland in the south (Fig. 3), the area could be expected to

yield a large and most interesting fauna of Trichoptera.

Rhyacophilidae and Limnephilidae

5

Table 1 . The family Limnephilidae in Alberta and eastern British Columbia.

Subfamily Tribe

Dicosmoecinae —

Apataniinae Apataniini

Neophylacinae —

Genus

Dicosmoecus

Onocosmoecus

Imania

Amphicosmoecus

Ecclisomyia

Apatania

Oligophlebodes

Neothremma

Pseudostenophylacinae —

Limnephilinae Limnephilini

Stenophylacini

Chilostigmini

Homophylax

Limnephilus

Grammotaulius

Nemotaulius

Anatolia

Asynarchus

Clistoronia

Philarctus

Arctopora

Lenar chus

Hesperophylax

Chyranda

Pycnopsyche

Philo casca

Glyphopsyche

Chilostigmodes

Psychoglypha

Phanocelia

MATERIALS

Total number of specimens

I examined 7,604 specimens of both sexes: 2,915 specimens of Rhyacophilidae; and

4,689 specimens of Limnephilidae. The total number of specimens of each species exam-

ined, by numbers per sex, is given at the end of its description in the text.

Sources of material

Most of the above material was collected by me during the summers of 1965, 1966,

6

Nimmo

Fig. 1. Alberta showing major highways, secondary roads, and collecting points. See also Fig. la and lb.

Rhyacophilidae and Limnephilidae

7

Fig. la. Outline map of the Forestry Trunk Road between Hinton and Nordegg (left), and Nordegg and Cochrane

(right). • - collecting points

8

Nimmo

\

Fig. lb. Forestry Trunk Road and connecting roads between Waterton and Banff (left); Banff-Jasper Highway between

Banff and Jasper (right); • - collecting points.

Rhyacophilidae and Limnephilidae

9

Fig. 2. Major physical features of Alberta and eastern British Columbia, showing rivers, lakes, and three levels of altitude.

10

Nimmo

Fig. 3. Forest Regions of Alberta and

eastern British Columbia (adapted from Rowe, 1959).

Rhyacophilidae and Limnephilidae

11

1967, and 1968. Some specimens were obtained from the collections of the Department

of Entomology, University of Alberta. A very small proportion was obtained on loan from

the following institutions: Canadian National Collection, Ottawa; Royal Ontario Museum,

Toronto; Illinois Natural History Survey, Urbana, Illinois, United States; and the United

States National Museum, Washington, D. C., United States. The source of the borrowed

material is given with species descriptions wherever applicable. Some material was obtained

from graduate students in the Department of Zoology, University of Alberta, who obtained

it in the course of their own studies, and passed it on to me for identification. The

material recorded from Great Slave Lake was the result of a collection made in the summer

of 1964 by Mr. D. J. Larson, and passed on to me. A small collection was obtained on loan

from the Banff National Park Museum, Banff, Alberta.

In the text, locality records have been condensed to map form. Lists of localities and

dates are given for new species only. A complete list of all such data is deposited in the

Department of Entomology, University of Alberta, Edmonton, and is available to inter-

ested workers.

Disposition of material

All borrowed material was returned to the lending institutions.

Type material. — All type material is deposited in the Canadian National Collection,

Ottawa, unless stated otherwise in the descriptions of new species. Borrowed material here

designated as type material was returned to the lending institution. Where sufficient para-

type material exists in unborrowed material, at least one of each sex is deposited in the

Strickland Museum, Department of Entomology, University of Alberta.

Other material. - Most of the remainder of the material, all of which was obtained in

the field in the course of this study, is deposited in the Strickland Museum of the Depart-

ment of Entomology, University of Alberta. All remaining material, with the agreement of

the Department of Entomology, University of Alberta, forms part of my own reference

collection, or was distributed to other institutions or workers who expressed an interest

in obtaining such material.

Determination of species present in the study area

It was too much to hope that all species known from the study area would be represented

in my field collections. Therefore recourse was had to the literature pertaining to North

American Trichoptera, and to Fischer’s ‘Trichopterorum Catalogus’ (1960, 1967, 1968)

in a search for species recorded from Alberta, but not represented in my collections. The

collections of the Canadian National Collection, Royal Ontario Museum, and the Illinois

Natural History Survey were searched for specimens from the area, both to add to my

records and to supplement the list of species. These methods proved most successful, and

the names of several species were added to the list in consequence.

METHODS

Collecting methods

Several methods were employed in collecting the adult specimens used in this study.

First was hand-netting, either of individuals in flight, or by sweeping vegetation adjacent

to bodies of fresh water. Specimens were collected from a variety of vegetation, including

trees overhanging the water, and sedges in the water. The next method involved searching

the undersides of bridges, or the interior surfaces of culverts under roads. This method was

very effective, but was of maximum use only after much practice.

12

Nimmo

The above methods were employed in daytime collecting. Collection of adult Trichoptera

is also possible at nighttime, by the use of light sources of various types. As the insects

generally land close to the light source, a white sheet is used below the light to render them

more conspicuous and thus easier to pick up. The best times and conditions for light trap-

ping are from twilight to about 1 or 1 Vi hours later, at air temperatures greater than 55 F, on

cloudy evenings with no wind other than the most gentle air movements (see Nimmo, 1966b).

The first light source was a kerosene pressure lamp. This method is of use on warm,

humid evenings only. Car headlights were also used. They were aimed toward the body of

water from which the insects were expected to arrive in flight. An electric lamp rich in

the ultra-violet wavelengths was especially productive, even on cooler evenings. It was most

reliable when connected to a mains supply of current. This, however, was rarely possible

and other sources were used, including a portable gasoline generator, and a portable 12-volt

battery and DC-AC rectifier. The generator and battery sources, however, were unreliable.

Preservation of material

I collected all specimens directly into 80% ethanol, which both killed and preserved the

specimens. While this preservative may fade some specimens it permitted me to manipulate

whole specimens under the microscope. Storage is also facilitated. Dried, pinned specimens

are difficult to handle, shrivel up on drying, and are much more liable to damage.

Sorting of collections

All material acquired at one time and locality was collected into a single vial and labelled

with pertinent information. In the laboratory the contents of each vial were sorted to

species and all specimens of each species, from each collecting episode, were placed collec-

tively and permanently in a new vial of 80% ethanol and labelled. After this initial sorting

the vials were sorted to groups, each of which contained specimens of one species.

Association of males and females. - On occasion it was difficult to correlate correctly the

specimens of the two sexes of a species. On the initial sorting of field collections this was

accomplished in one of three ways. Firstly, if both sexes were already described in the litera-

tures no problem was encountered. Secondly, if pairs in copula were taken in the field, each

pair was segregated immediately to a separate vial. Later examination in the laboratory pro-

vided the information required to correctly associate specimens of the two sexes of any one

species in mixed field collections, and specimens of each sex collected individually and in

separate vials. And finally, if the above two sources of information were not available, wing

colour patterns, venation, and various other body characters were used for associative pur-

poses. Frequently the general facies of the specimens was all that was required. Also, a

knowledge of the general facies of the genera involved assisted in narrowing the field. Rarely

were very closely related species taken together. This last method was, in retrospect, found

to have worked remarkably well; improper associations were rare and were later corrected.

Identification of material. - In identifying the material collected, the available literature

was consulted. The specimens of Rhyacophilidae were identifiable with the assistance of

Ross’ (1956) publication. For the Limnephilidae Schmid’s (1955) publication was used to

the generic level. Identification to species was then made with the aid of scattered minor

literature referred to by Schmid for each species. In case of doubt, material was forwarded

to F. Schmid, of the Entomology Research Institute, Ottawa, G. B. Wiggins, of the Royal

Ontario Museum, Toronto, H. H. Ross, of the Illinois Natural History Survey, Urbana,

Illinois, or to D. G. Denning, Moraga, California for identification. I also spent one week

at the Illinois Natural History Survey examining the collections, both for records, and for

purposes of identifying material.

Rhyacophilidae and Limnephilidae

13

Type material was not normally examined. Little such material was available at the

institutions which I visited. Many of my identifications were made from specimens iden-

tified by workers who had previously had access to such type material. For a very few

species type material was all that was available and this was either borrowed, or drawings

made from the type were obtained on loan from the original authors.

Preparation of material. - To identify the Trichoptera recorded as occurring in the study

area genitalic characters were used at the species level, and venational, genitalic, and other

non-genitalic characters were used at the supraspecific level. Material for identification and

drawing was prepared as set out below.

The wings. The wings of the right side of the thorax were illustrated. These were torn off

cleanly at their bases with stiff, needle-pointed forceps. They were then passed through

95% ethanol for washing and stiffening, and spread on a clean glass slide. A second slide,

with 2.0 mm wide strips of cellulose tape along each lower edge, was then placed on top of

the first and the whole assemblage placed in a small hand-sized press. After clamping the

slides into the press the long edges were united by an application of Lepage’s white ‘Bond-

fast’ glue, which was found effective in binding the two slides together on drying. The

cellulose tape prevented seepage of glue between the slides. The slides were then unclamped,

labelled, and filed for future use. The wings were taken from the males, unless sexual

dimorphism was evident on in situ examination, in which case the female wings were also

treated as above, and illustrated. When only the female was known, the wings of this sex

were illustrated.

The genitalia. The male and female genitalia were prepared for examination and illustra-

j tion by removing the entire abdomen of a specimen and boiling it in a very strong solution

of KOH, to dissolve the abdominal organs and tissues. The abdomen was then removed

immediately to glacial acetic acid for clearing. After using this procedure for some time, I

| discovered that, while in the acid, a vigorous evolution of gas occurred within the abdominal

contents; this gas was violently expelled on return to the boiling KOH, removing the

abdominal contents in large part. Several such transfers between the two solutions resulted

in swift removal of the abdominal contents. The genital capsule and abdomen together

| were then returned to the vial of 80% ethanol which contained the donor specimen.

Preparation of drawings

Wing drawings were made with a camera lucida mounted on a stereo binocular micro-

scope. Slides were made of the wings of all species recorded here, but not all are illustrated

as, in Limnephilus for example, the venation varied little between species.

Genitalic drawings were made using a square-grid eyepiece in a stereo binocular micro-

scope. The image was imparted to a segment of bristol board lined in pencil with a similar

squared grid. The size of the grid squares varied according to the size of the specimen, as it

was desired to produce drawings of similar sizes for all species. The genital capsule being

drawn was held steady, in a dish of 80% ethanol under the microscope, by a piece of wire

of sufficient weight inserted anteriorly into the abdominal cavity.

In the drawings of the male the genital segments (IX and X) are frequently extracted

from their normal position, retracted into segment VIII; this was done for greater clarity,

but was not possible with the specimens of certain species. In the drawings of the male

genital capsule of Rhyacophila spp. the lateral aspect shows the mesal face of the far (right

hand) side clasper as this is the face which bears important characters.

Measurements and scales

Wing length is used to indicate relative sizes of species to each other. It is the distance,

14

Nimmo

in millimeters, from the fore wing tip to the base at the costal edge where the wing folds

over at rest. Males were measured, unless only females were available. Scale bars are pro-

vided for the genitalic and wing drawings. The genitalic scale bars represent 0.5 mm or

0.25 mm, depending on the size of the specimen. The wing scale bars represent 4.0 mm.

All measurements and scale bars were obtained by use of a micrometer eyepiece in a stereo

binocular microscope. The scale bars for the genitalic drawings are immediately adjacent

to the lateral aspect of the male genital capsule. All drawings, male and female, with the

exception of wing drawings, which are located elsewhere, for any one species are to the

same scale. When only the female is known no scale is given. Drawings derived from sources

other than specimens available to me, have no scale, since the original sources had none.

Criteria employed

Inasmuch as this study is not a revision but a faunal survey, I restrict my remarks to taxa

at the species level. For higher taxa I have adhered to the work of Ross (1956) and Schmid

(1955). It is desirable to outline criteria at the species level as several new species are

described.

Characters used in distinguishing species. — In the study of Trichoptera at the species

level, with the exception of the species of a very few genera, the genitalic characters are of

prime importance. Species are segregated and recognised on the basis of differences in

genitalic characters. Other characters may be referred to in combination as distinguishing

one species from another. Such are, the colour pattern of the fore wing, if sufficiently

distinctive and constant, spur formula, coloration of the thorax, head or legs, and form and

setation of designated areas or parts of the body. But the use of such characters individually

is strictly subsidiary, as they are rarely sufficiently distinctive by themselves to provide a

basis for erection of species. On the other hand, such characters may be utilised in the

erection of higher taxa, when common to two or more species of still higher taxa. Wing

pattern is, however, of paramount importance in distinguishing species of certain genera

(e.g. Leptocella, of the Leptoceridae) where genitalic characters are highly variable and of

dubious utility.

In the study of genitalic structures at the species level, form, structure, setation or

spination are the important characters to be observed. Coloration, for example, is usually

held to be of no importance, varying with age of the specimens. While it is accepted,

indeed expected, that the characters in which interspecific differences may be detected

vary intraspecifically, in most species recorded here this variation is limited and can, with

practice, be recognised for what it is. In opposition to certain other groups of insects,

Trichoptera species may be distinguished by critical examination of the general facies of the

characters selected. No application of statistical techniques has yet been found necessary.

While supraspecific taxa are generally excluded from this discussion, it may be noted

that, in initially segregating species of higher taxa to membership in these taxa, the charac-

ters employed are frequently venational, or are a variety of general body characters, and are

frequently genitalic, involving considerations of characters less varied than at the specific

level, and more revealing of the broader evolutionary history of the group.

Criteria at the species level. — Simpson (1961) defines the genetical species as follows:

‘Species are groups of actually or potentially inter-breeding natural populations, which are

reproductively isolated from other such groups’. This biological species definition is the

ideal, which I accept, but it is normally impractical to use it as a working definition due to

lack of information. Simpson discusses this at some length. He also states that, in practice,

this definition employs morphological criteria, but without the adverse implications of the

‘morphospecies’.

Rhyacophilidae and Limnephilidae

15

In dealing with Trichoptera species only morphological and distributional data are avail-

able at present, as outlined above. The data are, however, applied with the intentions of the

above definition in mind; morphological differences are taken as evidence of reproductive

isolation.

In this study I employ the following definition: A species is that group of individuals

which is recognized as a unit by a multiplicity of characters, the nature of forms of which

are peculiar to, and constant within, the available specimens, and which are distinguished

from specimens of the presumed most closely related species by pronounced discontinuities

in any or all of the characters. Interspecific variation, as mentioned above, is taken into

account with regard to the constancy of characters within a species, and the discontinuities

between species.

While, in most species of the Alberta Rhyacophilidae and Limnephilidae, the interspecific

discontinuities are sufficiently evident as to require no comment, in some genera or species

groups the recognised species are very similar. This may lead to some confusion in sorting

if specimens of two similar species are collected simultaneously. With regard to such species,

particularly if described as new, I reserved judgement as to their separate identity until I

had sufficient information on which to base a decision. This information was acquired in

the form of collecting data. Specimens of the species concerned had to be collected sepa-

rately at different localities sufficiently often to instil confidence in regarding them as

separate entities.

Miscellaneous notes

Notes on the descriptions. — General body descriptions are derived from the male. Sexual

dimorphisms of the female are noted wherever applicable. Wing colours are of the male

unless stated otherwise; the hind wings are hyaline in most species. The costal area referred

to is that part of the wing between the costa and subcosta, extending from the humeral

cross-vein to the point at which the costa and subcosta meet distally.

Notes on the text. — Most species recorded herein are strictly nearctic in distribu-

tion. The literature for each of these species is, to the best of my knowledge, complete

to early 1970. However, for the holarctic species in the study area, only references to

nearctic literature are recorded here, and the reader is referred in appropriate cases to

Fischer’s ‘Trichopterorum Catalogus’ (1967; 1968) for a complete listing of palaearctic

literature. This system is adopted as the literature on these holarctic species is too ex-

tensive for complete inclusion here. Synonymies for each species are complete, however.

Type localities are named for each specific epithet, and are given in the citation for each

species.

The keys used here for supraspecific taxa are adapted in translation from Schmid (1955).

They are adapted by restricting them to the taxa recorded from the study area. At the

species level, keys have been constructed for the males, and females, of each genus, if

known. The character synopses for the supraspecific taxa of the Limnephilidae are adapted

in translation from Schmid (1955), and are greatly condensed.

Immediately following each species description is a short statement of the known, or

suspected, habitats, and biology, including notes on emergence if available and flying season,

of the species. This information was largely derived from personal notes.

Notes on distribution maps. — The range maps presented here give only an approximate

outline of collecting localities in the study area due to limitations of scale (see above, p. 1 1 ,

for information on exact listing of collecting localities for each species). The inset maps

of North America give only the nearctic distributions of each species. In the cases of

holarctic species the palaearctic ranges are described briefly in the text.

16

Nimmo

THE FAMILY RHYACOPHILIDAE STEPHENS

This family is represented in Alberta and eastern British Columbia by the genus Rhyaco-

phila Pictet only, belonging to the subfamily Rhyacophilinae Ulmer. Immediately following

are synopses of the familial characteristics of the Rhyacophilidae, as adapted from Mosely

(1939) and Mosely and Kimmins (1953), excluding the Glossosomatidae, and the subfamilial

characteristics of the Rhyacophilinae, as adapted from Mosely (1939) and Ross (1956). The

synopsis of characteristics for the genus Rhyacophila is adapted from Mosely (1939) and

Ross (1956). The grouping of species within the genus is adopted from Ross (1956).

Character synopsis of the Rhyacophilidae. — Ocelli three. Maxillary palpi each of five

articles in both sexes; basal articles short, remainder long, cylindrical. Metascutellum with

or without warts. In some genera the middle tibia is dilated considerably. Spurs 3, 4, 4 in

both sexes. Wings elongate, roughly parabolic, obliquely truncated apically. Hind wings

shorter, narrower than fore wings. Venation generally complete (Fig. 4). Fore wings with

apical cells fl-f5 present. Hind wings with only f2 and f5 present in some genera. Discoidal

cell open or closed on fore and hind wings, or lacking on hind wings. R1 of fore wings

forked apically or not. Thyridial cell present, sub-radial present or absent.

Character synopsis of the Rhyacophilinae. — Middle tibia not dilated. Fore wings with

apical cells fl-f5 present; hind wings with fl-f3, and f5, present. Discoidal cell of both fore

and hind wings open. Male genitalia with anal sclerite generally present, rarely absent.

The genus Rhyacophila Pictet

Synopsis of characters. — Antennae slender, shorter than wings; pedicel very short. Sec-

ond article of maxillary palpus short, globular. Tergum VIII of male rarely modified postero-

dorsally. Segment IX wide throughout, but varying in width. Claspers of two articles; of

varied size; large, conspicuous. Segment X with anal sclerite in most species. Female genitalia

long, tapered, slender, with pair of terminal cerci. Segments IX-XI membranous. Segment

VIII with basal portion sclerotized, in some species complexly, with attendant lobes.

The genus Rhyacophila in Alberta and eastern British Columbia. — As presented in this

study the genus is represented by 22 species, of which two are represented by unassociated

females. The species are primarily confined to the mountain and foothill areas, but several

range eastward to the plains, and two are transcontinental.

Following are separate keys to the males and females of the genus known to be present in

the area.

Key to the Males of the Alberta and eastern British Columbia species of Rhyacophila Pictet

la. Aedeagus simple, limnephiloid in general appearance, with paired lateral arms

and simple median shaft (Fig. 25, 29, 33). Segment X rather like bird’s head in

lateral aspect; the crown with multiple rounded depressions (Fig. 24, 28, 32)

2

lb. Aedeagus not limnephiloid, ranging from very small and simple (Fig. 91) to

large and complex, with as many as five lobes (Fig. 52) 4

2a.(la) Distal article of clasper very deeply cleft, with ventral lobe lanceolate, dorsal

lobe finger-like (Fig. 32). Lateral arms of aedeagus fringed distally with tuft of

fine hairs (Fig. 33) R. glaciera Denning, p. 21.

2b. Distal article of clasper not constructed 3

3a. (2b) Distal article of clasper with dorsal lobe directed strongly meso-anterad along

mesal face of basal segment (Fig. 24). Anal sclerite flanked by simple, laterally

Rhyacophilidae and Limnephilidae

17

3b.

4a. (lb)

4b.

5a. (4a)

5b.

6a. (4b)

6b.

7a.(6b)

7b.

8a. (7a)

8b.

9a.(8a)

9b.

10a.(8b)

10b.

1 la.(7b)

lib.

12a.(l la)

12b.

13a.(12a)

13b.

14a.(12b)

14b.

15a.(14b)

15b.

16a.(l lb)

uncleft, ventral lobes of segment X R. alberta Banks, p. 19.

Distal article of clasper with dorsal edge thick, fleshy; horizontally almost cylin-

drical. Anal sclerite flanked by two lobes of segment X on each side (Fig. 28)

R. tucula Ross, p. 20.

Postero-dorsal edge of tergum VIII with one or more lobes (Fig. 54, 86) 5

Postero-dorsal edge of tergum VIII unmodified 6

Tergum VIII with one small, rounded lobe (Fig. 54) . . . . R. rickeri Ross, p. 28.

Tergum VIII with three distinct lobes; lateral lobes large, rounded distally and

flanking curved, square tipped median lobe (Fig. 86, 87)

R. verrula Milne, p. 34.

Postero-dorsal edge of segment IX with distinct tuft of long, thick setae, direc-

ted posterad (Fig. 96) R. vemna Milne, p. 24.

Postero-dorsal edge of segment IX quite clear of setae or hairs 7

Postero-dorsal edge of segment IX developed posterad as distinct lobe pro-

jected well beyond segment (Fig. 39, 5 1, 89, 93) 8

Postero-dorsal edge of segment IX not so developed 11

Segment X with long, thin, curved and folded strap-like median lobe (Fig. 89,

93) 9

Segment X without strap-like median lobe 10

Lobe of segment IX bilobed distally, in dorsal aspect (Fig. 90)

R. vagrita Milne, p. 35.

Lobe of segment IX trilobed distally, in dorsal aspect; lateral lobes shorter than

median lobe (Fig. 94) R. milnei Ross, p. 36.

Segment X without anal sclerite. Aedeagus small, with median shaft short and

with two rounded lateral wings (Fig. 39); lateral arms ventral, membranous,

terminated by brush of stout, dark spines (Fig. 40)

R. acropedes Banks, p. 23.

Segment X with anal sclerite and large, dish-like tergal strap; segment X of two

thin, vertical plates (Fig. 51). Aedeagus huge, with membranous base, long

clavate lateral arms, hooded tip on median shaft and two lanceolate ventral

lobes (Fig. 52) R. hyalinata Banks, p. 27.

Distal article of clasper bilobed (Fig. 35, 57, 65, 69, 75, 79) 12

Distal article of clasper not bilobed 16

Distal article of clasper cleft apically, appearing scissors-like (Fig. 69, 75) . . . 13

Distal article of clasper not scissors-like 14

Anal sclerite small, with two button-like disto-lateral lobes (Fig. 70) curved

dorsad (Fig. 69) R. vaccua Milne, p. 31.

Anal sclerite larger, with two simple, rectangular distal lobes (Fig. 76) directed

ventrad from segment X (Fig. 75) R. chilsia Denning, p. 32.

Segment X massive, convoluted plate (Fig. 57, 58) longer than claspers. No

evident anal sclerite R. vepulsa Milne, p. 28.

Segment X shorter than claspers. Anal sclerite evident 15

Distal article of clasper with acuminate, triangular dorsal lobe (Fig. 35)

R. vofixa Milne, p. 22.

Dorsal lobe of distal article minute, hooked ventrad (Fig. 65)

R. pellisa Ross, p. 30.

Dorsal lobe of distal article large, rectangular (Fig. 79)

R. vobara Milne, p. 32.

Segment X long, plate-like, cleft deeply, in dorsal aspect (Fig. 63, 83) 17

18

Nimmo

16b.

17a.(16a)

17b.

18a.( 16b)

18b.

Key to the

Pictet

la.

lb.

2a.(la)

2b.

3a. (2a)

3b.

4a. (3a)

4b.

5a. (3b)

5b.

6a.(5a)

6b.

7a. (5b)

7b.

8a.(7b)

8b.

9a.(2b)

9b.

10a. (9a)

Segment X short, of two lateral plates flanking anal sclerite (Fig. 42, 47) ... 18

Aedeagus large plate with lateral edges curled dorsad, ejaculatory duct pro-

longed by slender, tapered, dorsally curved median tube (Fig. 62)

R. belona Ross, p. 29.

Aedeagus complex, with two very long, membranous lateral arms terminated

by spatulate, setose distal plate (Fig. 84) R. angelita Banks, p. 33.

Lateral plates of segment X simple (Fig. 42, 43) R. bifila Banks, p. 25.

Lateral plates of segment X bilobed (Fig. 47, 48)

R. coloradensis Banks, p. 26.

Females of the Alberta and eastern British Columbia species of Rhyacophila

Basal portion of segment VIII strongly sclerotized, clearly demarcated from

membranous distal portion (Fig. 26) 2

Basal portion of segment VIII weakly sclerotized, merged almost imperceptibly

with distal membranous portion (Fig. 85) 15

Sclerotized portion of segment VIII with distinct lobes or processes quite free

from main body of segment, except at bases (Fig. 26, 38, 45, 53, 65. 73) .... 3

Sclerotized portion of segment VIII without such processes (Fig. 30, 34, 41,

61,68,92, 100) 9

Distal portion of segment VIII, and segments IX-XI shortened, partly retracted

within sclerotized base of segment VIII (Fig. 45) 4

Distal portion of segment VIII, and segments IX-XI long, tapered gradually

distad 5

Ventral lobes of segment VIII long, thin, arcuate, in ventral aspect (Fig. 46)

R. bifila Banks, p. 25.

Ventral lobes of segment VIII short, rounded apically, on common pedicel

(Fig. 50) R. coloradensis Banks, p. 26.

Segment VIII with one medial ventral process, (Fig. 26, 27) 6

Segment VIII with two ventro-lateral processes (Fig. 38, 53, 65) 7

Ventral process acute-triangular, but with deep v-shaped distal cleft; process

extended well posterad under membranous portion of segment (Fig. 73, 74)

R. vaccua Milne, p. 31.

Ventral process short, triangular, with rounded apex (Fig. 27)

R. alberta Banks, p. 19.

Membranous distal portion of segment VIII flanked on either side, just distad

of sclerotized base by lightly sclerotized, oblong plate (Fig. 53)

R. hyalinata Banks, p. 27.

Membranous distal portion of segment VIII without sclerotized plates 8

Spermathecal sclerites long, slender, each with deep narrow hook at posterior

end (Fig. 38) R. vofixa Milne, p. 22.

Spermathecal sclerite long, triangular in lateral aspect (Fig. 64)

R. belona Ross, p. 29.

Posterior edge of sclerotized portion of segment VIII with dorsal portion offset

anterad, with dorsal and ventral edges joined by long sloping edge (lateral

aspect) (Fig. 61, 100) 10

Posterior edge not offset as above 11

Anterior sclerotized edge of segment VIII with retractor rod attached (Fig.

100) R. species 1, p. 37.

Rhyacophilidae and Limnephilidae

19

10b.

1 la.(9b)

lib.

12a.( 1 lb)

12b.

1 3a.( 1 2b)

13b.

14a.(13b)

14b.

14c.

15a.(lb)

15b.

16a.(15a)

16b.

1 7a.( 1 5b)

17b.

18a.(17a)

18b.

Anterior sclerotized edge of segment VIII without retractor rod attached (Fig.

61) R. vepulsa Milne, p. 28.

Membranous portion of segment VIII emergent from upper half of sclerotized

portion; lower half pinched off in form of thin, plate-like keel posteriorly

(Fig. 41) R. acropedes Banks, p. 23.

Membranous portion of segment VIII emergent from entire diameter of sclero-

tized portion 12

Dorsal surface of segment VIII immediately distad of sclerotized portion, mi-

nutely spinate (Fig. 30) R. tucula Ross, p. 20.

Membranous portion of segment VIII not spinate 13

Sclerotized portion of segment VIII with posterior edge indented ventrally,

laterally, and dorsally, as two pairs of lateral extensions (Fig. 81)

R. vobara Milne, p. 32.

Sclerotized portion of segment VIII with posterior edge circular, or only slight-

ly sinuate 14

No evident spermathecal sclerites (Fig. 34) R. glaciera Denning, p. 21.

Spermathecal sclerite long, narrow strap in ventral aspect, with slightly wider,

darker, posterior end and distinctly widened, pierced, anterior end (Fig. 68)

R. pellisa Ross, p. 30.

Spermathecal sclerite long, thin, irregular rod in lateral aspect; tip small, pick-

like (Fig. 92) R. vagrita Milne, p. 35.

Abdomen in area of segments VIII-X with one or more distinct annular swel-

lings (Fig. 88, 102) 16

No such annular swellings present 17

Tergum X clearly sclerotized R. verrula Milne, p. 34.

Tergum X not sclerotized. Spermathecal sclerite spindle shaped in lateral aspect,

with anterior end attenuated (Fig. 102) R. species 2, p. 37.

Spermathecal sclerite spindle shaped in lateral aspect (Fig. 56, 85) 18

Spermathecal sclerite relatively long, tapered posterad in ventral aspect; ante-

rior end cleft, with distinct median fissure (Fig. 99) R. vemna Milne, p. 24.

Spermathecal sclerite with both ends (in lateral aspect) attenuated (Fig. 85)

R. angelita Banks, p. 33.

Spermathecal sclerite (in lateral aspect) sigmoid in outline (Fig. 56)

R. rickeri Ross, p. 28.

The alberta group

Males of this group are characterised by large anal sclerites, simple tenth terga and,

especially, by simple, limnephiloid aedeagi, with simple lateral arms and median shaft. The

group contains four species of which three are found in Alberta.

Rhyacophila alberta Banks, 1918

(Fig. 4a, 4b, 24-27, 104)

Rhyacophila alberta Banks, 191 8; 21. (Type locality: Banff, Alberta). Banks, 1918:3b.

Dodds and Hisaw, 1925b:386. Ulmer, 1932:209. Betten, 1934: 135. Milne, 1936:98, 106,

110. Ross, 1944:291. Ross, 1950a:261. Ross, 1956:76, 1 16. Schmid, 1958: 13. Fischer,

1960:70. Ross, 1965:591. Schmid, 1970:55, 124.

Males of this species are distinguished from males of other species of the alberta group by

20

Nimmo

dorsal lobe of distal article of clasper directed meso-anterad (Fig. 24). The rounded triangu-

lar lobe on lower posterior edge of segment VIII is characteristic of females of this species

(Fig. 27).

Description. — Antennae very pale yellow. Vertex of head mottled red-brown. Fore wing

length of male 1 1 mm; pale yellowish brown, heavily irrorate, with dark areas concentrated

on veins. Stigma pale, opaque. Venation of fore and hind wings as in Fig. 4a, 4b.

Male genitalia. (Specimen from Gap, near Exshaw, Alberta). Segment IX rectangular

dorsally, tapered sharply antero-ventrad to lateral sutures then widened ventrally (Fig. 24).

Claspers each with massive rectangular basal article concave mesally, especially at base.

Distal article with long triangular ventral lobe, and dorsal lobe directed meso-anterad along

mesal face of basal article. Segment X with dorsal portion rather like skull and beak of bird,

with small rounded depressions at peak. Ventral part of segment cleft dorso-ventrally, flank-

ing divided anal sclerite. Aedeagus with median shaft long, thin, bulbous at mid point (Fig.

25); lateral arms long, with large, rounded, ventral lobe distally with five long, curved spines.

Female genitalia. (Specimen from Gap, near Exshaw, Alberta). Basal quarter of segment

VIII sclerotized, proximal edge slightly bulged and divided from distal area by thin annular

line; distal edge produced ventrally as short, stout, triangular tooth (Fig. 26, 27). Cerci

short, hyaline.

Notes on biology. — Adults of this species appears to emerge from streams ranging from

small mountain brooks to large, turbulent mountain torrents. The flying season extends

from August 12 to October 9.

Geographical distribution. — The known range of this species extends from Alaska to

Colorado and Utah (Fig. 104) apparently being confined largely to the Rocky Mountain

chain. In Alberta this species is found in the mountains and foothills at altitudes between

4,000’ and 6,500’. Dodds and Hisaw (1925b) recorded it from Colorado, between 9,000’

and 11,000’.

I have examined 174 specimens, 131 males and 43 females, from the study area.

Rhyacophila tucula Ross, 1950

(Fig. 5a, 5b, 28-31, 105)

Rhyacophila tucula Ross, 1950a:261. (Type locality: Gardner River, Yellowstone National

Park, Wyoming). Ross, 1952:45. Ross, 1956: 1 16. Ross, 1965:591. Anderson, 1967:508.

Smith, 196&658, 666-668, 673. Schmid, 1970:55, 124.

Males of this species are distinguished from males of other members of the alberta group

by bifid lateral flaps of segment X enclosing anal sclerite, and by long, rectangular dorsal

ridge or fold of distal article of clasper (Fig. 28). The spinate dorsal surface of membranous

portion of segment VIII (Fig. 30) is distinctive of females.

Description. — Antennae pale yellow. Vertex of head red-brown, with distinct cruciform

pattern posteriorly, formed from two intersecting sutures. Thorax straw yellow, to red-

brown dorsally. Spurs dark brown. Fore wing length of male 9.5 mm; pale red-brown,

irregularly irrorate with no distinct stigma. Venation of fore and hind wings as in Fig. 5a, 5b.

Male genitalia. (Specimen from Sundance Creek, west of Edson, Alberta). Segment IX

laterally rectangular but with large triangular bight in lower half of posterior edge (Fig. 28).

Claspers each with long, rectangular basal article channeled along entire length of mesal face.

Segment X laterally similar to birds head, with two bifid ventral lobes flanking anal sclerite.

Aedeagus with long, slender median shaft; distal portion of shaft very slender tube, basal

portion long slender bulb; lateral arms long, slender, smoothly expanded distally, each with

distal spine, and mesal edge with heavy, dark spines (Fig. 29).

Rhyacophilidae and Limnephilidae

21

Female genitalia. (Specimen from Alaska; in Illinois Natural History Survey). Segment

VIII with basal half sclerotized, tapered (Fig. 30); proximal half of sclerotized portion light

brown, remainder darker, clearly demarcated. Dorsal surface of membranous portion of

segment VIII minutely spinate. Spermathecal sclerite long, almost hyaline (Fig. 31); distal

end constricted as circular head (ventral aspect); lateral edges darker than remainder.

Notes on biology. — Smith (1968) records the larvae from . . small to medium streams

with mixed rubble bottoms”, in Idaho. My records indicate fairly fast, rock filled streams

to be the habitat of larvae of this species. Smith also records the adult flying season as

September-October. My records indicate a flying season of August 24 to October 12.

Geographical distribution. — The known range of this species extends from Alaska to

Wyoming and Oregon (Fig. 105). It extends east to the Rockies but appears to be centered

primarily in the Coast Ranges. In Alberta the species is known only from one locality in the

low eastern foothills near Edson. The two localities shown in eastern British Columbia are

also at low altitude (under 4,000’).

I have examined six males from the area.

Rhyacophila glaciera Denning, 1965

(Fig. 6a, 6b, 32-34, 106)

Rhyacophila glaciera Denning, 1965a: 263, 265. (Type locality: Glacier National Park, Mon-

tana). Schmid, 1970:55, 124.

Males of this species are distinguished from males of other species of the alberta group by

long, deeply cleft distal article of clasper. Females are distinguished by relatively unmodified

segment VIII: posterior rim of sclerotized portion only very shallowly indented laterally.

Description. — Antennae light yellowish brown. Vertex of head deep reddish brown,

except warts paler. Thorax brownish yellow, to red-brown dorsally. Spurs brown. Male fore

wing length 8.8 mm; clear mottled greyish brown; stigma evident but light. Venation of fore

and hind wings as in Fig. 6a, 6b.

Male genitalia. (Specimen from Mt. Edith Cavell, Jasper, Alberta). Segment IX with or

without small dorsal, hyaline fin (Fig. 32). Segment IX strongly pinched in at lateral sutures,

as in R. tucula. Claspers each with short basal article channeled on mesal face; distal article

large, with very deep u-shaped cleft dividing it to rounded, finger-like, dorsal lobe and knife-

like ventral lobe. Segment X with dorsal body only sparsely indented, forming rough semi-

circle. Aedeagus (Fig. 33) with median shaft in two distinct parts; distal part long, very thin;

proximal part much thicker, continued thus to base. Lateral arms clavate in dorsal aspect;

terminated by thick brush of setae.

Female genitalia. (Specimen from Mt. Edith Cavell, Jasper, Alberta). Basal part of seg-

ment VIII sclerotized, truncated, cone with slightly sinuate postero-lateral edges (Fig. 34).

No cerci evident.

A note on taxonomy. — In his original description Denning (1965a) states that this species

is not related to any other species of the genus. This may appear to be so, but an examina-

tion of the male genitalia, and a close comparison with the genitalia of R. alberta and tucula

places this species in the alberta group beyond doubt. The virtual identity of the aedeagi of

specimens of these species is of special importance in this connection.

Notes on biology. — This species frequents mountain streams ranging from small, turbu-

lent, rocky creeks, to very small alpine trickles. The adult flying season extends from August

19 to October 7. I have taken specimens of this species crawling about on one to two feet

of snow in October at the Mt. Edith Cavell alpine meadows in Jasper National Park. The

nearby stream was largely frozen, but with occasional open holes in the thin ice.

Nimino

Geographical distribution. - The known range of this species is very small. The species

was only recently described from Montana. It is known only from the Rocky Mountains

and Alberta. Altitude range is from 5,000’ to 7,000’.

Eight specimens, six males and two females, were examined from the study area.

The vofixa group

This group, represented here by only one species, is characterised by simple male geni-

talia in which the anal sclerite is divided into two large, ovate, lateral lobes. Aedeagus with

large, finger-like, dorsal process; median shaft flanked by two finely divided lateral lobes

like sheaves of spines. There are only two known species in this group, of which one occurs

in Alberta.

Rhyacophila vofixa Milne, 1936

(Fig. 7a, 7b, 35-38, 107)

Rhyacophila vofixa Milne, 1936:95, 102, 111. (Type locality: Edmonton, Alberta). Ross,

1944:291. Ross, 1956:80, 101, 116. Schmid, 1958:17. Fischer, 1960: 1 52. Smith, 1968:

672-673. Schmid, 1970:67, 126.

Males of this species are recognized by form of distal article of clasper, with acuminate

triangular dorsal lobe and thick, fleshy, triangular ventral lobe (Fig. 35). Females are recog-

nized by hooked spermathecal sclerites (Fig. 38).

Description. - Antennae yellow-brown. Vertex of head red-brown. Thorax light red-

brown, slightly darker dorsally. Spurs brown. Fore wing length of male 11.1 mm, deep

chocolate-brown with reddish tinge; scattered hyaline irrorations. Stigma dark, opaque,

clothed with fine mat of hairs.

Male genitalia. (Specimen from Rapids Creek, Gap, Alberta). Segment IX wide dorsally,

narrower in ventral third of total height. Extreme ventral area segregated by dark suture

lines (Fig. 35). Claspers each with large, trapezoidal, mesally concave, basal article. Distal

article bilobed; dorsal lobe acuminate triangular; ventral lobe triangular, larger than dorsal,

rounded distally, heavy. Segment X with concave dorsal plate; lateral walls sharp edged.

Anal sclerite of two rounded, mesally concave, lateral flaps located ventrad of distal tip of

segment X (Fig. 36). Aedeagus relatively small, with ejaculatory duct terminated on peculiar

pick-like, hyaline, median shaft; median shaft flanked basally by membranous lobes tipped

by numerous long, thin spines. Base of aedeagus surmounted by large, fleshy, dorsal lobe

terminated by dark, folded, sclerotized pocket (Fig. 37).

Female genitalia. (Specimen from Rapids Creek, Gap, Alberta). Basal portion of segment

VIII sclerotized in form of truncated cone terminated in two pairs of lateral lobes (Fig. 38).

Segment XI with pair of small, hyaline cerci. Spermathecal sclerites two, parallel to each

other, lightly sclerotized, tapered finely anterad, with heavier, deeply hooked posterior ends.

Notes on biology. - Adults emerge from non-turbulent, smoothly flowing, but swift,

mountain brooks, to smaller, but very turbulent mountain torrents. The flight season ex-

tends from July 15 to August 31.

Geographical distribution. — The known range of this species extends from Alaska to

Idaho and Washington (Fig. 107), apparently ranging throughout the northern Cordillera.

My collecting in Alberta indicates that the species is confined to the mountain and foothill

areas, between 3,000’ and 6,500’. The type locality is given as Edmonton by Milne (1936)

which is at 2,000’.

I have examined 255 specimens, 184 males and 71 females, from the study area.

Rhyacophilidae and Limnephilidae

23

The acropedes group

Males of this group, represented here by two species, are characterised by simple aedeagus

(Fig. 40) with lateral arms in form of membranous, extensible lobes with multi-spinate tips.

One branch of this group has segment X with anteriorly directed basal processes (Fig. 96).

In the second branch segment X is cleft dorsally down the middle. There are eight known

species in this group, of which one is known from Alberta.

Rhyacophila acropedes Banks, 1914

(Fig. 8a, 8b, 39-41, 108)

Rhyacophila acropedes Banks, 1914:201. (Type locality: Deer Creek, Provo Canyon, Utah).

Dodds and Hisaw, 1925b:386. Essig, 1926:176. Betten, 1934: 135. Milne, 1936:93, 102,

110. Ross, 1938b:4. Ross, 1941a:36. Ross, 1944:291. Denning, 1948a: 87. Leonard and

Leonard, 1949b:3. Morse and Blickle, 1953:69. Ross, 1956:75, 83, 84, 117. Fischer,

1960:69. Flint, 1962:479-480. Denning, 1963:244. Ross, 1965:591. Anderson, 1967:

508, 517. Smith, 1968:658, 660, 667. Schmid, 1970:87, 131.

Males of this species are distinguished from males of other species of the acropedes group

by lateral ‘wings’ on median shaft of aedeagus, and by acuminate lobe dorsad of segment X,

attached to segment IX. Females are distinguished by postero-ventral keel of segment VIII.

Description. — Antennae yellow-brown. Vertex of head uniformly deep red-brown. Thor-

ax deep yellow-brown, to deep red-brown dorsally. Spurs brown. Fore wing length of male

10.8 mm; colour grey-brown; costal area hyaline, stigma weak, indistinctly irrorate. Vena-

tion of fore and hind wings as in Fig. 8a, 8b.

Male genitalia. (Specimen from Cold Creek, Nojack, Alberta). Segment IX essentially

rectangular, with irregular posterior edge (Fig. 39); postero-dorsal edge produced posterad

as broad, triangular plate dorsad of segment X. Claspers each with basal article arcuate; basal

area laterally flattened, distally tubular. Heavy distal article bilobed, with small pyramidal

dorsal lobe, and massive ventral lobe roughly triangular in cross section. Segment X of two

approximately rectangular, vertical plates side by side; base of each plate with broad lateral

flange. Median shaft of aedeagus simple tapered tube (Fig. 40), flanked just distad of base

by two rounded lateral ‘wings’. Lateral arms large, membranous, tubes each terminated by

sheaves of long, dark spines.

Female genitalia. (Specimen from Cold Creek, Nojack, Alberta). Basal portion of segment

VIII heavily sclerotized; antero-lateral edges slightly depressed; ventro-posterior surface

keeled, tapered gradually to sharp posterior edge; membranous remainder of segment emer-

gent only above keel (Fig. 41). Spermathecal sclerites two, indistinct, simple, tapered.

Notes on biology. — This species emerges from a variety of stream types ranging from

torrential, rocky, mountain streams, to swift, smoothly flowing, shallow streams on pebble

beds, to very quiet, sluggish streams on earthen beds. Smith (1968) gives the adult flight

period in Idaho as late July to early August; peak emergence is in late July. My records give

the flight period in Alberta as July 1 to August 22. I have a record from Jaffray, in south

eastern British Columbia, dated May 10; seven of each sex were taken. Smith states that the

species overwinters in Idaho as third or fourth instar larvae; pupation occurs in late May and

June. He also adds that the adults are active in the afternoon, ceasing flight at dusk, and do

not come to light. My own observations confirm this, specimens usually being taken under

bridges or culverts. Both Smith (1968) and Denning (1948a) report that both sexes emit an

unpleasant odour when handled. I have not noticed this phenomenon.

Geographical distribution. — The known range of this species extends from British Colum-

24

Nimmo

bia in the west to Labrador in the east, and south as far as Colorado (Fig. 108). This is one of

the two transcontinental species of Rhyacophila known to occur in the study area. From the

map there is seen to be a gap in mid-continent; this may be genuine or an artifact of collect-

ing. In Alberta the species seems to be largely confined to the mountain and foothill regions.

However, two localities well away from the main Cordillera are recorded. One, near White-

court (Chickadee Creek), is on the south west edge of the Swan Hills, an isolated rise of land;

the other is at Cold Creek, Nojack, which is quite outside any unusual elevation of land, and

about 50 miles from the eastern extremity of the foothills. This stream is peculiar in afford-

ing a rather large selection of otherwise mountain caddis-flies while, at the same time, af-

fording a selection of plains species. This species ranges between 2,500’ and 7,000’ altitude.

I have examined 698 specimens, 449 males and 249 females, from the study area.

Rhyacophila vemna Milne, 1936

(Fig. 21a, 21b, 96-99, 120)

Rhyacophila vemna Milne, 1936:92, 102, 111. (Type locality: White River, Mt. Rainier,

Washington). Ross, 1944:291. Denning, 1948a: 105-106. Ross, 1956:84. Smith, 1965:

243. Schmid, 1970:86, 131.

Males of this species are distinguished from males of other species in the acropedes group

by long tuft of setae on posterior edge of tergum IX (Fig. 96), and by long, twisted, distal

article of clasper. Females are distinguished by open-ended, mesally fissured spermathecal

sclerite (Fig. 99).

Description. — Antennae red-brown. Vertex of head red-brown. Thorax light red to

yellow-brown, darker dorsally. Spurs brown. Fore wing length of male 18.1 mm; chocolate-

brown to reddish brown, heavily irrorate, with thick stigma. Hind wings with distal half

stained clear brown. Venation of fore and hind wings as in Fig. 21a, 21b.

Male genitalia. (Specimen from Gap, near Exshaw, Alberta). Segment IX with anterior

edges essentially straight, vertical; posterior edges with large, smooth, ventral indentation;

dorsal portion curved antero-dorsad to narrow dorsal ridge (Fig. 96). Clasper with distinct

mesal ledge basally on basal article; distal article twisted as if part of coil, with thick,

rounded, distal lip. Dorsal part of segment X of two thin, short, vertical plates; distally

with darkened, folded, deeply incised plate, denticulate on dorsal surface (Fig. 97). Median

shaft of aedeagus slender, tapered; lateral arms membranous, very thick, terminated by

heavy brushes of long, amber setae; these lobes surmount median shaft (Fig. 98).

Female genitalia. (Specimen from Gap, near Exshaw, Alberta). Basal half of segment VIII

simple, tapered, lightly sclerotized tube. Spermathecal sclerite in ventral aspect (Fig. 99)

spanner-like in appearance, with anterior end open; spindle shaped fissure located mesally.

Cerci minute.

Notes on biology. — This species inhabits small, swift, gravel bedded, mountain streams.

The flight period of the adults in Alberta extends from May 17 to July 7.

Geographical distribution. — The known range of this species extends from the Cascade

Mountains of Washington to the Rockies of Alberta.

I have examined 21 specimens, 12 males and nine females, from the study area.

The invaria group

Members of this group are characterised by essentially simple male genitalia. Anal sclerite

large, with deep root. Aedeagus with two lateral arms, complex median shaft, and specialised

ventral lobes. Tergal strap attached to aedeagal base; expanded into sclerotized apical band.

Rhyacophilidae and Limnephilidae

25

There are two main branches of this group (Ross, 1956): one is located in eastern North

America, the other in western North America. The western branch includes five species, of

which two are dealt with here.

Rhyacophila bifila Banks, 1914

(Fig. 9a, 9b, 42-46, 109)

Rhyacophila bifila Banks, 1914:201. (Type locality: Vernon, British Columbia). Essig,

1926:176. Betten, 1934:135. Milne, 1936:91, 110. Ross, 1944:291. Ross, 1947:127.

Denning, 1948a:98. Ross, 1952:43. Ross, 1956:88, 118. Fischer, 1960:76. Denning,

1963:244. Smith, 1965:242-243. Denning, 1965b:694. Smith, 1969:658,660-661,673.

Schmid, 1970:60, 125.

Males of this species are distinguished from males of other species in the invaria group by

long, thin, conical, very dark lateral arms of aedeagus; by heavy, blade-like ventral lobes.

Segment X is divided to two simple, lateral, tergal flaps (Fig. 42, 43). Females are distin-

guished by ventral lobes of segment VIII long, thin, arcuate, in ventral aspect (Fig. 46)

arising from separate bases.

Description. — Antennae brown; each article annulated dark brown distally. Vertex of

head very dark chocolate-brown. Thorax brown, to dark chocolate-brown dorsally. Legs yel-

low, distal ends of articles dark brown. Spurs brown. Fore wing length of males 10 mm; dark

chocolate-brown, irrorate; stigmatic area almost black. Venation of fore and hind wings as

in Fig. 9a, 9b.

Male genitalia. (Specimen from Canmore, Alberta). Tergum IX projected well posterad;

remainder of segment IX narrow, tapered slightly ventrad (Fig. 42). Clasper short, massive;

basal article roughly trapezoidal, mesal face occupied by two, almost contiguous, short

ridges. Distal article of clasper polygonal, distal edge slightly concave; mesal face ridged,

partly with fine setae. Segment X divided, as pair of flared, dorso-lateral plates; ventro-

laterad of each plate is single, large, rounded lobe which appears as dorsal extension of

tergal strap. Anal sclerite divided mesally (Fig. 43), arched dorsad. Median shaft of aedeagus

thick, with rounded dorsal lobe overhanging tip (Fig. 44); attached ventrad of median shaft

are two massive, folded, sclerotized lobes terminated by short, stout, acuminate plates with

tips directed dorsad. Lateral arms of aedeagus long, conical, attached at aedeagal base, with

distal quarter very finely attenuated as a hair.

Female genitalia. (Specimen from Canmore, Alberta). Segment VIII short, sclerotized,

with short, membranous distal portion enclosed by short, widely separated, blunt, dorsal

lobes, and blunt, vertically thin, ventral lobes attached to polygonal sternum (Fig. 45, 46).

Dorsal and ventral lobes connected by thin lateral band anterad of which segment VIII is

weakly sclerotized. Segments IX-XI short, stout. Cerci represented by two minute papillae.

Spermathecal sclerites absent.

Notes on biology. — This species emerges from a great variety of streams and rivers,

ranging from fast and turbulent to slow, from boulder strewn to pebble bottomed. Denning

(1965) records it from clear, cold mountain streams. Smith (1968) records the Idaho flight

period of the adults as late June to early September, peaking in late June. My records give

the Alberta flight period as May 22 to August 23. Smith (1968) states that the species is

crepuscular and its members are attracted to light.

Geographical distribution. — The known range of this species extends from British Colum-

bia and Alberta to California and Wyoming (Fig. 109). In Alberta this species is found in the

lower river courses, between 3,000’ and 5,000’, of the mountains and foothills.

I have examined 161 specimens, of which 102 were males, and 59 females.

26

Nimmo

Rhyacophila coloradensis Banks, 1 904

(Fig. 10a, 10b, 47-50, 110)

Rhyacophila coloradensis Banks, 1904a: Plate 1, Fig. 7. (Type locality: Las Vegas, New

Mexico). Banks, 1905:10. Banks, 1907a:41 . Ulmer, 1907a:210. Banks, 1911:354. Dodds

and Hisaw, 1925b:386. Essig, 1926:176. Muttkowski, 1929:192. Betten, 1934:135.

Milne, 1936:91, 111. Ross, 1938b:5. Ross, 1944:291. Denning, 1948a: 101. Ross, 1952:

43. Ross, 1956:88. 1 18. Schmid, 1958:12. Fischer, 1960:78. Denning, 1965b:691.

Smith, 1968:658, 661, 663, 673. Unzicker, 1868:4, 18, 44. Schmid, 1970:60, 125.

Rhyacophila stigmatica Banks, 1904a: 108. (Type locality: Las Vegas, New Mexico). Ulmer,

1905a:72. Banks, 1905:10. Fischer, 1960:78.

Rhyacophila anomala Banks, 1924:444. (Type locality: Tolland, Colorado). Dodds and

Hisaw, 1925b:386. Betten, 1934:135. Milne, 1936:111. Fischer, 1960:78.

Males of this species are distinguished from males of R. bifila Banks by bilobed, dorso-

lateral sclerites of segment X (Fig. 47, 48). Females are distinguished by paired ventral

lobes of segment VIII attached to common pedicel (Fig. 50) in R. coloradensis.

Description. — Antennae brown. Vertex of head virtually black. Thorax deep chocolate-

brown, to virtually black dorsally. Spurs dark brown. Fore wing length of male 10 mm; dark

chocolate-brown, irrorate, stigmatic area solid brown. Hind wings clear, dark brown distally,

stigma slightly darker. Venation of fore and hind wings as in Fig. 10a, 10b.

Male genitalia. (Specimen from Chancellor Peak, Yoho, British Columbia). Segment IX

virtually rectangular laterally, slightly wider dorsally than ventrally (Fig. 47), traversed by

lateral sutures. Basal article of clasper short, stout, almost square; mesal face with short

ventral ridge. Distal article of clasper massive, polygonal, with slightly concave mesal face

with distal half with short setae. Dorso-lateral sclerites of segment X mesally concave, with

dorsal hooks (Fig. 47, 48). Anal sclerite cleft mesally, arched dorsad, paralleling sclerites of

segment X. Tergal strap terminated as two small lobes laterad of anal sclerite. Median shaft

of aedeagus very similar to that of R. bifila Banks (Fig. 44, 49), but ventral lobes massive,

rounded, folded, sclerotized, not acuminate distally; lateral arms rod-like, tipped with mi-

nute spines (Fig. 49).

Female genitalia. (Specimen from Chancellor Peak, Yoho, British Columbia). Sclerotized

basal portion of segment VIII similar to that of R. bifila Banks laterally, except ventral

lobes horizontal, not vertical; in ventral aspect both lobes arise from common pedicel

(Fig. 50).

Notes on biology. — This species frequents both the fast, turbulent mountain creeks with

boulder beds, and the slower, smoother flowing, pebble-bottomed creeks. Smith (1969)

gives the Idaho flight period as March, April, May, and September, indicating two peaks of

emergence, one in Spring, the other in Fall. My records indicate only one period of emer-

gence in Alberta, albeit rather an extended one, from May to August, peaking in May (May

7 to August 30). I have one record, from Banff, Alberta, on April 17, 1915, by N. B. Sanson.

Smith (1968) indicates that the adults are most active at dusk in Idaho, but start flying in

the afternoon.

Geographical distribution. — The known range of this species extends from Alberta and

British Columbia to California, New Mexico, and all intervening states except Nevada (Fig.

1 10). It appears to be truly widespread throughout the Cordillera. In Alberta the species is

found in the foothills, but primarily in the mountains. In altitude it ranges from at least

3,000’ to about 6,000’. Dodds and Hisaw (1925b) recorded it from Colorado at between

6,000’ and 11,000’.

I have examined 268 specimens, 127 males and 141 females, from the study area.

Rhyacophilidae and Limnephilidae

27

The hyalinata group

Males of species in this group are characterised by wide apical band of tergal strap pro-

jecting beyond ventral corner of tergum X (Fig. 51); by truncate anal sclerite with definite

root, and by simple tergum X (Ross, 1956). There are four species in this group of which

one is known to occur in Alberta.

Rhyacophila hyalinata Banks, 1905

(Fig. 11a, lib, 51-53, 111)

Rhyacophila hyalinata Banks, 1905:10. (Type locality: southwestern Colorado). Banks,

1907a:41. Ulmer, 1907a:201. Banks, 1911:354. Dodds and Hisaw, 1925b:386. Essig,

1926:177. Betten, 1934: 135. Milne, 1936:96, 104, 111. Ross, 1938b:6. Ross, 1944:291.

Denning, 1948a: 101. Ross, 1956:88, 92, 118. Schmid, 1958:17. Fischer, 1960:96. Smith,

1968:658, 663-664, 673. Schmid, 1970:59, 124.

Males of this species are distinguished from those of other species by dorsally arched

postero-dorsal extension of segment IX; by shape of distal article of claspers, which are

parallel sided but bowed ventrad (Fig. 51), and by complex aedeagus (Fig. 52).

Description. — Antennae red-brown. Vertex of head deep chocolate. Thorax chocolate-

brown, to darker chocolate dorsally. Spurs brown. Fore wing length of male 13.7 mm; dark

purplish-brown, especially on veins; cells lighter, irrorate; stigma very thick. Venation of

fore and hind wings as in Fig. 11a, lib.

Male genitalia. (Specimen from Vicary Creek, north of Coleman, Alberta). Segment IX

wide; anterior edges bowed anterad, posterior edges angular. Postero-dorsal edge developed

as triangular, cup-like process overtopping segment X (Fig. 51). Basal article of clasper

short, rectangular. Distal article bowed ventrad; swollen tip with fine hairs. Segment X of

two large, warped, dorsal plates fused diagonally along mesal faces. Anal sclerite long, thin,

angular laterally, apical band of tergal strap engaged distally. Tergal strap horizontal, lateral

edges curled dorsad, apical band in form of recurved distal horn. Median shaft of aedeagus

heavily sclerotized (Fig. 52) with distal tip overhung by dorsally concave lobe; acute tri-

angular plate flanked basally by two lateral pads ventrad to ejaculatory duct. Lateral arms

attached to membranous bases; each clavate, long, terminated by small, acuminate spine.

Median shaft and lateral arms of aedeagus distad of massive membranous base overhung

basally by acuminate, ventrally concave plate.

Female genitalia. (Specimen from Vicary Creek, north of Coleman). Basal portions of

segment VIII sclerotized, flanked laterally by pair of lateral flaps, two acute triangular

lobes ventrad (Fig. 53). Two lightly sclerotized, dorso-lateral plates posterad. No evident

spermathecal sclerites, or cerci.

Notes on biology. — This species frequents the faster, more turbulent types of mountain

i stream with rocky beds; it is also, however, found in the faster, smoother flowing, pebble-

bottomed streams. Smith (1968) records the adult flying season as late June to early August

| in Idaho. My records give a total span from July 5 to September 12 in Alberta, with the

peak in later July.

Geographical distribution. — The known range of this species extends from Alberta and

| British Columbia to California and Colorado. The species is widespread in the Cordillera

(Fig. 111). In Alberta it appears to be confined to the mountain areas. In altitude it ranges

i between 3,500’ and 6,000’. Dodds and Hisaw (1925b) record it from Colorado at 9,000’ to

11,000’.

I have examined 428 specimens, 270 males and 158 females, from the study area.

28

Nimmo

The sibirica group

Males of this group are characterised by ventral fusion of lateral arms, forming ventral

membranous base terminated by ovate or elongate scoop with dorsal brush of hair (Fig. 54,

60) (Ross, 1956). Some species have lost this structure (Fig. 62). There are 26 species in this

group, of which four are known to occur in Alberta.

Rhyacophila rickeri Ross, 1956

(Fig. 12a, 12b, 54-56, 112)

Rhyacophila rickeri Ross, 1956:95, 98, 120. (Type locality: Babine River, 50 miles north

of Hazelton, British Columbia). Schmid, 1970:63, 126.

Males of this species are distinguished from males of other species of the sibirica group by

postero-dorsal lobe of tergum VIII, and similar lobe of tergum IX (Fig. 54). Tergal strap

wide, telescoped, with ventral aperture for aedeagus distinctive.

Description. — Antennae brown. Vertex of head dark brown anteriorly, lighter posteri-

orly. Thorax dark yellow-brown laterally, speckled dark brown on lighter ground. Spurs

yellow-brown. Fore wing length of male 1 1.2 mm; light greyish brown, transparent. Vena-

tion of fore and hind wings as in Fig. 12a, 12b.

Male genitalia. (Specimen from Mt. Edith Cavell, Jasper, Alberta). Tergum IX with bul-

bous postero-dorsal process dorsad of segment X (Fig. 54). Segment IX rectangular laterally,

with sinuate posterior edge. Basal article of clasper large, thick, directed postero-dorsad;

distal article long, tapered, setose. Segment X with two distinct cerci whose bases pass

mesad to two mesal protrusions (Fig. 55); thin flaps flank three-arched anal sclerite ventro-

laterad of cerci. Tergal strap prominent, dark, sinuate, with ventral aperture for passage of

aedeagus. Median shaft of aedeagus long, thin, with distinct basal piece (Fig. 54); attached

to large flared base, which flanks it; ventral process membranous, highly extensible, with

dorsally concave tip spinate internally.

Female genitalia. (Specimen from Mt. Edith Cavell, Jasper, Alberta). Proximal third of

segment VIII sclerotized; wide, shallow, longitudinal groove dorsally. Spermathecal sclerite

warped, dorsal surface concave, ventral convex, at posterior end, reversed at anterior end.

Laterally sclerite sigmoid in appearance (Fig. 56).

Notes on biology. — This species inhabits small, very shallow, cold alpine streams, in high

moraine country. These streams are little more than trickles but persist throughout the

year. I have taken specimens at Mt. Edith Cavell, on 1 to 2 feet of fresh snow in early

October.

Geographical distribution. — The known range of this species extends from Alaska to

Alberta and British Columbia (Fig. 1 1 2). I have taken specimens at only two localities, both

at about 7,000’ altitude.

I have examined 23 specimens, 20 males and three females, from the study area.

Rhyacophila vepulsa Milne, 1936

(Fig. 14a, 14b, 57-61, 113)

Rhyacophila vepulsa Milne, 1936:96, 102, 111. (Type locality: Salmon River, Lincoln

County, Oregon). Ross, 1944:291. Denning, 1948a: 106. Ross, 1952:45. Ross, 1956:97,

98, 120. Fischer, 1960:151. Denning, 1963:245. Anderson, 1967:508, 5 17, 5 18. Smith,

1968:658,670, 673. Thut, 1969:895,896, 897, 898. Schmid, 1970:125.

Males of this species are distinguished from males of other species of the sibirica group by

Rhyacophilidae and Limnephilidae

29

massive segment X (Fig. 57, 58) and by asymmetrical distal cup of ventral process of aedea-

gus (Fig. 60). Females are distinguished by angularly offset dorsal and ventral surfaces of

segment VIII (Fig. 61).

Description. — Antennae uniform chocolate-brown. Vertex of head dark chocolate. Tho-

rax dark reddish-brown dorsally, grey-brown laterally. Spurs dark brown. Fore wing length

of male 8.5 mm; uniform dull grey-brown; stigmatic area faint. Venation of fore and hind

wings as in Fig. 14a, 14b.

Male genitalia. (Specimen from Rowe Creek, Waterton National Park, Alberta). Segment

IX slightly narrower ventrally than dorsally (Fig. 57). Basal article of clasper with straight

dorsal edge, sinuate ventral edge. Distal article polygonal, slightly indented distally, ventral

lobe swollen. Segment X massive plaque (Fig. 57, 58) weakly bilobed distally; dorsal surface

irregular in lateral aspect, with distinct anterior hump; ventral surface concave anteriorly

and posteriorly, convex between. Median lobe of aedeagus long, thin, with thinner distal

quarter directed postero-dorsad (Fig. 59); attached ventrad of dorsally arched basal plate

with disto-ventral surface produced as dorsally directed, slender, rounded, distally spinate,

process. Proximad of basal complex of aedeagus is short, rounded, minutely spinate process

directed anterad above aedeagal base. Ventral process of aedeagus attached to membranous

base on slender pedicel; massive, ladle-like, setose around lateral walls; lateral walls asym-

metrical (Fig. 59, 60).

Female genitalia. (Specimen from Rowe Creek, Waterton National Park, Alberta). Basal

half of segment VIII simple, sclerotized, tapered cone; distal edge angular laterally (Fig. 61).

Notes on biology. — I have records of adults of this species taken in the vicinity of large,

turbulent rivers, and from small, pebbly slow streams. Smith (1968) records the larvae from

. . riffles of headwaters streams with compact, pebbly bottoms”, in Idaho. He also gives

the Idaho adult flight season as late July. My records for the study area are few, but they

indicate a flight season in early August (August 1-5). This is not a common species.

Geographical distribution. — The known range of this species extends from Alaska to

California and Montana (Fig. 1 13). In Alberta my records are too few to state with certainty

the range of the species, but it appears to be confined to the mountains. Altitudinally it

occurs from 4,000’ to 6,000’.

I have examined eight specimens, five males and three females, from three localities in

the study area.

Rhyacophila belona Ross, 1948

(Fig. 15a, 15b, 62-64, 112)

Rhyacophila belona Ross, 1948:19-20. (Type locality: East of Logan Pass, Glacier National

Park, Montana). Ross, 1956:97, 120. Schmid, 1970:65, 125.

Males of this species are distinguished from males of other species of the sibirica group by

long, thin, dorsally curved median shaft of aedeagus, with basal two-thirds flanked laterally

by thin up-curved wings (Fig. 62). Females are distinguished by lateral aspect of spermathe-

cal sclerite, which is triangular (Fig. 64).

Description. — Antennae almost black. Vertex of head black. Thorax black-brown, to

black dorsally. Spurs dark brown. Fore wing length of male 8.4 mm; very deep purplish

brown, with patches of slightly lighter purple-brown. Hind wings transparent chocolate-

brown. Venation of fore and hind wings as in Fig. 15a, 15b.

Male genitalia. (Specimen from Mt. Edith Cavell, Jasper, Alberta). Segment IX wide

dorsally, sharply narrowed ventrad (Fig. 62). Basal article of clasper straight, tapered distad;

distal article acute-triangular except for basal constriction, ventral edge thickened. Segment

30

Nimmo

X large, complexly folded, distally bilobed process (Fig. 62, 63) with roots well inside

segment IX. Ventrad of base of segment X is rugose, domed plate. Aedeagus simple, slender,

smoothly up-curved, tapered tube flanked along basal two-thirds by thin, dorsally curved,

lateral wings (Fig. 62).

Female genitalia. (Specimen from Mt. Edith Cavell, Jasper, Alberta). Basal portion of

segment VIII sclerotized, truncated-conical tube (Fig. 64); ventro-posterior edge with two

small processes; dorsad of these are two larger, triangular flaps. Cerci simple, papillate.

Notes on biology. — This species inhabits high alpine brooks in moraine topography.

The flight season of the adults extends from June 1 to July 21.

Geographical distribution. — This species is known only from Alberta and Montana

(Fig. 112). It ranges from 6,000’ to 7,000’.

I have examined 14 specimens, 10 males and four females, from the study area.

Rhyacophila pellisa Ross, 1938a

(Fig. 16a, 16b, 65-68, 114)

Rhyacophila pellisa Ross, 1938a: 118. (Type locality: Cascade Lodge, Rocky Mountain

National Park, Colorado). Ross, 1944:291. Denning, 1948a: 102. Ross, 1956:95, 97, 120.

Fischer, 1960:121. Denning, 1963:244. Anderson, 1967:508. Smith, 1968:666, 673.

Schmid, 1970:64, 125.

Rhyacophila doddsi Ling, 1938:61. (Type locality: Yellowstone Park, Wyoming). Ross,

1944:291.

Males of this species are distinguished from males of other species of the sibirica group by

very long claspers, very short segment X and simple aedeagus with long, thin, tapered

median shaft and simple, setose ventral process (Fig. 66). In dorsal aspect segment X is

simple, rounded, bilobed and deeply cleft (Fig. 64); another similar species which may

eventually be found in Alberta has similar segment X but with only very shallow cleft.

Females are distinguished by long, proximally pierced spermathecal sclerite (Fig. 68).

Description. — Antennae dark brown. Vertex of head almost black. Thorax dark choco-

late-brown, to almost black dorsally. Spurs dark brown. Fore wing length of male 8.4 mm;

uniform deep reddish brown; cells translucent, stigma prominent.

Male genitalia. (Specimen from Gap, near Exshaw, Alberta). Segment IX with robust,

rectangular dorsal area separated from partially keeled, narrower ventral area by fine suture

(Fig. 66). Basal article of clasper massive, tapered distad, variably channeled on mesal face;

distal article somewhat trapezoidal, with thick ventral lobe, minute ventrally hooked, dorsal

lobe. Segment X small, set dorsally on segment IX, deeply cleft mesally, with short, rounded

lobes (Fig. 64); merged ventrally with flat, square-cut plate dorsad of anus. Tergal strap and

apical band as smoothly curved distal hook. Median shaft of aedeagus long, thin, tapered,

recurved tube (Fig. 67); ventral process long, with membranous base, short sclerotized

pedicel, and slightly wider, setose, distal body.

Female genitalia. (Specimen from Gap, near Exshaw, Alberta). Basal half of segment VIII

strongly sclerotized (Fig. 68), dorsal surface slightly shorter than ventral. Spermathecal

sclerite in ventral aspect long, thin, clavate anteriorly; anterior end pierced; posterior end

slightly expanded, darker, with two-arched hyaline line near tip.

Notes on biology. — This species frequents streams ranging from small, pebble-bottomed

brooks, to larger, fast, rocky rivers. The Idaho flight season is given by Smith (1968) as late

July to early August. The Alberta flight season is from July 19 to August 31. Smith states

that daily flight is confined to late morning.

Geographical distribution. — The known range of this species extends from Alberta to

Rhyacophilidae and Limnephilidae

31

California and Colorado (Fig. 1 14). In Alberta it is largely confined to the mountain areas,

but appears to extend eastward in the foothills. Its altitudinal range is 3,500’ to 6,000’.

I have examined 142 specimens, 98 males and 44 females, from the study area.

The betteni group

This group is characterised, in males, by unusually long lateral arms of aedeagus; these are

ventral in position, and fused in some species. Of the eight known species of this group, two

are known to occur in the study area.

Rhyacophila vaccua Milne, 1936

(Fig. 13a, 13b, 69-74, 116)

Rhyacophila vaccua Milne, 1936:94-95, 102, 111. (Type locality: Cultus Lake, British

Columbia). Ross, 1944:291. Denning, 1948a:102. Ross, 1952:45. Schmid and Guppy,

1952:41. Ross, 1956:99, 100, 121. Fischer, 1960:150. Denning, 1963:245. Smith, 1968

658, 668, 673. Thut, 1969:895, 896, 897, 898. Schmid, 1970:68, 126.

Rhyacophila complicata Ling, 1938:60. Ross, 1944:291.

Rhyacophila bruesi Milne and Milne, 1940: 153-156. Ross, 1944:291 .

Males of this species are distinguished from males of other species of the betteni group by

abrupt ventral narrowing of segment IX (Fig. 69), by dorsally curved process of anal sclerite,

and by narrow tenth segment (Fig. 70). As the female of R. chilsia Denning is unknown I

cannot give comparative details within the group. However, females of this species are dis-

tinguished from all others by distinctive sternum of segment VIII (Fig. 73, 74), and by

complex spermathecal sclerites (Fig. 73).

Description. — Antennae dark brown. Vertex of head uniformly very dark brown. Thorax

dark grey-brown, mottled by almost white areas. Spurs dark brown. Fore wing length of

male 10.4 mm; dark grey -brown interspersed with hyaline areas. Venation of fore and hind

wings as in Fig. 13a, 13b.

Male genitalia. (Specimen from Red Earth Creek, Banff National Park, Alberta). Segment

IX with distinct, narrow, sternum (Fig. 69); tergum large, widened dorsally, dorsal surface

sloped antero-ventrad. Basal article of clasper rectangular laterally, with mesal shelf at base

joined medially with identical member of opposing clasper. Distal article bifid; dorsal lobe

acuminate, with basal median flange overlapped with basal lateral flange of thumb-like ven-

tral lobe. Segment X small, recessed dorsally into segment IX, channeled longitudinally along

dorsal surface (Fig. 70). Anal sclerite small, bilobed distally, curved dorsad. Tergal strap large,

curved, with apical band horizontal, attached to segment X disto-ventrally. Median shaft of

aedeagus long, thick, distal end cleft horizontally, with rounded dorsal lobe overhanging

ejaculatory duct (Fig. 71); median shaft completely shielded dorsally by thin, arched, dis-

tally acuminate plate (Fig. 71, 72); lateral arms fused except for heavily spinate distal lobes.

Female genitalia. (Specimen from Gap, near Exshaw, Alberta). Basal portion of segment

VIII heavily sclerotized, partly free distally from membranous portion, very deeply cleft

dorsally and ventro-laterally (Fig. 73, 74); sternum distinct from tergum, posterior edge

produced posterad as acuminate-triangular lobe, with v-cleft distally. Spermathecal sclerites

two long, slender, ventral rods, slightly bulbous posteriorly, set close together; dorsad of

these rods is located a large, dorsally arched structure tapered anterad; single unit with light

i and dark areas as in Fig. 73.

Notes on biology. — This appears to be rather a ubiquitous species, being taken in the

! vicinity of a great variety of streams. I have collected specimens in the vicinity of large and

32

Nimmo

small, fast and slow, rocky and pebbly streams. Smith (1968) records the larva from small to

medium, clear streams with mixed rubble bottoms, in Idaho. He also gives the adult flight

season in Idaho as September and October. The Alberta flight season is August 19 to

October 4.

Geographical distribution. - The known range of this species extends from Alberta and

British Columbia to California and Wyoming (Fig. 1 16). In Alberta it is confined largely to

the mountains, with a few records from the foothills. Most records are from south of Banff.

In altitude it ranges between at least 3,500’ and 6,000’.

I have examined 72 specimens, 47 males and 25 females, from the study area.

Rhyacophila chilsia Denning, 1950

(Fig. 75-78, 1 15)

Rhyacophila chilsia Denning, 1950:115-1 16. (Type locality: Maligne Canyon, Jasper, Al-

berta). Ross, 1956:100, 121. Schmid, 1970:68, 126.

Males of this species are distinguished from those of R. vaccua Milne by minute dorsal

lobe of distal article claspers, by segment X well separated dorsally from segment IX, and by

ventrally directed, angular, anal sclerite (Fig. 75).

The only known specimen of this species is a male, from Maligne Canyon, Jasper, Alberta.

Being thus unable to prepare my own drawings, Dr. D. G. Denning very kindly lent me his

original drawings of the species, which I have partly redrawn and present here (Fig. 75-78).

I also present Denning’s original description of the male, altering only his figure numbers to

mine.

Description. - ‘Length [?] 9.5 mm. Fore and hindwing fuscous, veins and pterostigma

somewhat darker. Body, head, palpi and antennae yellowish; legs and spurs luteous. Sixth

and seventh abdominal sternites with a short acute mesal spine.

Genitalia as in Fig. 75-78. Ninth segment gradually widened dorsally, meso-apical margin

of tergum in the form of a sub-acute projection. Tenth tergum narrow plate-like structure;

meso-dorsal margin projected caudad as acute process, pair of small acute mesal projections

close to ventral margin and best discernible from dorso-caudal aspect; mesal surface of

tergum concave; ventral process cleft nearly entire length (Fig. 75), and capable of only

slight dorso-ventral movement; dorsal aspect of tergum as in Fig. 77. Apical segment of

clasper gradually narrowed distally, distal margin cleft to form acuminate digitate process,

setation sparse. Structures in association with aedeagus as in Fig. 78; apex of lateral arms

somewhat asymmetrical, but each divided into four acute projections’ (Denning, 1950).

Notes on biology. — I do not know which part of Maligne Canyon is meant, but the

Maligne River in the area is fast, fairly smooth water on rocky, but not boulder, bottom; in

the canyon itself the river is considerably narrowed, deeper, with frequent pools and cas-

cades. Date of capture of the male was July 23.

Geographical distribution. — The position of the one known locality is indicated in Fig.

1 15. It is at an altitude of 3,800’.

The vobara group

Males of this group are distinguished by apical band of the tergal strap attached directly

to inner ends of anal sclerite (Fig. 79). One of the two known species occurs in Alberta.

Rhyacophila vobara Milne, 1936

(Fig. 17a, 17b, 79-81, 115)

Rhyacophilidae and Limnephilidae

33

Rhyacophila vobara Milne, 1936:94, 102, 111. (Type locality: Cultus Lake, British Colum-

bia). Ross, 1944:291. Denning, 1948a: 106. Ross, 1952:45. Ross, 1956:102, 116, 121.

Schmid, 1958:17. Fischer, 1960:152. Smith, 1968:672, 673. Schmid, 1970:67, 126.

Males of this species are distinguished by thin, high segment IX with sinuate edges (Fig.

79), by massive proximo-mesal swelling of basal article of claspers, and by dorsal toothed

plate at aedeagal base (Fig. 80).

Description. - Antennae dark brown. Vertex of head deep chocolate-brown, warts almost

white. Thorax dark brown. Spurs brown. Fore wing length of male 8.5 mm; pale to dark

chocolate-brown, with large irrorations, primarily at distal ends of peripheral cells. Stigma

weak. Venation of fore and hind wings as in Fig. 17a, 17b.

Male genitalia. (Specimen from Ranger Creek, Jasper Park, Alberta). Segment IX high,

narrow, pinched ventrad of mid-line (Fig. 79). Basal article of clasper sinuate, with massive

bulge on mesal face. Distal article almost square; distal edge irregular, with thick ventral

lobe. Segment X irregular longitudinally folded sclerite. Anal sclerite large, tapered basad,

with dark distal edge; articulated with ventral corner of segment X, which flanks it laterally.

Apical band of tergal strap curved, attached to segment X at ventral edge. Median shaft of

aedeagus short, thin, between two sclerotized, acuminate lateral arms; longer, thin, tubular

process dorsad of shaft (Fig. 80). Aedeagal base roofed over by heavy, dark, flat, plate with

large dorsal thorn; base ventrad of this plate enclosed in lightly sclerotized tube.

Female genitalia. (Specimen from Ranger Creek, Jasper Park, Alberta). Basal portion of

segment VIII sclerotized (Fig. 81) with two pairs of disto-lateral lobes; dorsal lobes rounded,

ventral lobes somewhat triangular.

Notes on biology. — Specimens of this species have been taken in the vicinity of small,

fast, turbulent mountain creeks, small, gravelly alpine trickles, and slow, deep streams

emerging from alpine peat bogs. The flight period of Alberta adults is from July 3 to

September 10.

Geographical distribution. — The known range of this species extends from the Yukon

Territory to Idaho. In Alberta my records are all from the mountains, except for one

locality in the northern foothills. In altitude this species ranges from at least 3,500’ to over

6,000’.

I have examined 36 specimens, 20 males and 16 females, from the study area.

The angelita group

Males of this group are characterised by large dorsal lobe of segment X (Fig. 82), by deep

root of anal sclerite, and by an extra pair of lateral lobes on aedeagus (Fig. 84). One of the

three known species in this group occurs in Alberta.

Rhyacophila angelita Banks, 1911

(Fig. 18a, 18b, 82-85, 117)

Rhyacophila angelita Banks, 191 1:352, 355. (Type locality: Pasadena, California). Essig,

1926:176. Betten, 1934:135. Milne, 1936:92, 103, 110. Ross, 1944:291. Denning,

1948a:97. Ross, 1952:43. Schmid and Guppy, 1952:41. Ross, 1956:102, 121. Schmid,

1958:12. Fischer, 1960:71. Denning, 1963:244. Smith, 1968:658, 660, 673. Schmid,

1970:62, 125.

Rhyacophila bipartita Banks, 1914:201-202. (Type locality: Banff, Alberta). Betten, 1934:

135. Milne, 1936:110. Fischer, 1960:71.

Rhyacophila species 1, Flint, 1962:478.

34

Nimmo

Males of this species are distinguished by large, elliptical bilobed dorsal lobe of segment X

(Fig. 83), and by structure of aedeagus (Fig. 84).

Description. — Antennae yellow. Vertex of head brownish yellow. Thorax brownish

yellow to straw. Spurs yellow-brown. Fore wing length of male 10.1 mm; hyaline, with very

faint yellow-brown pattern; veins dark red-brown. Stigma weak. Venation of fore and hind

wings as in Fig. 18a, 18b.

Male genitalia. (Specimen from Gap, near Exshaw, Alberta). Segment IX very wide dor-

sally; bowed, saddle-like; segment narrowed gradually ventrad; sternum curved anterad at

lateral sutures (Fig. 82). Basal article of clasper long, parallel-sided, sinuate. Distal article

mesally concave, with wide ventro-mesal ledge. Segment X small, enclosed plate, produced

postero-dorsad as massive dorsal plate cleft deeply mesally (Fig. 83). Anal sclerite large, with

deep root, flanking base of segment X. Tergal strap heavy, with laterally triangular apical

band. Median shaft of aedeagus small, tapered, thin (Fig. 84); flanked laterally by pair of

wide, dorsally hooked lobes. Two short, rectangular plates ventrad of median shaft. Lateral

arms long, with membranous, extensible bases and spatulate tips with mesal concavities

setose. Vertically bilobed prominence directed posterad on dorsum of aedeagal base.

Female genitalia. (Specimen from Gap, near Exshaw, Alberta). Basal portion of segment

VIII tapered, sclerotized, truncated cone merged distally with membranous portion (Fig.

85). Dorsal surface of segment VIII base with deep channel; distally extended into dorsal

swelling. Spermathecal sclerite simple, folded longitudinally, attenuated at each end.

Notes on biology. — This species appears to be cosmopolitan in its choice of habitat,

specimens being taken near almost every type of water course available in the moun-

tain area. The adult flight season extends in Alberta from July 7 to October 18. I have

records for May 23 and June 20 also, but most of my records are within the range stated

above.

Geographical distribution. — The known range of this species extends from the Yukon

Territory to California and Colorado (Fig. 1 17). The species has also been recorded from

the northern Appalachians of New Hampshire (Smith, 1968) which, in the present state

of knowledge of this species, is a very isolated record probably representing a post-glacial

remnant of a previously truly transcontinental species.

I have examined 530 specimens, 324 males and 206 females, from the study area.

The verrula group

This group contains only one known species of singular peculiarity. The detailed descrip-

tion following will serve to characterise the group.

Rhyacophila verrula Milne, 1936

(Fig. 19a, 19b, 86-88, 118)

Rhyacophila verrula Milne, 1936:90, 111. (Type locality: Cultus Lake, British Columbia).

Knowlton and Harmston, 1938:286. Ross, 1944:291. Denning, 1948a: 102. Denning,

1948b:22. Ross, 1952:45. Schmid and Guppy, 1952:41. Ross, 1956:108, 122. Schmid,

1958:12. Fischer, 1960:151. Denning, 1963:245. Smith, 1968:658, 671-672, 673. Thut,

1969:894, 895, 896, 897. Schmid, 1970:65, 126.

Rhyacophila oregonensis Ling, 1938:62. (Type locality: Corvallis, Oregon). Ross, 1944:291.

Males of this species are distinguished by trilobed postero-dorsal edge of tergum VIII

(Fig. 86), by fused postero-dorsal lobes of segment IX, hooked ventrad (Fig. 86, 87), and by

curious dorsal process of aedeagal base. Females are distinguished by sclerotized tergum X

Rhyacophilidae and Limnephilidae

35

and by two annular swellings of segment IX (Fig. 88).

Description. — Antennae dark brown, scapes yellow. Vertex of head dark brown antero-

mesally; remainder, and warts, yellow. Thorax mottled yellow and dark brown. Spur formu-

la of males 2, 4, 5; pro-thoracic spurs short, meso-apical spur of hind legs stout, long, dis-

tally bifid, in form of pincers; remainder normal. Spur formula of females 3, 4, 4; fore leg

spurs short, remainder normal. Fore wing length of male 12 mm; light yellow-brown; pattern

somewhat banded, colour alternated with hyaline areas; stigma opaque white. Venation of

fore and hind wings as in Fig. 19a, 19b.

Male genitalia. (Specimen from Gap, near Exshaw, Alberta). Postero-dorsal edge of ter-

gum VIII with projected median lobe dorsad of dorsal strap (Fig. 86, 87); flanked by two

flap-like lateral lobes. Segment IX with narrow, short, dorsal strap ventrad of which postero-

dorsal edges produced as two large, arched lobes fused distally but parted just at tips; each

lobe with blunt, ventral, process basally fused to equivalent member opposite. Postero-

lateral edges of segment IX rolled meso-anterad (Fig. 87). Basal article of clasper massive,

rectangular; distal article small, rounded, concave on mesal face. Segment X small, arched,

dorsal roof with two small distal lobes attached distally to ventral processes of lobes of seg-

ment IX. Anal sclerite as second, internal arch. Median shaft of aedeagus bulbous basally,

thin, tapered distally, in deep, thin walled, sclerotized trough (Fig. 86), connected with

stout, membranous, base; dorsal surface of base with long, slender, finger-like dorsal process,

sinuate, fitted distally with anal sclerite.

Female genitalia. (Specimen from Gap, near Exshaw, Alberta). Segment VIII long, tap-

ered, dilated annularly at distal extremity, lightly sclerotized for most of length (Fig. 88).

Segments IX and X with annular swelling at point of junction. Segment X with posterior

two-thirds of tergum sclerotized. Cerci small, membranous. No evident spermathecal sclerite.

Notes on biology. — Smith (1968) reports that the larvae of this species are totally phyto-

phagous, which sets the species apart from other species of the genus. He reports larvae from

small, cold, pebbled riffles of clear streams. I have usually taken specimens near larger, swift,

deep, boulder strewn mountain creeks, and occasionally from smaller, shallower, pebbly

creeks. Smith (1968) reports the Idaho adult flight period to be September to October,

peaking in September. My records indicate the Alberta flight season to range from August

20 to October 12.

Geographical distribution. — The known range of this species extends from Alaska to

California and Colorado (Fig. 118). In Alberta the species is found in the mountains and

high foothills, between 3,500’ and 4,000’.

I have examined 64 specimens, 47 males and 17 females, from the study area.

The vagrita group

Males of this group are characterised by prominent postero-dorsal, strap-like, lobes of

both segments IX and X (Fig. 89, 93), by small, very simple aedeagi (Fig. 91), and by inter-

locking of segment X, anal sclerite, and apical band of tergal strap; anal sclerite encloses

small, spherical, tip of segment X, and is itself flanked laterally by apical band (Fig. 86, 93).

Both species of this group are known from Alberta.

Rhyacophila vagrita Milne, 1936

(Fig. 20a, 20b, 89-92, 119)

Rhyacophila vagrita Milne, 1936:91-92, 105, 111. (Type locality: Cultus Lake, British

Columbia). Ross, 1944:291. Denning, 1948a: 105. Ross, 1950a:264. Ross, 1952:45. Ross,

36

Nimmo

1956:108, 122. Schmid, 1958:12. Fischer, 1960:150. Smith, 1968:658, 668, 670, 673.

Thut, 1969:895, 896, 897, 898. Schmid, 1970:45, 122.

Males of this species are distinguished from males of other species of the vagrita group by

distal article of claspers with acuminate dorsal lobe and thick, rounded, fleshy ventral lobe,

and by bilobed postero-dorsal process of segment IX (Fig. 89, 90). Only the female of this

species is known so no comparison can be made.

Description. — Antennae dark brown. Vertex of head dark brown anteriorly, laterally;

postero-mesally light. Thorax dark grey-brown, to richer reddish brown dorsally. Fore wing

length of male 9 mm; light grey-brown with scattered, rectangular, hyaline windows; stigma

distinct, brown. Venation of fore and hind wings as in Fig. 20a, 20b.

Male genitalia. (Specimen from Snaring River, Jasper National Park, Alberta). Segment IX

wide, with narrower dorsal strap peaked along anterior edge (Fig. 89); posterior edge of

dorsal strap developed as long, sinuate, distally bilobed, thin, strap-like process (Fig. 89, 90).

Basal article of clasper with very narrow base, approximately triangular, with mesal ledge

basally. Distal article of clasper lanceolate, curved, acuminate dorsally, rounded and fleshy

ventrally. Segment X with long, sinuate, dark brown dorsal process immediately ventrad of

similar process of segment IX (Fig. 89, 90); ventrad of base of lobe are two small, peg-like

cerci between which is located sclerotized, strap-like body of segment X, terminated be-

tween lateral horns of anal sclerite. Tergal strap sinuate, with apical band terminated at an-

tero-ventral corner of anal sclerite. Aedeagus minute; ejaculatory duct directed dorsad from

dorsal plate; dorsal process short, pick-like laterally; joined to sclerotized base (Fig. 91).

Female genitalia. (Specimen from Snaring River, Jasper National Park, Alberta). Segment

VIII with short sclerotized tube at base (Fig. 92), with sigmoid posterior edges. Spermathe-

cal sclerite long, thin, irregular, with minute, pick-like posterior tip; located centrally in

membranous internal tube. No evident cerci.

Notes on biology. — Little is known of this species. The one locality at which I have taken

it was adjacent to a wide, swift, smooth-flowing river with pebble and small boulder bottom.

Geographical distribution. — The known range of this species extends from Alberta and

British Columbia to Utah (Fig. 1 19). In Alberta the two known localities are at the bottoms

of major valleys at about 3,500’.

I have examined one specimen of each sex from the study area.

Rhyacophila milnei Ross, 1950

(Fig. 93-95, 119)

Rhyacophila milnei Ross, 1950a:264. (Type locality: Banff, Alberta). Ross, 1956:108, 122.

Schmid, 1970:45, 122.

Males of this species are distinguished from those of R. vagrita Milne by trilobed postero-

dorsal process of segment IX (Fig. 93, 94), by saddled dorsal area of segment IX, and by

parallelogram shaped distal articles of claspers.

Description. — Antennae brown. Vertex of head very deep brown, almost black. Thorax

deep reddish brown, to very dark brown dorsally. Spurs yellow-brown; spurs of middle and

hind legs long and heavy. Fore wing length of male 8 mm; translucent red-brown. Venation

of fore and hind wings identical to that of R. vagrita Milne.

Male genitalia. (Specimen from Banff, Alberta; paratype, in Illinois Natural History Sur-

vey). Segment roughly rectangular laterally (Fig. 93); dorsal surface longitudinally chan-

neled, with posterior edge of two lateral arches over minute, stubby, cerci; postero-dorsal

edge produced posterad as long, curved, trilobed strap; lateral lobes shorter than medial.

Basal article of clasper slightly narrowed basad, with thick, rounded dorsal edge, and thin,

Rhyacophilidae and Limnephilidae

37

ledged, ventro-mesal edge. Distal article of clasper fused to basal article; parallelogram-like,

with distinct acuminate dorsal lobe, and thick, rounded ventral lobe. Segment X with long,

thin, sinuate, dorsal process immediately ventrad of that of segment IX (Fig. 93, 94); ventral

surface concave. Ventral body of segment rounded, tubular, semi-circular laterally. Anal

sclerite large, enclosing tip of segment X, open dorsally. Aedeagus minute; ejaculatory duct

minute, directed postero-dorsad from membranous base below evenly rounded dorsal pro-

cess; dorsal groove extended from base of aedeagus to top of dorsal process (Fig. 95).

Female unknown.

Geographical distribution. — The known range of this species is presently restricted to the

type locality, which is simply ‘Banff, Alberta’ (Fig. 1 19).

Only one male of this species, a paratype, has been examined. The date of capture was

September 5.

Unassociated females

Rhyacophila species 1

(Fig. 23a, 23b, 100-101, 121)

Description. — Antennae dark brown. Vertex of head deep reddish brown. Thorax light

red-brown, to dark chocolate dorsally. Spurs brown. Fore wing length of female 8.0 mm;

pale, clear brown, veins dark, stigma brown; hind wings much the same colour as fore,

except anal area hyaline. Venation of fore and hind wings as in Fig. 23a, 23b.

Female genitalia. (Specimen from Lusk Creek, Kananaskis, Alberta). Basal half of seg-

ment VIII sclerotized; posterior edges angular, dorsal edge offset anterad (Fig. 100). Sper-

mathecal sclerite laterally spindle-like with long membranous sack attached anteriorly; distal

end and adjacent edges dark; ventrally tip square-cut; double hook pattern just anterad of

tip of dark coloration (Fig. 101), with shaft of hood faded anterad.

Notes on biology. — The two creeks from which I have taken specimens of this species are

shallow, slow riffled streams on small pebble beds. The dates of capture were May 18 in

southeastern British Columbia, and July 15 at Lusk Creek, in the Kananaskis valley of

Alberta.

Geographical distribution. — Only two records are available for this species at present

(Fig. 121). Both are from low altitudes, about 3,000’

Rhyacophila species 2

(Fig. 22a, 22b, 102-103, 121)

Description. — Antennae pale straw, scapes slightly darker. Vertex of head yellow, to

pale red-brown between ocelli. Thorax reddish straw, to pale red-brown dorsally. Spurs

brown. Fore wing length of female 10.4 mm; pale straw-yellow, no discernible pattern

except for opaque stigmatic area. Venation of fore and hind wings as in Fig. 22a, 22b.

Female genitalia. (Specimen from Gap, near Exshaw, Alberta). Basal two thirds of seg-

ment VIII very weakly sclerotized; segment long, tapered, terminated by annular swelling

(Fig. 102). Spermathecal sclerite laterally spindle shaped, with anterior end attenuated

(Fig. 102, 103); main body folded dorsad with thick lateral edges faded anterad.

Notes on biology. — The one known locality for this species is a fast, smooth mountain

creek on a medium sized boulder bed. Date of capture is September 25.

Geographical distribution. — The one locality is Rapids Creek, at the Trans-Canada High-

way, Gap, near Exshaw, Alberta (Fig. 121).

38

Nimmo

Fig. 4-10. Fore (a) and hind (b) wings of males of Rhyacophila. 4. R. alberta Banks. 5. R. tucula Ross. 6. R. glaciera

Denning. 7. R. vofixa Milne. 8. R. acropedes Banks. 9. R. bifila Banks, 10. R. coloradensis Banks.

Rhyacophilidae and Limnephilidae

39

16a

16b

Fig. 11-16. Fore (a) and hind (b) wings of males of Rhyacophila. 11. R. hyalinata Banks

vaccua Milne. 14. R. vepulsa Milne. 15. R. belona Ross. 16. R. pellisa Ross.

12. R. rickeri Ross. 13.

R.

40

Nimmo

Fig. 17-23. Fore (a) and hind (b) wings of males, except where otherwise stated, of Rhyacophila. 17. R. vobara Milne.

18. R. angelita Banks. 19. R. verrula Milne. 20. R. vagrita Milne. 21. R. vemna Milne. 22. R. species 2, female.

23. R. species 1, female.

Rhyacophilidae and Limnephilidae

41

Fig. 24-38. Genitalia of Rhyacophila species. R. alberta Banks, 24. Male, lateral aspect. 25. Aedeagus, lateral aspect.

26. Female, sclerotized base of segment VIII, lateral aspect. 27. Female, sclerotized base of segment VIII, posterior

edge, ventral aspect. R. tucula Ross, 28. Male, lateral aspect. 29. Aedeagus, lateral aspect. 30. Female, sclerotized base

of segment VIII, lateral aspect. 31. Spermathecal sclerite, ventral aspect. R. glaciera Denning, 32. Male, lateral aspect.

33. Aedeagus, dorsal aspect. 34. Female, sclerotized base of segment VIII, lateral aspect. R. vofixa Milne, 35. Male,

lateral aspect. 36. Segment X, dorsal aspect. 37. Aedeagus, lateral aspect. 38. Female, segment VIII, lateral aspect.

A. S. - anal sclerite. Cl. - Clasper. L. A. - lateral arm of Aedeagus. M. S. - median shaft of aedeagus. S. S. - sper-

mathecal sclerite. Scale bar for each species adjacent to lateral aspect of male genitalia; scale uniform for all drawings

of a species. No scale given when only female known.

42

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Fig. 39-53. Genitalia of Rhyacophila species. R. acropedes Banks, 39. Male, lateral aspect. 40. Aedeagus, lateral aspect.

41. Female, base of segment VIII, lateral aspect. R. bifila Banks, 42. Male, lateral aspect. 43. Segment X, dorsal aspect.

44. Aedeagus, lateral aspect. 45. Female, segments VIII-IX, lateral aspect. 46. Sclerotized base of segment VIII, ventral

aspect. R. coloradensis Banks, 47. Male, lateral aspect. 48. Segment X, dorsal aspect. 49. Aedeagus, lateral aspect.

50. Female, ventral lobes of segment VIII, ventral aspect. R. hyalinata Banks, 51. Male, lateral aspect. 52. Aedeagus,

lateral aspect. 53. Female, segment VIII, lateral aspect. A. B. - apical band of tergal strap. T. S. - tergal strap.

Rhyacophilidae and Limnephilidae

43

Fig. 54-68. Genitalia of Rhyacophila species. R. rickeri Ross, 54. Male, lateral aspect, including aedeagus. 55. Male,

posterior aspect. 56. Female, segments VIII-IX, lateral aspect. R. vepulsa Milne, 57. Male, lateral aspect. 58. Segment X,

dorsal aspect. 59. Aedeagus, lateral aspect. 60. Aedeagus, ventral process tip, dorsal aspect. 61. Female, segment VIII,

I base, lateral aspect. R. belona Ross, 62. Male, including aedeagus, lateral aspect. 63. Segment X, dorsal aspect. 64.

Female, segment VIII, base, lateral aspect. R. pellisa Ross, 65. Male, lateral aspect. 66. Segment X, dorsal aspect.

67. Aedeagus, lateral aspect. 68. Female, segment VIII, ventral aspect.

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Fig. 69-85. Genitalia of RhyacophUa species. R. vaccua Milne, 69. Male, lateral aspect. 70. Segment X, dorsal aspect

71. Aedeagus, lateral aspect. 72. Aedeagus, dorsal aspect. 73. Female, segment VIII, lateral aspect. 74. Sclerotized

base of segment VIII, ventral aspect. R. chilsia Denning, 75. Male, lateral aspect (redrawn from Denning). 76. Anal

sclerite, distal edge, dorsal aspect (after Denning). 77. Segment X, dorsal aspect (after Denning). 78. Aedeagus, lateral

aspect (after Denning). No scale given. R. vobara Milne, 79. Male, lateral aspect. 80. Aedeagus, lateral aspect. 81. Female,

segment VIII, base, lateral aspect. R. angelita Banks, 82. Male, lateral aspect. 83. Segment X, dorsal aspect. 84. Aedeagus,

lateral aspect. 85. Female, segment VIII, base, lateral aspect.

Rhyacophilidae and Limnephilidae

45

Fig. 86-103. Genitalia of Rhyacoplnila species. R. verrula Milne, 86. Male, lateral aspect. 87. Segments VIII, and IX,

dorsal aspect. 88. Female, segments VIII-XI, lateral aspect. R. vagrita Milne, 89. Male, lateral aspect. 90. Posterior

lobes, segments IX and X, dorsal aspect. 91. Aedeagus, lateral aspect. 92. Female, segment VIII, lateral aspect. R. milnei

Ross, 93. Male, lateral aspect. 94. Posterior lobes, segments IX and X, dorsal aspect. 95. Aedeagus, lateral aspect.

R. vemna Milne, 96. Male, lateral aspect. 97. Segment X, dorsal aspect. 98. Aedeagus, lateral aspect. 99. Female, segment

VIII, ventral aspect. R. species 1, 100. Female, segment VIII, base, lateral aspect. 101. Posterior end of spermathecal

sclerite, ventral aspect. R. species 2, 102. Female, segment VIII, lateral aspect. 103. Spermathecal sclerite, ventral aspect.

No scale given when only female known.

46

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Fig. 104-109. Maps of geographical distribution of Rhyacophila species in Alberta, and North America.

Rhyacophilidae and Limnephilidae

47

Fig. 110-115. Maps of geographical distribution of Rhyacophila species in Alberta, and North America.

48

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Fig. 116-121. Maps of geographical distribution of Rhyacopkila species in Alberta, and North America.

Rhyacophilidae and Limnephilidae

49

THE FAMILY LIMNEPHILIDAE KOLENATI

This family is represented in Alberta and eastern British Columbia by 9 1 species belong-

ing to 26 genera. The names of the genera are presented in Table 1 according to the scheme

used by Schmid (1955).

The species in the study area are discussed individually in the text. Within genera, divi-

sions to subgenera, or groups, follow Schmid (1955). Immediately following is a synopsis

of familial characteristics translated and greatly condensed from Schmid (1955). The synop-

ses of the subfamilial groups, including genera, are also derived and condensed from Schmid

(1955), as are the keys to all taxa to the generic level. Schmid (1955) should be consulted

for a complete exposition of the family and its constituent taxa. The keys include only

those groups found in the study area. Keys are provided to the males of the species of each

genus and are original unless stated otherwise. In the family as a whole the keys are

applicable primarily to the males, as male genitalic characters are used sufficiently often to

exclude the females. This situation is due to an insufficient knowledge of the females of the

various taxa within the family, only the males being known for many species. The situation

is aggravated by the relatively great homogeneity of body characters within the family,

other than in the genitalia. Keys are provided to the females of some genera, but at present

the best way to identify females is by association with males.

Some species are holarctic in distribution, ranging as far west as Europe. For such species

there is a very large European literature. In this work only the literature pertaining to such

species in North America is given in detail; the reader is referred, at the end of the literature

and synonymy list for each species, to the appropriate volume of Fischer’s ‘Trichopterorum

Catalogus’ for a complete listing.

Character synopsis of the Limnephilidae. — Ocelli three. Antennae as long as, or little

shorter than fore wings; thickened, not fine; scapes generally as long as head, cylindrical,

thickened. Maxillary palpi of straight, sub-cylindrical articles; with three articles in males,

five in females. Pronotum short. Dorsal line (i.e. pale coloured median stripe from inter-

ocellar space of head to metanotum) present or absent. Legs commonly long, heavy, not

hairy or silky but spinate; spines most abundant on the tibiae and tarsi, generally black.

Spurs yellowish, modified or not. Spur formula 1,3,4 to 1,1,1, variable intergenerically and

interspecifically. Femora and tarsi of fore legs of males of certain genera provided with

opposing brushes of short, stout, black spines. Fore wings of some genera basally narrow,

distally expanded at stigma, with widely rounded apex; hind wings much larger, with well

developed anal area (some dicosmoecine genera and all Limnephilinae). In all other genera

the hind wing is reduced to varying degrees of similarity to the fore wing by reduction of

anal area. Venation generally as in Fig. 122. Cross-vein R1-R2 of fore wing absent; f4 absent,

median cell open. R1 of fore wing with distal kink of varied intensity, followed by smooth

curve to encompass stigma, if present. Radial sector four branched, to encompass cells fl

and f2; discoidal cell longer than wide. Median and cubital veins each three branched to

delimit f3 and f5 respectively. Thyridial cell between M and Cul. All veins except Sc and R1

connected by cross-veins to form irregular, dispersed line known as anastomosis or chord.

Venation of hind wing almost identical to that of fore wing, except chord more dispersed.

Anal veins five in number. R1 without distal kink and bow, generally parallel to Sc.

Tergum VIII of male unmodified of variously developed and clothed with spines or hairs.

Segment IX single, rigid, heavily sclerotized tube of fairly uniform width all round, or of

varied widths; ventral area produced posterad to form shelf below aedeagus, or not pro-

duced; dorsal area reduced to strap of varied widths, or obsolete. Postero-ventral edges

recessed, or not. In posterior aspect segment IX divided to dorsal and ventral cavities by

50

Nimmo

mesally directed extensions of lateral walls, or not divided. Claspers movable or fused to

segment IX; composed of one or two articles, horizontal; if of two articles then curved,

pincer-like; if of one article then short, plaque-like. Segment X small or large, with or with-

out many lobes or branches.

Female genitalia less varied than those of males. Segment VIII unmodified or, in some

genera, with slight concavity in sternum. Segment IX short, cylindrical, cleft ventrally, or

lateral walls shortened to isolate ventral angles. In most genera segment IX of distinct terga

and sterna. Segment X more or less incised, sclerotized tube distinct from segment IX, or

fused to it. Supra-genital plate present except in a few genera, dorsad of genital cavity.

Vaginal aperture on sternum IX or between sterna VIII and IX. Vulval scale ventrad of

vaginal aperture, either as simple chitinous pad or strongly sclerotized, trilobed structure.

Key to the Subfamilies and certain Genera of Limnephilidae in Alberta and eastern British

Columbia

la. Discoidal cell of hind wing open distally, or R1 of fore wing united to Sc by cross-

vein terminated at wing edge (Fig. 129a) Apataniinae, p. 64.

lb. Discoidal cell of hind wing open basally; i.e. RS divided from wing base; or F3 of

hind wing absent, or both (Fig. 130, 131) Neophylacinae, p. 71.

lc. Discoidal cell of hind wing closed; f3 present 2

2a. (lc) Chord of fore wing a single, irregular line (Fig. 133-135)

Homophylax (Pseudostenophylacinae), p. 78.

2b. Chord of fore wing in two distinct lines (Fig. 138) 3

3a. (2b) Fore wing reddish, narrow. Male maxillary palpus very large

Chyranda (Limnephilinae), p. 143.

3b. Fore wing not narrowed. Maxillary palpus of normal size 4

4a.(3b) Fore wing reddish, large, rounded; fl with long common boundary with discoidal

cell (Fig. 128). Mesal face of male clasper with one or more sclerotized spines

(Fig. 195) Ecclisomyia (Dicosmoecinae), p. 61.

4b. Characters otherwise 5

5a.(4b) Clasper of male two-articled and movable. Vaginal aperture of female on segment

IX Dicosmoecinae, p. 50.

5b. Clasper of male of one article only, and fused to segment IX. Vaginal aperture of

female between segments VIII and IX Limnephilinae, p. 81.

The Subfamily Dicosmoecinae Schmid

Synopsis of characters. — Head very large; eyes large. Pronotum short; macrochaetae

well developed. Spur formula 1,3,3; 1,2,2; or 1,3,4. Fore wings medium to large, para-

bolic. Hind wings much larger than fore wings or only slightly larger; anal edge smoothly

convex. Venation of species in study area as in Fig. 122-128, basically simple and un-

modified.

Male genitalia complex, varied. There is feeble specialization of appendages, or strong

reduction in their numbers in some genera. Segment IX recessed into segment VIII in very

few genera; of fairly uniform width except for dorsal lobes in certain genera, such as

Dicosmoecus. Segment X not projected in most genera; large, roof-like over the aedeagal

recess. There are four pairs of appendages which are reduced or absent in some genera.

Claspers very large, pincer-like, two-articled, as in Dicosmoecus (Fig. 146), or smaller, with

articles tending to fuse, as in Amphicosmoecus (Fig. 164). In certain genera (e.g. Imania )

the claspers are complex, with distal article missing; not fused to segment IX. Aedeagus

Rhyacophilidae and Limnephilidae

51

highly varied in form and size; long and slender in some genera.

Segment IX of female genitalia of one piece, or of two distinct parts. Dorsal part well

developed, but reduced in certain genera; in form of long tube; appendages absent. Segment

X well developed, in form of simple cone (e.g. Onocosmoecus Fig. 162), bilobed dorsally

with ventral plate. Supra-genital plate present. Vaginal orifice on segment IX. Vulval scale

similar to that of other subfamilies but formed from posterior edge of segment IX, not VIII;

trilobed.

Key to the Genera of Dicosmoecinae in Alberta and eastern British Columbia

la. Fore wing reddish, large, rounded; fl with long common border with discoidal

cell (Fig. 128a) Ecclisomyia, p. 61.

lb. Fore wing with fl short, common border with discoidal cell (Fig. 122-127) .... 2

2a.(lb) Small insects, fore wing length less than 12 mm; fore wing very dark, brownish

black Imania, p. 56.

2b. Much larger insects, fore wing length over 17 mm; fore wing reddish brown or

grey to grey-black 3

3a.(2b) Clasper of male with both articles fused, not articulated. Lateral lobes of female

vulval scale stout, fleshy, with distal ends concave. Supra-genital plate arched

dorsad, projected prominently posterad, free of remainder of genitalia (Fig. 164,

168, 169) Amphicosmoecus, p. 55.

3b. Clasper of male with both articles articulated (Fig. 146). Median lobe of female

vulval scale projected free, between thin, placoid lateral lobes. Supra-genital plate

not as above; not projected freely and prominently (Fig. 151, 152, 156, 162,

163) 4

4a. (3b) Thorax abundantly clothed with silky hairs; large, black insects

Dicosmoecus, p. 5 1 .

4b. Thorax without many silky hairs; smaller, red-brown insects

Onocosmoecus, p. 53.

The Genus Dicosmoecus McLachlan

This genus is represented in the study area by two species.

Synopsis of characters. — Head very large; ocelli large, close-set. Spur formula 1,3,4.

Pleural sclerites, metanotum, and wing bases clothed with long, fine, silky hairs.

Male genitalia with segment IX narrow throughout, except for meso-ventral tongue which

encloses the clasper bases. Claspers very long, two-articled, in form of semi-circular pincer;

bases of claspers produced mesad as wide ledges divided by vertical ridge (Fig. 147); distal

article tapered sharply. Aedeagus long, thin; lateral arms slender, finely spinate; variably

fused to ensheath median shaft (Fig. 149).

Female genitalia with segment IX of two almost separate parts; tubular piece narrow,

projected. Segment X flared, narrowed basally. Supra-genital plate large, short, thick. Vulval

scale simple, pad-like, trapezoidal and convex (Fig. 151, 152).

Key to the Males of species of Dicosmoecus found in Alberta and eastern British Columbia

1 a. Mesal ridge of clasper bases in posterior aspect vertical, roughly rectangular in outline,

with smaller rectangular lateral lobe (Fig. 147) D. jucundus Banks, p. 52.

lb. Mesal ridge of clasper bases in posterior aspect not vertical, only outer edge visible;

long axis oriented obliquely dorso-ventrad ; lateral lobe small, acuminate (Fig. 154)

D. atripes (Hagen), p. 53.

52

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Key to the Females of species of Dicosmoecus found in Alberta and eastern British Co-

lumbia

la. Median lobe of vulval scale sub-equal in length to lateral lobes. Segment X projected

posterad, deeply cleft, the two halves thin, plate-like (Fig. 151)

D. jucundus Banks, p. 52.

lb. Median lobe of vulval scale only half as long as lateral lobes. Segment X short, cleft to

two short, rounded, fleshy lobes (Fig. 156) D. atripes (Hagen), p. 53.

Dicosmoecus jucundus Banks, 1 943

(Fig. 122a, 122b, 146-152,593)

Dicosmoecus jucundus Banks, 1943:358-359. (Type locality: Modoc County, California).

Ross, 1944:297. Schmid, 1955:36. Flint, 1966:376. Fischer, 1967:66.

Males of this species (which Flint, 1966, considers to be a synonym of D. atripes ) are

most easily distinguished from males of D. atripes by posterior aspect of basal ridge of

clasper (Fig. 147, 153), as defined in preceding key. Females are distinguished by length of

median lobe of vulval scale relative to lateral lobes (Fig. 151, 156).

Description. - Antennae dark brown; antero-mesal faces of scapes with longitudinal

glabrous, yellowish stripe. Vertex of head dark brown; warts yellowish. Thorax yellowish

laterally, yellow to red-brown dorsally in form of cruciform pattern on nota, with base of

cross directed posterad. Femora yellow, tibiae and tarsi chocolate-brown. Fore wing length

of male 25 mm; dark brownish grey, with prominent, almost black veins; costal area almost

hyaline and a clear area located at divergence of veins Ml+2 and M3. Venation of fore and

hind wings as in Fig. 122a, 122b.

Male genitalia. (Specimen taken 2 miles west of Hinton, Alberta). Segment IX high, very

narrow, of irregular width (Fig. 146). Sternum produced posterad as broad shelf ventrad of

clasper bases. Clasper massive, two-articled; distal article bowed slightly ventrad, with distal

tooth directed postero-ventrad. Basal article with meso-ventral edge developed mesad as

broad shelf (Fig. 147); shelf with two lobed vertical ridge extended meso-laterad; each lobe

rectangular in posterior aspect, lateral lobe smaller, almost at right-angles to mesal lobe (Fig.

148). Median lobes of segment X short, rounded, spinate and semi-membranous. Intermedi-

ate lobes of segment X long, finger-like, each flanked laterally by thin ridge (Fig. 146); lobes

connected by high, thin-ledged bridge with flat, plate-like crown (Fig. 147). Cercus long,

rounded-rectangular, setose distally. Aedeagus long, slender, head little wider than stem.

Lateral arms short, with thick basal half, needle-like distal half; four heavy, dark spines near

middle and tip with two or three spines in tight cluster (Fig. 149, 150). Mesal faces of

clasper bases and base of aedeagus joined by continuous, thin, sclerotized strap, looped

around aedeagal base (Fig. 147).

Female genitalia. (Specimen from 2 miles west of Hinton, Alberta). Posterior edge of

sternum VII with long, narrow fringe of short, hyaline hairs. Segment VIII with sternum

markedly narrowed by antero-mesal constriction (Fig. 151). Vulval scale massive; lateral

lobes around vaginal aperture; median lobe tapered slightly distad, tip rectangular, attached

completely distad by membrane. Segment IX small, notum massive, connected to vulval

scale by lateral lobes with broad ventral extremities (Fig. 152); lateral lobes darker than

notal area. Supra-genital plate small, membranous, located between vulval scale and segment

IX, in membranous sheet. Segment X of two large, convoluted lobes joined smoothly to

segment IX.

Notes on biology. — Specimens of this species appear to emerge from small to large,

smoothly flowing, pebbled streams and rivers. I have records of captures made along lake

Rhyacophilidae and Limnephilidae

53

edges, but it is possible that these are instances of individuals flying in from nearby streams.

Adults are found from the last week in July to the last week in August.

Geographical distribution. — The known range of this species extends from Alberta and

British Columbia to California (Fig. 593). In Alberta the species is confined to the low

mountain valleys and foothills at altitudes between 3,400’ and 5,450’.

I have examined 27 specimens, 20 males and seven females, from the study area.

Dicosmoecus atripes (Hagen), 1873

(Fig. 153-156, 592)

Platyphylax atripes Hagen, 1 873b:600-601 , 605, 606. (Type locality: Colorado Mountains,

Colorado). Putnam, 1876: 1 13. Banks, 1892:364. Ulmer, 1905a:21.

Dicosmoecus atripes; McLachlan, 1875:113. Banks, 1904a: 107. Ulmer, 1905b: 63-64. Ul-

mer, 1907a:60. Banks, 1907a:38. Martynov, 1914:243. Essig, 1926:176. Betten, 1934:

318. Milne, 1935:36, 50. Goodrich, 1935:57-64. Goodrich, 1937:243-248. Ross, 1938b:

30. Knowlton and Harmston, 1939:285. Goodrich, 1941:134-143. Ross, 1941a: 103.

Banks, 1943:358. Ross, 1944:297. Ross and Spencer, 1952:47. Schmid, 1955:36. Flint,

1960:4. Denning, 1963:259. Flint, 1966:376. Anderson, 1967:508. Fischer, 1967:64-65.

Specimens of this species are essentially identical to those of D. jucundus in the majority

of characters, but differ as outlined below.

Description. — Meso-basal ledge of male clasper with ridge; in posterior aspect (Fig. 154)

this ridge viewed along its crest; bi-partite, with larger lobe set at angle of about 40° along

its long axis; smaller lobe tooth-like, with distinct acuminate tip, located latero-anterad of

larger lobe. Distal article of clasper with distal tooth directed posterad from unbowed finger-

like extension of article (Fig. 153). Lateral arms of aedeagus each tipped by dense cluster of

spines; mid-point with four close-spaced lateral spines; basad with short single spine (Fig.

155).

Female genitalia essentially similar to that of D. jucundus in lateral aspect, but differing

markedly in ventral aspect (Fig. 156). Vulval scale massive, with short, tapered median lobe

square tipped; lateral lobes huge, bulbous, with distinct antero-lateral concavities. Supra-

genital plate minute, triangular, located in sheet of membrane. Vaginal aperture roofed over

by large sclerotized, straight-edged plate. Lateral lobes of segment X thick, fleshy, rounded,

completely separated ventrally.

Geographical distribution. — The known range of this species extends from Alberta (Bea-

ver Creek (Banks, 1943)) and British Columbia to California and New Mexico (Fig. 592).

I have no records from Alberta in my own collections. The drawings were taken from Utah

specimens in Illinois Natural History Survey.

The Genus Onocosmoecus Banks

This genus is represented by one species in the study area.

Synopsis of characters. - Spur formula 1,3,4. Pleural sclerites without silky hairs; macro-

chaetae less developed than in Dicosmoecus. Fore wings large, bluntly parabolic apically;

hind wings larger than fore wings, with posterior edge regularly convex. Chord of fore wing

markedly broken; posterior part slightly oblique to body axis. Chord of hind wing only

slightly disrupted; posterior part oblique to body axis.

Male genitalia much as in Dicosmoecus . Aedeagus relatively shorter; median shaft ex-

panded apically; lateral arms quite large, partly free, armed with few heavy spines (Fig.

159).

54

Nimmo

Female genitalia differing appreciably from Dicosmoecus. Segment IX massive, very

strong, of one piece; ventrally vulval scale flanked by two large lobes (Fig. 162, 163). Seg-

ment X conical. Supra-genital plate small. Vulval scale thick, trilobed; lateral lobes semi-

globose apically; small, tongue-like median lobe enclosed by lateral lobes (Fig. 163).

Onocosmoecus unicolor (Banks), 1897

(Fig. 157-163,594)

Anabolia unicolor Banks, 1897:27-28. (Type locality: Skokomish River, Washington). Ul-

mer, 1905a:20.

Dicosmoecus unicolor ; Banks, 1907a:38. Ulmer, 1907a:60. Essig, 1926:176. Betten, 1934:

318. Milne, 1935:36, 50. Ross, 1938b:30. Knowlton and Harmston, 1939:285. Ross,

1941a: 103. Ross, 1944:297.

Dicosmoecus (Onocosmoecus) unicolor ; Banks, 1943:361-362. Fischer, 1967:68.

Onocosmoecus unicolor\ Schmid, 1955:39. Lindroth and Ball, 1969:138.

Specimens of this species are distinguished from other limnephilids in the study area by

large size of body and wings, and by smoothly rounded parabolic form of wing tips. Speci-

mens of this species may be confused with the still larger specimens of Dicosmoecus in the

study area, and may be separated from them by general body coloration, which is red-brown

rather than grey to black as in Dicosmoecus.

Description. — Antennae straw-yellow; antero-mesal face of scapes setaless; each article

with distal half and anterior face depressed. Vertex of head dark straw-yellow. Thorax

generally yellow, to brownish dorsally except for yellowish warts. Spurs dark yellow. Fore

wing length of male 18.4 mm; warm orange-brown except for grey-brown areas at proxi-

mal ends of the four distal radial cells, and at first bifurcation of M. Anal area slightly

darker brown. Venation of fore and hind wings identical to that of Dicosmoecus (Fig.

122a, 122b).

Male genitalia. (Specimen from Gorge Creek, 20 miles west of Turner Valley, Alberta).

Segment IX high, narrow, sinuate (Fig. 157); with sparse fringe of setae round clasper base,

and denser patch ventrad of base. Clasper large, two-articled, with rather spindly distal

article black tipped. Basal article with ventro-lateral band of setae set in lighter coloured

area. Meso-basal ledge of clasper with transverse, rounded, ridge at postero-mesal corner

(Fig. 158). Median lobes of segment X long, narrow, partially fused; separated from ventral

portion of segment by wide membranous area. Ventral portion of segment X truncate-

triangular in dorsal aspect, with high sclerotized ridge on each side. Cercus large, blunt,

lanceolate in lateral aspect, with concave mesal face. Aedeagus large, simple, sinuate (Fig.

159); lateral arms heavy, short, armed with three to five straight, black spines of varied

lengths, with one dominantly large spine basally. Two large, black spines arise from dorsal

membranous area between median shaft and lateral arms of aedeagus (Fig. 160). Aedeagal

base flanked by two twisted, strap-like sclerites attached to antero-mesal edge of claspers

(Fig. 161).

Female genitalia. (Specimen from Gorge Creek, 20 miles west of Turner Valley, Alberta).

Posterior edge of sternum VII with wide fringe of fine, hyaline hairs, Vulval scale relatively

small, complex. Median lobe squat, triangular, rounded (Fig. 163); lateral lobes rounded,

spatulate on mesal faces, tending to enclose median lobe. Lateral lobes laterally enveloped

by two folds of membrane. Segment XI massive, tapered ventrad, with large lateral bulges

(Fig. 162, 163). No apparent supra-genital plate. Segment X large, tubular, merged imper-

ceptibly with segment IX; with large, projected, roof-like dorsal area, and shorter, hemi-

cylindrical ventral area with dentate posterior edge.

Rhyacophilidae and Limnephilidae

55

Notes on biology. — Larvae of this species apparently inhabit streams ranging from very

small to rivers, with no apparent preference as to nature of the bottom. There is some indi-

cation that larvae might also inhabit lakes. The adult flight season extends from early May

to late September, peaking in August.

Geographical distribution. — The known range of this species extends from Alaska to the

New England states and New Mexico (Fig. 594). In Alberta the species is commonest in the

low mountain valleys, but has been recorded well out in the plains, and is known from

Saskatchewan. In Alberta the known altitudinal range is from around 2,000’ to 5,450’.

I have examined 225 specimens, 154 males and 71 females, from the study area.

The Genus Amphicosmoecus Schmid

The single species of this genus is found in Alberta and eastern British Columbia.

Synopsis of characters. — Spur formula 1,2,4. Wings very large; fore wings bluntly

rounded apically; hind wings with large anal area and M2 present (Fig. 123b).

Male genitalia with narrow dorsal strap on segment IX; remainder of segment greatly ex-

panded ventrally (Fig. 164). Median lobes of segment X long, dorsally arched blades. Clasper

large, long, blunt, pincer-like (Fig. 165); two-articled, both articles fused. Aedeagus small,

housed in membranous pocket; lateral arms spiniform, armed with short, fine spines (Fig.

166).

Female with segment VIII unmodified. Vulval scale huge, massive; median lobe small,

located in deep cleft between lateral lobes which are fleshy, and enclose vaginal aperture

laterally (Fig. 168, 169). Segment IX large, of one piece, with no central lobes. Supra-genital

plate large, projected well posterad, sclerotized (Fig. 169). Segment X small, recessed into

segment IX; fused to segment IX; with long, thin median lobes.

Amphicosmoecus canax (Ross), 1947

(Fig. 123a, 123b, 164-169,595)

Dicosmoecus canax Ross, 1947:149-150. (Type locality: Logan Canyon, Utah). Fischer,

1967:65.

Amphicosmoecus canax; Schmid, 1955:50. Nimmo, 1965:787. Smith, 1965:243, 244.

Males of this species are distinguished from all others by large, sickle-shaped, median

lobes of segment X, and by claspers with fused articles. Females are distinguished by massive

vulval scale, with small, minutely trilobed median lobe, and by long, slender median lobes of

segment X.

Description. — Antennae orange-brown; scapes dark brown, with yellow, glabrous antero-

mesal faces. Vertex of head dark straw-yellow with light brown areas. Thorax yellow to

light brown especially laterad of terga. Spurs straw-yellow. Fore wing length of male 17.8

mm; pale greyish brown except for hyaline areas at extremity of Cu2, and initial bifurcation

of M. Anal area posterad of Cu2 darker, slightly irrorate. Venation of fore and hind wings as

in Fig. 123a, 123b.

Male genitalia. (Specimen from 2 miles east of Nordegg, Alberta). Segment IX with large

ventral body; dorsal strap narrow, heavy, bilobed (Fig. 165); slight concavities occur at

ventro-lateral faces of segment. Clasper large, two-articled, with articles fused; sharp, narrow,

ledge along ventro-mesal face, and distinct groove on dorso-lateral face of basal portion

thumb-like (Fig. 164, 165). Median lobes of segment X long, dorsally arched, thin blades;

sickle-like, with acuminate tips. Cercus spatulate, setose, attached to large, bowl-shaped,

lateral plates of segment. Aedeagus very simple (Fig. 166); tip slightly dilated, base slightly

56

Nimmo

wrinkled dorsally. Lateral arms very simple, plain rods with five to six simple, long spines

distally. Aedeagus connected basally to clasper bases by lateral straps bent at angle of about

60° at point of departure from aedeagal base; terminated by large, hollow, spherical struc-

tures (Fig. 167).

Female genitalia. (Specimen from 2 miles east of Nordegg, Alberta). Posterior edge of

tergum VII with short, narrow brush of short, yellow hairs. Vulval scale huge, with massive,

fleshy lateral lobes, and small, trilobed median lobe (Fig. 169). Segment IX composed of

single, trapezoidal tergum (Fig. 168). Supra-genital plate large, convoluted, with lateral

edges turned dorsad. Segment X small, complex, with two short, stout, lateral lobes and two

long, thin, hyaline, mesal lobes; bases of lobes produced ventrad as two broad, triangular

flaps fitted loosely into two dorsal channels of supra-genital plate. Segment X recessed into

segment IX.

Notes on biology. - Larvae of this species inhabit a wide variety of streams, from small,

quiet brooks, to large rivers and turbulent mountain streams. The adult flying season ex-

tends from mid-September to late October.

Geographical distribution. — The known range of this species extends from Alberta and

eastern British Columbia to Utah and California. Records are very scattered however (Fig.

595). In Alberta the species occurs mainly in the mountain and foothill areas. There is one

record from Cold Creek, Nojack, however, which is well outside of this area. My records

indicate an altitude range from at least 2,500’ to 4,200’.

I have examined 67 specimens, 51 males and 16 females, from the study area.

The Genus Imania Martynov

This genus is represented in the study area by four species, one of which is here described

as new. Ross (1950) divided the genus into four species groups, of which the tripunctata and

bifosa groups are known to occur in the area.

Synopsis of characters. — Form of head varied. Spur formula 1,3,4. Macrochaetae poorly

developed. Wings of different lengths; fore wings apically parabolic; hind wings slightly

larger than fore wings, hardly indented posterad of apex (Fig. 124-127). Venation complete;

fore wings unmodified, venation of hind wing slightly reduced. R1 of fore wing arched or

not at stigma, or joined to Sc by cross-vein. Chord of fore wing strongly disrupted, concave

to body. Hind wing with chord only slightly disrupted, posterior part oblique to body; Cula

absent. Frenulum of seven or eight strong, curved spines at costa base.

Male genitalia with segment IX narrow laterally, with wide dorsal process (Fig. 170).

Segment X elongate, slender; of two long, sclerotized, free or fused, portions (Fig. 171).

Clasper large, two-articled; not pincer-like, complex, directed dorso-posterad; basal article

long, subcylindrical, not baso-mesally enlarged, with baso-mesal area in some species devel-

oped as one or more sclerotized spines; distal article short, bilobed, with dorsal lobe with

heavy, short, black teeth. Aedeagus small; lateral arms slender, spiniform, free or basally

fused as sheath to median shaft (Fig. 175).

Female genitalia with segment IX of two parts not entirely separated; dorsal part simple,

narrow, quite long, Segment X of two very large dorsal plates, strongly sclerotized. Vaginal

aperture flanked by ventral lobes of segment X. Supra-genital plate large, not prominent.

Vulval scale reduced to one lobe, membranous, wrinkled (Fig. 176, 177).

Key to the Males of species of Imania found in Alberta and eastern British Columbia

la. Basal article of clasper with long, slender, acuminate spine attached to baso-mesal

face (Fig. 171, 174, 184) ( tripunctata group) 2a

Rhyacophilidae and Limnephilidae

57

Basal article of clasper without such spine (Fig. 179) /. bifosa Ross, p. 60.

Aedeagus with lateral arm bases fused to form sheath around aedeagal base (Fig.

175) /. cascadis Ross, p. 58.

Aedeagus without basal sheath (Fig. 171, 185) 3a

Median shaft of aedeagus tapered to tip (Fig. 172); segment X deeply cleft, of two

heavy, black, large spiniform lobes (Fig. 171) I. tripunctata (Banks), p. 57.

Median shaft of aedeagus not tapered distally (Fig. 185); segment X not deeply

cleft, with two short, acuminate distal lobes (Fig. 184)

I. hector Nimmo n. sp., p. 59.

Key to the Females of three species of Imania found in Alberta and eastern British Columbia

la. Single, median, lobe of vulval scale massive, rectangular, dorsally directed flap

(Fig. 176, 177) /. cascadis Ross, p. 58.

lb. Median lobe of vulval scale small, dorsally directed (Fig. 181, 186) 2a

2a.(lb) Ventral aspect of lateral lobes of segment X triangular; short, wide, rounded in

lateral aspect (Fig. 187) I hector Nimmo n. sp., p. 59.

2b. Ventral aspect of lateral lobes of segment X trapezoidal (Fig. 181); in lateral

aspect long, ventrally curved, slender (Fig. 182) I. bifosa Ross, p. 60.

The tripunctata group

There are three species of this group known from the study area, one of which is new.

Males of species in this group are characterised by long, sinuate, blade-like spines devel-

oped from baso-mesal face of clasper (Fig. 174). Females are recognizable by roughly tri-

angular ventral aspect of lobes of segment X (Fig. 176, 187).

Imania tripunctata (Banks), 1900

(Fig. 126a, 126b, 170-172,596)

Apatania tripunctata Banks, 1900b:472. (Type locality: Yakutat, Alaska). Ulmer, 1905a:

23. Banks, 1907a:41. Martynov, 1914:20. Essig, 1926:176.

Hypnotranus ? tripunctatus’, Ulmer, 1907a:72.

Allomyia tripunctata ; Banks, 1916:120. Dodds and Hisaw, 1925a:386. Ulmer, 1932:216,

217. Betten, 1934:52, 380. Milne, 1935:49. Ross, 1944:297. Denning, 1948c: 120.

Fischer, 1967:71-72.

Imania tripunctata ; Ross, 1950b:41 1, 412, 413. Ross and Spencer, 1952:47. Schmid, 1955:

43. Ross, 1965:590. Schmid, 1968:681.

Allomyia sty lata Denning, 1948c: 1 19-120. (Type locality: Snowy Range Mountains, Albany

County, Wyoming). Ross, 1950b:412. Fischer, 1967:71-72.

Males of this species are distinguished from males of other species of the tripunctata

group by massive, spiniform, heavily sclerotized distal lobes of segment X in ventral aspect

(Fig. 171), and by smoothly tapered median shaft of aedeagus (Fig. 172).

Description. — Antennae chocolate-brown. Vertex of head dark chocolate with reddish

tinges laterally. Thorax deep, warm red-brown laterally, darker dorsally. Spurs brown. Fore

wing length of male 1 1.4 mm; dull red-brown, with hyaline areas along veins. Venation of

fore and hind wings as in Fig. 126a, 126b.

Male genitalia. (Specimen from Cascade River, Banff National Park, Alberta). Posterior

edge of sternum VIII with long, single parallel row of heavy, hyaline setae (Fig. 170). Seg-

ment IX with very narrow ventral and lateral walls, broadened dorsad of clasper bases.

lb.

2a.(la)

2b.

3a.(2b)

3b.

58

Nimmo

Clasper massive, with high base, narrowed distad to uniform width along distal half of basal

article. Baso-mesal face of clasper with long, thin, sinuate black spine. Distally basal article

with flared fringe of long, stout setae. Distal article of clasper bilobed, claw-like, with dorsal

lobe flat ventrally, armed with many stout, short, black pegs. Ventral lobe rounded, short,

with some black spines distally. Segment X with median lobes overhung by large, rounded,

dorsal bulge. Median lobes long, thin, acuminate structures curved dorso-laterad distally.

Cercus long, almost rectangular plate (in Fig. 170 they are seen end-on, however, as they

project laterad). Lateral arms of aedeagus sclerotized, finely tapered, with curved base (Fig.

172). Median shaft tapered, with attenuated tip; ejaculatory duct opens on dorsal surface of

tip, between two lateral flanges.

Female genitalia. Unknown.

Geographical distribution. — The known range of this species extends from Alaska to

Washington and Colorado (Fig. 596). I have only one record of the species in Alberta, from

an altitude of 6,500’, several miles northeast of Banff, on the Cascade Fire Road.

I have examined a single male of this species, on loan from Illinois Natural History

Survey. The date of collection is May 30.

Imania cascadis Ross, 1950

(Fig. 124a, 124b, 173-177,597)

Imania cascadis Ross, 1950b:41 1, 412, 415. (Type locality: Steven’s Pass, nr. Berne, Wash-

ington). Schmid, 1955:43. Schmid, 1968:681.

Allomyia cascadis', Fischer, 1967:70.

Males of this species are distinguished from males of other species of Imania by basal

aedeagal sheath of lateral arm bases (Fig. 175); by foreshortened basal article of clasper

(Fig. 174); and by almost uncleft segment X, each half of which is more or less square cut

distally in ventral aspect. Females are distinguished by very wide, long median lobe of

vulval scale (Fig. 176).

Description. — Antennae reddish chocolate-brown; scapes with antero-mesal faces dark

brown, remainder pale yellow. Vertex of head deep chocolate-brown; warts pale. Thorax

deep red-brown laterally, almost black dorsally; warts pale. Fore wing length of male 10.4

mm; pale red-brown, lightly irrorate. Venation of fore and hind wings as in Fig. 124a,

124b.

Male genitalia. (Specimen from Spray River, at Banff, Alberta). Tergum VIII with single

row of long, thick setae parallel to posterior edge. Segment IX high, uniformly narrow,

bowed anterad (Fig. 173). Basal article of clasper with massive base and clearly delineated

distal clumps of setae originated from pale surficial areas. Baso-mesal face of clasper with

long, thin, sinuate black spine. Distal article of two short, stout lobes with teeth or pegs

only at distal edges. Segment X of two parallel, concave plates with distal edges thin, sharp,

directed dorsad (Fig. 173, 174). Cercus short, rounded lobe. Lateral arms of aedeagus arched

high over distal portion; attached to massive, plate-like bases (Fig. 175). Ejaculatory duct of

median shaft opening dorsally, well basad of cleft tip.

Female genitalia. (Specimen from Spray River, at Banff, Alberta). Tergum VIII with

straight, single row of long, stout setae parallel to posterior edge. Sternum VIII unpigmented

medially. Vulval scale of single, wide, dorsally curved median lobe (Fig. 176). Segments IX

and X imperceptibly fused (Fig. 177) as massive, sigmoid structure in lateral aspect, cleft

medially as two lateral plates abutted dorsally, each at angle of about 45°. Supra-genital

plate semi-circular, hyaline.

Geographical distribution. — The known range of this species extends from Washington to

Rhyacophilidae and Lirnnephilidae

59

Alberta (Fig. 597). In Alberta it is known only from the Spray River, Banff, at an altitude

of about 5,500’.

I have examined two specimens from the study area, one of each sex, in Illinois Natural

History Survey. The date given is May 30.

Imania hector Nimmo n. sp.

(Fig. 127a, 127b, 183-187, 597)

Males of this species are distinguished from males of other species of Imania by form of

segment X in ventral aspect (Fig. 184); segment cleft distally, two halves spread wide apart,

blade-like and acuminate. Median shaft of aedeagus short, blunt, with distinct distal head

(Fig. 185). Females are distinguishable by triangular form of lateral lobes of segment X

in ventral aspect (Fig. 186), and by long, narrow, dorsally directed median lobe of vulval

scale.

Description. — Antennae dark brown. Vertex of head dark brown to black. Thorax dark

red-brown. Femora of legs dark brown, remainder lighter brown. Fore wing length of male

7.6 mm; uniform dark brown except for slight irrorations along Rl, and three hyaline areas

at bases of F2 and F3, at bifurcation of Ml +2 and M3, and at end of Cu2. Venation of fore

and hind wings as in Fig. 127a, 127b.

Male genitalia. (Specimen from Sunshine Lodge, Banff National Park, Alberta). Tergum

VIII with single row of long setae parallel to posterior edge. Segment IX high, narrow,

essentially parallel sided (Fig. 183). Basal article of clasper massive, cylindrical throughout,

armed with long, light brown, sinuate spine on baso-mesal face (Fig. 183, 184). Distal article

bilobed; dorsal lobe parallel sided, moderately toothed ventrally. Segment X fused to seg-

ment IX dorsally. Median lobes fused at bases; distally separate, triangular, spinate. Main

body of segment X folded roof-like over anal aperture. Cercus long, narrow, in dorso-lateral

depression of segment X. Aedeagus with two pairs of lateral arms; dorsal pair short, heavy,

scythe-like; ventral pair long, straight, spatulate, in form of cylindrical sheath to mem-

branous median shaft (Fig. 185). Ejaculatory duct open dorsally between distal flanges.

Female genitalia. (Specimen from Sunshine Lodge, Banff National Park, Alberta). Poste-

rior edge of tergum VIII with single line of widely spaced hyaline hairs parallel to edge.

Vulval scale single, long, rectangular median lobe; distal half wrinkled, curved dorsad (Fig.

186, 187). Vulval scale and vaginal aperture flanked by two large, sclerotized bodies. Seg-

ments IX and X fused; comprising two large, rounded lobes with flat mesal faces sloped

meso-dorsad in form of roof-like structure; each lobe in ventral aspect triangular.

Notes on biology. — Individuals of this species appear to emerge from fast, turbulent

mountain creeks with boulder beds. The flight season extends from June 1 to July 9.

Geographical distribution. — The known range of this species includes three localities

close to the continental divide in Banff National Park, Alberta (Fig. 597). The altitudinal

range extends from around 6,000’ to 7,000’.

I have examined 13 specimens, nine males and four females, from the study area.

Holotype. - Male. Sunshine Lodge, Banff National Park, Alberta; July 9, 1962; W. R. M.

Mason.

Allotype. — Female. Same data as holotype.

Paratypes. — Same data as holotype; four males, one female. Sunshine Lodge, Banff

National Park, Alberta; July 9, 1962; K. C. Herrmann; four males. Hector Creek, at Banff-

Jasper Hwy. north of Lake Louise, Banff National Park, Alberta; June 15, 1967; A. Nimmo;

one female. Moraine Creek at Moraine Lake, Banff National Park, Alberta; June 1, 1958;

W. E. Ricker; one female.

60

Nimmo

All type material is in the Canadian National Collection, Ottawa (type number 10,583)

with the exception of the female from Hector Creek, which is in the Strickland Museum,

Dept, of Entomology, University of Alberta, Edmonton.

This species is not named for Hector Creek but for the character of the same name in

ancient Greek literature.

The bifosa group

One species of this group is known from the study area.

Males of the group are distinguished by absence of long, blade-like spine at baso-mesal

face of basal article of clasper (Fig. 179); by presence of a short, acuminate extension of

basal article of clasper, at meso-distal edge; and by presence of two pairs of long, curved,

blade-like lateral arms on aedeagus (Fig. 180). Females are distinguished by lack of medial

cleavage of segment X (Fig. 181); by trapezoidal outline in ventral aspect, of lateral bulges

of segment X; and by long, narrow, dorsally curved median lobe of vulval scale.

Imania bifosa Ross, 1950

(Fig. 125a, 125b, 178-182,597)

Imania bifosa Ross, 1950b:412, 415-417. (Type locality: Banff, Alberta). Ross and Spencer,

1952:47. Schmid, 1955:43. Schmid, 1968:681-682.

Allomyia bifosa ; Fischer, 1967:70.

Males of this species are distinguished from males of other species of the genus by absence

of baso-mesal spine on clasper base. Females are distinguished by trapezoidal lateral bulges

of segment X.

Description. — Antennae dark brown to black. Vertex of head black. Thorax black to

very dark brown. Fore wing length of male 8.5 mm; uniform dark chocolate-brown, appear-

ing black in living insects. Venation of fore and hind wings as in Fig. 125a, 125b.

Male genitalia. (Specimen from Mt. Edith Cavell, Jasper National Park, Alberta). Tergum

VIII with single line of long setae parallel to posterior edge. Segment IX very narrow, bowed

anterad in lateral aspect. Clasper large, with massive basal article of uniform width (Fig.

178) . Basal article with meso-distal edge produced posterad as short, acuminate spine (Fig.

179) . Distal article bilobed; ventral lobe thin, parallel sided in lateral aspect; dorsal article

irregular, arched dorsad, with heavily toothed ventral surface. Segment X with short, bul-

bous cercus; median lobes triangular in lateral aspect, curved postero-laterad; intermediate

lobes small, located on ventral edges of median lobes. Segment X roof-like, with carinate

dorsal ridge. Aedeagus with median shaft curved ventrad; ejaculatory duct terminated in

dorsal groove. Two pairs of lateral arms; basal pair long, slender, arched dorsad; distal pair

similar but shorter and wider (Fig. 180).

Female genitalia. (Specimen from Mt. Edith Cavell, Jasper National Park, Alberta). Poste-

rior edge of tergum VIII with single line of well spaced setae. Vulval scale with single,

median lobe originated from fold of membrane, curved dorsad. Segment IX with semi-

cylindrical dorsum almost completely dissociated from two ventro-lateral lobes around

genital cavity (Fig. 182). No evident supra-genital plate. Segment X fused completely with

segment IX and cleft mesally very shallowly (Fig. 181).

Notes on biology. - Individuals of this species emerge from cold, alpine streams which

originate primarily as glacial melt-water and flow over moraine debris. The adult flight

season extends from June 29 to July 22.

Geographical distribution. - The known range of this species is restricted to southern

Rhyacophilidae and Limnephilidae

61

Alberta and British Columbia (Fig. 597). In Alberta the species is known only from high

mountain regions close to the continental divide, at altitudes between 6,000’ and 7,500’.

I have examined 1 16 specimens, 71 males and 45 females, from the study area.

The Genus Ecclisomyia Banks

Three species of this genus are known from the study area. Of these one is unidentified as

only a single female is known.

Synopsis of characters. - Spur formula 1,3,4. Wings of normal size, elongate; fore wing

rather narrowly parabolic; hind wing little larger. Venation complete, almost unmodified.

Fore wing with long, narrow, discoidal cell; chord strongly disrupted, posterior part oblique

to body. Hind wing with very long discoidal cell, chord not so disrupted, and posteriorly

oblique to body (Fig. 128a, 128b).

Male genitalia with lateral walls of segment IX very wide; segment greatly narrowed

dorsally and ventrally (Fig. 188, 194). Cercus very large, long, mesally concave. Median

lobes of segment X fused in form of long process, concave basally, flattened distally. Clasper

small, not recessed into segment IX, of one article, conical, with one or more stout, or long

and thin, spines on baso-mesal face (Fig. 189, 195).

Female genitalia with segment IX very large, massive, of one piece. Segment X conical.

Supra-genital plate short. Vaginal aperture along entire length of segment IX, partially

obstructed by single, median, membranous vulval scale (Fig. 192, 193).

Key to the Males of two species of Ecclisomyia found in Alberta and eastern British Colum-

bia

la. Colour pale yellow-red. Clasper of genitalia with two pairs of long, fine, baso-

mesal spines (Fig. 188, 189). Size smaller E. maculosa Banks, p. 61.

lb. Colour dark purple-brown. Clasper of genitalia with single, massive, black, heavily

sclerotized baso-mesal spine (Fig. 195). Size larger . . . E. conspersa Banks, p. 62.

Key to the Females of species of Ecclisomyia found in Alberta and eastern British Columbia

la. Colour pale yellow-red. Single median lobe of vulval scale massive, with distinct

v-notch on distal edge (Fig. 192); supra-genital plate very large, very deeply cleft

mesally. Size smaller E. maculosa Banks, p. 61.

lb. Colour dark red-, or purple-brown. Vulval scale and supra-genital plate not as

above. Size larger 2a

2a.(lb) Vulval scale very wide, long, rectangular, point of origin in membrane between

segments VIII and IX (Fig. 200) E. sp. 1 , p. 63.

2b. Vulval scale with rectangular tip, membranous, originated imperceptibly from

membrane between segments VIII and X (Fig. 199) . . E. conspersa Banks, p. 62.

Ecclisomyia maculosa Banks, 1 907

(Fig. 128a, 128b, 188-193,598)

Ecclisomyia maculosa Banks, 1907a: 123-124. (Type locality: Boulder, Colorado). Banks,

1907b:40. Dodds and Hisaw, 1925b:386. Essig, 1926:176. Betten, 1934:356. Milne,

1935:37, 50. Ross, 1938b:31. Ross, 1944:300. Denning, 1948b: 18. Ross, 1950b:423-

425. Denning, 1951:161. Ross and Spencer, 1952:50. Schmid, 1955:59. Anderson, 1967:

508. Fischer, 1967:77-78.

Males of this species are distinguished from males of other species of Ecclisomyia by two

62

Nimmo

pairs of long, fine spines on each clasper (Fig. 188, 189). Females are distinguished by large,

distally notched, median lobe of vulval scale. Specimens may be distinguished with the

naked eye simply by their small size, and pale yellowish red coloration with scattered

irrorations.

Description. — Antennae light yellow-brown; antero-mesal face of scapes paler, glabrous.

Vertex of head with antero-mesal area brown bounded by yellow. Thorax pale reddish or

yellowish brown. Lateral faces of maxillary palpi black or otherwise; mesal faces light brown

to yellow. Lateral faces of front and middle tibiae black. Fore wing length of male 7.9 mm;

very light reddish or yellowish brown, uniformly irrorate, except for hyaline costal area.

Venation of fore and hind wings as in Fig. 128a, 128b.

Male genitalia. (Specimen from Lake Agnes, Lake Louise, Alberta). Segment IX with

wide lateral walls, and narrower but strong dorsal strap (Fig. 188, 189). Clasper of one

article, large, longer than thick, with two pairs of long, dark spines on baso-mesal face; basal

pair about three times longer than distal pair (Fig. 188). Median lobe of segment X skittle-

like in dorsal aspect (Fig. 189), with small dilated tip; laterally fringed with sparse, short,

sharp setae. Intermediate lobes short, blunt. Cercus long, rounded, mesally concave. Aedea-

gus very simple, with dorsally directed, scoop-like tip extended over ejaculatory duct as

hood; ejaculatory duct opening in padding of hyaline membrane on ventral surface of dorsal

hood (Fig. 190, 191). Distal edge of hood dark brown, with u-shaped mesal notch.

Female genitalia. (Specimen from Lake Agnes, Lake Louise, Alberta). Vulval scale large,

with median ridge between two rounded, ventral concavities; sides roughly parallel, poste-

rior edge with shallow, wide v-notch (Fig. 192). Segment IX high, narrow, of one piece

(Fig. 193). Tufts of long, yellow setae laterally, about mid-point. Supra-genital plate large,

long, rectangular, divided in two rounded, rectangular lobes by very deep v-cleft. Segment X

of two rounded, conical lobes joined dorsally toward base; each lobe with mesally directed

flanges on basal half of ventral edges (Fig. 192).

Notes on biology. — Individuals of this species emerge from small, riffled mountain

creeks, running gently over fine gravel bottoms. The adult flying season extends, in the

study area, from July 14 to August 14.

Geographical distribution. — The known range of this species extends from Alberta and

British Columbia to Oregon and Colorado (Fig. 598). In Alberta it is confined to the moun-

tains and foothills, ranging in altitude from 4,500’ to 7,000’.

I have examined 59 specimens, 54 males and five females, from the study area.

Ecclisomyia conspersa Banks, 1907

(Fig. 194-199, 599)

Ecclisomyia conspersa Banks, 1907b: 123. (Type locality: Olympia, Washington). Banks,

1907a:40. Essig, 1926:176. Neave, 1929:189. Betten, 1934:356. Milne, 1935:37, 50.

Ross, 1 94 la: 1 14. Ross, 1944:300. Ross, 1950b:423, 425. Denning, 1951:162. Schmid

and Guppy, 1952:42. Ross and Spencer, 1952:50. Schmid, 1955:59. Flint, 1960:24.

Denning, 1963:263. Fischer, 1967:76. Lindroth and Ball, 1969:138.

Males of this species are distinguished by massive, black, heavily sclerotized spine on

baso-mesal face of clasper (Fig. 195). Females are distinguished by membranous, tapered,

rectangular tipped median lobe of vulval scale (Fig. 199).

Description. — Antennae brown; scapes dark brown, antero-mesal faces white in female,

pale brown in male. Vertex of head dark brown, posterior edge and angles paler. Thorax

light to dark chocolate-brown; warts of terga yellowish. Spurs dark brown. Fore wing length

of male 13.8 mm; dark reddish brown, uniformly coloured costal area. Venation as in E.

Rhyacophilidae and Limnephilidae

63

maculosa.

Male genitalia. (Specimen from Kicking Horse Campground, Yoho National Park, British

Columbia). Segment IX almost equilateral-triangular in lateral aspect (Fig. 194). Dorsal

strap wide, short. Basal half of anterior edges banded with light and dark brown; remainder

pale to hyaline. Clasper large, stubby; two articles fused but traces of suture evident. Setae

on ventral, disto-ventral, and distal surfaces. Baso-mesal face occupied by massive, black,

strongly sclerotized spine (Fig. 195), hooked laterad at tip; base of spine hemispherical.

Median lobe of segment X acute-triangular in ventral aspect (Fig. 195), with disto-ventral

hook. Intermediate lobes small, setose, placoid. Cercus large, rectangular, distally rounded,

slightly concave on mesal face. Aedeagus large, with ejaculatory orifice on large, fleshy lobe

between two stout, dark brown, distal spines (Fig. 196). Duct ventral on aedeagus, flanked

by two lateral folds. Aedeagal strap roughly triangular (Fig. 197) at attachment point, with

two thin straps connected to clasper bases.

Female genitalia. (Specimen from Kicking Horse Campground, Yoho National Park, Brit-

ish Columbia). Sternum VIII divided mesally by band of membrane of two lateral sternites

(Fig. 199). Vulval scale rectangular distally, with rippled surface; attached to tapered fleshy

extension of sternum VIII. Segment IX massive, trapezoidal in lateral aspect, fused almost

imperceptibly to segment X, except ventrally (Fig. 198). Supra-genital plate cleft distally in

form of two lobes; plate arched laterally, flanged to form partly enclosed passage to vagina.

Segment X large, triangular in lateral aspect, with small notch distally. Ventral edges arched

over and beyond anal aperture.

Notes on biology. — Individuals of this species emerge from small to large mountain

creeks flowing turbulently over small stone or boulder bottoms. I have observed pupae of

this species emerge from Whitehorse Creek, Cadomin, Alberta, at an altitude of about

6,500’, on May 21. The pupae crawled out of the water onto streamside boulders or ice

indiscriminately, remained still for about 10 minutes, then started to emerge as adults from

the pupal skin, which required about 5 minutes. Upon completion of emergence the adults

were active, but did not attempt to fly for about another 10-15 minutes. A total of 33 males

and 17 females were collected on this occasion. The adult flight season extends from May

17 to September 12 in the study area.

Geographical distribution. — The known range of this species extends from Alaska to

California and New Mexico (Fig. 599). In the study area the species is confined to the

mountain and foothill areas, ranging in altitude from 2,600’ to 7,300’.

I have examined 180 specimens, 131 males and 49 females, from the study area.

Ecclisomyia species 1

(Fig. 200-201, 600)

Only a single female is known to me, which is distinguished from the females of the other

species of Ecclisomyia treated here by very large, rectangular, median lobe of vulval scale.

This lobe has definite point of origin from membrane between segments VIII and IX, and is

concave dorsally, convex ventrally (Fig. 200, 201).

Description. — Antennae yellow-brown; scapes darker, with antero-mesal faces almost

white, glabrous. Vertex of head dark brown centrally, lighter round edges. Thorax deep

yellow-brown laterally, to dark brown dorsally. Spurs brown. Fore wing length of female

15.8 mm; red-brown, heavily irrorate, except for clear costal area. Venation identical to that

of is. maculosa.

Female genitalia. Vulval scale single, large, rectangular, median lobe (Fig. 200), concave

dorsally, convex ventrally, i.e. spatulate. Segment IX massive, trapezoidal in lateral aspect

64

Nimmo

(Fig. 201). Supra-genital plate short, wide, bilobed. Segment X relatively large, closed dor-

sally, deeply cleft ventrally; very slightly bilobed distally, with lateral walls constricted

antero-ventrad.

Geographical distribution. — The single known female specimen was taken under a road

bridge over the Miette River, about 1 mile south of Jasper, Alberta, on highway 93a (Fig.

600), on July 6. It was already dead and trapped in cobwebs.

The Subfamily Apataniinae Ulmer

Head rather long, both sides convex; eyes, ocelli and cephalic warts small. Pronotum

short; macrochaetal development slight. Spur formula varied, identical in both sexes of the

same species. Wings constant, identical in both sexes; fore wing elongate, obliquely para-

bolic; hind wing hardly larger than fore, with convex trailing edge; clearly indented at

termination of Cu2. Frenulum of three strong spines at base of costa of hind wing; spines

curved basally, sharp, flattened distally. Venation complete, somewhat modified. R1 of fore

wing of most species arched distally, united to Sc by cross-vein. Chord single line, irregularly

disposed. Hind wing with distally open discoidal cell, very short f 1 and four anal veins. See

Fig. 129a, 129b.

Male genitalia with segment IX lengthened longitudinally slightly; clasper bases enclosed

or not. Dorsally segment IX only weakly narrowed, with or without two basal lobes. These

lobes various in size, separate or fused together, or fused to median lobes of segment X.

Segment X sclerotized, annular, around anus; slightly developed and inconspicuous; with or

without three pairs of appendages. Clasper always large, two-articled, movable, pincer-like in

certain groups. Distal article various, little reduced. Aedeagus with median shaft and two

lateral arms, freely movable.

Female genitalia with segment IX of one piece; wide dorsally, latero-ventrally attenuated

with large, conical, ventral lobes. Segment X slender, poorly sclerotized; in some taxa long,

tubular, with lateral concavities. Supra-genital plate present in most taxa; membranous or

concave, rigid. Vulval scale membranous lobe of sternum VIII or with small lateral lobes.

Vaginal aperture along length of sternum IX.

In the study area the subfamily Apataniinae is represented by one genus, Apatania

Kolenati, of the tribe Apataniini (Martynov). In consequence, no outline of characters of

the subfamily or genus is given. The species of Apatania are distinguished by the key to sub-

families on page 50. Schmid (1953, 1954a, 1955) presents a detailed account of the sub-

family and its constituent taxa.

The Genus Apatania Kolenati

This genus is represented in the study area by five species belonging to four groups. One

of these species is new.

Key to the Males of species of Apatania found in Alberta and eastern British Columbia

la. Intermediate lobes of segment X with tips hooked antero-laterad, darker than

remainder of genitalia (Fig. 206, 207) 2a

lb. Intermediate lobes of segment X not so hooked, not necessarily darker than

remainder of genitalia (Fig. 202, 221) 4a

2a.(la) Lateral arms of aedeagus attached to fleshy dorsal lobes at base; long, laminate

blades (Fig. 208,213) 3a

Attachment of lateral arms not as above; median shaft of aedeagus thick, short,

2b.

Rhyacophilidae and Limnephilidae

65

with bulbous tip (Fig. 218) A. crymophila McLachlan, p. 69.

3a. (2a) Median shaft of aedeagus strongly curved ventrad to form half circle (Fig. 208)

A. shoshone Banks, p. 68.

3b. Median shaft slightly curved ventrad, of irregular outline (Fig. 213)

A, stigmatella (Zetterstedt), p. 67.

4a.(lb) Clasper very long, slender (Fig. 221) A. alberta Nimmo n. sp., p. 70.

4b. Clasper very short; distal article bilobed (Fig. 202)

A. zonella (Zetterstedt), p. 65.

Key to the Females of species of Apatania found in Alberta and eastern British Columbia

la. Segment X separate and distinct from segment IX (Fig. 219, 224) 2a

lb. Segments X and IX fused solidly together (Fig. 204, 210, 214) 3a

2a.(la) Median lobe of vulval scale abruptly widened distally to rectangular tip (Fig. 225)

A. alberta Nimmo n. sp., p. 70.

2b. Median lobe short, simple (Fig. 220) A. crymophila McLachlan, p. 69.

3a.(lb) Segment X large, visible, fused solidly to segment IX; with acuminate distal lobes

(Fig. 204, 214) 4a

3b. Segment X minute, concealed by posterior lobes of segment IX; segment IX

massive, formless, rounded A. shoshone Banks, p. 68.

4a. (3a) Segments IX and X fused (Fig. 214) A. stigmatella (Zetterstedt), p. 67.

4b. Ventro-lateral lobes of segment IX present, distinct, separated by broad band of

membrane (Fig. 204) A. zonella (Zetterstedt), p. 65.

The fimbriata group

One species of this group is presently known from the study area.

Synopsis of characters. — Fore wing sexual dimorphism strong in certain species; male

fore wing stigma well marked or not; C thickened. R1 of hind wing arched at stigma or not,

in contact with Sc.

Male genitalia with cerci and median lobes only of segment X present, free (except in

A. kyotensis), large in most species. Cerci long, simple, like slender pegs in most species.

Claspers medium sized; distal article distinct; small, bilobed in certain species. Aedeagus

spinate; lateral arms laminate blades.

Female genitalia with dorsum of segment IX not prominent; simple and convex; short-

ened in some species. Supra-genital plate sclerotized, fused solidly to segment X. Segment X

short, small, as large as segment IX.

Apatania zonella (Zetterstedt), 1840

(Fig. 129a, 129b, 202-205, 600)

Phryganea stigmatella var. zonella Zetterstedt, 1840:1066. (Type locality; Lapland).

Apatelia zonella ; (Literature Palaearctic; see Fischer, 1967: 125-126).

Apatidea zonella’, (Literature Palaearctic; see Fischer, 1967:126).

Apatania zonella’, Schmid, 1953: Fig. 1. Schmid, 1954a:30-34. Schmid, 1955:82. Flint,

1960:26. Corbet, 1966:981. (For Palaearctic literature see Fischer, 1967:126).

Goniotaulius arctica Boheman, 1865. (Literature Palaearctic; see Fischer, 1967:127-129).

Goniotaulius arcticus’, (Literature Palaearctic; see Fischer, 1967: 127).

Apatelia arctica’, (Literature Palaearctic; see Fischer, 1967:127-128).

Apatidea arctica ; (Literature Palaearctic; see Fischer, 1967: 129).

66

Nimmo

Radema arctica\ Ross, 1944:297.

Apatidea auricula not Forssiund; (Literature Palaearctic; see Fischer, 1967: 129).

Apatania groenlandica Kolbe, 1912. See Fischer, 1967:129.

Apatelia groenlandica ; See Fischer, 1967: 129.

Radema groenlandica, Ross, 1944:297.

Apatania inornata Wallengren, 1886. See Fischer, 1967:129.

Apatelia inornata-, See Fischer, 1967: 129.

Apatania palmeni Sahlberg, 1894. See Fischer, 1967: 129-130.

Apatidea palmeni-, See Fischer, 1967:130.

Apatelia palmeni-. See Fischer, 1967:130.

Apatania stigmatella not Zetterstedt; See Fischer, 1967:130.

Apatania zonella (Zetterstedt) var. dalecarlica Forssiund; See Fischer, 1967: 130.

Males of this species are distinguishable from males of other species of Apatania by very

short claspers, with bilobed distal article (Fig. 202). Females are distinguishable by acumi-

nate postero-ventral lobes of segment X and presence of ventro-lateral lobes of segment IX

(Fig. 204).

Description. — Antennae dark brown; scapes with antero-mesal faces glabrous. Vertex of

head black. Thorax very dark brown. Femora of legs irregularly patterned with dark and

lighter brown. Spurs yellow-brown. Fore wing length of male 8.5 mm; translucent dark

brown, with thick, irregular stigma. Venation of fore and hind wings as in Fig. 129a, 129b.

Stigma of female fore wing weaker than male.

Male genitalia. (Specimen from Lake Hazen, Ellesmere Island, Northwest Territories).

Tergum VIII with irregular single row of long setae parallel to posterior edge. Segment IX

with narrow dorsal strap pinched in at each side as distinct crown; segment gradually ex-

panded ventrad (Fig. 202). Clasper stout, short, with basal article laterally flattened, nar-

rowed basally, disto-ventrally with long setae. Distal article bilobed; ventral lobe flattened

dorso-ventrally, directed mesad. Segment X with irregularly dentate, pedicilate cerci. Median

lobes large, hooked ventrad distally, with smooth dorsal edges and irregularly dentate ventral

edges. Intermediate lobes fused in form of small, ventrally hooked lobe ventrad of median

lobes. Aedeagus simple, with large, warped, sword-like lateral arms attached to membranous

base dorsad of median shaft (Fig. 203). Median shaft arched dorsad, distally acuminate,

bilobed, with cluster of basally directed spines disto-ventrally.

Female genitalia. (Specimen from Vermilion Lakes, Banff, Alberta). Posterior edge of

sternum VII with narrow band of short, hyaline hairs. Vulval scale with single, median lobe

(Fig. 205), slightly rugose distally. Segment IX small, parallel-sided in lateral aspect (Fig.

204); with large, irregular latero-ventral lobes separated by band of membrane. Supra-genital

plate not evident. Segment X larger than IX with wide, smooth, warped ventral surface.

Dorsal surface concave laterally, with median ridge.

Notes on biology. — This, species is known to me in the study area from only two locali-

ties of the most opposite characteristics. The first is Lake Agnes, at Lake Louise, in Banff

National Park, Alberta, at an altitude of 6,885’, in the alpine meadows. The second is

Vermilion Lakes just west of Banff, Alberta, at an altitude of 4,538’. This locality is a dense,

valley bottom swamp in the Bow River valley. The collecting dates were July 21, and July 5

respectively.

Geographical distribution. — The known range of this species is Holarctic, extending in

North America, from northernmost Canada (Ellesmere Island) to British Columbia, Mon-

tana, and Minnesota (Fig. 600).

I have examined three females of this species from the study area, and one male and seven

females from Lake Hazen, Ellesmere Island, Northwest Territories.

Rhyacophilidae and Limnephilidae

67

The stigmatella group

Two species belonging to this group are known from the study area.

Synopsis of characters. - Cerci and intermediate lobes of male segment X large, massive,

concave mesally. Median lobes of segment X on mesal faces of intermediate lobes (Fig. 207);

very short and slender. Female genitalia with segment IX very long, without latero-ventral

lobes (Fig. 210). Supra-genital plate poorly developed. Segment X very small.

Ap at ania stigmatella (Zetterstedt), 1840

(Fig. 211-215,602)

Phryganea stigmatella Zetterstedt, 1840:1066. (Type locality: Lapland).

Limnephilus stigmatellus ; Walker, 1852:50.

Apatania stigmatella', Ulmer, 1907a:76. Banks, 1907a:41. Banks, 1908b:61, 64. Dodds and

Hisaw, 1925b:386. Ulmer, 1932:215. Betten, 1934:379-380. Schmid, 1953: Fig. 4m.

Schmid, 1954a: 13-14. Schmid, 1955:82. Flint, 1960:28. Fischer, 1967:1 18-121. Smith,

1969:48.

Parapatania stigmatella ; See Fischer, 1967 : 1 19.

Apatelia stigmatella', Milne, 1935:25, 49. Ross, 1938b:29.

Radema stigmatella', Ross, 1944:297. Kimmins and Denning, 1951:121-122. Unzicker,

1968:4, 20, 54.

Apatania pallida Hagen, 1861:270. Hagen, 1864:805. Banks, 1892:364. Ulmer, 1905a:23.

Ulmer, 1907a:76. Banks, 1907a:41. Thienemann, 1926:274. Betten, 1934:379. Milne,

1935:49. Ross, 1938b:29.

Apatania frigida McLachlan, 1867:57-58. Brauer, 1876:287. Milne, 1935:49. Kimmins and

Denning, 1951:121-122.

Males of this species are distinguished by irregularly sinuate median shaft of aedeagus

(Fig. 213), and by distal tooth set dorsally on intermediate lobes of segment X (Fig. 211).

Females are distinguished by absence of latero-ventral lobes of segment IX, by segment X

visible in lateral aspect (Fig. 214), and by simple, narrow, median lobe of vulval scale (Fig.

215).

Description. — Antennae brown; scapes white, with antero-mesal faces brown, glabrous.

Vertex of head black, warts white. Thorax dark brown to almost black dorsally. Femora

blotchy brown, light brown, hyaline. Spurs brown. Fore wing length of male 9.6 mm; light,

clear yellow-brown, no pattern. Venation identical with that of A. zonella.

Male genitalia. (Specimen from Simpson Islands, Great Slave Lake, Northwest Territories).

Tergum VIII with single row of long, well spaced setae parallel to posterior edge. Segment

IX roughly rectangular, wider laterally (Fig. 211); dorsal strap segregated by two lateral

grooves, bulged slightly dorsad. Clasper massive, with cylindrical, fluted basal article; distal

article semi-circular, claw-like, fringed internally with long setae (Fig. 211, 212). Segment

X with large, flared, triangular cerci. Median lobes long, thin, merged basally with cerci.

Intermediate lobes complex, dark, with dorsal processes closely associated with cerci and

laterad of them; with distal lateral and dorsal teeth. Aedeagus with median shaft irregularly

sinuate (Fig. 213); aperture of ejaculatory duct disto-dorsad, between lateral flaps; lateral

arms mounted dorsad of median shaft, long, slender, laminate blades.

Female genitalia. (Specimen from Simpson Islands, Great Slave Lake, Northwest Terri-

tories). Posterior edge of sternum VII with wide band of short, hyaline hairs. Vulval scale

single, strongly dorsally curved median lobe (Fig. 214, 215). Segment IX large, irregular,

with large postero-ventral cavity. Segment X small, with anterior edges slanted from vertical

68

Nimmo

(Fig. 214); anal cavity distinct.

Geographical distribution. — The known range of this species is Holarctic; in North Amer-

ica it extends from Alaska to Colorado, in the south, and Newfoundland, in the east (Fig.

602). The Colorado record is curious and, I suspect, open to some doubt. The species has

not yet been recorded from Alberta but is included here as there is a high probability of it

occurring in the far north of the province, which is similar to the area surrounding Great

Slave Lake, and less than 100 miles distant.

I have examined a single specimen of each sex, from Great Slave Lake. These were taken

on August 29.

Apatania shoshone Banks, 1 924

(Fig. 206-210, 601)

Apatania shoshone Banks, 1924:442. (Type locality: Yellowstone National Park, Wyoming).

Schmid, 1954a: 15-16. Schmid, 1955:82. Fischer, 1967: 1 17. Smith, 1969:48.

Apatania shoshone ; Betten, 1934:380.

Apatelia shoshone; Milne, 1935:25, 49. Ross, 1938b:29.

Radema shoshone ; Ross, 1944:297.

Males of this species are distinguished by long, slender distal hooks of intermediate lobes

of segment X (Fig. 206), and by strongly ventrally curved median shaft of aedeagus (Fig.

208). Females are distinguished by massive, formless segment IX, and minute segment X

(Fig. 209, 210).

Description. — Antennae dark brown; antero-mesal face of scapes yellow, glabrous. Ver-

tex of head very dark brown. Thorax dark brown, with interspersed lighter areas. Spurs

yellow. Fore wing length of male 8.3 mm; pale, clear brown. Cross-vein Cu-Rl white line

across stigma. Venation identical to that of A. zonella.

Male genitalia. (Specimen from Waterton National Park gate, Hwy. 5, Alberta). Tergum

with single line of long, well spaced setae parallel to posterior edge. Segment IX of roughly

uniform width throughout, sinuate (Fig. 206). Clasper massive, with cylindrical basal article,

and smaller distal article strongly hooked mesad, with ventral tooth (Fig. 206, 207). Seg-

ment X with short, cylindrical median lobes, projecting, triangular cerci, and large, dark,

trifid intermediate lobes hooked antero-laterad distally (Fig. 206, 207). Median shaft of

aedeagus stout, curved strongly ventrad; ejaculatory pore in membranous area between dis-

tal flaps (Fig. 208). Lateral arms long, each laminate blade on single membranous dorsal

process of aedeagal base.

Female genitalia. (Specimen from Cameron Lake, Waterton National Park, Alberta). Vul-

val scale of single median lobe; rectangular except for slightly expanded tip; distal end of

lobe rugose ventrally (Fig. 209). Segment IX massive, of indefinite shape (Fig. 210); open

ventrally and posteriorly with two short lateral lobes laterad of segment X and posterior

opening. Supra-genital plate small, membranous, ventrad of postero-dorsal opening of seg-

ment IX. Segment X minute, bilobed, between lateral lobes of segment IX.

Notes on biology. - I am uncertain as to the possible emergence sites of this species. The

single female from the Waterton Park gates may have emerged from the adjacent Waterton

River, or flown in from the nearby Maskinonge Lake. The remainder of my records are from

locations adjacent to large lakes. The adult flight season extends from July 23 to August 19.

Geographical distribution. — The known range of this species extends from Alberta to

Colorado (Fig. 601). In Alberta I have records from the extreme south west corner of the

province, at altitudes between 4,189’ and 5,445’.

I have examined 19 specimens, 14 males and five females, from the study area.

Rhyacophilidae and Limnephilidae

69

The wallengreni group

One species belonging to this group is known to occur in the study area.

Synopsis of characters. - Fore wing stigma usually weak; costal vein not thickened. Sc

and R1 of hind wing parallel throughout.

Male genitalia with segment IX narrow. Cerci setose, slightly concave mesally; triangular

and projected well posterad (Fig. 216). Median lobes of segment X very slender; separate,

or fused. Claspers unmodified, elongated, not thick, with basal article cylindrical; distal

article as long as basal or shorter, sickle shaped in most species.

Female genitalia with dorsal part of segment IX characteristic, with pits and ridges, large;

ventral lobes prominent <Fig. 219). Segment X long, slender; 1/3-1 /4 times diameter of seg-

ment IX; quite divided to dorsal and ventral parts in certain species. Supra-genital plate tri-

angular, shorter than segment X, weakly sclerotized.

Apatania crymophila McLachlan, 1880

(Fig. 216-220, 601)

Apatania crymophila McLachlan, 1880:44. (Type locality: Northwest Siberia). Ulmer,

1905a:23. Ulmer, 1907a:76. Schmid, 1954a:5-6. Schmid, 1955:81. Fischer, 1967:102-

103. (For references to Palaearctic literature see Fischer, 1967).

Apatelia aenicta Ross, 1 938c: 1 62-163. (Type locality: Churchill, Manitoba). Schmid, 1954a:

5.

Radema aenicta', Ross, 1944:297.

Males of this species are distinguished from males of other species of Apatania by dilated,

heavily setose, distal article of clasper (Fig. 216), and by short, crooked, distally bulbous,

median shaft of aedeagus (Fig. 218). Females are distinguished by simple median lobe of

vulval scale (Fig. 220), and segment X distinctly separated from segment IX (Fig. 219).

Description. - Antennae dark brown to almost black. Vertex of head black. Thorax

black. Legs black to deep brown. Spurs red-brown. Fore wing length of male 9.8 mm; uni-

form greyish brown, with large, opaque, stigmatic area. Venation identical to that of A.

zonella.

Male genitalia. (Specimen from Simpson Islands, Great Slave Lake, Northwest Territories).

Postero-dorsal edge with single line of long setae. Segment IX high, narrow (Fig. 216); with

row of about six short, heavy setae near clasper base. Clasper massive, with laterally com-

pressed distal article heavily setose. Median lobes of segment X sinuate, bilobed; cerci at-

tached laterad of median lobes, distally splayed. Intermediate lobes massive, hooked plates

(Fig. 216, 217). Median shaft and lateral arms of aedeagus attached to common base (Fig.

218); lateral arms dorsad of median shaft. Median shaft distally bulbous, tip directed ven-

trad. Lateral arms as long as median shaft, thin, recurved.

Female genitalia. (Specimen from Simpson Islands, Great Slave Lake, Northwest Terri-

tories). Sternum VII with posterior edge clothed by broad band of scattered, fine, hyaline

hairs. Vulval scale small, short, rectangular, disto-ventrally rugose, median lobe (Fig. 220).

Segment IX high, narrow, parallel-sided, with broad, short, lateral lobes laterad of segment

X (Fig. 219). No apparent supra-genital plate. Segment X with bilobed portion, and fused,

plate-like, median lobes with common central spur (Fig. 219).

Geographical distribution. — The known range of this species is Holarctic; in North Amer-

ica it extends from Alaska to Manitoba (Fig. 601). It is not yet recorded from Alberta, but

occurs in Great Slave Lake, Northwest Territories, which is so close to the northern bound-

ary of Alberta that it is reasonable to expect that it occurs in the Province.

70

Nimmo

I have examined 71 specimens, 43 males and 28 females, from Simpson Islands, Great

Slave Lake.

The complexa group

One species belonging to this group is known from the study area, and is here described

as new.

Synopsis of characters. — Male genitalia slightly enlarged. Segment IX with very long,

slender, paried, dorsal lobes (Fig. 221); fused with marked median channel or not. Cerci

small, oval. Intermediate lobes of segment X very long, slender. Median lobes varied in size,

small in some species, slender; interlocked with lobes of segment IX, shorter. Claspers varied.

Basal article long, cylindrical. Distal article much shorter, spinate; in some species long,

spiniform. Aedeagus short, stout.

Female genitalia with segment IX long throughout (Fig. 224). Segment X long, divided

to dorsal and ventral parts. Supra-genital plate very short, thick, unsclerotized, fused to

segment IX.

Apatania alberta Nimmo n. sp.

(Fig. 221-225,602)

Males of this species are distinguished from males of other species oi Apatania by aedea-

gus (Fig. 221) with lanceolate lateral arms attached laterad of aedeagal base at point where

aedeagus bends sharply ventrad. Females are distinguished by single, median lobe of vulval

scale abruptly broadened distally into rectangular head (Fig. 225).

Description. — Antennae very dark brown to black. Vertex of head black, setae hyaline.

Thorax quite black. Spur formula 1,2,4; brown. Fore wing length of male 8.5 mm; uniform

black. Cross-vein C-Rl white. Venation identical to that of A. zonella.

Male genitalia. (Specimen from Rapids Creek, Gap, Alberta). Tergum VIII with distinct

postero-distal ridge with single row of long setae. Segment IX with high-peaked dorsal strap

(Fig. 221); gradually widened ventrad, with small postero-ventral process. Clasper long,

slender, with tip of distal article with short, stout setae. Base of proximal article with slight

lateral depression; distal portion of article with long, slender setae. Segment IX with paired

postero-dorsal processes, long, tapered, acuminate blades, fused basally. Median lobes of

segment X directed postero-laterad, bilobed, with ventral lobe smaller; dark brown. Inter-

mediate lobes ventrad of median lobes, with bifid base recessed into segment IX, and long,

rounded, distally slightly dilated process in lateral aspect (Fig. 221). Intermediate lobes

acute triangular plates in dorsal aspect (Fig. 223); abruptly pinched off at tip. Cercus small,

dark, triangular, dorso-laterad of median lobe bases. Aedeagus with semi-cylindrical median

shaft; open on dorsal surface (Fig. 221, 222). Lateral arms angular, directed dorso-laterad,

attached laterad of aedeagal base.

Female genitalia. (Specimen from Rapids Creek, Gap, Alberta). Sternum VII with poste-

rior edge clothed with short, hyaline hairs. Vulval scale with single, massive, median lobe

(Fig. 224, 225); distal portion abruptly expanded laterally, rectangular, ventrally rugose.

Segment IX large, semi-rectangular in lateral aspect (Fig. 224), with slightly expanded ven-

tral portion. No evident latero-ventral lobes, or supra-genital plate. Segment X distinct from

segment IX, rounded rectangular in lateral aspect, with disto-lateral clefts; with membranous

ventral lobe dorsad of vaginal orifice; segment X completely open ventrally.

Notes on biology. — Adults of this species are usually associated with small, turbulent

creeks, but, on occasion, some are near large, smooth-flowing rivers. A flight season from

Rhyacophilidae and Limnephilidae

71

May 5 to June 22 is indicated.

Geographical distribution. — To date this species is known only from the Banff area of

Alberta (Fig. 602). All records are from areas at about 4,250’ to 4,550’ altitude.

I have examined 17 specimens, four males and 13 females, from the study area.

Holotype. - Male. Rapids Creek, Gap, Alberta; June 15, 1967; A. Nimmo.

Allotype. — Female. Same data as holotype.

Paratypes. — Same data as holotype; four females. Forty Mile Creek at Trans-Canada

Hwy., Banff National Park, Alberta; June 15, 1967; A. Nimmo; five females. Bow River at

Trans-Canada Hwy., Canmore, Alberta; A. Nimmo; one male, three females. As previous

record; May 23, 1967; A. Nimmo; one male. Road to Sundance Canyon, Banff, Alberta;

June 22, 1962; G. B. Wiggins; one male.

The type series has been assigned the type number 10,584 in the Canadian National

Collection, Ottawa.

The holotype, allotype, and seven female paratypes are in the Canadian National Collec-

tion. One male and two female paratypes are in the Royal Ontario Museum, Toronto,

Ontario. One male and two female paratypes are in the United States National Museum,

Washington. The remaining male and female paratypes are in the collection of the Strickland

Museum, Dept, of Entomology, University of Alberta, Edmonton, Alberta.

This species is named for the Province of Alberta.

The Subfamily Neophylacinae Schmid

Head short, large, eyes prominent. Ocelli small, placed well anterad. Pronotum short,

chaetose. Spur formula 1,2,2; 1,2,3; 1,2,4; 1,3,3; or 1,3,4. Meso-apical spur of male hind leg

modified in certain taxa. Wings rather variable, not reduced. Fore wings narrow, but greatly

widened at chord in some taxa. Hind wings shorter than fore. Fore wings irrorate in certain

taxa, hind wings darkened. Frenulum large; subcosta of hind wing with three to four very

long, basal spines. Fore wing venation complete; hind wings rather incomplete, with strong

sexual dimorphism. Fore wing R1 strongly arched at stigma, in some taxa united to Sc by

cross-vein. Discoidal cell long, narrow; chord strongly disrupted. Hind wings of Oligophle-

bodes and Neothremma with much reduced but constant venation coupled with sexual

dimorphism.

Male genitalia of short pieces in form of peculiar massive ensemble. Segment VIII seti-

ferous. Segment IX very large, enclosing remainder of genitalia, especially claspers; strongly

convex ventrally in form of plate fused to claspers; segment with pronounced lateral relief

(Oligophlebodes) or large appendage ( Neothremma ); dorsally very narrow, indistinguishable

from segment X. Dorsal lobes absent. Segment X visible only by three pairs of appendages.

Claspers small, not prominent, well recessed into segment IX; two articled, with basal article

reduced. Aedeagus, except in Neothremma, emergent from centre, or higher, of segment IX;

slender, barely movable; lateral arms present or absent.

Female genitalia with segments IX and X very small, closely fused. Segment IX of two

distinct parts; dorsal part short, or vestigial. Segment X fairly large, prominent, of two large,

lateral, quite distinct pieces; narrowly cleft dorsally, quite open ventrally. Vaginal aperture

wide and open on segment IX. Vulval scale simple or bilobed, attached to segment IX.

Ventral lobes of segment IX more or less fused to vulval scale.

Key to the Genera of Neophylacinae in Alberta and eastern British Columbia

la. Segment X of male two blunt lobes and two protuberant appendages (Fig. 226).

Clasper small, recessed into segment IX. Ventral lobes of female segment IX large,

72

Nimmo

fused anterad of vulval scale (Fig. 230) Oligophlebodes, p. 72.

lb. Segment X of male two very long blades strongly hooked ventrad (Fig. 244). Mesal

face of segment IX with long, bilobed spine. Claspers almost entirely fused. Ventral

lobes of female segment IX quite free (Fig. 248) Neothremma, p. 75.

The Genus Oligophlebodes Ulmer

This genus is represented in the study by three species, one of which is described as

new.

Synopsis of characters. — Spur formula 1,3,3. Apico-mesal spur of hind tibia thickened

basally, bristled on mesal face. Fore wing obliquely parabolic apically; hind wing slightly

larger, notched apically (Fig. 131). Venation constant, sexually dimorphic on hind wings.

R1 of fore wing joined to Sc by short cross-vein; chord disrupted little; posteriorly oblique

to body. R5 absent in male hind wing; only f2 present. Female hind wing with fl, f2, and f5

present.

Male genitalia with segment IX strongly developed, with pronounced lateral process

attached to median lobes of segment X. Median lobes of segment X rounded, concave,

separated by wide space. Aedeagus very small, embedded in membranous mass, very high in

segment IX. Claspers small, recessed in fissure of segment IX.

Female genitalia with ventral lobes of segment IX transverse plates, massive, not prom-

inent, weakly sclerotized, fused to ventral surfaces of vulval scale; also fused to sternum

VIII.

Key to the Males of species of Oligophlebodes found in Alberta and eastern British Columbia

la. Distal spine of clasper, short, curved dorsad (Fig. 238)

O. zelti Nimmo n. sp., p. 74.

lb. Distal spine of clasper straight (Fig. 226), or with disto-ventral tooth (Fig. 232)

2a

2a.(lb) Distal spine of clasper long, slender, straight (Fig. 226) . ... O. ruthae Ross, p. 72.

2b. Distal spine of clasper short, stout, with disto-ventral tooth (Fig. 232)

O. sierra Ross, p. 73.

Key to the Females of species of Oligophlebodes found in Alberta and eastern British

Columbia

la. Segment X in ventral aspect deeply cleft, lateral lobes roughly triangular (Fig.

231) O. ruthae Ross, p. 72.

lb. Segment X in ventral aspect not deeply cleft, or, if so, not widely separated or

triangular (Fig. 237, 243) 2a

2a.(lb) Lateral lobes of segment X closely appressed, vulval scale latero-ventrally exca-

vated (Fig. 237) O. sierra Ross, p. 73.

2b. Lateral lobes of segment X in ventral aspect bilobed (Fig. 243), with semi-circular

dorsal lobes O. zelti Nimmo n. sp., p. 74.

Oligophlebodes ruthae Ross, 1944

(Fig. 130a, 130b, 131a, 131b, 226-231,603)

Oligophlebodes ruthae Ross, 1944:283, 285, 300. (Type locality: Roe’s Creek, Glacier

National Park, Montana). Ross, 1949b: 127-128. Ross and Spencer, 1952:50. Schmid,

1955:99. Fischer, 1967:155. Schmid, 1968:687.

Rhyacophilidae and Limnephilidae

73

Males of this species are distinguished from males of other species of Oligophlebodes by

long, straight distal spine of clasper (Fig. 226). Females are distinguished by deeply divided,

well separated, triangular lateral lobes of segment X in ventral aspect (Fig. 231).

Description. — Antennae dark brown; scapes somewhat darker. Vertex of head uni-

formly dark brown. Thorax more or less uniform dark brown. Spur formula 1,2,2, in

both sexes in specimens examined here. Male only with mesal spur of hind leg swollen

basally; curved, with short, thin process on inner face of curve. Fore wing length of male

7.4 mm; uniform light yellow to yellow-brown. Venation of male and female wings as in

Fig. 130-131.

Male genitalia. (Specimen from Cameron Creek, Waterton National Park, Alberta). Ter-

gum and sternum VIII each with single, distinct row of long setae parallel to posterior edge.

Segment IX with high, thin dorsal strap and small, thin, irregular ventral body (Fig. 226).

Ventral area produced posterad as long, narrow, triangular plate, ventrad of claspers (Fig.

227). Clasper fused to segment IX, large, complex, partly recessed into segment IX; with

distinct, basally directed, long, dark, distal tooth; in ventral aspect tooth hooked mesad

(Fig. 226, 227). Segment X with cerci trapezoidal, small. Median lobes well separated, with

setose posterior edges. Lateral arms of aedeagus close to base of ejaculatory duct projected

below them (Fig. 228, 229); distal end of lateral arms curved mesad.

Female genitalia. (Specimen from Cameron Lake, Waterton National Park, Alberta).

Posterior edge of sternum VII with narrow fringe of short, fine setae. Vulval scale small,

semi-circular, dark, median lobe partly enclosed ventrally by ventro-lateral lobes of segment

IX (Fig. 231). Segment IX rectangular in ventral aspect, internally hollow, with ventral cleft;

in effect large hood. Supra-genital plate not evident. Segment VIII with simple invagination

on postero-lateral edge. Segment X bilobed (Fig. 231); lobes connected only basally; fused

almost imperceptibly to segment IX (Fig. 230).

Notes on biology. — The adult flight season extends from July 3 to August 29. The adults

appear to emerge from the mountain creek type of stream, ranging from relatively slow,

gentle, gravel-bottomed foothills creeks to the more torrential, boulder strewn creeks.

Geographical distribution. - The known range of this species extends from Alberta and

British Columbia to Oregon and Utah (Fig. 603). In Alberta it is found only in the mountain

and foothill areas, ranging in altitude from about 5,000’ to 6,000’.

I have examined 52 specimens, 35 males and 17 females, from the study area.

Oligophlebodes sierra Ross, 1944

(Fig. 232-237, 604)

Oligophlebodes sierra Ross, 1944:283, 284-285, 300. (Type locality: Dana Fork, Tolumne

River, Yosemite National Park, California). Ross, 1949b: 127. Ross and Spencer, 1952:50.

Schmid, 1955:99. Flint, 1960:32. Denning, 1963:261-262. Smith, 1965:244. Fischer,

1967:144. Schmid, 1968:685, 687.

Males of this species are distinguished from males of other species of Oligophlebodes by

short, stout distal spine of claspers, with disto-ventral tooth (Fig. 232). Females are distin-

guished by closely appressed lateral lobes of segment X, and by ventro-lateral concavities of

vulval scale.

Description. — Antennae light brown. Vertex of head uniform reddish brown. Thorax

uniformly yellow or pale purplish brown. Spur formula 1,3,3; meso-apical spur of male hind

tibia swollen basally, with short, thin process on inner edge of curve. Fore wing length of

male 7.6 mm; uniform pale brown, almost hyaline; no pattern except for pale grey stigma.

Venation identical to that of O. ruthae.

74

Nimmo

Male genitalia. (Specimen from Athabasca River, Entrance, Alberta). Tergum and sternum

of segment VIII with single rows each of setae parallel to posterior edges. Segment IX with

small ventral body; high, very thin dorsal strap (Fig. 232). Clasper relatively large, fused to

segment IX, distally black, with small disto-ventral tooth; with distinct, mesally directed,

horizontal hooks (Fig. 233). Segment X with median lobes well separated, with irregularly

dentate edges, and with distinct medially directed process (Fig. 233). Cercus large, rounded

triangular, attached to horizontal dorsal edge of segment IX. Aedeagus minute; ejaculatory

duct syringe-like, with bulbous inner end at attachment of membranous sperm duct (Fig.

234, 235); otherwise straight tube. Lateral arms dorsad and posterad of duct, overhung by

large fold of hyaline membrane.

Female genitalia. (Specimen from Idaho; in Illinois National History Survey). Tergum

VIII very large, separated from small, ventral sternum by wide band of folded membrane

(Fig. 236); with row of strong setae parallel to posterior edge. Vulval scale single, median,

dark, ventro-laterally concave lobe (Fig. 237). Segment IX fused almost indistinguishably to

segment X; antero-laterally invaginated anterad into segment VIII. Segment X of two closely

appressed lateral lobes.

Notes on biology. — My only record of this species in the study area is a single male taken

at black-light about 100’ above the Athabasca River, at the point where the Forestry Trunk

Road crosses, at Entrance, Alberta. At that point the river is quite turbulent, with an obvi-

ously rocky bed. The date was July 25.

Geographical distribution. — The known range of this species extends from Alberta and

British Columbia to California and Colorado (Fig. 604). The only Alberta record as given

above was at an altitude of about 3,270’.

Oligophlebodes zelti Nimmo n. sp.

(Fig. 238-243, 604)

Males of this species are distinguished from males of other species of Oligophlebodes by

short, black, dorsally curved distal lobes of claspers (Fig. 238), which are more widely sepa-

rated than in the similar O. sigma (Fig. 239). Also ventral plate of segment IX prominent,

horizontal and projected further posterad than in O. sigma. Females are distinguished by

bilobed lateral lobes of segment X in ventral aspect (Fig. 243).

Description. — Antennae light to dark brown. Vertex of head dark brown. Thorax uni-

form dark reddish brown. Spur formula 1,2,2; meso-apical spurs of male hind legs basally

enlarged, claw-like, with small spine basally, on inside of claw. Fore wing length of male 8.4

mm; pale yellowish brown, no pattern. Venation identical to that of O. ruthae.

Male genitalia. (Specimen from South Creek, Forestry Trunk Road, 20 miles south of

Nordegg, Alberta). Segment VIII with single band of long, fine setae each side of tergum,

parallel to posterior edge. Segment IX with high, narrow dorsal strap, with large, rounded

triangular ventral body, and with short, dark, triangular plate projected posterad, ventrad of

claspers (Fig. 238, 239). Clasper distally black, with distal process curved dorsad; in ventral

aspect tip hooked mesad, otherwise squat triangular. Segment X with membranous median

lobe and large, rectangular, setose, lateral lobes. Cercus small, dark, trapezoidal. Aedeagus

with short, stout ejaculatory tube surmounted by two dark, bowed, lateral arms slightly

bulbous distally (Fig. 240, 241). Median shaft of aedeagus surmounted by rectangular, mem-

branous mass lightly spinate dorsally. Sperm duct passed anterad by way of large, oval aper-

ture situated ventrally on membranous aedeagal sheath.

Female genitalia. (Specimen from South Creek, at Forestry Trunk Road, 20 miles south

of Nordegg, Alberta). Posterior edge of sternum VII with narrow band of short, fine well

Rhyacophilidae and Limnephilidae

75

spaced setae. Vulval scale dark, table-like on median stalk (Fig. 243). Segment IX trape-

zoidal, with antero-lateral invaginations hooked ventrad (Fig. 242). Ventro-lateral lobes of

segment IX square in lateral aspect, triangular in ventral aspect. Segment X fused smoothly

to segment IX, except dorsally, where slight demarcation is noticeable; roughly triangular in

lateral aspect; cleft dorsally. Segments IX and X together form large, cavernous hood,

entirely open ventrally.

Notes on biology. — Individuals of this species emerge from small, turbulent, boulder

or coarse gravel mountain streams. The adult flying season extends from July 14 to Au-

gust 19.

Geographical distribution. — This species is presently known only from the mountains

and foothills of Alberta (Fig. 604) between 4,700’ and 5,500’ in altitude.

I have examined 55 specimens, 16 males and 39 females, from the study area.

Holotype. — Male. South Creek, Forestry Trunk Road, 20 miles south of Nordegg,

Alberta; August 8, 1965; A. Nimmo.

Allotype. — Female. Same data as for holotype.

Paratypes. — Same data as for holotype; 36 females. Same data as for holotype, except

July 14, 1967; one male. Red Earth Creek, Trans-Canada Highway, Banff National Park,

Alberta; July 30, 1967; A. Nimmo; one male. Helen Creek, Banff-Jasper Hwy., north of

Lake Louise, Alberta; August 10, 1967; A. Nimmo; one female. Rowe Brook, Waterton

National Park, Alberta; August 19, 1965; A. Nimmo; one male. Lynx Creek, Forestry Trunk

Road, north of Clearwater River, Alberta; July 14, 1967; A. Nimmo; five males, one female.

South Creek, Forestry Trunk Road, 20 miles south of Nordegg, Alberta; August 12, 1968;

A. Nimmo; five males, one female. Wampus Creek, Cadomin, Alberta; August 18, 1967;

K. Zelt; one male.

The type series has been assigned the type number 10,589 in the Canadian National

Collection.

The holotype, allotype, and three male and 36 paratypes are in the Canadian National

Collection. The Lynx Creek paratypes are in the Strickland Museum, Dept, of Entomology,

University of Alberta, Edmonton, Alberta. One male and one female paratype are in the

United States National Museum.

This species is named for Ken Zelt, a graduate student in the Dept, of Zoology, University

of Alberta, who collected a single male near Cadomin.

The Genus Neothremma Banks

Two species of this genus are known from the study area.

Synopsis of characters. — Lateral ocelli almost immediately posterad of exceptionally

large anterior cephalic warts. Spur formula 1,3,4. Maxillary palpus of male with long brush

of distally hooked setae (Fig. 249). Fore wing venation unmodified; hind wing venation

much reduced, with some sexual dimorphism. Fore w'ing f3 tapered proximally to point;

chord strongly irregular (Fig. 132a). Hind wing with minute discal cell, fl with long proxi-

1 mal stem; male Ml+2, M3+4, Cul , Cu2 all separate; with three anal veins. In the female hind

wing M similar but Cu of three veins.

Male genitalia with segment IX well developed, not enclosing any appendages; laterally

with very long, bifid, sclerotized lobe directed postero-mesad. Segment X much smaller than

IX, well separated; median lobes long, simple, postero-ventrally arched blades, well separated

: basally by membrane of anus (Fig. 244). Clasper along ventral edges of segment IX; two-

| articled, with basal article of each clasper fused mesally as ventral plate; distal article baso-

dorsad on basal article; large, fused to segment IX, covered with distinct tubercles.

76

Nimmo

Female genitalia with segment IX of two pieces (Fig. 247). Segment X of two large dorsal

pieces, and ventral scale. Vulval scale large, bifid (Fig. 248). Ventral lobes of segment IX

large, slightly concave mesally, laterad of vulval scale.

Key to the Males of species of Neothremma found in Alberta and eastern British Columbia

la. Distal tooth of second article of clasper dorso-anterad of distal extremity (Fig. 244);

intermediate lobes of segment X roughly rectangular in lateral aspect

N. alicia Banks, p. 76.

lb. Distal tooth of second article of clasper a postero-dorsal continuation of article (Fig.

591a); intermediate lobes of segment X trifid in lateral aspect

N. laloukesi Schmid, p. 77.

Neothremma alicia Banks, 1930

(Fig. 132a, 132b, 244-249, 605)

Neothremma alicia Banks, 1930a: 229-230. (Type locality: Tolland, Colorado). Dodds and

Hisaw, 1925a: 127-129. Dodds and Hisaw, 1925b:386. Neave and Bajkov, 1929:202.

Betten, 1934:413. Milne, 1936:116, 123. Balduf, 1939:122. Ross, 1944:300. Ross,

1949a:92. Schmid, 1955:101. Flint, 1960:5, 31 . Denning, 1966:233. Fischer, 1967:145.

Unzicker, 1968:4, 20, 52.

Neothremma alicea; Ross, 1938b: 45.

Males of this species are distinguishable by high-arched, blade-like median lobes of seg-

ment X; by lateral processes of posterior edges of segment IX; and by fused basal articles of

claspers. Females are recognizable by massiveness of genitalia (Fig. 247), with huge, rounded,

latero-ventral lobes of segment IX.

Description. — Antennae dark brown, scapes paler, six times longer than pedicel, with

long bunch of hyaline hairs on mesal faces. Vertex of head red-brown. Frons with scattered,

spatulate-tipped hairs. Maxillary palpus of male with article III cylindrical, fringed on poste-

rior face with distinct, even brush of long, black, distally hooked hairs (Fig. 249). Thorax

light yellow to red-brown. Spurs yellow-brown. Fore wing length of male 7.7 mm; light red-

brown; no pattern. Venation as in Fig. 132a, 132b.

Male genitalia. (Specimen from Banff, Alberta). Tergum VIII with single line of long,

slender setae parallel to posterior edge. Segment IX with narrow, distinct, dorsal strap;

wide ventrally, with small, rounded, ventro-posterad process (Fig. 244, 245). Clasper mas-

sive, articulated to segment IX, with disto-dorsal teeth; two-articled, with distal article

dorsad of basal article, rounded, setose. Claspers fused ventrally together at basal articles.

Segment IX with long, distally bifid latero-posterad processes. Median lobes of segment X

long, dorsally arched, acuminate blades. Intermediate lobes roughly rectangular, flared

ventro-laterad. Aedeagus with massive membranous base (Fig. 246) connected to base of

proximal clasper articles; each clasper base with short, stout, black spine at point of fusion.

Median shaft of aedeagus tapered, sclerotized, scoop-shaped structure with ejaculatory duct

projected dorsad as thin, isolated tube.

Female genitalia. (Specimen from Bow River, at Trans-Canada Hwy., west of Lake Louise,

Alberta). Tergum VIII with single, dorsal line of long setae. Vulval scale with squat, tapered,

distally bifid median lobe (Fig. 248). Segment IX small, rectangular, with massive, rounded

latero-ventral lobes (Fig. 247). Segment X with large, triangular lateral lobes laterad of mesal

structure like inverted bowl. Vaginal orifice flanked by two lateral plates of segment X.

Notes on biology. - Specimens of this species emerge from small to large mountain

streams, usually of the less turbulent variety. Adult flight season extends from June 22 to

Rhyacophilidae and Limnephilidae

77

August 13.

Geographical distribution. — The known range of this species extends from Alberta and

British Columbia to Oregon, Utah and Colorado (Fig. 605). In Alberta it is restricted to the

mountain areas close to the continental divide, ranging in altitude from 4,500’ to 7,000’.

I have examined six specimens, two males and four females, from the study area.

Neothremma laloukesi Schmid, 1968

(Fig. 591a, 591b, 605)

Neothremma laloukesi Schmid, 1968:692-693. (Type locality: Lake Louise, Alberta).

Males of this species are distinguished from males of N. alicia by the characters presented

in the key to males of Neothremma known from the study area.

The only known specimen of this species is a male, from Lake Louise, Alberta. Being thus

unable to prepare my own drawings, Dr. F. Schmid very kindly lent me his original drawings

of the species, which are presented here (Fig. 591a, 591b). I also present Schmid’s original

description of the male, in translation from the French, altering only his figure numbers to

mine.

Description. — ‘Fore wings uniformly golden brown. Head abundantly clothed by very

long, golden hairs which are slightly denser between the scapes of the antennae. Male maxil-

lary palpi short, thick, with dense clusters of black hairs on mesal faces, as in N. alicia Banks.

Venation similar to alicia. Wing expanse of male 13 mm.

Male genitalia. Very similar to alicia, but basal article of clasper distinctly longer and

narrower (Fig. 591a). Distal article of clasper also longer in lateral aspect, with clear outline

gradually tapered from base to apex; distally curved dorsad at obtuse angle; regularly trape-

zoidal in ventral aspect (Fig. 591b), tapered distad.’

Female genitalia. Unknown.

Notes on biology. — Date of capture of the single known male was June 7.

Geographical distribution. — The single locality is indicated in Fig. 605.

The Subfamily Pseudostenophylacinae Schmid

Head short, very large; eyes large. Ocelli large, protuberant. Thoracic macrochaetae long,

dense. Spur formula 1,2,2; 1,3,3; or 1,3,4. Wings large; fore wing of varied sizes, apically

elliptical. Hind wing not larger than fore wing except when anal area well developed; anal

edge convex or not; anal area sexually dimorphic in some taxa. Certain taxa with fore wings

strongly granular, densely clothed with fine, bristly hairs; hind wings similar. Fore wings

brown, irrorate. Venation complete. R1 of fore wing not arched at stigma; discoidal cell

very long; chord irregularly disrupted, markedly curved in most taxa; f5 and thyridial cell

sessile. Hind wing chord similar, but more oblique to body.

Male genitalia segment VIII with strongly marked, spinate zone, or not. Segment IX very

short in lateral aspect, or longer. Cerci small, lateral, fused to intermediate lobe bases; inter-

mediate lobes large, massive, generally paired, or as single horizontal plate, or paired plates

elongated meso-dorsad. Claspers one-articled, reduced in certain taxa to simple buttons

along edge of segment IX. Aedeagus very large in certain taxa; voluminous but short; mem-

branous or sclerotized. Lateral arms enormous in most taxa, varied, membranous.

Female genitalia well sclerotized, loosely connected. Segment IX of two separate parts;

dorsal part simple, without appendages. Segment X not prominent, of two lateral lobes.

Ventro-lateral lobes of segment IX small, well sclerotized, well separated; meso-ventral part

small, membranous, attached to lateral lobes. No supra-genital plate. Vaginal aperture be-

78

Nimmo

tween segments VIII and IX. Vulval scale small, strongly sclerotized; lobes lightly connected,

movable; median lobe very small, narrow, very thick in some taxa; lateral lobes strongly

sclerotized, voluminous, very thick.

This subfamily is represented in the study area by one genus, Homophylax.

The Genus Homophylax Banks

This genus is represented in the study area by three species, of which one is described as

new. Females of only two species are known.

Synopsis of characters. — Spur formula 1,3,4. Wings large; fore wing markedly widened at

chord, bluntly parabolic apically; hind wing blunted apically, scarcely larger than fore wing.

Venation complete, modified, with strong sexual dimorphism in hind wings. Fore wing

chord single line, slightly oblique to body anteriorly, not disrupted. Male hind wing with

small, narrow discoidal cell; R and M crowded toward C; fl petiolate, f2 tapered proxi-

mad to point, f3 with long petiole. Hind wing of female with very large, triangular, dis-

coidal cell.

Male genitalia with sclerotized, smooth, posterior bulge to tergum VIII. Segment IX very

wide laterally, ventrally. Median lobes of segment X in form of two symmetrical, sclerotized

cavities dependent from very thin dorsal strap; fused mesally as strongly sclerotized mesal

ridges with dark, dorsal and ventral paired lobes. Intermediate lobes stout, twisted, lobed

plates fused to segment IX; interlocked with meso-dorsal blades of claspers. Claspers with

small, ventral plates; antero-mesal angles produced postero-dorsally as long, thin, acuminate,

strongly sclerotized, dark blades. Cerci large, somewhat oval lobes in lateral aspect (Fig.

250); fused at bases with sclerotized cavities of intermediate lobes. Aedeagus very small,

stout.

Female genitalia with dorsum of segment IX small. Segment X rather narrow, long tube

(Fig. 253). Ventral lobes of segment IX large, prominent, vertical, separate plates. Vulval

scale blunt, recurved; two or three lobed. Supra-genital plate very small.

Key to the Males of species of Homophylax found in Alberta and eastern British Columbia

la. Meso-dorsal lobes of clasper long, narrow (Fig. 250); median processes of segment

X with flat-topped dorsal spines with lateral teeth distally (Fig. 251)

H. crotchi Banks, p. 78.

lb. Meso-dorsal lobes of clasper short, wide, heavy (Fig. 225, 261). Dorsal spines of

median lobe of segment X without disto-lateral tooth 2a

2a. (lb) Median lobe dorsal spines fused basally, parallel sided (Fig. 256), with ventrally

directed distal hook (Fig. 255) H. acutus Denning, p. 79.

2b. Median lobe dorsal spines well separated basally, tapered, directed postero-laterad

(Fig. 262) H. baldur Nimmo n. sp., p. 80.

Key to the Females of two species of Homophylax found in Alberta and eastern British

Columbia

la. Segment X in ventral aspect (Fig. 254) narrow, with deep ventral cleft, shallow

dorsal cleft H. crotchi Banks, p. 78.

lb. Segment X in ventral aspect (Fig. 258) wide, with dorsal cleft at least as deep as

ventral H. acutus Denning, p. 79.

Homophylax crotchi Banks, 1920

(Fig. 133a, 133b, 250-254, 606)

Rhyacophilidae and Limnephilidae

79

Homophylax crotchi Banks, 1920:345-346. (Type locality: Victoria, British Columbia).

Betten, 1934:364. Milne, 1935:23, 51. Ross, 1938b:33. Ross, 1944:300. Ross and Spen-

cer, 1952:50. Schmid, 1955:114. Denning, 1964:253, 254. Fischer, 1967:156.

Males of this species are distinguished from males of other species of Homophylax by

small, laterally toothed dorsal spine of median lobes (Fig. 251), and by long, thin meso-

dorsal lobe of clasper. Females are distinguished by narrow segment X in ventral aspect (Fig.

254), with unequal dorsal and ventral clefts.

Description. - Antennae light red-brown. Vertex of head red-brown. Thorax warm light

red-brown. Spurs brown. Fore wing length of male 16.4 mm; light brownish yellow, in-

terspersed with slightly darker, irregular areas. Venation as in Fig. 133a, 133b. Without

basal fore wing pouch and longitudinal hind wing fold between Rs and M (see Fig. 134,

135).

Male genitalia. (Specimen from Washington State, United States; in United States Na-

tional Museum, Washington, D. C.). Tergum VIII with postero-dorsal area with distinct

tooth close to posterior edge; with distinct lateral concavities (Fig. 250). Segment IX

with dorsal strap narrowed laterally to junction with main body, expanded evenly ventrad.

Claspers with ventral lobes slightly separated (Fig. 251), short, rounded; lateral lobes finger-

like, thick, fleshy; median lobes long, narrow blades with slightly thickened black tips.

Median lobes or plates of segment X large, cupped laterad, with small, spiniform, recurved

ventral lobes, larger, flat-topped, dorsal lobes each with distinct disto-lateral tooth. Inter-

mediate lobes extended laterad from anal membrane, curved posterad above base of clasp-

ers. Cercus large, arched slightly dorsad; short, rounded distally. Aedeagus with large, mem-

branous, dorsal lobe dorsad of median shaft (Fig. 252); like inverted trough, with concave

ventral surface.

Female genitalia. (Specimen from Banff, Alberta). Sterna IV, V, and VI each traversed by

thin dark line parallel to posterior edge interrupted mesally by roughly triangular tooth.

Vulval scale with median lobe of two inconspicuous, small protuberances of posterior edge

(Fig. 254). Lateral lobes curved dorsad to enclose two sclerites projected from vagina. Seg-

ment IX with laterally triangular, shell-like lateral lobes suspended from short, narrow

dorsum. No supra-genital plate evident. Segment X long, tubular, of two lateral lobes distally

projected well posterad of all other structures (Fig. 253).

Geographical distribution. — The known range of this species extends from Vancouver

Island and Washington State to Alberta. In Alberta I have only two records of the species:

I Banff, at about 5,000’ on Sulphur Mountain; and Lost Lake, Waterton National Park, at

! 5,500’.

Dates of capture were September 10 and August 17 respectively. I have not taken any

specimens of this species myself.

I have examined one male and two females of this species; only the females are from the

study area, however.

Homophylax acutus Denning, 1964

(Fig. 134a, 134b, 255-259, 606)

Homophylax acutus Denning, 1964:254, 256-258. (Type locality: Wallace, Idaho). Nimmo,

1965:787.

Males of this species are distinguished from males of other species of Homophylax by

short, wide, heavy mesal lobes of claspers, and by large, basally fused, ventrally hooked

dorsal spines of median lobe of segment X (Fig. 255). Females are distinguished by large,

short segment X, with equal dorsal and ventral clefts (Fig. 258).

80

Nimmo

Description. — Antennae brownish yellow. Vertex of head pale yellow. Fore wing length

of male 16.4 mm; dull brownish yellow, with darker areas especially in posterior parts of

wing. Venation as in Fig. 134a, 134b. Hind wing with large, longitudinal fold posterad of

Rs; fold with numerous scales, especially basally. Male fore wing with soft, membranous

fold basally; female without fold.

Male genitalia. (Specimen from Mt. Edith Cavell, Jasper National Park, Alberta). Segment

IX with thread-like dorsal strap; main body of segment wide ventrally, tapered abruptly

dorsad, with slight concavities ventrad of peak (Fig. 255). Clasper with mesal lobe short,

wide, bowed mesad in dorsal aspect (Fig. 256); lateral lobe bifid distally, separated by deep

mesal cleft. Segment X with median processes of two toothed plates flared laterad in form

of deep, sclerotized concavities; dorsal hooks only slightly parted, square tipped; ventral

hooks directed dorsad, recurved. Intermediate lobes triangular, acuminate in lateral aspect.

Cercus with straight ventral edge, dorsal edge curved gradually ventrad. Aedeagus with

smooth, sclerotized, basal sheath followed by high, peaked dorsal lobe; median lobe sclero-

tized distally.

Female genitalia. (Specimen from Moraine Lake, Banff National Park, Alberta; in Cana-

dian National Collection). Similar to H. crotchi, but lateral lobes of vulval scale larger;

median lobe more pronounced (Fig. 259); vaginal sclerites more globose, with concave

mesal faces. Ventro-lateral lobes of segment IX triangular but more rounded than in H.

crotchi. Segment X heavier, larger, shorter; dorsal cleft deeper than ventral (Fig. 258).

Notes on biology. — This species is recorded from only two localities in Alberta to date.

One locality is a deep, morainic mountain lake; the second is located in high, alpine mead-

ows, with shallow pools and small water trickles. Altitudes are 6,200’ and 7,000’ respective-

ly. Dates of capture were August 6 and 22.

Geographical distribution. — The known range of this species extends from Idaho to

Alberta (Fig. 606). In Alberta it appears to be confined to high altitude creeks or pools.

I have examined two males and one female from the study area.

Homophylax baldur Nimmo n. sp.

(Fig. 135a, 135b, 260-263, 607)

Males of this species are distinguished by spiniform, widely separated dorsal processes of

median lobes of segment X (Fig. 262) and by small membranous dorsal lobe of aedeagus

(Fig. 263).

Description. — Antennae light yellow-brown; scapes with antero-mesal faces devoid of

long setae. Vertex of head yellowish to reddish brown, with red-brown band between lateral

ocelli. Thorax uniformly reddish yellow. Spurs red-brown. Fore wing length of male 17.3

mm; light yellowish brown, with slightly darker bands mesally and posteriorly. Anal flap of

fore wings with white scales. Venation as in Fig. 135a, 135b. Fore wing with basal flap or

pouch (Fig. 135a, 260). Hind wing with pronounced fold anterad of Cula; fold internally

with hyaline scales; scales commonest in basal area of wing.

Male genitalia. (Specimen from Cameron Lake, Waterton National Park, Alberta). Postero-

dorsal edge of tergum VIII triangular, raised plaque; in lateral aspect an overhanging tooth

(Fig. 261). Segment IX with anterior edges produced anterad; posterior edge continuous to

peak of dorsal strap. Dorsal strap virtually non-existent. Clasper with heavy, black, meso-

dorsal process directed postero-dorsad; with straight dorsal edges, sinuate ventral edges; base

warped at 90° to meet lateral lobe of clasper (Fig. 262). Median lobe of clasper fused ven-

trally, weakly divided. Segment X with two vertical median plates; plates toothed dorsally

and ventrally; produced laterad as sclerotized concavities; dorsal teeth separated, acuminate;

Rhyacophilidae and Limnephilidae

81

ventral teeth small, button-like. Cercus almost triangular, with rounded tips. Aedeagus with

small, membranous, dorsal process originated from membranous middle portion; bilobed in

dorsal aspect.

Female genitalia. Not known.

Notes on biology. - The one Alberta record of this species is a high (5,445’), large moun-

tain lake. The Utah record listed below is situated at 9,700’. Dates of capture were August

19, and September 17 respectively.

Geographical distribution. — The known range of this species is restricted to two localities

(Fig. 607): one in extreme south west Alberta, the other in Utah.

I have examined 23 specimens from the study area, all males, and one other male, from

Utah.

Holotype. — Male. Cameron Lake, Waterton National Park, Alberta; August 19, 1965;

A. Nimmo.

Paratypes. — Same data as for holotype; 22 males. La Baron Lake, Circleville Mountain,

15.9 miles west Junction, Pinto County, Utah, United States; September 17, 1967; G. E.

Ball; one male.

The type series has been assigned the type number 10,585 in the Canadian National

Collection, Ottawa. The holotype and 19 paratypes are in the Canadian National Collection;

the Utah specimen is in the Strickland Museum, Dept, of Entomology, University of Alberta,

Edmonton, Alberta; and one paratype each are in the Royal Ontario Museum, Toronto and

the United States National Museum, Washington.

This species is named for Baldur, a character of Norse mythology encountered in my

reading.

The Subfamily Limnephilinae Ulmer

Synopsis of characters. — Spur formula 1,3,4, but the following combinations are also

found: 1,1,1; 0,2,2; 1,2,2; 2,2,2; 0,3,3; or 1 ,3,3. Wings very varied in size, from very large to

little more than scales, dicosmoecine in shape in certain genera; hind wings with well devel-

j oped anal area; fore wing colour highly varied; hind wings hyaline in most genera. Frenulum

barely evident, of some long setae at extreme base of subcosta. Venation complete, only

I feebly varied except in Enoicyla and Phanocelia ; identical in both sexes. Fore wing dis-

coidal cell one to three times longer than its own stem; thyridial cell pedicillate in very

I few taxa. Hind wing chord more or less broken, generally very oblique posterad; with five

; anal veins.

Male genitalia simple, with three pairs of appendages. Tergum VIII with spinate or setose

postero-dorsal process or not. Segment IX constant; more or less elongate laterally, some-

what shortened ventrally; in most genera very reduced dorsally. Segment as whole deeply

recessed into segment VIII. Cerci rounded lobes; concave mesally, unarmed or not, with or

without teeth, ridges, or crenulations; small to very large, strongly sclerotized. Intermediate

lobes of segment X sclerotized, not very varied in form but varied in size; between cerci, not

fused. Lateral angles of segment IX tapered or not, curved mesad to effect a certain amount

of separation between anal and aedeagal cavities. Clasper one-articled, fused to segment IX;

generally comparable in form to very oblique cone, with apex directed dorso-posterad.

Aedeagus highly variable in form and size, long or not, slender, with distal spines; membran-

ous basally or distally; lateral arms generally large, with all degrees of reduction among taxa.

Female genitalia with segment IX of two parts. Dorsal part large or not, conical, tapered

i to segment X. Segment X variable; tubular, cylindrical or conical; cleft dorsally, ventrally,

or even laterally, various parts reduced to independent scales in certain genera. Ventral parts

82

Nimmo

of segment IX of two lobes and median part. Supra-genital plate present or absent. Vaginal

aperture between segments VIII and IX. Vulval scale trilobed or not; thickened, fleshy;

three lobes fused basally or not, but in the Limnephilini lateral lobes not entirely fused to

median; relative proportions of lobes highly variable among taxa.

Following is a key to tribes of Limnephilinae, using males only. The females proved in-

tractable in the attempt to discover cohesive key characters and are best identified in associ-

ation with the male, or by comparison with drawings. This key is good only for the study

area.

Key to the Males of the Tribes of Limnephilinae found in Alberta and eastern British

Columbia

la. Lateral arms of aedeagus simple, spiniform, or apparently so (Fig. 542), with no

accessory spines or lobes; originated dorsad or ventrad of aedeagal base (Fig. 531,

547, 568, 580), not laterad 2a

lb. Lateral arms clavate, multi-lobate, multi-spinate, parti-membranous, or combina-

tion of these (Fig. 473, 486, 527); or if spiniform (Fig. 367), attached laterad of

aedeagal base Limnephilini, p. 82.

2a.(la) Ejaculatory duct terminated at extreme tip of median shaft of aedeagus, including

lateral lobes or processes, excepting lateral arms (Fig. 531, 536, 547)

Stenophylacini, p. 142.

2b. Ejaculatory duct terminated basad of extreme tip of median shaft, on dorsal

surface of shaft (Fig. 550, 553), or between longer distal lobes (Fig. 565)

Chilostigmini, p. 148.

The Tribe Limnephilini Schmid

Character synopsis of the Limnephilini. — Pronotum more developed than in other tribes

of subfamily. Base of pro-femur and apex of opposing tibia with or without black brushes.

Spur formula varied. Wings medium or small, not reduced, similar in both sexes. Fore wing

evenly strap-like, little wider at stigma, with oblique, truncated apex. Hind wings much

larger than fore. Fore wing coloration strongly contrasted; with large clear streak in mid-

wing, second at proximal end of apical cells, third at distal end of M4+5, and fourth in

thyridial cell. Chord of fore wing very oblique to body, narrowly broken. Hind wing chord

parallel to body, zigzagged regularly, strongly accentuated.

Male genitalia with posterior edge of tergum VIII finely, not densely, spinate. Segment IX

with postero-lateral edges convex or not, as supported to cerci. Cerci very varied in size and

form. Intermediate lobes of segment X varied, pincer-like in opposition to neighbouring

cerci or not; with lateral teeth or not. Claspers varied, base button-like, to almost vestigial;

free part slender, directed almost horizontally. Aedeagus strong, large; median shaft very

simple, unarmed, folded and extensible at base or not. Lateral arms with slender base and

expanded, spinate tip.

Female genitalia with segment IX of two parts in most taxa, well developed, close set.

Dorsal part prominent or not, without prominent lobes. Appendages present or not; large,

free; fused solidly, either to segment IX, or segment X. Segment X much smaller than

segment IX; cylindrical, slender, deeply cleft or not. Segment X large, with thick, fleshy

walls, hardly cleft at all; extended as one piece, not separate scales as in Stenophylacini.

Ventral lobes of segment IX large, convex, in contact ventrally. Supra-genital plate large,

free, prominent, ogival. Vulval scale trilobed, incompletely fused to sternum VIII, inter-

vening sutures clearly visible.

Rhyacophilidae and Limnephilidae

83

Key to Genera and Subgenera of Liinnephilini found in Alberta and eastern British Columbia

la. Apex of fore wing notched (Fig. 141a) . . .Nemotaulius ( Macro taulius), p. 123.

lb. Apex of fore wing smooth, without indentations 2a

2a.(lb) Apical spur of fore tibia large, triangular Philarctus, p. 132.

2b. Spurs normal 3a

3a. (2b) Fore wing with one or more longitudinal, median, silver lines bordered with

black Hesperophylax, p. 139.

3b. Fore wings without such lines 4a

4a. (3b) R4+5 of hind wings strongly tinted brown Gr ammo taulius, p. 122.

4b. R4+5 of hind wing not so coloured. Spur formula 1 ,3,4; 1,2,3; or 1 ,2,2 .... 5a

5a.(4b) Dorsal strap of segment IX well developed 6a

5b. Dorsal strap of segment IX very narrow, recessed into segment VIII 8a

6a.(5a) Wing span less than 20 mm; fore wing weakly irrorate Arctopora, p. 133.

6b. Wing span greater than 25 mm; fore wing strongly irrorate 7a

7a.(6b) Dorsal strap of segment IX of male large plate overhanging remainder of geni-

talia (Fig. 495) Lenarchus (Lenarchus), p. 135.

7b. Dorsal strap of segment IX of male quite short, but prolonged by large plate

formed by fused cerci (Fig. 501) Lenarchus (Paralenarchus), p. 136.

8a.(5b) Cercus of male strongly toothed, segment IX narrow throughout (Fig. 440)

Clistoronia (Clistoroniella), p. 120.

8b. Characters otherwise 9a

9a.(8b) Sc. of hind wing turned sharply anterad distally (Fig. 138b)

Limnephilus, p. 83.

9b. Sc. of hind wing only slightly turned anterad (Fig. 142b) 10a

10a.(9b) Intermediate lobes of male segment X much smaller than cerci, button-like

Asynarchus , p. 128.

10b. Intermediate lobes of male segment X plate-like, spiniform, or reduced ... 11a

1 la.(10b) Fore wing reddish, uniformly irrorate, or with regularly spaced minute patches

of brown Anabolia, p. 124.

lib. Fore wings otherwise colored; patterned with bars or large patches of colour,

ranging from black to almost hyaline Limnephilus, p. 83.

The Genus Limnephilus Leach

This genus is represented in the study area by 33 species, of which two are new, and one

is known only from the female. In 1955 Schmid arranged the species of the genus in species

groups. Sixteen groups are known from the study area. Besides these are three species which

he did not classify, even though he placed other single species in groups of their own. In this

study these three single species are placed in monotypic groups. According to Schmid

(1955) the characters of this genus are the same as for the tribe.

Key to the Males of species of Limnephilus from Alberta and eastern British Columbia

la. Postero-dorsal edge of tergum VIII spinate or setose (Fig. 370, 402) 2a

lb. Postero-dorsal edge of tergum VIII not spinate or setose (Fig. 309) 23a

2a.(la) Spinate area of tergum VIII produced posterad, or ventrad, to varying degrees

(Fig. 326, 342,385,427) 3a

2b. Spinate area of tergum VIII not produced (Fig. 273, 370, 421) 21a

3a.(2a) Spinate area of tergum VIII light or heavy bulbous lobe projected well posterad

of membranous connection to segment IX (Fig. 291, 390, 402, 408) 4a

84

Nimmo

3b.

4a. (3a)

4b.

5a. (4a)

5b.

6a. (5a)

6b.

7a. (6a)

7b.

8a. (7a)

8b.

9a. (7b)

9b.

10a.(6b)

10b.

1 la.(5b)

lib.

12a.(l lb)

12b.

13a.(4b)

13b.

14a.(13a)

14b.

15a.(14a)

Spinate area of tergum VIII not projected well posterad of membranous con-

nection to segment IX; not bulbous or spinate 18a

Mesal face of cercus with one or more black, strongly sclerotized teeth (Fig.

270b, 292,321) 5a

Mesal face of cercus without such teeth 13a

Teeth basad of distal edge of cercus in single, dorso-ventral row (Fig. 265, 278,

301,320) 6a

Teeth of cercus not arranged thus 11a

Lateral arms of aedeagus distally as acuminate, meso-dorsal, sclerotized teeth

flanked by at least partly membranous, extensible ventro-lateral lobe (Fig. 266,

280, 293) 7a

Lateral arms of aedeagus, if distally divided, with ventro-lateral lobes sclero-

tized, rigid, not membranous or extensible (Fig. 299, 322) 10a

Median lobes of segment X long, laminate, narrow, dorsally curved blades (Fig.

264, 291) 8a

Median lobes of segment X not as above; short, wide basally, tapered distally

(Fig. 270b, 278) 9a

Median lobes of segment X parallel almost to tips (Fig. 264); spinate dorsal

process of tergum VIII large, globose L. sublunatus Provancher, p. 89.

Median lobes of segment X tapered evenly and gradually distad (Fig. 291);

spinate postero-dorsal process of tergum VIII long, tapered, directed ventrad

L. susana Nimmo n. sp., p. 93.

Distal process of clasper long, thin, tapered (Fig. 270a); spinate process of

tergum VIII large, globose L. sansoni Banks, p. 90.

Distal process of clasper short, stout, blunt (Fig. 278); spinate process of

tergum VIII small, thumb-like L. hageni Banks, p. 91.

Tip of clasper black, strongly sclerotized, with dorsally directed tooth (Fig.

320); median lobes of segment X hooked postero-ventrad

L. externus Hagen, p. 99.

Tip of clasper not so armed, blunt (Fig. 298); median lobes of segment X

tapered, directed directly postero-dorsad, with no hook

L. indivisus Walker, p. 95.

Median lobes of segment X directed directly postero-dorsad (Fig. 326); spinate

lobe of tergum VIII directed ventrad, located between cerci

L. sericeus (Say), p. 100.

Median lobes of segment X wide basally, tapered abruptly to thin tooth curved

dorso-anterad (Fig. 385, 402); spinate process of tergum VIII directed back-

wards over genitalia 12a

Clasper with wide base, tapered gradually postero-dorsad (Fig. 385)

-. L. perpusillus Walker, p. 112.

Clasper originated abruptly from base, almost rectangular in lateral aspect,

divided shallowly distally (Fig. 402) L. labus Ross, p. 115.

Median lobes of segment X with distal portion curved dorsad (Fig. 305, 365,

408) 14a

Median lobes of segment X otherwise 17a

Median lobes of segment X long, narrow, evenly tapered throughout length

(Fig. 304, 390) 15a

Median lobes of segment X otherwise 16a

Clasper with wide base, abruptly narrowed to finger-like distal portion (Fig.

Rhyacophilidae and Limnephilidae

85

15b.

16a. (14b)

16b.

17a. (13b)

17b.

18a.(3b)

18b.

19a.(18a)

19b.

20a.(18b)

20b.

21a.(2b)

21b.

22a. (2 lb)

22b.

23a. (lb)

23b.

24a. (23a)

24b.

25a.(24a)

25b.

26a. (24b)

26b.

27a.(23b)

27b.

28a. (27a)

28b.

304) L. infernalis (Banks), p. 96.

Clasper short, blunt, with short base (Fig. 390) .... I. argenteus Banks, p. 113.

Clasper base very short; cercus narrow, long, dark, strongly sclerotized on distal

edge (Fig. 408, 409) L. minusculus (Banks), p. 116.

Clasper with long base; cercus with wide base, short, rounded (Fig. 365)

L. spinatus Banks, p. 108.

Both cerci and median lobes of segment X short, strongly sclerotized, massive

(Fig. 433, 434) L. canadensis Banks, p. 120.

Both cerci and median lobes of segment X long, slender, tapered, strongly

sclerotized (Fig. 332, 333) L. femoralis (Kirby), p. 102.

Lateral arms of aedeagus distally bilobed (Fig. 286, 429); cercus relatively

short, broad 19a

Lateral arms of aedeagus simple (Fig. 344, 350); cercus very long, slender ....

20a

Cercus long, parallel-sided; clasper short (Fig. 427) . .L. rhombicus (L.), p. 118.

Cercus short, triangular; clasper with long, narrow distal process (Fig. 284). . . .

L. partitus Walker, p. 92.

Postero-dorsal edges of segment IX distinctly concave; clasper long, acuminate

(Fig. 350) L. valhalla Nimmo n. sp., p. 106.

Postero-dorsal edges of segment IX not concave; clasper short, blunt (Fig. 342)

L. moestus Banks, p. 104.

Clasper massive, with distinct dorsal tooth (Fig. 421)

L. nigriceps (Zetterstedt), p. 117.

Clasper otherwise (Fig. 370) 22a

Median lobes of segment X long, with irregular edges; disto-lateral tooth present

(Fig. 273, 274); no teeth on mesal face of cercus. . . L. extractus Walker, p. 91.

Median lobes of segment X short, with smooth edges; directed postero-mesad

to teeth of mesal faces of cerci (Fig. 370, 371) .... L. hyalinus Hagen, p. 109.

Clasper with long, narrow base (Fig. 375, 414) 24a

Clasper with short base (Fig. 337, 355, 360) 27a

Lateral arm of aedeagus expanded distally; blade-like, fringed peripherally with

spines or setae (Fig. 3 1 1 , 3 1 7) 25a

Lateral arm not expanded distally (Fig. 377, 416) 26a

Cercus deeply cleft distally, with distal ends of each lobe black, strongly sclero-

tized (Fig. 309, 310) L. ornatus Banks, p. 97.

Cercus not cleft, trapezoidal in lateral aspect (Fig. 315); meso-distal edge with

distinct, regular, black tooth or spine (Fig. 316)

L. picturatus McLachlan, p. 98.

Lateral arm of aedeagus reduced; simple, distally spinate, membranous lobe

(Fig. 377); cercus triangular, vertically high (Fig. 375)

L. secludens Banks, p. 110.

Lateral arm of aedeagus long, thin, with three or four distal spines across

median shaft (Fig. 416, 417); clasper large, thin plate directed mesad, with

black, dentate, dorsal edge (Fig. 414, 418) L. kennicotti Banks, p. 116.

Tips of median lobes of segment X directed dorso-laterad (Fig. 355, 380). . 28a

Tips of median lobes of segment X not directed dorso laterad (Fig. 337, 347.

360, 395) 29a

Clasper with disto-lateral tooth (Fig. 380) L. janus Ross, p. 111.

Clasper without such tooth; much smaller than median lobes of segment X

86

Nimmo

29a. (27b)

29b.

30a.(29a)

30b.

31a.(29b)

31b.

Key to the

la.

lb.

2a.(la)

2b.

3a.(2b)

3b.

4a.(3b)

4b.

5a. (4a)

5b.

6a. (5a)

6b.

7a. (6a)

7b.

8a. (6b)

8b.

9a. (5b)

L. lopho Ross, p. 107.

Lateral arm of aedeagus curved sharply dorsad; not bilobed (Fig. 347, 398)

30a

Lateral arm of aedeagus straight or, if turned dorsad, not sharply; not bilobed

distally (Fig. 339, 362) 31a

Cercus short, wide, thick (Fig. 395, 396) L. alberta Denning, p. 114.

Cercus long, narrow, thin, with concave mesal face (Fig. 347, 349)

L. cockerelli Banks, p. 105.

Cercus long, narrow, tapered, with meso-distal tooth; median lobes of segment

X set higher than cerci, arched, lyre-shaped in dorsal aspect (Fig. 337, 338)

L. nogus Ross, p. 103.

Cercus short, squat, distally rounded, with concave mesal face; median lobes of

segment X set low, between cercus, lamellar, parallel (Fig. 360, 361)

L. parvulus (Banks), p. 107.

Females of species of Limnephilus from Alberta and eastern British Columbia

Segment X distinct from segment IX, either by distinct suture line or abrupt

decrease in size, or both. Segment X partly recessed into segment IX or not

(Fig. 268, 295, 331, 406, 431) 2a

Segment X fused to segment IX; no suture lines or abrupt decrease in size;

demarcated by slight declivity or not (Fig. 296, 318, 359, 363, 400, 419, 426)

17a

Segment IX with ventro-lateral lobes separate, distinct; demarcated by sutures

(Fig. 271, 290, 324, 431, 438) 13a

Segment IX with ventro-lateral lobes, or not; if present, an integral part of seg-

ment, not separated by sutures (Fig. 268, 331, 340, 369, 388, 406) 3a

Segment IX with distinct dorsal and ventral portions joined by laterally con-

stricted strap; segment X flanked dorsally and ventrally, not laterally (Fig. 268,

295,388) 11a

Segment IX not constricted laterally, or segment X not flanked dorsally, or

ventrally (Fig. 336, 369, 406, 412) 4a

Segment X with dorso-lateral lobes (Fig. 313, 331, 336, 383, 406, 412) ... . 5a

Segment X without dorso-lateral lobes (Fig. 340, 369) 10a

Opposing edges of median and lateral lobes of vulval scale very close, at least at

base; if close only at base, in form of v-pattem (Fig. 330, 384, 407, 413) .. . 6a

Opposing edges of vulval scale lobes markedly separated (Fig. 314, 336). ... 9a

Dorso-lateral lobes of segment X tapered in lateral aspect (Fig. 383, 412) ... 7a

Dorso-lateral lobes of segment X not tapered; blunt or rounded distally (Fig.

331,406) 8a

Lateral lobes of vulval scale with concave ventral faces in lateral aspect (Fig.

383) L. janus Ross, p. 111.

Lateral lobes of vulval scale in lateral aspect with ventral faces not concave,

median lobe markedly longer than lateral lobes (Fig. 412)

L. minusculus (Banks), p. 116.

Meso-dorsal lobes of segment X, immediately dorsad of anus, black, long, very

thin, acuminate in lateral aspect (Fig. 331) L. sericeus (Say), p. 100.

Meso-dorsal lobes of segment X immediately dorsad of anus short, triangular,

blunt in lateral aspect (Fig. 406) L. labus Ross, p. 115.

Dorso-lateral lobes of segment X long, thin, finger-like (Fig. 313)

Rhyacophilidae and Limnephilidae

87

9b.

10a.(4b)

10b.

1 la.(3a)

lib.

12a.(l la)

12b.

13a.(2a)

13b.

14a.(13a)

14b.

15a.(14a)

15b.

16a.(14b)

16b.

17a.(lb)

17b.

18a.(17a)

18b.

19a.(18a)

19b.

20a.(19a)

20b.

2 1 a.( 1 9b)

L. omatus Banks, p. 97.

Dorso-lateral lobes of segment X short, squat, triangular (Fig. 335, 336)

L. femoralis (Kirby), p. 102.

Segment IX with distinct, trapezoidal, lateral lobes projected postero-laterad of

segment X (Fig. 340) L. nogus Ross, p. 103.

Segment X without such lobes, with minute dorsal portion (Fig. 369)

L. spinatus Banks, p. 108.

Cercus short, squat, rounded, appressed to top surface of segment X (Fig. 295,

388) 12a

Cercus long, lamellar, attached to segment IX basally, otherwise free (Fig. 268)

* L. sublunatus Provancher, p. 89.

Median lobe of vulval scale approximately equal to lateral lobes (Fig. 389);

segment X oriented vertically in lateral aspect (Fig. 388)

L. perpusillus Walker, p. 112.

Median lobe of vulval scale projected well beyond lateral lobes, up to twice

their length (Fig. 294); segment X oriented antero-posterad (Fig. 295)

L. susana Nimmo n. sp., p. 93.

Genitalia with cercus or cercus-like lobes dorsad of segment X (Fig. 271, 290,

324,431) 14a

Genitalia without cerci; segment X conical in ventral aspect (Fig. 437)

L. canadensis Banks, p. 120.

Vulval scale small, shallowly recessed into sternum VIII (Fig. 290, 325); cercus

in lateral aspect not divergent from segment X (Fig. 290, 325) 15a

Vulval scale large, deeply recessed into sternum VIII (Fig. 272, 432); cercus in

lateral aspect divergent widely from segment X (Fig. 271, 431) 16a

Ventro-lateral lobe of segment IX divided as two sclerites (Fig. 290)

L. partitus Walker, p. 92.

Ventro-lateral lobe of segment IX undivided, of one piece (Fig. 324)

L. externus Hagen, p. 99.

Base of cercus dorsad of segment X (Fig. 271), squarely cleft disto-laterally

L. sansoni Banks, p. 90.

Base of cercus antero-dorsad of segment X (Fig. 431); segment X not disto-

laterally cleft L. rhombicus (L.), p. 118.

Ventro-lateral lobe of segment IX separate and distinct from dorsal part of seg-

ment by suture or membrane (Fig. 307, 3 18, 363, 419) 18a

Ventro-lateral lobe of segment IX not distinguished from dorsal part of seg-

ment by suture (Fig. 282, 346, 378, 400) 25a

Segment X with pair of dorso-lateral lobes or cerci (Fig. 302, 307, 318, 374)

19a

Segment X without dorso-lateral lobes (Fig. 363, 394, 419, 426) 22a

Distal end of segment X pair of meso-dorsal lobes or scales and single ventral

lobe (Fig. 308,319) 20a

Distal end of segment X without such lobes (Fig. 303, 373) 21a

Ventro-lateral lobes of segment IX roughly rectangular in lateral aspect, ori-

ented vertically (Fig. 318), the two lobes meeting but not fused ventrally (Fig.

319) L. picturatus McLachlan, p. 98.

Ventro-lateral lobes of segment IX triangular in lateral aspect (Fig. 307); not

meeting ventrally (Fig. 308) L. infernalis (Banks), p. 96.

Dorso-lateral lobes of segment X flanked laterally by rectangular walls (Fig.

88

Nimmo

21b.

22a. ( 18b)

22b.

23a. (22b)

23b.

24a.(23b)

24b.

25a. (17b)

25b.

26a. (25a)

26b.

27a.(26a)

27b.

28a.(26b)

28b.

29a.(25b)

29b.

30a. (29a)

30b.

31a.(29b)

31b.

373, 374) L. hyalinus Hagen, p. 109.

Dorso-lateral lobes of segment X free, not flanked laterally (Fig. 302)

L. indivisus Walker, p. 95.

Segment X composed of massive, mesally completely cleft, postero-dorsal plate

(Fig. 419, 420) L. kennicotti Banks, p. 116.

Segment X otherwise 23a

Segment X indistinguishable from segment IX; massive, blunt, with small, sim-

ple anal aperture at extreme posterior end (Fig. 425, 426)

L. nigriceps (Zetterstedt), p. 117.

Segment X not massive; deeply cleft mesally, tapered posterad in lateral aspect

(Fig. 364, 394) 24a

Ventro-lateral lobes of segment IX high, narrow, oriented vertically (Fig. 394)

L. argenteus Banks, p. 113.

Ventro-lateral lobes of segment IX almost as high as wide, with no special

orientation (Fig. 363) L. parvulus (Banks), p. 107.

Ventro-lateral part of segment IX produced posterad as narrow, tapered lobe

(Fig. 282, 296, 346, 353) 26a

Ventro-lateral part of segment IX produced posterad at most as very broad-

based, abruptly tapered lobe (Fig. 276, 359, 378, 400) 29a

Dorso-lateral lobes, or cerci, of segment X long, thin, well separated from

remainder of vulval scale much longer than lateral lobes 27a

Dorso-lateral lobes, or cerci, of segment X short, rounded distally, lamellar (Fig.

346, 354); median lobe of vulval scale no longer than lateral lobes 28a

Segment X cleft laterally (Fig. 296) L. species 1, p. 94.

Segment X not cleft laterally (Fig. 282) L. hageni Banks, p. 91.

Segment X with dorsal lobes located laterally (Fig. 354)

L. valhalla Nimmo n. sp., p. 106.

Segment X with dorsal lobes more mesally located (Fig. 345)

L. moestus Banks, p. 1 04.

Segment X cleft laterally (Fig. 276, 378) 30a

Segment X not cleft laterally (Fig. 359, 400) 31a

Dorso-lateral lobes, or cerci, of segment X small, short, reaching no more than

halfway to extremities of segment (Fig. 378) . . . L. secludens (Banks), p. 110.

Dorso-lateral lobes, or cerci, of segment X large, triangular, equally as long as

segment (Fig. 276) L. extractus Walker, p. 91.

Segment X deeply cleft dorsally (Fig. 401) L. alberta Denning, p. 114.

Segment X not deeply cleft dorsally (Fig. 358) L. lopho Ross, p. 107.

The subcentralis group

Members of this group are recognizable in the field by wing pattern. This consists of a

series of dark brown bars on lighter background; initially with median longitudinal band

in vicinity of subradial cell, terminated at chord; distad of chord are two shorter bands, one

between fl and f2, the other on f4; remainder of wing irrorate to very irregularly patterned.

Males are distinguished by thin, slightly concave mesal cerci, usually with short or long row

of black teeth basad of distal ends; and by lamellate, tapered, median lobes of segment X

(Fig. 273, 278, 291). Females are distinguished by presence of cerci or cercus-like postero-

dorsal lobes of segment X, and by segment X being separated from remainder of genitalia

by suture line (Fig. 268, 271, 290).

Rhyachophilidae and Limnephilidae

89

Limnephilus sublunatus Provancher, 1877

(Fig. 138a, 138b, 264-269, 607)

Limnophilus sublunatus Provancher, 1877:243. (Type locality: Quebec). Provancher, 1878a:

146. Provancher, 1878b: 131-132. Schmid, 1955:135.

Limnephilus sublunatus ; Banks, 1907a:37. Betten, 1926:529. Betten, 1934:333-334. Milne,

1935:44, 52. Milne, 1936:59. Ross, 1938b:34. Betten and Mosely, 1940:129. Banks,

1943:341-342. Ross, 1944:298. Ross and Merkley, 1952:448. Robert, 1960:59. Wiggins,

1961:700-701. Fischer, 1968:331-332.

Limnophilus americanus Banks, 1900a:253. (Type locality: Idaho). Ulmer, 1905a: 19, Ul-

mer, 1907a:43. Essig, 1926:175. Fischer, 1968:331.

Limnephilus americanus Banks, 1907a:36. Betten, 1934:320. Milne, 1935:52. Ross, 1938b:

34. Fischer, 1968:331.

Limnephilus macgillivrayi Banks, 1908a: 263. (Type locality: Axton, New York). Banks,

1930a:226. Milne, 1935:44, 51. Milne, 1936:59. Fischer, 1968:331-332.

Males of this species are distinguished from males of other species of the group by long,

smoothly curved, lamellate cerci (Fig. 264), and by similar median lobes of segment X

which are curved dorsad and rather blunt distally. Females are distinguished by thin, deli-

cate cerci of segment X, which are apparently attached to segment IX; segment X recessed

into segment IX, long, tapered, thin walled, in lateral aspect (Fig. 268).

Description. — Antennae pale yellow. Vertex of head brown-yellow, except for paler

warts. Thorax pale yellow, with light brown patches on dorsal areas of pleura. Brush of

anterior femur of male half length of femur; sparse, with short, black spines. Fore wing

length of male 13.9 mm; orange-brown, clear areas hyaline; costal margin clear for two-

thirds of length. Stigma light brown. Venation as in Fig. 138a, 138b.

Male genitalia. (Specimen from Canmore, Alberta). Postero-dorsal edge of tergum VIII

produced posterad as large spinate bulb; spines black, sparse. Dorsal strap of segment IX of

uniform width, overhanging main body of segment at angle of about 45° (Fig. 264). Main

body of segment IX almost square; posterior edge directed mesad under postero-dorsal

corner, which has finger-like aspect laterally. Clasper long, with wide base tapered quickly

to finger-like dorsal portion. Segment X with median lobes long, thin, lamellate; distal

third of dorsal edges black; lobes slightly up-turned in lateral aspect, warped in dorsal aspect

(Fig. 265). Cercus long, narrow, of uniform width, with distal end black-toothed; with

slightly darker dentate ridge two-thirds of distance from base (Fig. 265). Lateral arms of

aedeagus bipartite distally; dorsal lobe sclerotized, dorsally directed spine (Fig. 266); ventral

lobe membranous except for clear, sclerotized tip with fringe of brown setae.

Female genitalia. (Specimen from Canmore, Alberta). Vulval scale with slight concavities

at posterior edges of lateral lobes; median lobe distally blunt, longer than lateral lobes (Fig.

269). Segment IX with lateral faces deeply depressed, recessed anterad as flanges. Ventral

area directed posterad under segment X as acuminate plates (Fig. 268). Supra-genital plate

large, semicircular, with hyaline basal area; with rectangular, membranous, basal area. Seg-

ment X very deeply cleft mesally (Fig. 269), with acuminate distal tips to lateral lobes; in

lateral aspect segment tapered gradually posterad to tips. Cercus long, narrow, distal half

clothed with short, fine, hyaline setae; cercus slightly longer than segment X, apparently

attached to segment IX.

Notes on biology. — Specimens of this species appear to emerge from pools, sloughs or

lakes fringed with horse-tails, or sedges. The flight period extends from July 1 to October 3.

My records are insufficient to define a peak.

Geographical distribution. — The known range of this species extends from British Co-

90

Nimmo

lumbia, Idaho and Colorado to New Hampshire and Quebec (Fig. 607), with a large, mid-

continental blank. In Alberta it is confined to the passes, and low valleys, of the mountains

and foothills. In altitude it ranges from about 4,100’ to 6,878’. I have taken specimens at

the upper limit, but on the whole it is usually taken at the lower altitudes.

I have examined 45 specimens, 17 males and 28 females, from the study area.

Limnephilus sansoni Banks, 1918

(Fig. 270a-272, 608)

Limnephilus sansoni Banks, 1918:19. (Type locality: Banff, Alberta). Dodds and Hisaw,

1925b:386. Betten, 1934:337. Milne, 1935:44, 51. Ross, 1938b:39. Ross, 1944:298.

Ross and Merkley, 1952:448. Fischer, 1968:295.

Limnophilus sansoni ; Schmid, 1955:135.

Males of this species are distinguished from males of other species in the group by long,

thin, tapered, dorsal process of clasper; by laterally toothed tip of median lobes of segment

X; and by row of heavy, black teeth on mesal face of cercus (Fig. 270b). Females are dis-

tinguished by large, isolated cerci of segment X, and by segment X cleft squarely on latero-

mesal faces (Fig. 271). Venation identical to that of L. sublunatus.

Description. — Antennae dark brown; scapes darker, with antero-mesal faces pale, gla-

brous. Vertex of head almost black in ocellar triangle; remainder brown, with warts yellow.

Thorax generally dark brown. Body of female generally rich red-brown, with dark brown

only dorso-laterally. Brushes of male anterior femora of strong, black spines on basal half

only. Spurs yellow. Fore wing length of male 14.0 mm; light brown; costal area clear to

distal end of sub-costa. Stigma present. Clear areas other than costal margin slightly cloudy

brown to clear yellow.

Male genitalia. (Specimen from Totem Creek area, Banff-Jasper Hwy., north of Lake

Louise, Alberta). Postero-dorsal edge of tergum VIII prolonged posterad as large, black,

spinate bulb. Segment IX with very narrow dorsal strap directed postero-dorsad at about

45°. Main body of segment square; posterior edge indented at clasper bases as wide, thumb-

like process dorsally (Fig. 270a). Clasper with long, narrow base and long, thin, finger-like

dorsal process. Median lobes of segment X with small disto-lateral tooth; dorso-lateral edges

black, denticulate; attached to mesal edges of small, basal plates. Cercus triangular, with

posterior edges black, dentate; with strongly sclerotized, dorso-ventral, black, dentate ridge

(Fig. 270b). Aedeagus essentially identical to that of L. sublunatus, except for slightly

broader dorsal lobe.

Female genitalia. (Specimen from Totem Creek area, on Banff-Jasper Hwy., north of

Lake Louise, Alberta). Vulval scale median lobe blunt, little tapered, longer than lateral

lobes (Fig. 272). Ventro-lateral lobes of segment IX square, with acuminate, concave pro-

cess directed posterad (Fig. 271). Supra-genital plate attached ventrad of segment IX, wide,

with slightly flanged lateral edges, flanges folded mesad. Segment X rectangular in lateral

aspect, with square lateral clefts; in ventral aspect wide, triangular basally; cleft ventrally

only slightly; dorsal cleft complete to cereal bases. Cercus located on posterior half of

segment X, projected posterad beyond segment for half its length.

Notes on biology. — Adults of this species are collected almost exclusively in the vicinity

of sedge ponds or sloughs. The adult flight season extends from July 25 to October 3 with

an apparent peak of emergence from mid-August to mid-September.

Geographical distribution. — The known range of this species extends from Alaska to

Colorado, though records are few (Fig. 608). In Alberta it is confined to the mountain and

foothill areas, ranging in altitude from 3,700’ to 5,350’.

Rhyacophilidae and Limnephilidae

91

I have examined 276 specimens, 103 males and 173 females, from the study area.

Limnephilus extractus Walker, 1852

(Fig. 273-277, 609)

Limnephilus extractus Walker, 1852:34. (Type locality: St. Martin’s Falls, Albany River,

Hudson’s Bay). McLachlan, 1863:157, 161. Hagen, 1864:835. Banks, 1892:363. Banks,

1907a:36. Banks, 1908a: plate 18, Fig. 6-7. Betten, 1934:324. Milne, 1935:44, 51.

Milne (D. J.), 1943:192. Ross, 1944:298. Ross and Merkley, 1952:449. Morse and

Blickle, 1953:98. Fischer, 1968:101-102.

Goniotaulius extractus ; Hagen, 1864:815.

Limnophilus (Goniotaulius) extractus’, Hagen, 1861:260.

Limnophilus extractus’, McLachlan, 1876b:7. Ulmer, 1905a: 19. Ulmer, 1907a:44. Betten

and Mosely, 1940:135-136. Schmid, 1955:135.

Males of this species are distinguished from males of other species of the group by long,

slender, black, median lobes of segment X; by inconspicuous posterior lobe of tergum VIII;

and by short, trapezoidal claspers (Fig. 273). Females are distinguished by almost complete-

ly cleft segment X (Fig. 277); by lack of separate and distinct ventro-lateral lobe of segment

I IX (Fig. 276); and by laterally cleft segment X.

Description. — Antennae yellow to pale brown. Vertex of head yellow, to very pale

brown in patches. Thorax pale yellow with slight local darkenings to pale brown. Male with

brush of fore leg spinate, black, occupying only basal half of mesal face, in narrow band.

Spurs yellow. Fore wing length of male 12.4 mm; pale brown to yellow; costal margin

clear to wing tip and beyond. Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from Cold Lake, Alberta). Postero-dorsal margin of tergum

VIII lightly setose; bounded laterally by depressions; setae pale. Segment IX with short

dorsal strap; strap with mesal process directed ventrad (Fig. 273). Main body of segment

abruptly expanded ventrad of dorsal strap, rectangular and featureless. Clasper small, blunt,

I attached to lower half of posterior edge of segment IX; trapezoidal in lateral aspect. Median

lobes of segment X finger-like in lateral aspect; projected well posterad of cerci, black, each

with small, distinct disto-lateral tooth (Fig. 274). Cercus triangular, concave on mesal face,

not projected posterad of segment IX. Aedeagus simple; lateral arms originated dorsad of

; median shaft (Fig. 275); tips fringed with heavy setae along dorsal edges only; setae increase

! in length distally. Median shaft attached to distinct basal, sclerotized collar.

Female genitalia. (Specimen from Flatbush, Alberta). Vulval scale large, lobes well spaced;

median lobe longer than lateral lobes, square-tipped, slightly tapered from base (Fig. 277);

lateral lobes curved mesad distally. Segment IX large, higher than wide (Fig. 276); with

i' slight latero-posterior lobes. Supra-genital plate semi-circular. Segment X fused to segment

( IX; of two pairs of lobes; cerci broad, triangular in lateral aspect; main body of segment

cleft completely in vertical plane (Fig. 277); cleft slightly in horizontal plane.

Notes on biology. — This is a lake species. Adults are collected near sedge and other water

weed-choked ponds and lakes. The adult flight period extends from May 22 to July 11.

Geographical distribution. - The known range of this species is primarily boreal, ex-

; tending from Great Slave Lake and central Alberta to the northern New England States

1 (Fig. 609). In Alberta it is confined to the central and northeast lowlands.

I have examined 27 specimens, 25 males and two females, from the study area.

Limnephilus hageni Banks, 1930

(Fig. 278-283,610)

92

Nimmo

Limnephilus hageni Banks, 1930a:226. (Type locality: Fort Resolution, Great Slave Lake).

Betten, 1934:336. Milne, 1935:44, 51. Ross, 1938b:36. Ross, 1944:298. Ross and

Merkley, 1952:448. Fischer, 1968:173.

Limnophilus hageni ; Schmid, 1955:135.

Males of this species are distinguished from males of other species in the group by high,

narrow clasper bases, with thick, blunt dorsal process (Fig. 278); by row of black teeth ex-

tended only half way across cerci; and by dorso-laterally directed tips of median lobes of

segment X. Females are distinguished by long, thin, cerci extended more than half their

length beyond segment X; by absence of discrete ventro-lateral lobes of segment IX; and by

very long median lobe of vulval scale (Fig. 282, 283).

Description. — Antennae light yellowish brown. Vertex of head with diamond-shaped

black area with two comers occupied by lateral ocelli. Thorax light yellowish brown, with

darker areas on thorax. Brush of male fore leg of single row of black spines extended along

basal half of femur. Spurs yellow-brown. Fore wing length of male 1 1.8 mm; light brown,

clear areas hyaline. Costal area quite clear; stigma slight. Venation identical to that of

L. sublunatus.

Male genitalia. (Specimen from Forestry Trunk Road in area of Kananaskis Lakes, Alber-

ta). Postero-dorsal edge of tergum VIII produced as short, black-spinate bulb. Segment IX

with long, tapered dorsal strap; main body of segment stout, rectangular, with postero-

dorsal angle directed mesad, furrowed (Fig. 278). Clasper short, blunt distally; base high,

narrow. Median lobes of segment X short, stout in lateral aspect, black distally, with tips

curved dorso-laterad (Fig. 278, 279). Cercus large, with dorsal edge curved gently distad;

posterior edge black, irregularly dentate; mesal face with short, curved dentate line from

dorsal edge. Median shaft of aedeagus with distinct distal head; opening of ejaculatory duct

at extreme tip. Lateral arms large, each bifid, ventral lobe membranous, extensible, distally

fringed with long setae; dorsal lobe sclerotized, toothed distally, with tooth directed dorso-

mesad (Fig. 281).

Female genitalia. (Specimen from Forestry Trunk Road in area of Kananaskis Lakes,

Alberta). Vulval scale with long, narrow, median lobe (Fig. 283); lateral lobes small, widely

spaced, lateral edges slightly flanged. Segment IX large, rectangular, with small ventro-lateral

lobes (Fig. 282). Supra-genital plate large, semi-circular. Segment X indistinct from segment

IX; deeply cleft dorsally. Cerci long, thin, well separated from segment X.

Notes on biology. — Adults of this species are found in smaller ponds or sloughs with

dense sedge and horse-tails present. The adult flight period extends from July 15 to Septem-

ber 22; my records are insufficient to indicate a peak period.

Geographical distribution. — The known range of this species extends from British Colum-

bia to Ontario and Great Slave Lake (Fig. 610). In Alberta it is confined to the mountains

and foothills, ranging in altitude from 4,600’ to 5,350’.

I have examined 26 specimens, 16 males and 10 females, from the study area, and one

(male) from Great Slave Lake.

Limnephilus partitus Walker, 1852

(Fig. 139a, 139b, 284-290, 611)

Limnephilus partitus Walker, 1852:32. (Type locality: St. Martin’s Falls, Albany River,

Hudson’s Bay). McLachlan, 1863:157, 161. Banks, 1907a:37. Betten, 1934:330. Milne,

1935:20, 47. Milne, 1936:60. Ross, 1944:298. Ross and Merkley, 1952:448. Etnier,

1965:149. Fischer, 1968:252-253.

Limnophilus partitus ; Betten and Mosely, 1940:127-129.

Rhyacophilidae and Limnephilidae

93

Males of this species are distinguishable by irregular posterior edges of segment IX (Fig.

284); by high, narrow-based claspers, with dorsal process long, thin, directed dorso-posterad;

by very wide dorso-posterior process of tergum VIII (Fig. 285); and by small, sclerotized

ventral lobe of lateral arm (Fig. 286). Females are distinguished by divided ventro-lateral

lobes of segment IX (Fig. 290).

Description. — Antennae black. Vertex of head black, warts dark brown. Thorax black in

male, brown in female. Spurs hght brown. Fore wing length of male 12.3 mm; brown, with

large clear areas. Costal margin clear for about four-fifths of length. Stigma present, with

distinct, white area immediately basad. Venation as in Fig. 139a, 139b; both fore and hind

wings with cell between f2 and f3 pedicellate.

Male genitalia. (Specimen from pond 2 miles east of Nordegg, Alberta). Tergum VIII

black with pale ventro-lateral edges; postero-dorsal edge produced posterad as black-spinate

bulge dorsad of genitalia. Dorsal strap of segment IX hyaline, discrete from remainder of

segment (Fig. 284); narrow in posterior aspect (Fig. 285). Main body of segment squat, with

distinct bulge on dorsal edge. Clasper with base coincident with entire posterior edge of

segment IX; dorsal process finger-like. Median lobes of segment X attached to mesal edges

of lateral, concave lobes; massive, directed dorso-posterad; quite black. Cercus with distal

edge black, dentate; with thin, weak line of teeth on mesal face. Lateral arms of aedeagus

bilobed (Fig. 286); dorsal lobe heavily sclerotized, distally spiniform, directed dorsally;

ventral lobe hyaline, spatulate, fringed distally with setae.

Female genitalia. (Specimen from pond 2 miles east of Nordegg, Alberta). Vulval scale

wide (Fig. 289); lateral arms with concave distal faces; median lobe wide basally, gradually

tapered distad, rounded. Segment IX with small dorsal body, divided to ventro-lateral lobes

(Fig. 290). Supra-genital plate large, projected well posterad of vulval scale. Segment X com-

pletely cleft dorsally; anal opening wide, flared. Cerci short, acute-triangular in ventral

aspect, held close to segment X.

Notes on biology. — Adults of this species are collected near ponds or sloughs thickly

vegetated with sedges, or sedges and horse-tails. The adult flight period extends from August

7 to October 3. Peak emergence appears to be from mid-August to mid-September.

Geographical distribution. — The known range of this species extends from British Colum-

j bia and Great Slave Lake, to Newfoundland (Fig. 611). In Alberta it is confined to the

mountains and foothills, ranging in altitude from 3,800’ to 5,200’.

I have examined 77 specimens, 49 males and 28 females, from the study area and Great

| Slave Lake.

Limnephilus susana Nimmo, n. sp.

(Fig. 291-295, 612)

Males of this species are distinguished from males of other species in the group by evenly

tapered, dorsally directed median lobes of segment X (Fig. 291); and by tapered, ventrally

curved lobe of tergum VIII. Females are distinguished by short, stout cerci closely appressed

to segment X and of equal length (Fig. 295); and by vulval scale median lobe twice as long

as lateral lobes (Fig. 294).

Description. — Antennae red-brown. Vertex of head reddish to yellowish brown except

for dark, chocolate-brown between ocelli, in cruciform pattern in male. Thorax very dark

brown laterally and dorsally. Brush of male fore leg of scattered short, black spines on basal

half of femur. Spurs light red-brown. Fore wing length of male 14.6 mm; chocolate-brown,

with lighter, reddish brown, areas. Costal area almost hyaline. Venation identical to that of

L. sublunatus.

94

Nimmo

Male genitalia. (Specimen from Forestry Trunk Road, at Pembina River, Alberta). Pos-

tero-dorsal lobe of tergum VIII broad, somewhat flattened, tapered, curved ventrad. Seg-

ment IX high, narrow, with main body nearly rectangular in lateral aspect (Fig. 291); with

deep furrow at postero-dorsal angle. Dorsal strap deep, vertical band. Clasper large, with

long, sinuate base; dorsal process of uneven width, directed dorsad distally, twisted slightly

to form ridge at right angles to body axes. Median lobes of segment X long, narrow, evenly

tapered, dorsally upturned blades; with disto-lateral tooth (Fig. 292) and large concave

plates basally. Cercus massive, curved, of approximately uniform width; distally with row of

dark teeth; second series of three to four dark teeth two-thirds of distance from base (Fig.

292) . Median shaft of aedeagus stout, dorsally curved; lateral arms massive, bilobed (Fig.

293) . Dorsal lobe heavy, distally toothed, sclerotized; ventral lobe membranous but with

lightly sclerotized distal edges with long, stout setae dorsally.

Female genitalia. (Specimen from Forestry Trunk Road, at Pembina River, Alberta).

Vulval scale large, with long, rectangular, median lobe, rectangular, almost square, lateral

lobes (Fig. 294); all lobes well separated; median lobe twice as long as lateral lobes. Supra-

genital plate large, pentagonal in ventral aspect. Segment IX with two thin, ventral lobes,

and triangular dorsal portion, all connected by narrow lateral straps (Fig. 295). Segment X

robust, terminated distally by two pairs of lobes; dorsal lobes thin, triangular; ventral lobes

vertically thin, wide in lateral aspect. Cerci short, closely attached to dorsal surface of seg-

ment X; rounded, spindle-shaped.

Notes on biology. — Known only from a single slough with thick growth of horse-tails,

and a fine-bladed sedge; water shallow. Date of capture was September 22.

Holotype. — Male. Forestry Trunk Road about 1 mile north of crossing of Pembina River,

Alberta (Fig. 612); September 22, 1968. A. Nimmo.

Allotype. — Female. Same data as holotype.

Paratypes. — Same data as for holotype; one male, two females.

The holotype, allotype, and one female paratype are in the Canadian National Collection,

Ottawa, where they have been assigned the type series number 10,586. The remaining male

and female paratypes are in the Strickland Museum. Dept, of Entomology, University of

Alberta, Edmonton, Alberta.

This new species is named for my wife, Susan.

Limnephilus species 1

(Fig. 296-297,612)

Females of this species are distinguished from females of other species in the group by

large dorsal body to segment IX, by long, distally up-turned segment X, and by long,

slender, tapered cerci not appressed dorsally to segment X (Fig. 296).

Description. — Antennae yellow-brown. Vertex of head deep red-brown. Thorax yellow-

brown. Spurs dark brown; formula 1,3,3. Fore wing length of female 1 1.8 mm; red-brown,

with hyaline areas typical of the pattern for the subcentralis group, except no evident longi-

tudinal bars at wing tip. Venation identical to that of L. sublunatus.

Male genitalia. Unknown.

Female genitalia. (Specimen from 2 miles north of Athabasca Falls, on Hwy. 93a, Jasper

National Park, Alberta). Median lobe of vulval scale much longer than lateral lobes, with

square tip, gradually tapered. Lateral lobes sub-triangular (Fig. 297). Segment IX large, with

two ventro-lateral, rounded, not separate, lobes (Fig. 296). Dorsal portion convex, bounded

by lateral declivities which result in concave sides of segment. Segments IX and X partly

separated by faint suture. Supra-genital plate relatively small, convex ventrally; polygonal in

Rhyacophilidae and Limnephilidae

95

ventral aspect. Segment X cylindrical, deeply cleft mesally; not so deeply cleft laterally;

dorsal lobes thus formed small, triangular in lateral aspect; ventral lobes larger, wider. Cerci

long, tapered, thin, free from dorsal surface of segment X.

The single specimen of this species known to me was taken from a small muskeg lake 2

miles north of Athabasca Falls, on Hwy. 93a, Jasper National Park, Alberta, September 10,

1966 (Fig. 612).

The stigma group

Males of this group are characterised by relatively small claspers, with high, ventrally

tapered bases, and short, blunt dorsal processes; by small postero-dorsal lobes of tergum

VIII projected well posterad from membranous connection to segment IX (Fig. 304). Fe-

males are characterised by presence of separate and discrete ventro-lateral lobes of segment

IX (Fig. 302).

Limnephilus indivisus Walker, 1852

(Fig. 298-303, 614)

Limnephilus indivisus Walker, 1852:34. (Type locality: Nova Scotia). McLachlan, 1863:

157, 161. Hagen, 1864:836. Banks, 1892:363. Banks, 1899:208. Simpson, 1903:98-100.

Banks, 1907a:36. Lloyd, 1915:205-208. Krafka, 1923: plate 9, Fig. 58. Betten, 1926:

529. Ricker, 1932:132. Ricker, 1934:54. Betten, 1934:325-326. Milne, 1935:46, 51.

Mickel and Milliron, 1939:575-579. Ross, 1941:109. Ross, 1944:5, 185, 186, 189, 191,

298. Proctor, 1946:211. Leonard and Leonard, 1949a: 18. Schmid and Guppy, 1952:42.

Ross and Spencer, 1952:48. Ross and Merkley, 1952:448. Morse and Blickle, 1953:98.

Flint, 1960:5, 47-49, 51, 53, 57. Robert, 1960:59. Etnier, 1965:149. Fischer, 1968:

196-198. McConnochie and Likens, 1969:150.

Limnophilus indivisus’, Hagen, 1861:260-261. Ulmer, 1905a: 19. Ulmer, 1907a:44. Siltala,

1907a: 17. Siltala, 1907b:329, 486. Lloyd, 1921 :41, 47-5 1 . Branch, 1922:256, 257, 258,

259, 263, 265, 266, 267, 270. Sibley, 1926:107, 191, 193, 194. Muttkowski and Smith,

1929:259. Handlirsch, 1936: Fig. 1601, 1608. Needham and Lloyd, 1937:345. Balduf,

1939:144, 151, 161, 179. Betten and Mosely, 1940:129-132. Schmid, 1955:136.

| Limnephilus subguttatus Walker, 1852:34-35. Ross, 1944:191, 298. Fischer, 1968:197-198.

ii Limnophilus subguttatus ; Hagen, 1861:261. Hagen, 1873a:295. McLachlan, 1875:59. Pro-

vancher, 1877:243. Provancher, 1 878b: 131. Provancher, 1878a: 146. Betten and Mosely,

1940:129, 132. Ross, 1944:191, 298.

Males of this species are distinguished from males of other species of the group and genus

: by distally flared or expanded cerci (Fig. 298); and by short, narrow, tapered, median lobes

of segment X. Females are distinguished by right-angled, ventro-lateral lobes of segment IX

(Fig. 302); and by long, thin, acute-triangular cerci of segment X closely appressed to the

dorso-lateral surface of segment X.

Description. — Antennae pale yellow; scapes darker, with longitudinal band of fine, short

setae on antero-mesal faces. Vertex of head reddish brown, warts paler. Thorax yellow,

tending to light reddish brown dorsally. Spurs dark reddish yellow. Fore wing length of

I male 14.9 mm; yellow to pale brown. Costal area clear to distal end of costa. Venation

identical to that of L. sublunatus ; with short area of short, black hairs on vein R2 of

i male hind wing (as in Fig. 140b); absent in female. Abdominal sterna VI and VII of male

! with short, triangular lobes in middle of posterior edges; similarly in female, but on sterna

V and VI.

96

Nimmo

Male genitalia. (Specimen from Gorge Creek, 20 miles west of Turner Valley, Alberta).

Postero-dorsal process of tergum VIII with short, black spines. Dorsal strap of segment IX

not arched highly, directed postero-dorsad, with distinct, spatulate ventro-mesal process

(Fig. 298, 301) connected to segment X. Body of segment IX wide dorsally, narrowed

ventrad; with distinct black band of varied width along anterior edges. Clasper short, blunt,

with high, narrow base. Median lobes of segment X conical, short, thin, tapered distad,

black; attached to mesal edges of basal plates; directed slightly dorso-laterad. Cercus broad-

ened distad; short, with distal rim black, dentate; with row of heavy, black teeth vertically

on mesal face (Fig. 301); with tufts of black setae at distal corners. Lateral arms of aedeagus

bilobed, with bifid bases (Fig. 299); ventral lobe rounded, spatulate, with fringe of heavy,

dark setae on dorsal edge; dorsal lobe directed dorso-mesad (Fig. 300), with black, spiniform

tip and hyaline lateral flange; posterior edge with four very heavy, black spines.

Female genitalia. (Specimen from Gorge Creek, 20 miles west of Turner Valley, Alberta).

Vulval scale with two triangular lateral lobes, uniformly tapered median lobe. Ventro-lateral

lobes of segment IX distinct, discrete (Fig. 302), bent at right angles around base of segment

X. Dorsal portion of segment IX small, almost indistinguishable from segment X. Supra-

genital plate small, smoothly curved (Fig. 303). Segment X deeply cleft dorsally; conical in

ventral aspect, situated between cerci. Cerci closely appressed to dorso-lateral surfaces of

segment X, tapered to finger-like tips.

Notes on biology. — I have frequently taken adults of this species in the vicinity of small

lakes or sloughs which are thickly bordered by cattail rushes, or coarse sedges. The adult

flight period extends from July 14 to September 30; records are insufficient to determine a

peak.

Geographical distribution. - The known range of this species extends from British Colum-

bia to Illinois, Nova Scotia and Quebec (Fig. 614). In Alberta it is found in the plains, foot-

hills, and low mountain passes.

I have examined 25 specimens, 15 males and 10 females, from the study area.

Limnephilus infemalis (Banks), 1914

(Fig. 304-308,613)

Anisogamus infemalis Banks, 1914:154-156. (Type locality: Pinnacle Mountain, Fulton

County, New York).

Stenophylax infemalis ; Betten, 1926:529. Betten, 1934:343-344.

Limnephilus infemalis ; Milne, 1935:46, 51. Milne (D. J.), 1943:192, 194, 195. Ross, 1944:

298. Leonard and Leonard, 1949a: 18. Ross and Merkley, 1952:448. Etnier, 1965:149.

Nimmo, 1965:783-786. Fischer, 1968:198. Clifford, 1969:582.

Limnophilus infemalis ; Schmid, 1955:136.

Males of this species are distinguished by antero-dorsally directed median lobes of seg-

ment X (Fig. 304); by large, mesally concave cerci heavily clothed on mesal faces with long

setae; and by meso-ventral process of dorsal strap of segment IX connected to segment X

(Fig. 305). Females are distinguished by large ventro-lateral lobes of segment IX (Fig. 307);

and by large, triangular cerci fused to posterior edges of segment IX.

Description. — Antennae uniformly pale yellow. Vertex of head yellow except for darker

area mesad of lateral ocelli. Thorax pale yellow, to brownish yellow. Brush of male fore leg

extended barely half length of femur; composed of very stout, black spines.

Male genitalia. (Specimen from Sturgeon River, St. Albert, Alberta). Postero-dorsal lobe

of tergum VIII small, bulbous distally; with short, stout, sparsely scattered, black spines.

Segment IX with very high, thin dorsal strap; strap with large, ventro-mesally directed horn

Rhyacophilidae and Limnephilidae

97

connected to segment X; main body of segment small, irregular (Fig. 304, 305). Clasper

thumb-like, widened slightly at high, very narrow base. Median lobes of segment X attached

to mesal edges of slightly concave, basal plates; curved antero-dorsad, with slight distal

swelling. Cercus very large, concave mesally; mesal face thickly clothed with long setae.

Aedeagus simple; ejaculatory duct terminated at extreme tip, lateral arms attached to dorsal

membranous base of aedeagus, with slightly dilated, setose tips (Fig. 306).

Female genitalia. (Specimen from Sturgeon River, St. Albert, Alberta). Vulval scale with

short, tapered, blunt median lobe (Fig. 308); lateral lobes deeply channelled on mesal faces.

Ventro-lateral lobes of segment IX triangular in lateral aspect (Fig. 307); main body of seg-

ment relatively large, not distinct from segment X. Segment X inconspicuous, ventrad of

massive, triangular cerci; of simple, triangular dorso-lateral lobes fused to ventral surfaces of

cerci (Fig. 308), and ventral, concave, triangular plate ventrad of anal orifice. Supra-genital

plate visor-like, arched dorsad.

Notes on biology. — Adults of this species emerge from lakes, sloughs and quiet streams

bordered with thick growths of sedges and cattail reeds. The adult flight period extends

from August 14 to September 22.

Geographical distribution. — The known range of this species extends from Alaska to

, Alberta, Michigan and Maine (Fig. 613). In Alberta it is confined to the plains, low foothills,

and low mountain passes.

I have examined 256 specimens, 206 males and 50 females, from the study area and

Great Slave Lake.

The ornatus group

Males of the single species of this group are characterised by massive, distally bifid cerci

(Fig. 309); by claspers with high, narrow bases and short, stubby dorsal processes; and by

| simple aedeagus (Fig. 311). Females are characterised by massive segment IX without ventro-

lateral lobes (Fig. 313); by segment X which is discrete from segment IX; and by long,

tapered cerci well separated from segment X.

Limnephilus ornatus Banks, 1897

(Fig. 309-314, 615)

| Limnephilus ornatus Banks, 1897:27. (Type locality: Sherbrook, Canada). Banks, 1899:

208. Banks, 1900b:467. Banks, 1907a:37. Banks, 1908b:63. Betten, 1926:529. Banks,

1930b: 128. Neave, 1934:167. Betten, 1934:329-330. Milne, 1935:46, 51. Ross, 1944:

186, 189-190, 298. Proctor, 1946:21 1 . Leonard and Leonard, 1949a: 18. Ross and Merk-

ly, 1952:441. Morse and Blickle, 1953:98. Robert, 1960:59. Wiggins, 1961:700. Etnier,

1965:149. Nimmo, 1966a:692. Nimmo, 1966b:224. Fischer, 1968:249-250. Lindroth

and Ball, 1969:138. McConnochie and Likens, 1969:150.

Limnophilus ornatus-, Betten, 1901:573. Ulmer, 1905a: 19. Ulmer, 1907a:44. Nakahara,

1915:95. Sibley, 1926:107, 191.Essig, 1926: 176. Forsslund, 1932:57-59. Mosely, 1932:

573. Ulmer, 1932:213. Carpenter, 1938:530, 534. Henricksen, 1939:23. Fristrup, 1942:

20. Schmid, 1955:136. (See Fischer, 1968:249-250, for Palaearctic literature).

w Limnophilus elegans Mosely (not Curtis), 1929:502, 504, 507. Fischer, 1968:250.

Description. — Antennae yellow. Vertex of head uniformly yellow. Thorax pale yellow to

1 light reddish brown. Brush of male fore leg narrow row of black setae along mesal face of

< basal half of femur. Spurs yellow. Fore wing length of male 14.3 mm; pale yellow to light

reddish brown, with large hyaline areas. Costal area clear except for opaque area in distal

98

Nimmo

third of stigma. Venation identical to that oi L. sublunatus.

Male genitalia. (Specimen from Indian Head, Saskatchewan). Tergum VIII without poste-

ro-dorsal process. Segment IX with short, thin dorsal strap. Main body with large, blunt lobe

at postero-dorsal angle; with distinct row of heavy setae along ventral half of posterior edge

(Fig. 309). Clasper with high, narrow base and short, blunt, thumb-like dorsal process.

Median lobes of segment X attached to complexly folded baso-lateral plates; ventral edges

and lateral faces minutely dentate; lobes black distally. Cercus large, bifurcated distally;

distal lobes black, heavily sclerotized at tips (Fig. 310). Lateral arms of aedeagus slightly

expanded distally (Fig. 311); distal half warped, concave mesally (Fig. 312) with fringe of

setae along posterior edge.

Female genitalia. (Specimen from Indian Head, Saskatchewan). Vulval scale with short,

rectangular median lobe; lateral lobes narrow, distinctly separated distally from median lobe

(Fig. 314). Segment IX without discrete latero-ventral lobes (Fig. 313); curved slightly to

embrace segment X. Supra-genital plate small, rounded, almost elliptical, set into ventral

depression of segment IX. Segment X of inter-leaved dorsal and ventral parts distally; dorsal

lobes darker, fused basally to ventral trough. Cerci long, thin, tapered, well separated from

dorsal surface of segment X.

Notes on biology. — I have not taken individuals of this species in the study area, but the

records available indicate that they inhabit sloughs and small lakes. The adult flight period

extends from July 13 to September 10.

Geographical distribution. — The known range of this species extends from Alaska to

Illinois and Greenland (Fig. 615) in North America. It is actually a Holarctic species with

records extending west to Europe and Iceland. In Alberta it is confined to the plains or long,

low mountain valleys leading to the plains.

I have examined 1 1 specimens, three males and eight females, from the study area.

The picturatus group

Males of the single species are characterised by large, mesally concave claspers with disto-

mesal spine (Fig. 315, 316); by absence of postero-dorsal process of tergum VIII; and by

short, relatively wide dorsal strap dorsad of large, trapezoidal segment IX. Females are dis-

tinguished by massive ventro-lateral lobes of segment IX (Fig. 318); by complete lack of

separation of segments IX and X dorsally; and by minute, triangular cerci closely appressed

to dorso-lateral surfaces of segment X.

Limnephilus picturatus McLachlan, 1875

(Fig. 315-319,616)

Limnophilus picturatus McLachlan, 1875:78-79. (Type locality: Europe). Brauer, 1876:286.

Ulmer, 1905a: 19. Ulmer, 1907a:42. Ulmer, 1932:213. Schmid, 1955:136. (See Fischer,

1968:256-259, for Palaearctic literature).

Limnephilus picturatus ; Ross and Merkley, 1952:449, 454-455. Fischer, 1968:256-259.

(See Fischer, 1968:256, 258, for Palaearctic literature).

Limnophilus exulans McLachlan, 1876, (See Fischer, 1968:258).

Limnophilus miyadii Tsuda, 1924, (See Fischer, 1968:258).

Phryganea notatus Zetterstedt, 1840, (See Fischer, 1968:258-259).

Description. — Antennae dark brown; scapes and pedicels black; yellowish brown through-

out in females. Head entirely black. Thorax black; brush of male fore leg on basal two-thirds

of femur; opposing face of tibia minutely spinate. Spurs yellow. Fore wing length of male

Rhyacophilidae and Limnephilidae

99

9.1 mm; pale brownish yellow; costal area clear; not patterned. Venation identical to that of

L. sublunatus.

Male genitalia. (Specimen from Bow Pass, Banff National Park, Alberta). Segment IX with

short dorsal strap; main body robust (Fig. 315). Clasper with long, narrow base; dorsal pro-

cess short, with irregular edges, directed dorso-posterad. Median lobes of segment X narrow

basally, widened at mid-point, narrowed distally to dorso-lateral hooks (Fig. 315, 316);

black throughout. Median lobes attached to simple latero-basal plates. Lateral arms of aedea-

gus simple, gradually widened distad to narrow, acuminate tips with several tufts of setae or

spines (Fig. 317).

Female genitalia. (Specimen from Sunwapta Pass, Jasper National Park, Alberta). Vulval

scale triangular in general outline (Fig. 319); median lobe blunt, almost rectangular; lateral

lobes semi-triangular, with straight posterior and mesal edges. Segment IX with massive,

rectangular, ventro-lateral lobes (Fig. 318); dorsal body of segment small, indistinguishable

from segment X. Supra-genital plate small, crescentic. Segment X small, with two minute

dorsal lobes and single ventral lobe. Cerci small, triangular, immovable, closely appressed to

dorso-lateral surfaces of segment X.

Notes on biology. — Adults of this species are found near shallow, sedge-fringed sloughs

or small pools. The adult flight period extends from August 5 to September 22. There

appears to be a peak at about the end of August and beginning of September.

Geographical distribution. — The nearctic range of this holarctic species extends from

Alaska to Hudson’s Bay, and to Colorado (Fig. 616). In Alberta it is confined to the moun-

tains and foothills, ranging in altitude from about 5,000’ to 6,880’.

I have examined 71 specimens, 42 males and 29 females, from the study area.

The externus group

Males of the single species of this group present in the study area are distinguishable by

small, triangular cerci (Fig. 320); by black-spinate postero-dorsal bulb of tergum VIII; and

by small, vertical, posteriorly hooked median lobes of segment X. Females are distinguished

by massive ventro-lateral lobes of segment IX (Fig. 324); and by large, roughly triangular

cerci of segment X very close to segment X.

Limnephilus externus Hagen, 1861

(Fig. 320-325, 617)

Limnophilus externus Hagen, 1861:257. (Type locality: North Red River, Canada). Ulmer,

1905a: 19. Ulmer, 1907a:44. Ulmer, 1932:212. Schmid, 1955:137. (See Fischer, 1968:

99, for Palaearctic literature).

Limnephilus externus’, Hagen, 1864:835. Banks, 1892:636. Banks, 1907a:36. Betten, 1934:

152. Neave, 1934:167. Milne, 1935:44, 51. Ross, 1938b:36. Knowlton and Harmston,

1939:286. Banks, 1943:342. Ross, 1944:298. Leonard and Leonard, 1949a: 17. Schmid

and Guppy, 1952:42. Ross and Merkley, 1952:447. Ross and Spencer, 1952:48. Flint,

1960:5. Wiggins, 1961:701. Denning, 1963:261. Etnier, 1965:149. Fischer, 1968:99-101.

(See Fischer, 1968:99-100, for Palaearctic literature).

Limnophilus congener McLachlan, 1875:56-57. (See Fischer, 1968:100, for Palaearctic

literature).

Limnephilus congener’, (Palaearctic; see Fischer, 1968:100).

Limnephilus luteolus Banks, 1899:207, 208. (Type locality: Washington State). Banks,

1907a:36. Milne, 1935:44, 51. Fischer, 1968:101.

100

Nimmo

Limnophilus luteolus Ulmer, 1905a: 19. Ulmer, 1907a:44. Essig, 1926:176.

Limnephilus oslari Banks, 1907b: 121-122. (Type locality: Colorado). Banks, 1907a:37.

Dodds and Hisaw, 1925b:286. Betten, 1934:337. Milne, 1935:51. Ross, 1938b:38.

Fischer, 1968:101.

Limnophilus oslari', Essig, 1926:176.

Limnephilus tersus Betten, 1934:334. (Type locality: Old Forge, New York). Milne, 1935:

48, 52. Milne, 1936:62. Ross, 1944:298. Ross and Merkley, 1952:447. Fischer, 1968:

101.

Description. — Antennae uniform yellowish brown. Vertex of head black immediately

mesad of lateral ocelli, otherwise brown. Thorax light reddish brown. Brush of male fore leg

single row of black spines on basal half of mesal face of femur. Spurs hyaline to yellow.

Fore wing length of male 21.6 mm; greyish brown, with hyaline areas. Costal area clear to

distal end of sub-costa. Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from Wapta Lake, Kicking Horse Pass, Yoho National Park,

British Columbia). Postero-dorsal process of tergum VIII distinctly bulbous, with mesal

band of dark spines. Dorsal strap of segment IX arched posterad, with ventro-mesal process

(Fig. 320). Main body of segment rectangular, slightly curved, with distinct groove at

postero-dorsal angle. Clasper short, slender, with virtually non-existent base; dorsal process

with single, dorsally directed distal tooth. Median lobes of segment X black distally, each

with two small distal teeth, dorsally and ventrally; lobes vertical, hooked posterad; bases

bifid. Cercus triangular, with dorsal angles produced posterad; distal edges with row of

prominent, black teeth (Fig. 321). Lateral arms of aedeagus distally fringed with dorsal row

of setae directed anterad (Fig. 322, 323).

Female genitalia. (Specimen from Wapta Lake, Kicking Horse Pass, Yoho National Park,

British Columbia). Vulval scale with evenly tapered median lobe; lateral lobes widely spaced

(Fig. 325). Segment IX with massive, triangular ventro-lateral lobes in contact ventrad of

segment X (Fig. 324, 325); dorsal portion of segment small, triangular in dorsal aspect.

Supra-genital plate straight-edged in ventral aspect, with triangular distal edge. Segment X

small, discrete from segment IX, with deep dorsal cleft, sinuate ventral edge. Cerci large,

triangular, close to segment X.

Notes on biology. — Adults of this species are associated with smaller, sedge-fringed

ponds and sloughs. The adult flight period extends from July 24 to October 4; I have

one record from June 15, also. The peak appears to occur from mid-August to mid-Sep-

tember.

Geographical distribution. - The nearctic range of this holarctic species extends from

Great Slave Lake to California and Newfoundland in North America (Fig. 617). In Alberta

it occurs in the mountains and plains. It ranges in altitude from 2,000’ to 5,340’.

I have examined 276 specimens, 108 males and 168 females, from the study area.

The sericeus group

Males of species of this group are characterised by high, narrow claspers with little or no

dorsal process; by long, acute-triangular median lobes of segment X; and by meso-ventrally

directed, relatively long, tubular, postero-dorsal process of tergum VIII (Fig. 326). Females

of this group are characterised by segment IX without discrete ventro-lateral lobes; by very

acuminate dorsal blade of segment X; and by rectangular, free cerci (Fig. 330, 331).

Limnephilus sericeus (Say), 1824

(Fig. 326-331,618)

Rhyacophilidae and Limnephilidae

101

Phryganea sericea Say, 1824:309. (Type locality: Northwest Territories). Harris, 1835:582.

Say, 1859:207.

Limnophilus sericeus’, Hagen, 1861 :256. Ulmer, 1905a: 19. Ulmer, 1907a:44. Schmid, 1955:

137.

Limnephilus sericeus ; Hagen, 1864:839. Banks, 1892:363. Banks, 1907a:37. Betten, 1934:

337. Milne, 1935:21. Milne, 1936:61. Ross, 1941a: 110. Ross, 1944:186, 192, 298.

Leonard and Leonard, 1949a: 19. Ross and Merkley, 1952:443. Ross and Spencer, 1952:

48. Schmid and Guppy, 1952:42. Morse and Blickle, 1953:98. Flint, 1960:51-52. Robert,

1960:59. Etnier, 1965: 149. Flint, 1966:380. Fischer, 1968:297-298.

Anabolia decepta Banks, 1899:208-209. (Type locality: Washington State).

Description. — Antennae yellow-brown. Vertex of head dark chocolate-brown, except for

lighter warts. Thorax yellow-brown, with darker areas. Male with very small brushes on ante-

rior femora. Spurs yellow. Fore wing length of male 10.6 mm; light brown, clear areas hya-

line. Costal area clear for most of distance to end of costa. Venation with humeral cross-vein

either missing in both wings or reduced, otherwise as in L. sublunatus.

Male genitalia. (Specimen from Crimson Lake, Alberta). Postero-dorsal process of tergum

VIII lengthened to cylindrical column directed meso-ventrad between cerci. Dorsal strap of

segment IX narrow, high; main body of segment very slim in lateral aspect, with postero-

dorsal angles blunt, curved mesad (Fig. 326). Clasper very acute-triangular with no dorsal

process. Median lobes of segment X long, narrow, acuminate blades with dorsal edges den-

tate, black; attached to mesal edges of basal, convex plates which extend laterad to cereal

bases. Cercus trapezoidal in lateral aspect; mesal edge massively dentate, black. Median shaft

of aedeagus short, stout, tapered; distal aspect of tip of median shaft as in Fig. 329. Lateral

arm with sclerotized, dorsally rugose base, membranous distal part (Fig. 327); tips lanceo-

late, fringed with short setae.

Female genitalia. (Specimen from Crimson Lake, Alberta). Vulval scale long, tapered,

distally rounded median lobe (Fig. 330); lateral lobes small, with concave ventral faces

(Fig. 331). Segment IX without discrete ventro-lateral lobes; larger ventrally, extending

postero-ventrad under segment X (Fig. 331). Segment X with tubular body; dorsal edge

of anal orifice extended posterad as very thin, black blade (Fig. 331); slightly cleft dis-

tally (Fig. 330). Cerci long, quite free from body of segment X; parallel sided in lateral

aspect.

Notes on biology. — Adults of this species emerge from a variety of habitats, ranging from

ponds, lakes or sloughs fringed with sedges, cattail rushes, or horse-tails, to small or large

streams and rivers of a quiet or smooth flowing nature. The flight season extends from June

15 to October 3, with no evident peak.

Geographical distribution. — The known range of this species extends from Alaska to

Oregon, and east to Quebec and the New England States (Fig. 618). In Alberta it is found

indiscriminately in the mountains, foothills and plains. In the mountain areas it is found to

altitudes of 6,675’.

I have examined 8 1 specimens, 44 males and 37 females, from the study area.

The luridus group

Males of this group are characterised by minute claspers; by long, narrow cerci; by

massive postero-dorsal bulbs of tergum VIII; and by ventrally narrow segment IX (Fig. 332).

Females are characterised by massive segment IX without discrete ventro-lateral lobes; by

small segment X; and by small, triangular cerci (Fig. 336).

102

Nimmo

Limnephilus femoralis Kolenati, 1848

(Fig. 332-336, 620)

Limnephilus nebulosus Kirby, 1837:253. (Preoccupied). (Type locality: British America).

Walker, 1852:50. McLachlan, 1863:157, 161. Banks, 1899:208. Banks, 1900b:469.

Banks, 1907a: plate 18, Fig. 9. Banks, 1908b:61, 63. Betten, 1934:327-328. Milne,

1935:46, 51. Ross, 1944:298. Ross and Merkley, 1952:448, 453-454. Wiggins, 1961:

701. Fischer, 1968: 1 13-1 15. Lindroth and Ball, 1969: 1 38. (See Fischer, 1968:1 13-115,

for Palaearctic literature).

Limnophilus nebulosus ; Kolenati, 1848:26. Hagen, 1861:259. McLachlan, 1876b:6. Brauer,

1876:286. Meinert, 1897:160-161. Jacobsen, 1898:215. Ulmer, 1905a: 19, 80. Ulmer,

1907a:42, 45. Kolbe, 1912:41. Essig, 1926:176. Mosely, 1929:502-507. Forsslund, 1932:

59. Ulmer, 1932:212, 217. Betten and Mosely, 1940:121-127. Schmid, 1955:138. (See

Fischer, 1968: 113-115, for Palaearctic literature).

Goniotaulius nebulosus ; Kolenati, 1859a: 157, 173. Hagen, 1864:816. Banks, 1892:363.

(See Fischer, 1968:113, for Palaearctic literature).

Limnophilus femoralis Kolenati, 1848:26. Hagen, 1861:260. Ulmer, 1905a: 19. Ulmer, 1932:

214, 217. (See Fischer, 1968: 1 12-1 13, for Palaearctic literature).

Limnephilus femoralis'. Walker, 1852:50. Hagen, 1864:835. Banks, 1892:363. Ulmer, 1907a:

42, 44. Banks, 1907a:36. Betten, 1934:324. Milne, 1935:20. Milne, 1936:61 . Betten and

Mosely, 1940:121, 122-126. Fischer, 1968:112-1 15. (See Fischer, 1968: 1 12-1 13, for

Palaearctic literature).

Goniotaulius femoralis', Kolenati, 1859a: 157, 173. Hagen, 1864:815. Banks, 1892:863. (See

Fischer, 1968: 1 12, for Palaearctic literature).

Limnephilus perforatus Walker, 1852:33. (Type locality: McKenzie and Slave Rivers area of

Arctic America). Johnson, 1927:49. Betten and Mosely, 1940:121, 123, 126. Fischer,

1968:115.

Limnephilus stipatus Walker, 1852:29. (Type locality: McKenzie and Slave Rivers area).

Betten and Mosely, 1940:121, 123, 126. Ross and Merkley, 1952:448. Fischer, 1968: 115.

Limnophilus stipatus;?ro\anchQT, 1877:243, 244. Provancher, 1878b: 131, 132. Provancher,

1878a: 146. Ulmer, 1907a:42.

Limnophilus subpunctulatus Hagen, 1861:261. (Type locality: Canada). Packard, 1869:161.

Packard, 1876:617-618. Fischer, 1968:1 15.

Description. — Antennae light brown; scapes and pedicels darker. Vertex of head brown,

warts paler. Thorax light to dark brown. Spurs yellow. Fore wing length of male 13.5 mm;

light brown, irrorate, Costal area entirely clear. Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from Simpson Islands, Great Slave Lake, Northwest Territories).

Postero-dorsal edge of tergum VIII developed as massive, globular bulb clothed distally with

small, stout, black spines. Dorsal strap of segment IX short, arched posterad; main body of

segment roughly triangular in lateral aspect, with very narrow sternum (Fig. 332). Clasper

minute, triangular. Median lobes of segment X black, long, acute-triangular, with rugose

lateral faces; intermediate lobes of segment X small, cylindrical, black, rugose pegs laterad of

median lobes (Fig. 333). Cercus long, narrow, distally rounded, with distal half of mesal

faces black, rugose. Lateral arms of aedeagus expanded distally (Fig. 334), with toothed

dorsal edges with four heavy setae; distal part fringed with short, dark setae, with thicker

setae on mesal faces.

Female genitalia. (Specimen from Simpson Islands, Great Slave Lake, Northwest Terri-

tories). Vulval scale broad, triangular, with triangular lateral lobes well separated from blunt,

tapered, median lobe (Fig. 335). Segment IX large, curved in lateral aspect; without discrete

Rhyacophilidae and Limnephilidae

103

ventro-lateral lobes (Fig. 336). Supra-genital plate short, narrow, lunate in ventral aspect.

Segment X small, with two triangular dorsal lobes well separated from ventral channel of

anal orifice. Cerci short, triangular, closely parallel to dorso-lateral surfaces of segment X;

segment X as a whole discrete from segment IX.

Notes on biology. — The only collection made by myself was at the Vermilion Lakes,

Banff National Park. These lakes are broad, shallow, sedge-fringed, swampy bodies of water.

Adult flight season in July (July 1-19).

Geographical distribution. - The nearctic range of this holarctic species extends from

Alaska to Greenland, Newfoundland, and Washington (Fig. 620), in North America. In

Alberta the species appears to be a plains species found in the mountains only in the larger,

low valleys. However, records are presently too few to allow proper conclusions.

I have examined 1 1 specimens, five males and six females, from the study area.

The nogus group

Males of the single known species of this group are characterised by wide dorsal strap in

lateral aspect (Fig. 337); by narrow ventral region; by median lobes of segment X set dorsad

of cerci; and by short claspers with short bases. Females are characterised by lack of discrete

ventro-lateral lobes of segment IX (Fig. 340); and by curious triangular ventral lobe formed

by fusion of segment IX ventrad of segment X (Fig. 341).

Limnephilus nogus Ross, 1944

(Fig. 337-341, 619)

Limnephilus nogus Ross, 1944:281-282, 298. (Type locality: McMinnville, Oregon). Ross,

1947:152. Schmid and Guppy, 1952:42. Ross and Spencer, 1952:48. Ross and Merkley,

1952:445. Denning, 1963:261 . Fischer, 1968:246-247.

Limnophilus nogus', Schmid, 1955: 139.

Description. — Antennae uniform deep red-brown; antero-mesal faces of scapes without

stout, heavy setae. Vertex of head uniform dark red-brown. Thorax dark chocolate-brown.

Brush of male fore leg on basal third of femur; of short, black spines. Fore wing length of

male 13.3 mm; dark red-brown, irregularly irrorate, with scattered larger hyaline areas.

Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from Oregon; Illinois Natural History Survey). Main body of

segment IX robust, narrowed gradually dorsally and ventrally (Fig. 337); dorsal strap very

thin only at meso-dorsal portion (Fig. 338). Clasper small, fused to segment IX; with short

base and body; with disto-mesal tooth. Median lobes of segment X long, parallel-sided in

lateral aspect; lyre-shaped in dorsal aspect, with distal dilations; attached to rectangular,

sclerotized bases dorsad of cerci. Cercus long, narrow, slightly tapered, with disto-mesal

tooth. Lateral arms of aedeagus bilobed distally (Fig. 339); dorsal edges of each lobe setose.

Female genitalia. (Specimen from Hosmer, British Columbia). Vulval scale roughly quad-

rangular (Fig. 341); with acute-triangular lateral lobes, long, rectangular median lobe. Seg-

ment IX of single unit; main body vertically narrow; with large, trapezoidal lateral lobes

laterad of segment X (Fig. 340); with large, dorsally curved ventro-mesal processes fused

ventrad in form of large triangular lobe (Fig. 341). Supra-genital plate wide, very small,

short. Segment X massive, with lateral edges folded dorso-mesally to partly enclose dorsal

surface of segment; in ventral aspect broad, rectangular plate with sinuate lateral edges and

short, mesal cleft (Fig. 341) continued as ventro-mesal channel.

Notes on biology. — I have not collected specimens of this species. The only record within

104

Nimmo

the study area is a single female from Hosmer, British Columbia. The other records available

to me from elsewhere indicate a flight season divided into two parts; the earliest flight

occurs in May, the second in September and October.

Geographical distribution. — The known range of this species extends from British Colum-

bia to California (Fig. 619). In the study area the single record is from an altitude of about

3,000’.

I have examined 18 specimens, two males and 16 females, from British Columbia. Seven-

teen of these specimens are from southwestern British Columbia and Oregon.

The sitchensis group

Males of this group are characterised by long, narrow cerci (Fig. 342); by short, wide-

based, placoid claspers; by long, wide-based, triangular median lobes of segment X; and by

aedeagi with tips of median shaft and lateral arms curved markedly dorsad; tip of median

shaft flattened ventrally, occupied by ejaculatory duct pore (Fig. 344). Females are charac-

terised by segment IX with ventro-lateral lobes distinct but not discrete from dorsal body;

by large, thumb-like cerci fused solidly to segment IX, directed ventro-posterad along sides

of segment X; and by thin walled, tubular segment X (Fig. 346, 353).

Limnephilus moestus Banks, 1 908

(Fig. 342-346, 621)

Limnephilus moestus Banks, 1908b:61, 62. (Type locality: Grand Lake, Newfoundland).

Johnson, 1927:107. Banks, 1930b: 128. Betten, 1934:327. Milne, 1935:46, 51. Ross,

1938b:37. Ross, 1944:6, 14, 186, 189, 191-192, 298. Proctor, 1946:211. Leonard and

Leonard, 1949a: 18. Ross and Merkley, 1952:445. Morse and Blickle, 1953:98. Robert,

1960:59. Wiggins, 1961 : 701 . Etnier, 1965: 149. Nimmo, 1966a:692. Nimmo, 1966b:224.

Fischer, 1968:236-237. McConnochie and Likens, 1969:150.

Limnophilus hingstoni Mosely, 1929:502, 504-507. Forsslund, 1932:56. Ulmer, 1932:213,

217. Ross, 1944:191, 298. Fischer, 1968:236-237.

Males of this species are distinguished by exceptionally long cerci (Fig. 342); by massive

claspers deeply indented into segment IX; and by presence of lightly setose, incipient

postero-dorsal process of tergum VIII. Females are distinguished by cerci projected well

posterad of ventral extremity of segment X (Fig. 346); and by short tip to ventro-lateral

extension of dorsal body of segment IX.

Description. — Antennae light brown; scapes dark brown, with antero-mesal faces paler,

glabrous. Vertex of head dark brown. Thorax light brown, darker dorsally. Spurs yellow.

Fore wing length of male 1 1.6 mm; medium brown. Costal area clear to mid-point of wing.

Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from Lethbridge, Alberta). Postero-dorsal edge of tergum VIII

slightly protuberant, lightly setose. Segment IX overall rather narrow (Fig. 342); dorsal strap

relatively thick, with distinct ventro-mesal process; ventral edge of segment with several

stout setae. Clasper large, fused to posterior edge of segment IX; directed postero-mesad,

with slight basal declivity; with three very distinct, long, black setae on ventral edge. Median

lobe of segment X large, triangular (Fig. 342), with swollen bases (Fig. 343); black distally,

light brown basally; minutely dentate disto-lateraily; attached to mesal edges of basal plates

which fit closely with bases of claspers and cerci. Cercus very long, narrow, tapered very

gradually distad; mesally concave, with narrow meso-basal shelf (Fig. 343); distal half of

mesal surface tuberculate. Median shaft of aedeagus without discrete head (Fig. 344); curved

Rhyacophilidae and Limnephilidae

105

dorsad, with three minute spines in orifice of ejaculatory duct. Lateral arms bifid basally;

mesal branches fused together dorsally across base of median shaft; arms dilated slightly dis-

tally, curved dorsad, fringed ventrally and distally with seate.

Female genitalia. (Specimen from Bow Pass, Banff National Park, Alberta). Vulval scale

with short, squat, slightly tapered median lobe; lateral lobes each rectangular (Fig. 345).

Dorsal body of segment IX small, indistinguishable from segment X (Fig. 346); divided by

lateral grooves from much larger, roughly triangular, ventro-lateral processes, with posterior

process short. Supra-genital plate very wide, short, with straight posterior edge. Segment X

small, tubular, darker than remainder of genitalia; deeply cleft ventrally, with short, v-cleft

dorsally (Fig. 345); thin walled. Cerci relatively long, finger-like, dark brown, directed

ventro-posterad, beyond ventral extremity of segment X.

Notes on biology. — Specimens of this species emerge from ponds or sloughs dominated

by sedges. The adult flight period extends from July 19 to September 12.

Geographical distribution. — The known range of this species extends from British Colum-

bia and Utah to Greenland and Newfoundland (Fig. 621). In Alberta most specimens were

collected in the mountains, but I have one record from Lethbridge, well out in the plains.

In the mountains it may attain altitudes of up to 6,878’.

I have examined 44 specimens, 35 males and nine females, from the study area and the

Northwest Territories.

Limnephilus cocker elli Banks, 1900

(Fig. 347-349, 622)

Limnophilus cockerelli Banks, 1900c: 124. (Type locality: Sapello, New Mexico). Ulmer,

1905a: 19. Ulmer, 1907a:44. Essig, 1926:175. Schmid, 1955:137.

Limnephilus cockerelli ; Banks, 1904a: 107. Betten, 1934:335. Milne, 1935:46, 51. Ross,

1938b:34-35. Ross, 1941a: 109. Ross, 1944:298. Ross and Merkley, 1952:445. Fischer,

1968:65.

Males of this species are distinguished by wide dorsal strap of segment IX (Fig. 347); by

short, parallel-sided, bilobed claspers, with dorsal lobe thin, sharp, and ventral lobe thick,

rounded; by short, triangular median lobes of segment X; and by relatively short aedeagus

(Fig. 348).

Description. — Antennae brown; scapes darker except for paler, glabrous antero-mesal

| faces. Vertex of head mottled dark and light brown. Thorax light red-brown; brushes of

male fore leg of thin, black spines along basal third of anterior edge of femur. Spurs brown.

Fore wing length of male 14.9 mm; light yellow brown, irrorate; anterior half of wing

virtually hyaline. Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from Eisenhower Junction, Banff National Park, Alberta).

Segment IX high, narrow, with wide dorsal strap (Fig. 347); narrow ventrally; postero-

lateral angles of segment folded, directed mesad as finger-like process. Clasper articulated

to segment IX; distally bilobed, with acuminate dorsal lobe, full, rounded, ventral lobe;

setose. Median lobes of segment X black, with dentate disto-lateral faces (Fig. 349); short,

triangular in lateral aspect (Fig. 347); attached to lateral edges of two deeply concave

basal plates. Cercus long, narrow, parallel-sided, with concave, dentate mesal face. Median

shaft of aedeagus curved sharply dorsad to large, slanted tip; lateral arms similarly curved,

with slightly expanded tips fringed with setae (Fig. 348) extended part way across lateral

faces.

Female genitalia. Unknown.

Geographical distribution. — The known range of this species extends from Alberta, south

106

Nimmo

along the Rocky Mountains to New Mexico. In Alberta it is known from a single locality,

at Eisenhower Junction, Banff National Park, at an altitude of 4,600’ (Fig. 622).

I have examined a single male specimen from the study area, in the Canadian National

Collection, Ottawa.

Limnephilus valhalla Nimmo n. sp.

(Fig. 350-354, 622)

Males of this species are distinguished by exceptionally long cerci, bowed ventrad half

way from base (Fig. 350); by distinct spinate process of tergum VIII; and by massive, tri-

angular median lobes of segment X. Females are distinguished by distinct posterior process

of ventro-lateral body of segment IX (Fig. 353); and by high, narrow, rectangular dorsal

body of segment IX.

Description. — Antennae brown; scapes darker, with antero-mesal faces lighter, glabrous.

Vertex of head black except for postero-lateral angles. Thorax light red-brown. Brush of

male fore leg of thin, dark spines on basal third of anterior face of femur. Spurs yellow.

Fore wing length of male 1 1.6 mm; dark brown, clear areas hyaline. Costal area clear. Vena-

tion identical to that of L. sublunatus ; costa hyaline basad of humeral cross-vein.

Male genitalia. (Specimen from Mt. Edith Cavell, Jasper National Park). Postero-dorsal

process of tergum VIII small, hemispherical, with short black spines. Dorsal strap of segment

IX high, narrow. Main body of segment small, with postero-dorsal angle directed posterad;

with fringe of setae on postero-ventral edge (Fig. 350). Clasper small, fused to segment IX,

with several long, irregular setae on posterior edge. Median lobes of segment X with distal

halves black; massive, triangular, thin plates. Cercus long, narrow, bowed ventrad at mid-

point; slightly concave mesally, with black mesal faces on distal halves (Fig. 351). Median

shaft of aedeagus curved strongly dorsad; lateral lobes similar, with distal portion at right

angles to basal arm, fringed with setae (Fig. 352).

Female genitalia. (Specimen from Mt. Edith Cavell, Jasper National Park, Alberta). Vulval

scale with very broad, short, slightly tapered median lobe (Fig. 354); lateral lobes rectangu-

lar, with concave distal faces. Segment IX high, narrow, with acute-triangular processes at

postero-ventral angle (Fig. 353). Supra-genital plate short, wide, rectangular. Segment X

relatively large, with scalloped posterior edges; lateral edge with large tooth dorsally; deeply

cleft dorsally and ventrally (Fig. 354). Cerci short, thumb-like, directed postero-ventrad

beside segment X.

Notes on biology. — Adults of this species were collected from small, sedge-lined, moun-

tain lakes or pools.

Geographical distribution. — This species is known from only two localities; one in Alber-

ta, at an altitude of 6,800’; the second is in British Columbia, at 5,220’, 1 mile west of the

continental divide (Fig. 622).

Holotype. — Male. First pools on trail to Mt. Edith Cavell alpine meadows, Jasper Nation-

al Park, Alberta; July 25, 1966; A. Nimmo.

Allotype. — Female. Same data as holotype.

Paratypes. — Same data as holotype; one male, one female. Wapta Lake beaver pond,

Kicking Horse Pass, Yoho National Park, British Columbia; August 10, 1967; A. Nimmo;

one female.

The type series is in the Canadian National Collection, Ottawa, and has been assigned the

number 10,587.

This species is named for ‘Valhalla’, home for those fallen in battle, as noted in my read-

ings in Norse mythology.

Rhyacophilidae and Limnephilidae

107

Limnephilus lopho Ross, 1 949

(Fig. 355-359, 623)

Limnephilus lopho Ross, 1949b: 1 19-120. (Type locality: Hood River Meadows, Mt. Hood,

Oregon). Schmid and Guppy, 1952:42. Ross and Merkley, 1952:445. Ross and Spencer,

1952:48. Fischer, 1968:201.

Limnophilus lopho \ Schmid, 195 1 : 2-6. Schmid, 1955:137.

Description. - Antennae with basal two-thirds brown; distal third yellow; scapes, pedicels

dark brown except for yellow, glabrous, antero-mesal faces of scapes. Vertex of head dark

brown between lateral ocelli. Thorax black or dark brown; brush of male fore leg of long,

fine, black spines on basal third of femur. Spurs yellow. Fore wing length of male 12.7 mm;

brown to dark reddish brown. Costal area clear. Venation identical to that ofZ,. sublunatus.

Male genitalia. (Specimen from Mt. Edith Cavell, Jasper National Park, Alberta). Dorsal

strap of segment IX high, thick (Fig. 355). Main body of segment roughly triangular, with

narrow sternum; anterior edges with distinct flanges. Clasper short, broad, thin plate with

rounded tips. Median lobes of segment X large, triangular, black on distal halves, with tips

directed dorso-laterally. Intermediate lobes small, rugose, bulbous pegs; black, heavily sclero-

tized. Cercus thumb-like in lateral aspect, with tip minutely dentate mesally, black (Fig.

356). Median shaft and lateral arms of aedeagus curved sharply dorsad (Fig. 357); lateral

arms fringed distally, with short fringe on meso-ventral edges.

Female genitalia. (Specimen from Mt. Edith Cavell, Jasper National Park, Alberta). Vulval

scale almost square (Fig. 358); lateral lobes rectangular; median lobe short, narrow, well

separated from lateral lobes. Dorsal body of segment IX small, rectangular, separated dor-

sally from segment X by slight declivity (Fig. 359); ventro-lateral extremities triangular in

lateral aspect, fused ventrally. Supra-genital plate wide, very short, sinuate. Segment X in

lateral aspect plough-like; deeply cleft ventrally; shallowly but widely cleft dorsally (Fig.

358). Cerci short, broad, rounded lobes located baso-laterad to segment X.

Notes on biology. — Adults of this species are found adjacent to small alpine pools or

seepage slopes heavily clothed with sedges. The flight period extends from July 25 to

August 5.

Geographical distribution. — The known range of this species extends from Oregon to

British Columbia and Alberta (Fig. 623). I have taken it at altitudes of 6,600’ and 6,800’, on

the east slope of Tonquin Valley, and at Mt. Edith Cavell, both in Jasper National Park.

The assimilis group

There are two species of this group present in the study area (see Fig. 360, 365). In geni-

talic characters these two species are so divergent that I cannot provide a list of common

characters. Each species is characterised below.

Limnephilus parvulus (Banks), 1905

(Fig. 360-364, 623)

* Stenophylax ? parvulus Banks, 1905:9-10. (Type locality: New Hampshire).

Parachiona parvulus', Banks, 1907a: 39.

Hypnotranus ? parvulus', Ulmer, 1907a:72.

Algonquina parvula', Banks, 1916:121, 122. Betten, 1934:370. Milne, 1935:31,49.

Limnephilus parvulus', Ross, 1938b:39. Ross, 1944:298. Ross and Merkley, 1952:449. Morse

and Blickle, 1953:98. Etnier, 1965: 149. Fischer, 1968:253-254.

108

Nimmo

Limnophilus parvulus ; Schmid, 1955:141.

Apolopsyche pallida Banks, 1924:442. (Type locality: Manitoba). Betten, 1934:370.

Anolopsyche pallida ; Criddle, 1925: 16.

Limnephilus pallida'. Banks, 1930a:226-227. Milne, 1935:49. Ross, 1938b:38-39.

Limnephilus pallidus', Fischer, 1968:254.

Limnephilus roberti Banks, 1930a: 226-227. (Type locality: Lake Winnipeg, Manitoba).

Betten, 1934:337. Neave, 1934:167. Milne, 1935:46, 5 1 . Ross, 1938b:39. Fischer, 1968:

254.

Males of this species are distinguished from males of other species of Limnephilus by

massive, distally toothed claspers; by thin-bladed, black median lobes of segment X which

are curved slightly ventrad (Fig. 360); and by lack of postero-dorsal process on tergum VIII.

Females are distinguished by discrete ventro-lateral lobes of segment IX; and by small,

tapered, thin segment X (Fig. 363).

Description. — Antennae light brown; scapes brown, with paler antero-mesal faces. Vertex

of head with two dark areas between lateral ocelli. Thorax medium to light brown. Male

fore leg with very slight brush of short, brown spines at base of femur. Spurs pale yellow.

Fore wing length of male 10.5 mm; light yellowish brown; clear areas smoky white; coloured

areas closely following venation; irrorate. Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from Great Slave Lake, Northwest Territories). Dorsal strap of

segment IX wide, with acuminate meso-ventral spine (Fig. 360). Main body of segment large,

tapered slightly ventrad, with straight posterior edge, sinuate dorsal edge. Clasper with disto-

dorsal tooth; mesal face of tooth concave, fringed with small, black dentations; base of

clasper short. Median lobes of segment X long, tapered, bowed slightly ventrad; lateral faces

black, ventral edges minutely dentate; lobes thin, with disto-lateral teeth (Fig. 361). Cercus

short, sub-triangular in lateral aspect, fringed distally with long setae. Lateral arms of aedea-

gus hardly expanded distally, fringed along dorsal edges with setae (Fig. 362).

Female genitalia. (Specimen from Great Slave Lake, Northwest Territories). Vulval scale

with distinct, straight base (Fig. 364); lateral lobes simple, curved mesad; median lobe rec-

tangular. Dorsal body of segment IX small, tapered abruptly at junction with segment X

(Fig. 363); ventro-lateral lobes large. Supra-genital plate with simple, curved posterior edge;

small. Segment X small, long, tapered; deeply cleft mesally (Fig. 364). Cerci acute-triangular

in lateral aspect; closely appressed to lateral surfaces of segment X.

Notes on biology. — Adults of this species are associated with lakes and sloughs with a

heavy growth of sedges or Typha. The flight season extends from May 25 to July 5.

Geographical distribution. — The known range of this species extends from Alberta and

Great Slave Lake to Quebec and New Hampshire (Fig. 623). In Alberta it is found largely in

the plains, and in the low, major mountain valleys.

I have examined 38 specimens, 27 males and 1 1 females, from the study area and Great

Slave Lake.

Limnephilus spinatus Banks, 1914

(Fig. 365-369, 624)

Limnephilus spinatus Banks, 1914: 149-150. (Type locality: Vineyard, Utah). Betten, 1934:

337. Ross, 1938b:40. Ross, 1944:299. Ross and Merkley, 1952:451. Denning, 1963:261.

Fischer, 1968:314.

Limnophilus spinatus\ Essig, 1926: 176. Schmid, 1955:141.

Anabolina spinata; Milne, 1935:41, 49.

Males of this species are distinguished from males of other species of Limnephilus by very

Rhyacophilidae and Limnephilidae

109

long base of the clasper (Fig. 365); by high, thin, dorso-anteriorly hooked, acuminate median

lobes of segment X; and by presence of small, black-spinate, postero-dorsal lobe of tergum

VIII. Females are distinguished by very massive ventro-lateral lobe of segment IX (Fig. 369);

by minute dorsal body of segment IX; and by postero-ventrally slanted segment X.

Description. — Antennae dark brown; scapes dark chocolate-brown, with antero-mesal

faces lighter, setaless. Vertex of head uniform dark brown. Thorax light brown, with darker

areas. Brush of male fore leg well developed, of short, stout, black spines along entire length

of femur. Spurs yellow. Fore wing length of male 10.8 mm; warm chocolate-brown. Costal

area entirely clear. Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from Hound Creek, 25 miles southeast of Cascade, Montana).

Tergum VIII with small, black-spinate, postero-dorsal process. Dorsal strap of segment IX

directed postero-dorsad (Fig. 365). Main body of segment of irregular outline, narrowed

ventrad. Clasper with very high, narrow base, with long, thin, dorso-posteriorly directed

dorsal process. Median lobes of segment X square in lateral aspect, with postero-dorsal

angles produced as long, anteriorly curved, acuminate spines; lobes attached to mesal edges

of rectangular, basal plates (Fig. 366). Cercus squat, distally rounded, with concave mesal

face. Lateral arms of aedeagus very long, slender, spiniform (Fig. 367).

Female genitalia. (Specimen from Crowsnest Pass, 8 miles west of Coleman, Alberta).

Vulval scale with broad, abruptly tapered median lobe (Fig. 368); lateral lobes with blunt

lateral horns. Segment IX with minute dorsal body (Fig. 369), massive ventro-lateral lobes

of indeterminate form. Segment X large, of two sets of sclerites; basal set longer, rectangu-

lar; distal set short, each roughly triangular in lateral aspect. Supra-genital plate wide, short,

with almost straight posterior edge.

Notes on biology. — Adults of this species were found near small ponds or sloughs, and

slow streams. The flight period extends from July 25 to August 30.

Geographical distribution. — The known range of this species extends from Alberta to

California and Colorado (Fig. 624). In Alberta specimens have been collected in the plains

and low mountain valleys of the southwestern comer of the province.

I have examined 22 specimens, 21 males and one female, from the study area and

Montana (one male).

The incisus group

Four species of this group are known from the study area. As with the assimilis group

there appears to be no significant community of characters to define the group (Fig. 370-

398).

Limnephilus hyalinus Hagen, 1861

(Fig. 370-374, 627)

Limnophilus hyalinus Hagen, 1861:258. (Type locality: North Red River, Canada). McLach-

lan, 1876b:7. Ulmer, 1907a:44. Schmid, 1955:140.

Limnephilus hyalinus ; McLachlan, 1863:157. Banks, 1892:363. Banks, 1899:208. Banks,

1907a:36. Betten, 1934:336. Neave, 1934:167. Milne, 1935:5 1 . Ross, 1938b:36. Banks,

1943:342. Ross, 1944:186, 189, 191, 298. Leonard and Leonard, 1949a:18. Ross and

Merkley, 1952:441. Ross and Spencer, 1952:48. Morse and Blickle, 1953:98. Robert,

1960:59. Wiggins, 1961 : 701. Etnier, 1965: 149. Nimmo, 1966a:692. Nimmo, 1966b:224.

Fischer, 1968:179-180.

Limnephilus species 1; Betten, 1934:335. Ross, 1938b:36. Fischer, 1968:180.

110

Nimmo

Males of this species are distinguished by close conjunction between distal edge of each

median lobe of segment X and black line of teeth on mesal face of each cercus (Fig. 370,

371). Females are distinguished by scoop-shaped ventro-median lobes of segment IX (Fig.

373) , which are discrete from dorsal body of segment; by large, triangular dorsal body of

segment IX; and by long, thin cerci flanked laterally by latero-dorsal lobes of segment X.

Description. — Entire body and wings very pale, straw-yellow; warts, and costal area of

fore wings almost white. Fore wing length of male 9.8 mm. Venation identical to that of

L. sublunatus.

Male genitalia. (Specimen from Vermilion, Alberta). Postero-dorsal edge of tergum lightly

setose. Dorsal strap of segment IX relatively short, thick. Main body of segment relatively

narrow, slightly narrowed ventrad (Fig. 370). Clasper short, with wide base; triangular in

lateral aspect. Median lobes of segment X short, distal halves black, tapered slightly distad

(Fig. 371); distal edges vertical, straight, coincident with vertical line of black teeth on mesal

faces of cerci. Cercus widened distad, slightly concave mesally, with vertical row of black

teeth at mid-point of mesal face. Lateral arms of aedeagus long, expanded distad (Fig. 372),

each with dorsal spine basad of membranous, irregular distal half.

Female genitalia. (Specimen from Manyberries, Alberta). Vulval scale almost circular,

with curved, parallel-sided lateral lobes (Fig. 373), longer, tapered, median lobe. Segment IX

with large, triangular dorsal body, larger, scoop-shaped, discrete, ventro-lateral lobes (Fig.

374) . Supra-genital plate small, evenly rounded distally. Segment X composed of two dorso-

lateral, rectangular lobes laterad of long, thin cerci, with small ventral lobe.

Notes on biology. - Adults of this species are found near sloughs and lakes which are

thickly vegetated with sedges and cattail rushes. The flight season extends from July 5 to

September 24, with a possible peak in August.

Geographical distribution. — This species is transcontinental, extending from Alaska to

Oregon, Colorado, Illinois, and Newfoundland (Fig. 627). In the study area it occurs

throughout the plains regions, and through the mountains along the low, major valleys

reaching altitudes of 5,200’.

I have examined 91 specimens, 48 males and 43 females, from the study area and Great

Slave Lake.

Limnephilus secludens Banks, 1914

(Fig. 375-379, 625)

Limnephilus secludens Banks, 1914:152. (Type locality: Penticton, British Columbia).

Betten, 1934:337. Milne, 1935:50. Ross, 1938b:39. Ross, 1944:299. Ross and Spencer, 3

1952:48. Ross and Merkley, 1952:436, 450. Denning, 1963:261. Denning, 1965b:697.

Etnier, 1965:149. Fischer, 1968:297.

Limnophilus secludens', Essig, 1926:176. Schmid, 1955:140.

Colpotaulius secludens', Milne, 1936:60. Schmid and Guppy, 1952:48.

Males of this species are distinguishable by very short, membranous, distally spinate

lateral arms of aedeagus (Fig. 377). Females are distinguished by short, tubular segment X

with short, peg-like cerci solidly fused to dorso-lateral surfaces of segment (Fig. 378, 379).

Description. — Antennae brown; scapes lighter, setaless on antero-mesal faces. Vertex of

head dark brown; lighter laterally. Thorax dark reddish brown; brush of male fore leg of

short, black spines along full length of femur; opposing face of tibia with corresponding :

brush. Spurs very long, yellow-brown. Fore wing length of male 14.2 mm; light yellow- d

brown with darker areas; costal area clear except for slight distal darkening. Venation

identical to that of L. sublunatus.

Rhyacophilidae and Limnephilidae

111

Male genitalia. (Specimen from Lethbridge, Alberta). Dorsal strap of segment IX short,

with ventro-mesal process connected to segment X (Fig. 375, 376). Main body of segment

tall, narrow, rectangular in lateral aspect. Clasper high, narrow, tapered, ventrad, acute-

triangular. Median lobes of segment X short, narrow, black; tips directed dorso-laterad.

Cercus short, irregular, deeply concave mesally. Median shaft of aedeagus with straight,

thick basal portion (Fig. 377); distal third turned sharply dorsad, thin, narrow, acuminate.

Lateral arms short, stubby, membranous; each with cluster of short, black spines distally.

Female genitalia. (Specimen from Lethbridge, Alberta). Vulval scale with median lobe

longer than lateral lobes, tapered from wide base (Fig. 379); lateral lobes concave ventrally;

Mesal edges sinuate, black. Dorsal body of segment X distinguished from segment X by

posterior declivity (Fig. 378); ventro-lateral lobes almost separated from dorsal body; large,

with straight, angular edges. Supra-genital plate squat, triangular. Segment X large, tubular,

with minor dorsal and ventral projections. Cerci short, finger-like, fused solidly to dorso-

lateral faces of segment X.

Notes on biology. - Adults of this species are found near lakes and sloughs with heavy

growths of sedges and cattail reeds. The flight period extends from June 3 to September 23,

with a possible peak in late July and August.

Geographical distribution. — The known range of this species extends from Great Slave

Lake to Wisconsin, New Mexico, and California (Fig. 625). In Alberta it seems to be restric-

ted to the plains, with occasional records from the foothills.

I have examined 372 specimens, 178 males and 194 females, from the study area and

Great Slave Lake.

Limnephilus janus Ross, 1938

(Fig. 380-384, 626)

Colpotaulius minusculus Banks, 1924:439. (Preoccupied). (Type locality: Tolland, Colo-

rado). Betten, 1934:339. Milne, 1935:42, 50. Milne, 1936:60. Ross, 1938b:37. Fischer,

1968:199.

Limnephilus janus Ross, 1938b: 37. (Type locality: Tolland, Colorado). Ross, 1944:299.

Ross and Merkley, 1952:451. Etnier, 1965:1.

Limnophilus janus ; Schmid, 1955:140.

Males of this species are distinguished by high, very thin dorsal strap of segment IX (Fig.

380); by large, relatively long cerci with concave mesal faces and irregular dorso-mesal edges

(Fig. 381); and by long, distally thin and acuminate median lobes of segment X. Females are

distinguished by trilobed dorsal shield of segment X; and by absence of discrete ventro-

lateral lobes of segment IX (Fig. 383).

Description. — Antennae pale yellow. Vertex of head pale yellow-brown, with dark areas

adjacent to ocelli. Thorax pale yellow with some slightly darker areas. Brush of male fore

leg composed of black spines along basal two-thirds of femur. Spurs pale yellow. Fore wing

length of male 7.2 mm; greyish white, with scattered brown areas tending to follow veins.

Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from 6 miles south of Elk Point, Alberta). Dorsal strap high,

narrow, with sharp dorsal edge (Fig. 380). Main body of segment IX wide, with distinct

antero-ventral bulge. Clasper short, with short base; almost rectangular with dorso-distal

tooth. Median lobes of segment X long, thin, each with wide base abruptly narrowed to thin

distal process with tip directed dorso-laterad (Fig. 381). Cercus long, with disto-dorsal edge

curled mesad, irregular, black. Aedeagus simple, with short, tapered median lobe, and long,

dorsally curved, distally expanded, acuminate lateral arms, clothed on ventral edges with

112

Nimmo

setae (Fig. 382).

Female genitalia. (Specimen from 6 miles south of Elk Point, Alberta). Vulval scale with

wide based, strongly tapered median lobe (Fig. 384); lateral lobes large, with concave lateral

faces. Segment IX large, with no separate ventro-lateral lobes (Fig. 383). Supra-genital plate

rectangular, set well ventrad of segment IX to which it is connected by band of folded mem-

brane. Segment X large, separate from segment IX, with cylindrical ventral portion sur-

mounted by trilobed dorsal roof, with lateral lobes.

Notes on biology. — I have collected specimens of this species only in the vicinity of large

or small, shallow, sedge-edged lakes, ponds or sloughs. The flight season extends from July

12 to September 13, with peak about mid-August.

Geographical distribution. - This species is known from Alberta, Saskatchewan, Min-

nesota, and Colorado (Fig. 626). In Alberta it is a species of the plains and low foot-

hills.

I have examined 40 specimens, 20 males and 20 females, from Alberta and Saskatchewan.

Limnephilus perpusillus Walker, 1852

(Fig. 385-389, 628)

Limnephilus perpusillus Walker, 1852:35. (Type locality: St. Martin’s Falls, Albany River,

Hudson’s Bay). Hagen, 1861:254. Ross, 1941a:107. Ross, 1944:298. Ross and Merkley,

1952:436, 450. Etnier, 1965:149. Fischer, 1968:255-256. Clifford, 1969:582.

Colpotaulius perpusillus ; McLachlan, 1863:161. Hagen, 1864:809. McLachlan, 1874:36.

Banks, 1892:363. Ulmer, 1905a: 18. Ulmer, 1907a:37. Banks, 1907a:37. Banks, 1908b:

61. Kolbe, 1912:42. Milne, 1935:42, 50. Milne, 1936:60.

Limnophilus perpusillus', Packard, 1876:617. Schmid, 1955:140.

Anabolina perpusillus ; Betten, 1934:338.

Anabolina perpusilla \ Betten and Mosely. 1940:140-142.

Colpotaulius rhaeus Milne, 1935:42, 50. (Type locality: Guelph, Ontario). Milne, 1936:60.

Ross, 1941a: 107. Ross, 1944:298. Fischer, 1968:256.

Limnephilus merinthus Ross, 1938c: 166. (Type locality: Churchill, Manitoba). Ross, 1941a:

107. Ross, 1944:298. Fischer, 1968:256.

Males of this species are distinguished by small postero-dorsal, spinate lobe of tergum

VIII (Fig. 385); by square median lobes of segment X with postero-dorsal angles developed

as anteriorly hooked spines; and by large, black spine on dorso-mesal edge of cerci (Fig.

285, 386). Females are distinguished by massive, fleshy ventral body of segment IX (Fig.

388); and by vertically oriented segment X.

Description. — Antennae light reddish brown; scapes and pedicels darker, antero-mesal

faces setaless. Vertex of head with circular dark brown area between lateral ocelli. Thorax

light brown, with darker areas. Brush of male fore leg composed of short, fine black spines

along entire length of femur. Spurs yellow. Fore wing length of male 7.9 mm; light yellow-

ish brown; very lightly irrorate in posterior regions. Costal area entirely clear. Venation

identical to that of L. sublunatus.

Male genitalia. (Specimen from camp 3 miles east of Nordegg, Alberta). Postero-dorsal

edge of tergum VIII with small, free, black-spinate lobe. Dorsal strap of segment IX narrow

(Fig. 385). Main body of segment narrow, tapered ventrad; tapered dorsad to merge gradu-

ally with dorsal strap. Clasper with very long, narrow base produced dorso-posterad as

finger-like dorsal process. Median lobes of segment X with square bases; postero-dorsal

angles produced dorso-anterad as short, slender, hooked spines. Cercus short, triangular in

lateral aspect, with dorso-mesal edge curled ventrad with single large spine (Fig. 385, 386).

Rhyacophilidae and Limnephilidae

113

Median shaft of aedeagus simple, short, tapered; lateral arms curved dorsad, each with

distally expanded, acuminate tip fringed along ventral edge with setae (Fig. 387).

Female genitalia. (Specimen from pond 10 miles south of McLeod River, Forestry Trunk

Road, Alberta). Vulval scale with squat, short median lobe tapered distad (Fig. 389). Lateral

lobes almost rectangular, with deep channel on mesal edge of each lobe. Dorsal body of

segment IX small; ventro-lateral extensions massive, fused ventrally (Fig. 388, 389). Supra-

genital plate membranous, lunate, of two wide lateral plates fused ventrally to form channel,

with short, narrow, distal extension. Cerci short, thin, almost rectangular lobes attached

between lateral plates of segment X.

Notes on biology. — Specimens of this species are collected near ponds with large quanti-

ties of sedge or bullrushes, or smaller creeks. The flight season extends from July 2 to

September 6.

Geographical distribution. — The known range of this species extends from Alberta to

Colorado and Newfoundland. In Alberta it occurs primarily in the plains and low foothills

(Fig. 628).

I have examined 1 2 specimens, three males and nine females, from the study area.

The argenteus group

Males of the single species in this group are characterised by presence of spinate postero-

dorsal lobe of tergum VIII; by large segment IX with short dorsal strap; by short, broad

claspers; and by long, narrow, evenly tapered median lobes of segment X (Fig. 390). Females

are characterised by discrete ventro-lateral lobes of segment IX; by segments IX and X being

fused; and by minute, thin cerci (Fig. 394).

Limnephilus argenteus Banks, 1914

(Fig. 390-394, 629)

Limnephilus argenteus Banks, 1914:152. (Type locality: Nipigon, Ontario). Betten, 1934:

320. Milne, 1935:46, 51. Ross, 1938b:34. Ross, 1944:14, 298. Ross and Merkley, 1952:

447. Fischer, 1968:25. Lindroth and Ball, 1969:138.

Limnophilus argenteus ; Schmid, 1955:143.

Description. — Antennae brown; scapes with antero-mesal faces lighter, almost glabrous.

Vertex of head with two dark brown areas mesad of lateral ocelli. Thorax brown with

darker areas. Spurs light brown. Fore wing length of male 16.8 mm; light to dark brown,

irrorate, clear areas very pale brown, not hyaline. Costal area sparingly irrorate. Venation

identical to that of L. sublunatus.

Male genitalia. (Specimen from Vermilion Lakes, Banff). Postero-dorsal edge of tergum

VIII produced posterad as small, black-spinate bulb. Dorsal strap of segment IX short, with

sharp dorsal edge (Fig. 390). Main body of segment massive, irregular. Clasper short, broad,

very stout, with short base. Median lobes of segment X long, narrow at bases, tapered evenly

to acuminate tips; curved dorso-laterad (Fig. 391). Cercus short, almost square in lateral

aspect, with wide baso-mesal extension dorsally; concave mesally. Median shaft of aedeagus

attached to folded, membranous base, curved dorsad (Fig. 392); lateral arms straight basally,

curved dorsad distally to expanded, flared, mesally concave tips.

Female genitalia. (Specimen from Worsley, Alberta). Vulval scale large, trapezoidal in

general outline (Fig. 393); median lobe long, gradually tapered; lateral lobes thin, arched

laterad. Dorsal body of segment IX small, fused imperceptibly to segment X (Fig. 394);

ventro-lateral lobes discrete from dorsal body, high, narrow. Supra-genital plate large, evenly

114

Nimmo

rounded distally. Segment X small, distally dark; deeply narrowly cleft mesally (Fig. 393).

Cerci two minute, thin, papillate structures attached to dorsal surface of segment X.

Notes on biology. — The single record which I have of this species from the study area is

from large, sedge marshes just west of Banff, Alberta. The dates of capture in Alberta range

from July 1 to July 12.

Geographical distribution. — The known range of this species extends from Alaska to

Illinois and Ontario (Fig. 629).

I have examined 10 specimens, two males and eight females, from the study area and

Great Slave Lake.

The alberta group

The single species known in this group is characterised by high, narrow segment IX (Fig.

395), with virtually no dorsal strap; by short, slender, tapered claspers; and by massive,

thick, heavily sclerotized cerci (Fig. 396). Females are characterised by high, narrow seg-

ment IX (Fig. 400); and by vulval scale almost completely demarcated from sternum VIII

(Fig. 401).

Limnephilus alberta Denning, 1958

(Fig. 395-401,629)

Limnephilus alberta Denning, 1958:96. (Type locality: Pipestone River, Lake Louise, Alber-

ta). Denning, 1965b:698.

Description. — Antennae dark brown; scapes with pale, glabrous antero-mesal faces.

Vertex of head very dark brown, warts paler. Thorax dark brown; warts very pale. Spurs

yellow. Fore wing length of male 14.9 mm; light brown, with darker areas; clear areas

mostly hyaline. Costal area clear on basal two-thirds. Venation identical to that of L.

sublunatus.

Male genitalia. (Specimen from 1 1 miles west of Banff, on Trans-Canada Hwy.). Dorsal

strap of segment IX virtually non-existent (Fig. 395). Main body of segment sinuate,

widened gradually dorsad from narrow ventral area. Clasper small, narrow, tapered, finger-

like, with very short base. Median lobes of segment X small, trapezoidal, slanted dorso-

mesad (Fig. 396). Cercus rectangular in lateral aspect, tapered basad from flat, black distal

face, in dorsal aspect (Fig. 397). Median shaft of aedeagus short, stout, with large, discrete,

distal head (Fig. 398). Lateral arms curved smoothly dorsad, not widened, each fringed

along mesal face with long setae (Fig. 398, 399).

Female genitalia. (Specimen from 1 1 miles west of Banff, on Trans-Canada Hwy.). Vulval

scale with angular anterior edge, almost separate from sternum VIII (Fig. 401); median lobe

long, acute-triangular; lateral lobes stout, close to median lobe. Segment IX high, narrow,

with slightly wider ventroTateral area (Fig. 400). Supra-genital plate short, broad, suspended

from segment IX by wide, wrinkled membrane. Segment X with hemi-cylindrical ventral

portion short, and dorsal, mesally flattened, dorsally deeply cleft portion (Fig. 401) much

longer, tapered. No evident cerci.

Notes on biology. — Specimens of this species are found near sedge lakes, ponds, and

sloughs. The flight period extends from July 5 to September 10.

Geographical distribution. — The known range of this species is restricted largely to the

continental divide area of Alberta and British Columbia (Fig. 629). In altitude it ranges from

about 3,700’ to 5,350’.

I have examined 28 specimens, eight males and 20 females, from the study area.

Rhyacophilidae and Limnephilidae

115

The asiaticus group

Males of this group are characterised by presence of postero-dorsal lobe on tergum VIII;

by large, smooth tooth on dorso-mesal edge of cercus; and by wide-based claspers, with

short, stout dorsal processes (Fig. 402). Females are distinguished by massive segment IX of

single piece, without ventro-lateral lobes; and by massive, short lobed vulval scale (Fig. 406,

407).

Limnephilus labus Ross, 1941

(Fig. 402-407, 630)

Limnephilus labus Ross, 1941a: 105-106. (Type locality: Wood Lake, Boulder, Colorado).

Ross, 1944:298. Ross and Merkley, 1952:436, 451. Flint, 1960:5. Fischer, 1968:200.

Limnophilus labus\ Schmid, 1955:141.

Description. — Antennae reddish brown, scapes darker. Vertex of head dark brown

between ocelli and posterad to posterior edge of cranium. Thorax reddish brown; brush

of male fore leg composed of dense, stout, black setae along entire length of femur.

Spurs reddish. Male fore wing length 9.5 mm; grey-brown, with darker areas close to

posterior edge. Costal area clear to distal end of subcosta. Venation identical to that of

L. sublunatus.

Male genitalia. (Specimen from Waterton, Alberta). Postero-dorsal edge of tergum VIII

with small, thin, spinate lobe. Dorsal strap of segment IX short, wide (Fig. 402); main body

of segment high, narrow, with anterior lobes; with short, blunt peg at postero-dorsal angles.

Clasper with rectangular dorsal process and high, narrow base. Median lobes of segment X

with broad bases tapered abruptly to small, acuminate postero-dorsal hooks. Cercus with

very concave mesal face, with dorsal edge arched meso-ventrad (Fig. 403); with large,

smooth, black spine. Median shaft of aedeagus long, irregular, curved dorsad (Fig. 404);

lateral arms long, each curved sharply dorsad at mid-point, with expanded, acuminate distal

lobe fringed ventrally with long setae (Fig. 404, 405).

Female genitalia. (Specimen from Waterton, Alberta). Vulval scale with massive, short

lobes (Fig. 407); lateral lobes rectangular; median lobe very broad, blunt, with very little

space between it and lateral lobes. Segment IX massive, of one rectangular piece (Fig. 406);

segment X completely enclosed. Supra-genital plate very wide, short. Segment X distinct

from segment IX; dorsal surface steep, terminated in two small, black-tipped dorsal lobes.

Cerci large, stout, attached to dorsal surface of segment X; short, blunt.

Notes on biology. — Adults of this species are found near sedge or cattail ponds, lakes or

sloughs. A very few (three) records indicate that this species may also inhabit rivers and

smaller creeks. The flight season extends from July 25 to September 13, with a possible

peak in late August.

Geographical distribution. — The known range of this species extends from Alberta and

Saskatchewan to Idaho and Colorado (Fig. 630). In Alberta it is confined to the southern

and central plains and lower foothills.

I have examined 40 specimens, 23 males and 17 females, from the study area.

The fenestratus group

Diagnostic genitalic characters are not available because of a too great diversity. Two

species of the group are known from the study area.

116

Nimmo

Limnephilus minus cuius (Banks), 1907

(Fig. 408-413, 631)

Stenophylax minusculus Banks, 1907b: 120. (Type locality: Olympia, Washington). Banks,

1907a:39.

Apolopsyche minusculus ; Banks, 1916:121. Betten, 1934:370.

Algonquina minuscula ; Milne, 1935:31, 49.

Limnephilus minusculus ; Ross, 1938b:37. Ross, 1944:298. Ross and Merkley, 1952:445.

Fischer, 1968:234. Clifford, 1969:582. Lindroth and Ball, 1969:138.

Limnophilus minusculus’, Schmid, 1955:139.

Males of this species are distinguished from males of other species of Limnephilus by

presence of postero-dorsal lobe of tergum VIII; by warped cerci with mesal projection and

cross-axial ventral ridge (Fig. 408, 409); and by form of aedeagus. especially lateral arms

(Fig. 410). Females are distinguished by single unit segment IX; by discrete segment X; and

by double-lunate groove ventrad of segment X as seen in ventral aspect (Fig. 412, 413).

Description. — Antennae very dark brown; scapes darker. Vertex of head black. Thorax

very dark brown; brush of male fore leg sparse, only on basal third of femur. Spurs dark

yellow. Fore wing length of male 8.4 mm; pale chocolate-brown; costal area clear. Venation

identical to that of L. sublunatus.

Male genitalia. (Specimen from Flatbush, Alberta). Postero-dorsal edge of tergum VIII

with prominent, lightly spinate lobe. Dorsal strap of segment small, relatively narrowed,

especially ventrally. Clasper short, with very short base; tapered evenly to rounded tip;

directed postero-dorsad. Median lobes of segment X small, thin plates, with black tips; with

wide bases, narrowed abruptly to small dorsal hooks; basal plates square, concave (Fig. 409).

Cercus small, slender in lateral aspect, with tip at right angles, directed mesad, with black,

dentate ventral edge (Fig. 409). Median shaft of aedeagus short, stout, tapered, with slightly

dilated tip (Fig. 410, 411); lateral arms long, with very slender, sclerotized basal halves;

distal halves membranous, extensible, with zigzag at bases; fringed distally, mostly on dorsal

edges, with long setae (Fig. 410).

Female genitalia. (Specimen from Flatbush, Alberta). Vulval scale large, with short, stout

median lobe, rectangular lateral lobes with groove along each median face (Fig. 413). Seg-

ment IX of one piece, roughly rectangular (Fig. 412); with slight postero-ventral lobes.

Supra-genital plate membranous, wide, short, rectangular except for posteriorly indented

edge. Segment X very small, discrete from segment IX in lateral aspect; tubular, short. Cerci

short, dorso-ventrally flattened, rounded in ventral aspect; fused solidly to segment X.

Notes on biology. — Adults of this species are found associated with sedge or cattail

reed sloughs or small lakes, and small streams or rivers on gravel beds. The flight season

extends from May 25 to July 16, with peak in late June and early July.

Geographical distribution. — The known range of this species extends from Alaska to

Washington, Colorado, New York, and Labrador (Fig. 631). In Alberta it is confined to

the plains, low foothills, and major, low mountain valleys, up to about 4,500’ altitude.

I have examined 106 specimens, 93 males and 13 females, from the study area.

Limnephilus kennicotti Banks, 1920

(Fig. 414-420, 632)

Limnephilus kennicotti Banks, 1920:344. (Type locality: Great Slave Lake, Northwest

Territories). Betten, 1934:336. Milne, 1935:44, 5 1 . Milne (D. J.), 1943:298. Ross, 1944:

298. Ross and Merkley, 1952:441. Ross, 1953: 155. Wiggins, 1961 : 70 1 . Ross, 1965:593.

Rhyacophilidae and Limnephilidae

117

Fischer, 1968:199.

Limnophilus kennicotti; Schmid, 1955: 139.

Males of this species are distinguished by massive claspers with very long bases (Fig. 414).

mesally directed, thin, body (Fig. 418), and black-toothed dorsal edges. Females are distin-

guished by massive, mesally cleft dorsal body of segments IX and X combined (Fig. 419,

420).

Description. — Antennae dark brown; scapes darker, with pale, setaless, antero-mesal

faces. Vertex of head very dark brown, warts paler. Thorax very dark brown, warts paler.

Spurs brownish yellow. Fore wing length of male 8.9 mm; light brown; costal margin clear

to end of subcosta; clear areas white. Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from Canmore, Alberta). Dorsal strap of segment IX equal in

height to main body; wide. Main body of segment very wide dorsally, narrowed ventrad

(Fig. 414); with slight concavities at postero-ventral margins. Clasper massive, thin, directed

mesad; with dorsal edges black-toothed (Fig. 418). Median lobes of segment X large, long,

with wide bases tapered distad to rounded tips; very wide in dorsal aspect, with low, black-

toothed ridge just basad of tips (Fig. 415). Cercus large, long, arched dorsad in lateral aspect;

with slight declivity on mesal face, produced mesad as triangular ledge (Fig. 415). Median

shaft of aedeagus long, slender; lateral arms shorter, slender, sclerotized, attached to dorsal

surface of aedeagus; each with cluster of three spines distally, spines crossed over dorsal

surface of aedeagus (Fig. 416, 417).

Female genitalia. (Specimen from Lethbridge, Alberta). Vulval scale small, with short,

distally rounded median lobe (Fig. 420); lateral lobes very small, irregular, narrow, directed

postero-laterad. Segment IX with discrete ventro-lateral lobes (Fig. 429); dorsal body in-

distinguishable from segment X. Supra-genital plate large, rectangular, visor-like in lateral

aspect. Segment X massive, solidly fused to segment IX; rectangular in lateral aspect, with

meso-dorsal hooks distally; in ventral aspect of two large, quite separate plates (Fig. 420).

Notes on biology. - Adults of this species are found near lakes and rivers. The flight

period extends from June 24 to October 3, with a possible peak in August and September.

Geographical distribution. — The known range of this species extends from British Co-

lumbia to Newfoundland, and north to Greenland (Fig. 632). In Alberta it is found both on

the plains, and in the foothills and mountains, to altitudes of about 6,675’.

I have examined 45 specimens, 10 males and 35 females, from the study area.

The nigriceps group

Males of the single species of this group are characterised by massive segment IX with

short, narrow, dorsal strap (Fig. 421); by massive claspers with large, heavily sclerotized,

black, dorsal tooth; and by small, light-coloured median lobes of segment X (Fig. 422).

Females are characterised by discrete ventro-lateral lobes of segment IX (Fig. 426); and by

solid, large segment X and dorsal body of segment IX fused.

Limnephilus nigriceps (Zetterstedt), 1840

(Fig. 140a, 140b, 421-426,633)

Phryganea nigriceps Zetterstedt, 1840: 1066. (Type locality: Lapland).

Limnophilus nigriceps', Ross and Merkley, 1952:443, 453. Schmid, 1955:142. (See Fischer,

1968:237, 245, for Palaearctic literature).

Limnephilus nigriceps', Betten, 1934:56, 87. Fischer, 1968:237-246. (See Fischer, 1968:

238, for Palaearctic literature).

118

Nimmo

Limnophilus affinis Hagen (not Curtis), 1858. (See Fischer, 1968:245, for Palaearctic liter-

ature).

Limnephilus affinis ; Fischer, 1968:245.

Pryganea pilosula (See Fischer, 1968:245).

Limnephilus pilosulus', (See Fischer, 1968:245).

Chaetotaulius striola Kolenati. (See Fischer, 1968:245, 246, for Palaearctic literature).

Limnophilus striola', (See Fischer, 1968:246, for Palaearctic literature).

Limnephilus striola ; (See Fischer, 1968:245-246, for Palaearctic literature).

Limnephilus forcipatus Banks, 1924:439-440. Criddle, 1925:16. Betten, 1934:336. Milne,

1935:46, 51. Milne (D. J.), 1943:192, 194, 195. Ross, 1944:298. Ross and Merkley,

1952:443. Fischer, 1968:139.

Description. — Antennae dark brown; scapes black in males, pale yellow in females;

antero-mesal faces paler, setaless. Vertex of head black except for paler warts. Thorax deep

black; brush of male fore leg composed of short, stout, black spines on basal half of femur.

Spurs yellow. Fore wing length of male 1 1.6 mm; pale brown; anterior half of wing clear,

posterior half irrorate. Venation of fore and hind wings as in Fig. 140a, 140b; R2 of hind

wing with row of short, black hairs on ventral surface.

Male genitalia. (Specimen from Exshaw, Alberta). Postero-dorsal edge of tergum VIII

lightly setose. Dorsal strap of segment IX short, curved posterad (Fig. 421). Main body of

segment massive, widened ventrad, with postero-dorsal angle rounded, large. Clasper massive,

short, wide, with large, black, heavily sclerotized disto-dorsal tooth. Median lobes of seg-

ment X small, directed dorso-posterad, with slight lateral hooks at tips (Fig. 422). Cercus

large, parabolic, with large, black, heavily sclerotized disto-mesal tooth (Fig. 422, 423).

Lateral arms of aedeagus long, slender, sclerotized, spinate distally, curved sharply dorsad

(Fig. 424); large, wing-like, spinate, lobe on disto-ventral margin.

Female genitalia. (Specimen from Exshaw, Alberta). Vulval scale relatively small, with

short, slender, tapered median lobe (Fig. 425); lateral lobes large, roughly triangular. Ventro-

lateral lobes of segment IX discrete, large, irregular in outline (Fig. 426). Dorsal body of

segment indistinguishable from massive segment X. Supra-genital plate triangular, large. No

evident cerci.

Notes on biology. — Adults of this species are found near small sedge ponds or sloughs.

The flight season is from August 23 to September 22.

Geographical distribution. — The nearctic range of this holarctic species extends from

Alaska to Alberta and Manitoba (Fig. 633). In Alberta I have collected specimens in the

foothills and mountains, in the low valleys.

I have examined 20 specimens, 15 males and five females, from the study area.

The rhombicus group

Males of this group are characterised by postero-dorsal lobe on tergum VIII; by large,

curved claspers with black, heavily sclerotized distal teeth; and by relatively small median

lobes of segment X. Females are characterised by long distinct cerci; discrete ventro-lateral

lobes of segment IX; and by separate segment X.

Limnephilus rhombicus (L.), 1758

(Fig. 427-432, 634)

Phryganea rhombica Linnaeus, 1758:548. (Type locality: Sweden). (See Fischer, 1968:

270-294, for Palaearctic literature).

Rhyacophilidae and Limnephilidae

119

Phryganea rhomboides', (See Fischer, 1968:271).

Friganea rhomboea', (See Fischer, 1968:271).

Chaetotaulius rhombicus ; (See Fischer, 1968:272-273, for Palaearctic literature).

Limnephila rhombica ; (See Fischer, 1968:272).

Limnophila rhombica ; (See Fischer, 1968:282, 283).

Leimnephila rhombica ; (See Fischer, 1968:274).

Limnophilus rhombicus', Hagen, 1861:254. Kolbe, 1912:41. Sibley, 1926:107, 191, 194.

Ulmer, 1932:212. Betten and Mosely, 1940:132-134. Schmid, 1955:134. (See Fischer,

1968:270-294, for Palaearctic literature).

Limnephilus rhomnicus; (See Fischer, 1968:278).

Limnephilus rhombicus ; Walker, 1852:22. Vorhies, 1909:661-666. Betten, 1926:529. Raw-

son, 1930:46. Betten, 1934:331-333. Ross, 1944:8, 14, 185, 186, 189, 190, 198. Leon-

ard and Leonard, 1949a: 19. Flint, 1960:5, 49. Robert, 1960:59. Wiggins, 1961:701.

Denning, 1965b:698. Etnier, 1965:149. Fischer, 1968:270-294. (See Fischer, 1968:270-

294, for Palaearctic literature).

Goniotaulius combinatus Walker, 1852:28-29. (Type locality: St. Martin’s Falls, Albany

River, Hudson’s Bay). Hagen, 1864:815, 834. Banks, 1892:363. Banks, 1899:208. Banks,

1907a:36. Ulmer, 1907a:44. Lloyd, 1915:203-205. Johnson, 1927:49. Brues, 1930:393.

Banks, 1930b: 128. Milne, 1935:44, 47, 51. Elkins, 1936:668-669. Betten and Mosely,

1940:132, 134. Ross, 1944:190, 298. Proctor, 1946:211. Fischer, 1968:294.

Limnophilus combinatus', Hagen, 1861:255. McLachlan, 1875:50. Ulmer, 1905a: 19. Lloyd,

1921:44-47. Muttkowski and Smith, 1929:259. Balduf, 1939:172, 174, 178. Pennak,

1953: Fig. 361b, 36 Id. Fischer, 1968:294.

Description. — Antennae yellow-brown. Vertex of head uniformly deep red-brown. Tho-

rax uniform reddish brown; brush of male fore leg composed of fine, reddish hairs along

basal half of anterior face of femur. Fore wing length of male 17 mm; reddish brown, partly

irrorate, with large, irregular, hyaline areas. Vein A3 of fore wing dark brown, with heavy

vestiture of stout, brown setae. Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from 2 miles west of Lake Louise, Alberta). Postero-dorsal

edge of tergum VIII extended sharply ventrad as rectangular, slightly bulbous, black-spinate

screen (Fig. 427, 428). Dorsal strap of segment low, wide vertically; main body of segment

stout, narrowed ventrad. Clasper small, with high, narrow base; dorsal process triangular.

Median lobes of segment X small, short, black, with square-cut tips directed postero-laterad.

Cercus massive, long, arched dorsad, with parallel sides; distal edge straight in lateral aspect,

with ventral edge black, heavily sclerotized, dentate (Fig. 428). Lateral arms attached dorsal-

ly on aedeagal base; distally bifid; ventral lobe large, flap-like, fringed dorsally (Fig. 429)

with long spines; dorso-median lobes folded mesad, armed with several stout, black, distal

teeth (Fig. 429, 430).

Female genitalia. (Specimen from Whitehorse. Yukon; in Canadian National Collection,

Ottawa). Vulval scale large, deeply imbedded in sternum VIII (Fig. 432); median lobe short,

truncate-triangular; lateral lobes clavate in ventral aspect. Ventro-lateral lobes of segment IX

large, roughly triangular in lateral aspect (Fig. 431), discrete from dorsal body of segment;

dorsal body small, triangular in dorsal aspect. Supra-genital plate large, semi-circular, hya-

line. Segment X separable from segment IX; long, tubular, shallowly cleft mesally; with

shallow groove on dorso-lateral surfaces, ventrad of cereal bases. Cerci long, with wide bases

each abruptly tapered at mid-point.

Notes on biology. — The single male specimen examined was probably taken in the sedge

sloughs just west of Lake Louise townsite, in Banff National Park. Date of capture was July

20.

120

Nimmo

Geographical distribution. — The nearctic range of this holarctic species extends from

Yukon Territory to Colorado, Illinois, Newfoundland and Greenland (Fig. 634).

The diver sus group

Males of this group are characterised by black-spinate, postero-dorsal lobes of tergum

VIII; by short, stout, triangular, thick cerci and median lobes of segment X; and by wide-

based claspers with long, distally heavily sclerotized, black-toothed tips (Fig. 433). Females

are characterised by discrete, ventro-lateral lobes of segment X; and by segment X separate

from segment IX (Fig. 438).

Limnephilus canadensis Banks, 1908

(Fig. 433-438, 635)

Limnephilus canadensis Banks, 1908a:264. (Type locality: Laval County, Canada). Betten,

1934:321. Ross, 1938b: 34. Denning, 1941b:201-202. Milne (D. J.), 1943:192, 193, 194,

195. Ross, 1944:299. Ross and Merkley, 1952:450. Flint, 1960:5. Etnier, 1965:148.

Fischer, 1968:53. Clifford, 1969:582.

Anabolina canadensis’, Milne, 1935:41, 49. Denning, 1937:23.

Limnophilus canadensis', Schmid, 1955:142.

Description. — Antennae dark brown; scapes darker, with antero-mesal faces yellow,

glabrous. Vertex of head dark brown, faded to brownish yellow laterad of lateral ocelli.

Thorax dark brown; brush of male fore leg along entire length of femur; anterior tibiae with

similar brushes on opposing faces. Spurs yellow, to brown. Fore wing length of male 7.0

mm; light reddish brown; costal area clear to end of subcosta; clear areas white; dark areas

irrorate. Venation identical to that of L. sublunatus.

Male genitalia. (Specimen from Nordegg, Alberta). Postero-dorsal edge of tergum VIII

with slender, prominent, black-spinate lobe. Dorsal strap of segment IX short, sharp edged

dorsally (Fig. 433); main body of segment wide, narrowed ventrad, with sinuate dorsal

edges. Clasper with high, narrow base; dorsal process long, directed dorsad, with distal edge

black, heavily sclerotized, with two distinct teeth. Median lobes of segment X small, short,

triangular, wide (Fig. 433, 434). Cercus similar, with dorsal angle setose, with long setae.

Median shaft of aedeagus long, tapered, with distinct head; lateral arms shorter, expanded

distally, acuminate, with disto-ventral edges fringed with short setae (Fig. 435, 436).

Female genitalia. (Specimen from Athabasca River, Grosmont, Alberta). Vulval scale rec-

tangular (Fig. 437); median lobe short, tapered from wide base; lateral lobes longer, rectan-

gular, each with slight distal widening. Dorsal body of segment IX small, high, narrow, with

postero-dorsal process (Fig. 438); ventro-lateral lobes discrete, large, irregular. Supra-genital

plate roughly rectangular, with squat triangular distal edge. Segment X large, tubular, coni-

cal in ventral aspect (Fig. 437), black disto-dorsally. No evident cerci.

Notes on biology. — Adults of this species are found near lakes, and slower, smoothly

flowing rivers. The flight period extends from June 9 to August 12.

Geographical distribution. — The known range of this species extends from Alberta to

Maine (Fig. 635). In Alberta it is confined to the plains or low foothills.

I have examined 12 specimens, five males and seven females, from the study area.

The Genus Clistoronia Banks

One species of this genus is known from the study area; it belongs to the subgenus

Rhyacophilidae and Limnephilidae

121

Clistoronia Banks.

Synopsis of characters. — Cephalic macrochaetae short, fine, sparse; pronotal macrochae-

tae very strong, abundant. Spur formula 1,3,3. Wings large; fore wings clearly widened at

chord, truncated apically; hind wings rather narrow. Fore wing coloration varied, quite

patchy. Fore wing venation much as in Limnephilus spp. but chord weakly disrupted,

almost straight, oblique anteriorly; f3 narrowed basally. Hind wing chord strongly disrupted,

distinctly concave to body.

Male genitalia with tergum VIII non-spinate. Segment IX very short, high (Fig. 439);

postero-lateral angles prominent, support to segment X mesally. Cerci large, strong, heavily

sclerotized; in lateral aspect triangular, bilobed distally. Median lobes of segment X spur-

like, horizontal. Clasper weakly developed; distal portion long, conical, simple, directed

mesad. Aedeagus large; median shaft much folded basally (Fig. 442); lateral arms spinate,

each simple or bilobed.

Female genitalia with dorsal body of segment IX vestigial. Ventro-lateral lobes of segment

IX quite large, convex plates, in close contact ventrally (Fig. 443, 444). Supra-genital plate

large, ogival, prominent. Segment X varied, short, long, slender. Cerci or cercus-like lobes

large, strongly flattened dorso-ventrally, fused to segment IX, or segment X, which is imme-

diately ventrad. Median lobe of vulval scale varied in size; lateral lobes sub-quadrangular or

rounded (Fig. 444).

Qistoronia magnifica (Banks), 1899

(Fig. 439-444, 635)

Halesus magnifica Banks, 1899:209. (Type locality: Olympia, Washington). Banks, 1907a:

38.

Halesus magnificus’, Ulmer, 1905a:21. Ulmer, 1907a:56. Essig, 1926:176.

Clistoronia magnus\ Banks, 1916: 1 19.

Qistoronia magnifica, Betten, 1934:371. Banks, 1943:343. Ross, 1944:297. Ross and Merk-

ley, 1952:436. Schmid and Guppy, 1952:42. Schmid, 1955:156. Fischer, 1969:55-56.

Stenophylax magnificus’, Milne, 1935:32, 52.

Limnephilus magnificus ; Ross and Merkley, 1952:441 , 45 1 .

Anabolia caroli Denning, 1941b: 196-197. (Type locality: Robson, British Columbia).

Limnephilus caroli ; Ross, 1944:298.

Males of this species are distinguished from males of other species of Limnephilidae by

large, triangular cerci (Fig. 439); by fused median lobes of segment X (Fig. 440); and by

mesally directed black teeth on mesal faces of cerci. Females are distinguished by long, thin,

acuminate dorsal lobes of segment X, ventro-mesad of cerci (Fig. 443).

Description. — Antennae yellow-brown; antero-mesal faces of scapes with few setae.

Vertex of head deep red-brown. Thorax deep reddish brown. Fore wing length of male 18.1

mm; light chocolate-brown, irrorate, with larger hyaline areas. Venation identical to that of

Limnephilus spp.

Male genitalia. (Specimen from Waterton, Alberta). Dorsal strap of segment IX very slim,

incomplete dorsally (Fig. 439). Main body of segment vertically narrow, moderately ex-

panded laterally, with postero-dorsal angles produced mesad as long, narrow processes. Clas-

per with narrow base, dorsal process long, tapered, acute-triangular in lateral aspect. Median

lobes of segment X fused throughout length (Fig. 440), conical, rounded. Cercus massive,

thick, triangular in lateral aspect, with short ventral tooth; mesal face with black, heavily

sclerotized tooth. Median shaft of aedeagus short, stout, curved gently dorsad (Fig. 442);

lateral arms parallel to median shaft, greatly widened distally; each distally bilobed, with

122

Nimmo

dorsal lobe longer, narrower, more acuminate than ventral. Both lobes fringed with setae.

Female genitalia. (Specimen from Waterton, Alberta). Vulval scale with squat, conical

median lobe; lateral lobes triangular distally, well clear of median lobe (Fig. 444). Dorsal

body of segment IX small, fused to segment X (Fig. 443); ventro-lateral lobes large, triangu-

lar in lateral aspect, fused ventrad of segment X; not discrete from dorsal body. Supra-

genital plate small, semi-circular. Segment X large, tubular, deeply cleft laterally, not so

deeply in vertical plane; dorsal lobes long, narrow; ventral lobes shorter, wide, darker. Cerci

short, acuminate in lateral aspect, wide in ventral aspect.

Notes on biology. — Adults of this species are collected near sedge lakes or ponds.

The flight period extends from June 11 to August 16, with a peak in mid-July.

Geographical distribution. — The known range of this species extends from Alberta and

British Columbia to Utah (Fig. 635). In Alberta it is confined strictly to the area of the

continental divide, but in the lower major valleys.

I have examined 17 specimens, 13 males and four females, from the study area.

The Genus Grammotaulius Kolenati

This genus is represented in the study area by a single species.

Synopsis of characters. — Head somewhat elongated, eyes small. Posterior cephalic warts

very large, oval. Posterior ocelli surrounded by many hairs. Spur formula 1,3,4. Pronotum at

least half as long as head. Wings large; fore wings uniformly, or patchily coloured. R5 of fore

wing darkened in some species; always so on hind wing. Chord of fore wings very oblique

anteriorly to body, not strongly disrupted. Hind wing chord parallel to body, strongly

disrupted.

Male genitalia rigid, very large. Tergum VIII not spinate. Segment IX long longitudinally

and vertically. Cerci very thick, to very large, slender; in some species short, massive, thick;

dentate. Claspers poorly developed, not projected. Aedeagus very thick, similar to that of

Limnephilus', lateral arms not erectile or setose, but simple or bilobed.

Female genitalia without discrete ventro-lateral lobes; dorsal body of segment IX narrow,

elongated, with two free appendages. Segment X tubular, very large, short, sclerotized,

barely cleft mesally. Supra-genital plate small. Vulval scale large, with three large, sub-equal,

projected lobes in most species.

Grammotaulius in terregationis (Zetterstedt), 1840

(Fig. 445-450, 636)

Phryganea interregationis Zetterstedt, 1840:1063-1064. (Type locality: Greenland). Kolen-

ati, 1848:25, 26, 30, 40-41. (See Fischer, 1967:212-213, for Palaearctic literature).

Limnephilus interregationis', Walker, 1852:19.

Limnophilus interregationis', Hagen, 1861:254-255. (See Fischer, 1967:213, for Palaearctic

literature).

Grammotaulius interregationis', Banks, 1892:363. Banks, 1907a:36. Ulmer, 1907a:38. Mose-

ly, 1929:502, 507, 508-509. Ulmer, 1932:212, 217. Betten, 1934:316. Milne, 1935:27,

50. Denning, 1941a:232-233. Ross, 1944:185, 297. Schmid, 1950a:348-35 1 . Ross and

Spencer, 1952:47. Schmid, 1955:145. Schmid, 1964:914, 917.Etnier, 1965: 148. Fisch-

er, 1967:212-214. Clifford, 1969:582. (See Fischer, 1967:213, for Palaearctic literature).

Grammotaulius praecox Hagen, 1873c:45 1-452. (Type locality: British America). Banks,

1892:363. Jacobson, 1898:215. Ulmer, 1905a: 18. Ulmer, 1907a:39. Banks, 1907a:36.

Kolbe, 1912:42. Betten, 1934:316. Milne, 1935:50. Ross, 1938b: 32. Fischer, 1967:214.

Rhyacophilidae and Limnephilidae

123

Limnophilus praecox ; Ross, 1941a: 1 10.

Grammotaulius sibiricus Mosely, 1929:502, 507.

Description. — Antennae light brown; scapes cream-white with glabrous antero-mesal

faces. Vertex of head with black mesal diamond limited by lateral ocelli and median warts;

remainder cream-white. Thorax black, shaded in areas to dark brown. Spurs brown. Fore

wing length of male 20.9 mm; chocolate-brown to light reddish brown. Hind wing stigma

light brown; R5 set in light brown band. Venation essentially identical to that of Limne-

philus sublunatus.

Male genitalia. (Specimen from Wapta Lake, Kicking Horse Pass, Yoho National Park,

British Columbia). Segment IX massive, rectangular; dorsal strap short, narrow to point of

extinction dorsally (Fig. 445). Clasper small, blunt, short. Median lobes of segment X very

small, short, with meso-dorsal black edges (Fig. 446). Cercus huge, thick, armed with heavily

sclerotized teeth along distal edge; with ventral, black tooth directed postero-mesad. Median

shaft of aedeagus simple. Lateral arms long, expanded slightly distally, each with two distal

lobes (Fig. 447, 448); basal or dorsal lobe minute, with distal tuft of spines; ventral lobe

large, lanceolate, fringed dorsally with short spines.

Female genitalia. (Specimen from Banff, Alberta). Vulval scale massive, triangular (Fig.

450); median lobe short, parallel-sided, lateral lobes with narrow bases, widened distally.

Segment IX large, inseparably fused to segment X dorsally; ventral lobes much larger, with

straight edges (Fig. 449). Supra-genital plate short, broad, with evenly curved posterior

edge. Segment X relatively small, short, with pair of meso-lateral lobes. No evident cerci.

Notes on biology. — Adults of this species are found near sedge ponds, lakes or sloughs,

and mountain streams of a turbulent and rocky nature. Flight season of the adults extends

from July 4 to September 27.

Geographical distribution. — The nearctic range of this holarctic species extends from

Yukon Territory to British Columbia, Minnesota, Nova Scotia, and Greenland (Fig. 636). In

Alberta it is confined to the foothills and mountains, ranging in altitude from 4,500’ to

6,800’.

I have examined 25 specimens, 13 males and 12 females, from the study area.

The Genus Nemotaulius Banks

A single species, of the subgenus Macrotaulius, is known from the study area.

Synopsis of characters. — Eyes quite small. Head quite elongate, with toothed ridge along

occipital border. No cephalic warts present. Pronotum twice length of head in certain spe-

cies. Spur formula 1,3,4. Fore wings coriaceous, apically indented (Fig. 141a). Hind wings

very large, slightly indented. Fore wings very reddened, not irrorate. Venation of fore wings

with very large discoidal cell; chord feebly disrupted, hardly oblique to body; hind wings

similar.

Male genitalia with postero-dorsal edge of tergum VIII developed as large, black-spinate

bulb. Segment IX usually large, well developed laterally. Claspers of medium size, massive in

appearance, thick, concave, unarmed, not close to border of segment IX. Aedeagus large,

strong, similar to that of Limnephilus spp.; lateral arms non-erectile, simple, fringed distally

with setae or spines.

Female genitalia with dorsal part of segment IX narrowed, triangular, weakly developed;

no free appendages. Segment X continuous with segment IX, tubular; short, massive, poorly

cleft mesally. Ventro-lateral lobes of segment IX huge, largely continguous ventrally. Supra-

genital plate small. Vulval scale very large, plate-like, entirely flanked by sternum VIII;

median lobe very small, sunk between two sub-quadrangular lateral lobes.

124

Nimmo

Nemotaulius hostilis (Hagen), 1873

(Fig. 141a, 141b, 451-455,637)

Glyphotaelius hostilis Hagen, 1873c:444-446. (Type locality: Saskatchewan). Banks, 1892:

363. Ulmer, 1905a: 18. Ulmer, 1907a:40. Banks, 1907a:36. Lloyd, 1921:41, 43-44.

Dodds and Hisaw, 1925a: 124-125. Dodds and Hisaw, 1925b:386. Betten, 1926:529.

Sibley, 1926:107, 193. Berry, 1927:3. Banks, 1930b: 127. Betten, 1934:313. Milne,

1935:28, 50. Ross, 1938b:32. Balduf, 1939:122. Ross, 1941a: 1 13. Ross, 1944:183, 297.

Leonard and Leonard, 1949a: 16. Ross and Spencer, 1952:47. Morse and Blickle, 1953:

97. Pennak, 1953:580. Robert, 1960:59. Wiggins, 1961:700. Etnier, 1965:148.

Nemotaulius ( Macro taulius) hostilis; Schmid, 1952a:229-23 1 . Schmid, 1955:149. Flint,

1960:5, 54, 55-56. Nimmo, 1966a:691. Fischer, 1967:250-251. Clifford, 1969:583.

Lindroth and Ball, 1969:138.

Description. — Antennae brown; scapes darker, with antero-mesal faces pale, almost

glabrous. Vertex of head deep red-brown; flat, evenly covered with large pustulate bumps.

Thorax pale yellow, with slightly darker areas laterally; dorsal areas flat, pustulate. Spurs

yellow. Fore wing length of male 22.8 mm; grey-brown to almost black in mixture of solid

blocks of colour, and irrorate areas. Costal area hyaline except for slight basal irroration.

Venation as in Fig. 141a, 141b; distal edge of fore wing crenulate.

Male genitalia. (Specimen from George Lake, near Busby, Alberta). Postero-dorsal edge

of tergum VIII developed as large, black-spinate bulb. Dorsal strap of segment IX short;

complex, of sclerotized flanges merged gradually with main body (Fig. 451). Main body of

segment high, spindle-shaped, not wide. Clasper with long, narrow base; dorsal process

massive, short, thick; claw-like distally. Median lobes of segment X small, short, dorso-

laterally hooked spines (Fig. 451, 452). Cercus small, wide, lobe without mesally concave

face. Aedeagus large, with simple median shaft (Fig. 453); lateral arms long, with wide

bases and distal tips; tips acuminate, fringed on edges with long setae.

Female genitalia. (Specimen from George Lake, near Busby, Alberta). Vulval scale rela-

tively small, with very short, rounded, median lobe, massive, triangular lateral lobes (Fig.

455). Dorsal body of segment IX and segment X solid tapered tube, deeply cleft dorsally

(Fig. 454). Ventro-lateral lobes of segment IX huge, discrete from dorsal body, irregular,

fused solidly ventrally. Supra-genital plate small, triangular, membranous with sclerotized

edges.

Notes on biology. — Adults of this species are found near lakes and sloughs overgrown

with sedges or, particularly, cattail reeds. The flight period extends from June 3 to August

10.

Geographical distribution. — The known range of this species extends from Alaska to

Colorado and Newfoundland (Fig. 637). In Alberta it is confined to the plains region.

I have examined 39 specimens, 28 males and 1 1 females, from the study area.

The Genus Anabolia Stephens

This genus is represented in the study area by three species.

Synopsis of characters. — Cephalic warts small, pronotum somewhat elongate. No dorsal

line. Spur formula 1,3,4. Fore wings large, slightly truncated, rounded, or parabolic apically;

hind wings variable in size. Coloration of wings characteristic, fairly constant; fore wings

reddish brown, with no hyaline or darker areas; uniformly coloured or minutely irrorate.

Venation somewhat variable; chord rectilinear, anteriorly oblique, weakly disrupted, con-

cave to body; hind wing chord posteriorly oblique, disrupted.

Rhyacophilidae and Limnephilidae

125

Male genitalia with tergum VIII non-spinate. Segment IX generally well developed later-

ally, rigid, sclerotized; dorsal strap slender, with blunt elongate, median process in some

species, dentate in most species. Median lobes of segment X more or less triangular, sclero-

tized lamellae. Claspers with slender, pad-like basal part fused to segment IX, sometimes

twisted helically; dorsal process horizontal, slender, cylindrical. Aedeagus large; median

shaft short, thick, folded at base; lateral arms slender, normally bilobed, spinate.

Female genitalia very stubby, not projected. Segment IX of two pieces in lateral aspect;

dorsal part small, with appendages which are free, or fused at base of segment X. Segment X

small, conical, cleft vertically. Ventro-lateral lobes of segment IX very large, massive, con-

tiguous or separated ventrally. Supra-genital plate small. Vulval scale variable.

Key to the Males of species of Anabolia found in Alberta and eastern British Columbia

la. Median shaft of aedeagus very short, stubby, with distal head recessed into basal

part (Fig. 458) A. consocia (Walker), p. 125.

lb. Median shaft of aedeagus long, slender (Fig. 463, 468) 2a

2a.(lb) Clasper with minute dorsal process and high, very slender base (Fig. 461 )

A. ozburni (Milne), p. 126.

2b. Clasper with large, conical, black-tipped dorsal process (Fig. 466)

A. bimaculata (Walker), p. 127.

Key to the Females of species of Anabolia found in Alberta and eastern British Columbia

la. Segment X with cerci or cercus-like lobes; vulval scale with median lobe longer

than lateral lobes (Fig. 459, 460, 469, 470) 2a

lb. Segment X without such lobes; median lobe of vulval scale not longer than lateral

lobes (Fig. 464, 465) A. ozburni (Milne), p. 126.

2a.(la) Ventro-lateral lobes of segment IX massive, fleshy, with triangular posterior edge

(Fig. 459) A. consocia (Walker), p. 125.

2b. Ventro-lateral lobes of segment IX small, roughly rectangular in lateral aspect,

except for triangular postero-ventral lobe projected well posterad under segment

X (Fig. 469) A. bimaculata (Walker), p. 127.

Anabolia consocia (Walker), 1852

(Fig. 456-460, 638)

Limnephilus (Goniotaulius) consocius Walker, 1852:33. (Type locality: North America).

Limnephilus consocius ; Ross, 1944:185, 186, 189, 190-191, 298. Sprules, 1947:30, 78.

Leonard and Leonard, 1949a: 17. Ross and Merkley, 1952:449. Morse and Blickle, 1953:

98. Pennak, 1953: Fig. 361. Etnier, 1965:148. McConnochie and Likens, 1969:150.

Stenophylax consocius ; Hagen, 1859:134. Hagen, 1864:885. Ulmer, 1905a:21.

Goniotaulius consocius-, Hagen, 1864:815.

Anabolia consocia ; Hagen, 1861:264. McLachlan, 1863:157, 161. Hagen, 1864:803. Banks,

1892:363. Banks, 1897:28. Banks, 1905:9. Ulmer, 1905a:20. Schmid, 1950b:3 19-322.

Schmid, 1955:150. Flint, 1960:5, 56, 58. Fischer, 1969:13-15.

Limnephilus consocia', Proctor, 1946:211. Wray, 1950:21.

Colpotaulius consocia; Banks, 1905:9. Banks, 1907a: 37. Martynov, 1914:221. Banks, 1916:

122. Johnson, 1927:49.

Arctoecia consocia-, Ulmer, 1907a:47. Lloyd, 1921:53-56. Banks, 1923:146. Betten, 1926:

528. Sibley, 1926:190. Berry, 1927:3. Banks, 1930b: 127. Ulmer, 1932:214. Betten,

1934:90, 91, 314-315. Milne, 1935:39-49. Balduf, 1939:152. Betten and Mosely, 1940:

126

Nimmo

111-113. Proctor, 1946:210.

Arctocia consocia; Sibley, 1926: 107.

Arctoecia consicia', Muttkowski and Smith, 1929:259.

Colpotaulius medialis Banks, 1905:8. (Type locality: Ontario). Banks, 1907a: 37. Banks,

1908c: 154. Martynov, 1914:221. Brimley, 1938:253.

Arctoecia medialis', Ulmer, 1907a:47. Krafka, 1915: plate 7. Lloyd, 1921:42. Krafka, 1923:

plate 31. Milne, 1935:49.

Limnephilus medialis', Ross, 1938b:36. Ross, 1944: 190, 298.

Males of this species are distinguished from males of other species of the genus by short,

blunt median shaft of aedeagus (Fig. 458) with distal head recessed into basal portion; by

distally flared, black-edged claspers with deeply concave mesal faces (Fig. 456, 457); and by

rectangular median lobes of segment X. Females are distinguished by massive, fleshy, ventro-

lateral lobes of segment IX (Fig. 459), with triangular posterior edge; by minute dorsal body

of segment IX; and by median lobe of vulval scale slender, tapered, much longer than lateral

lobes (Fig. 460).

Description. — Antennae yellowish brown. Vertex of head red-brown, with mesal longi-

tudinal stripe of dark brown. Thorax pale yellow; with mesal longitudinal stripe of dark

brown along terga. Thoracic warts fragmented to individual setal bases in some specimens.

Male anterior femur without brush. Spurs pale yellow. Fore wing length of male 14.1 mm;

light brownish yellow; faintly irrorate basally; veins irregularly darkened. Venation identical

to that of Limnephilus spp.

Male genitalia. (Specimen from Cold Creek, Nojack, Alberta). Dorsal strap of segment IX

with dorso-mesal process lightly setose (Fig. 456). Main body of segment wide dorsally,

tapered gradually ventrad. Clasper small, with high, narrow base; dorsal process short, origi-

nated gradually from base, with distally black, clawed tip. Median lobes of segment X large,

rectangular in lateral aspect, thick in dorsal aspect (Fig. 457), with distal third black; basal

plates large, complexly folded. Cercus massive, distal edge black, with disto-ventral angle

attenuated as black tooth (Fig. 457). Median shaft of aedeagus short, stout, with wrinkled

base (Fig. 458); distal head recessed into basal body. Lateral arms stout, curved dorsad,

tapered slightly distad; distally fringed on dorsal and ventral edges with short setae.

Female genitalia. (Specimen from Chicago, Illinois; in Illinois Natural History Survey).

Vulval scale large; median lobe longer than lateral lobes (Fig. 460), narrow, evenly tapered

distad; lateral lobes curved, widened distad, with concave mesal faces. Dorsal body of seg-

ment IX minute, narrow, strap-like (Fig. 459); ventro-lateral lobes massive, fleshy, with tri-

angular posterior edges, discrete from dorsal body. Supra-genital plate small, wide, tapered

posterad, truncated distally; convex ventrally. Segment X completely cleft ventro-mesally,

dorsally two edges joined by sheet of membrane (Fig. 460); triangular in lateral aspect. Cerci

roughly triangular, held closely to dorso-lateral faces of segment X; fused solidly to segment

X.

Geographical distribution. — The known range of this species extends from Alberta to

Virginia and Quebec (Fig. 638). The two records from Alberta available to me are from the

plains region, at about 2,100’.

I have examined two specimens of this species from the study area, one adult and one

pupal male. I have also examined a female from Illinois.

Anabolia ozbumi (Milne), 1935

(Fig. 142a, 142b, 461-465, 639)

Arctoecia ozbumi Milne, 1935:39, 49. (Type locality: Guelph, Ontario).

Rhyacophilidae and Limnephilidae

127

Anabolia ozburni', Ross, 1944:298. Schmid, 1950b:325-327. Schmid, 1955:151. Nimmo,

1966a:692. Nimmo, 1966b:224. Fischer, 1969:34-35.

Limnephilus ozburni ; Leonard and Leonard, 1949a: 18. Leonard and Leonard, 1949b:6.

Ross and Merkley, 1952:450. Morse and Blickle, 1953:98. Etnier, 1965:149.

Males of this species are distinguished from males of other species of Anabolia by massive,

thick cerci (Fig. 461); by minute dorsal process of claspers; and by dorso-mesal process of

dorsal strap large, projected well posterad (Fig. 462). Females are distinguished by large seg-

ment IX fused solidly to segment X dorsally (Fig. 464); and by lack of cerci or cercus-like

lobes.

Description. — Antennae dark brown; scapes very dark, with antero-mesal faces lighter,

glabrous. Vertex of head very dark brown. Thorax dark to very dark brown. Spurs yellow.

Fore wing length of male 10.8 mm; light reddish brown, irrorate; costal area clear. Venation

as in Fig. 142a, 142b; leading edge of fore wing truncate and R1 straight to edge, not bowed.

Hind wing with deep indentation at Cu.

Male genitalia. (Specimen from Cold Lake, Cold Lake, Alberta). Dorsal strap of segment

IX laterally narrow, dorsally expanded to large triangular, horizontal plate with single seta

in distal clear area. Main body of segment robust, narrowed abruptly ventrad (Fig. 461).

Clasper with high, very narrow base surmounted by minute dorsal process. Median lobes of

segment X massive, triangular, with distal halves black; tips hooked laterad as small teeth

(Fig. 462). Cercus massive, rectangular in lateral aspect, with distal face wide, black, fringed

with setae; attached to wide internal bases. Median shaft of aedeagus long, narrowed slightly

at mid-point (Fig. 463), expanded to folded distal head. Lateral arms each of uniform width

for two-thirds length, then widened slightly to thin, spatulate tip cleft distally as two acumi-

nate spines; dorsal and ventral edges of tips setose; mesal faces with pair of heavy spines.

Female genitalia. (Specimen from Michigan; in Illinois Natural History Survey). Vulval

scale small, with rectangular median lobe (Fig. 465); lateral lobes with concave mesal edges

curved, lateral edges distinctly angular. Dorsal body of segment IX large, wide, fused to seg-

ment X dorsally, but divided from it ventrally by suture (Fig. 464). Ventro-lateral lobes of

segment IX massive, almost square in lateral aspect, discrete from dorsal body of segment.

Supra-genital plate minute, short, not projected laterally beyond distal extremities of lateral

lobes of vulval scale. Segment X cylindrical internally, triangular in lateral aspect, with

dorsal edges of mesal cleft sinuate; median cleft deep, with dorsal cleft angular, ventral cleft

rounded (Fig. 465).

Geographical distribution. — The known range of this species extends from Alberta to

Quebec, New Hampshire and Michigan (Fig. 639). In Alberta it is known only from Edmon-

ton and Cold Lake to the northeast; both are plains localities, and Cold Lake is in the boreal

forest region.

I have examined two males from the study area, and one female from Michigan. Dates of

capture of the Alberta specimens were July 1 1 and 17.

Anabolia bimaculat a (Walker), 1852

(Fig. 466-470, 640)

Limnephilus (Goniotaulius) bimaculatus Walker, 1852:30. (Type locality: St. Martin’s Falls,

Albany River, Hudson’s Bay). Ross, 1941a: 110. Milne (D. J.), 1943:192, 194, 196. Ross,

1944:186, 189, 298. Leonard and Leonard, 1949a: 17. Ross and Spencer, 1952:48. Ross

and Merkley, 1952:449. Morse and Blickle, 1953:98. Pennak, 1953: Fig. 362R. Robert,

1960:59. Etnier, 1965:148.

Limnephilus (Desmotaulius) bimaculatus ; Hagen, 1861 : 263.

128

Nimmo

Anabolia bimaculata', McLachlan, 1863:157, 161. Hagen, 1864:803. Banks, 1892:363.

Banks 1897:28. Ulmer, 1905a:20. Ulmer, 1905b:61-62. Ulmer, 1907a:46. Banks, 1907a:

37. Martynov, 1914:209. Krafka, 1923: plate 8. Dodds and Hisaw, 1925a: 124. Dodds

and Hisaw, 1925b:386. Betten, 1926:529. Banks, 1930b: 128. Ricker, 1932:132. Ricker,

1934:54. Betten, 1934:353-354. Neave, 1934:167. Milne, 1935:43, 49. Denning, 1937:

34-37. Betten and Mosely, 1940:142-144. Denning, 1941b: 195. Schmid, 1950b:88-95.

Schmid, 1955:151. Flint, 1960:5, 56, 57, 114. Fischer, 1969:4-5.

Desmotaulius bimaculatus ; Hagen, 1864:81 1.

Anabolia maculata\ Dodds and Hisaw, 1925a: Fig. 1. Balduf, 1939:121.

Males of this species are distinguished from males of other species of Anabolia in the

study area by conical dorsal processes of claspers with black tips; by dorsal strap of segment

IX without dorso-mesal process; and by small, triangular median lobes of segment X (Fig.

466). Females are distinguished by relatively small ventro-lateral lobes of segment IX with

small, triangular ventro-posterior lobes; by ventro-lateral lobes in contact mesally (Fig. 470);

and by small, low, dorsal body of segment IX (Fig. 469).

Description. — Antennae brown; antero-mesal faces of scapes paler, glabrous. Vertex of

head dark brown. Thorax dark brown dorsally but with paler meso-longitudinal line; lateral-

ly pale yellow. Spurs pale brown. Fore wing length of male 15.8 mm; deep chocolate-brown

to pale yellowish brown; lightly irrorate; costal area clear, pale yellow. Venation not signifi-

cantly different from that of Limnephilus spp.

Male genitalia. (Specimen from George Lake, near Busby, Alberta). Dorsal strap of seg-

ment IX short, slender, directed anterad (Fig. 466). Main body of segment roughly inverted-

triangular. Clasper with long, narrow base; dorsal process short, conical, heavily sclerotized,

black distally. Median lobes of segment X short, triangular, black distally, each with minute

disto-lateral tooth (Fig. 467). Cercus of medium size, rounded-triangular distally; with con-

cave mesal face, thick base. Median shaft of aedeagus slender, tapered evenly distad, with

distinct distal head small (Fig. 468); lateral arms shorter than median shaft, each basally

curved sharply dorsad, divided distally to long, acuminate dorsal lobe and small, triangular

ventral lobe; short setae located only along edge between lobes.

Female genitalia. (Specimen from George Lake, near Busby, Alberta). Vulval scale with

three lobes well separated (Fig. 470); median lobe longer than laterals, conical in ventral

aspect, rugose at base; lateral lobes with channelled mesal faces. Dorsal body of segment IX

small, with short, triangular, dorsal extension; main part of dorsal body fused to segment X

(Fig. 469). Ventro-lateral lobes of segment IX relatively small, rectangular in lateral aspect

except for triangular postero-ventral angles; discrete from dorsal body. Segment X of irregu-

lar outline in lateral aspect; distally acuminate, deeply cleft mesally. Cerci short, fused at

bases to dorso-lateral surfaces of segment X. Supra-genital plate small, rectangular.

Notes on biology. — Adults of this species are found near cattail sloughs, lakes, or ponds,

or smaller, quietly flowing, rivers and creeks. The flight period extends from June 1 1 to

August 29 with a possible peak at end of July and beginning of August.

Geographical distribution. — The known range of this species extends from Great Slave

Lake to British Columbia, New Mexico, Michigan, and Maine (Fig. 640). In Alberta it is

fairly ubiquitous throughout the plains and lower major mountain valleys.

I have examined 417 specimens, 274 males and 143 females, from the study area.

The Genus Asynarchus McLachlan

This genus is represented in the study area by three species belonging to two groups.

Synopsis of characters. — Head slightly elongate; eyes not prominent. Dorsal line barely

Rhyacophilidae and Limnephilidae

129

visible. Spur formula 1,3,4. Pronotum short. Wings of normal size, shortened in female of

some species and cold adapted forms. Fore wings much as in Limnephilus spp. but more ex-

panded at chord; hind wings somewhat larger than fore. Fore wings basically brown, with

small irrorations and larger clear areas at thyridial cells, chord, and distally along M4+5.

Male genitalia with tergum VIII non-spinate. Segment IX laterally strongly convex and

sclerotized. Cerci medium to large, varied; in certain species slender, strongly concave; distal

edge heavily sclerotized, mesal faces with massive tooth or ridge in some species. Median

lobes of segment X small, ventrad of cerci. Claspers with prominent basal pieces fused to

segment IX; distal processes bifid, pincer-like, black. Aedeagus large, quite similar to that of

Limnephilus spp.; lateral arms slender, bilobed.

Female genitalia with dorsal body of segment IX short, inconspicuous; with two large,

laterally flattened, appendages fused basally to each other in some species. Segment X ven-

trad of these appendages; with thin, hardly cleft walls. Ventro-lateral lobes of segment IX

huge, high, massive; in most species contiguous meso-ventrally. Supra-genital plate large.

Vulval scale partly recessed into sternum VIII; lateral lobes long, oblique straps.

Key to the Males of species of Asynarchus found in Alberta and eastern British Columbia

la. Clasper very large, blade-like, triangular (Fig. 471); median lobes of segment X

large, triangular; dorsal strap relatively wide A. mutatus (Hagen), p. 129.

lb. Claspers small, distally bifid (Fig. 476, 482); median lobes of segment X small

hooks; dorsal strap very narrow 2a

2a.(lb) Cercus hooked meso-ventrad distally, with second, ventral, black tooth distinct

(Fig. 482) A. aldinus (Ross), p. 131.

2b. Disto-dorsal angle of cercus triangular in lateral aspect (Fig. 476), not divided

from disto-ventral angle A. curtus (Banks), p. 130.

Key to the Females of species of Asynarchus found in Alberta and eastern British Columbia

la. Segment IX a single, massive unit, grossly enlarge ventrad (Fig. 474)

A. mutatus (Hagen), p. 129.

lb. Segment IX with clear, almost discrete ventro-lateral lobes (Fig. 480)

A. curtus (Banks), p. 130.

A. aldinus (Ross), p. 131.

The contumax group

A single species of this group is known in the study area.

Males of this group are distinguished by short, thin-walled, mesally concave cerci; and by

massive median lobes of segment X. Females are distinguished by very large ventro-lateral

lobes of segment IX; and by narrow, oblique lateral lobes of vulval scale.

Asynarchus mutatus (Hagen), 1861

(Fig. 471-475, 641)

Hallesus mutatus Hagen, 1861:267. (Type locality: Labrador). Hagen, 1864:818. Banks,

1892:364. Ulmer, 1905a:21. Ulmer, 1907a:56. Banks, 1907a:38. Ross, 1938b:32.

Stenophylax mutatus ; Betten, 1934:345.

Anabolia mutata ; Milne, 1935:43, 49.

Limnephilus mutatus ; Ross, 1938b:37-38. Ross, 1944:298. Ross and Merkley, 1952:449.

Etnier, 1965:149.

130

Nirnmo

Asynarchus mutatus ; Schmid, 1954b:65-67. Schmid, 1955:153. Fischer, 1969:50-51.

Males of this species are distinguished from males of other species of Asynarchus by large,

triangular, blade-like claspers (Fig. 471); by high, short, triangular median lobes of segment

X; and by short, irregular cerci. Females are distinguished by massive, unit segment IX (Fig.

474) which completely encircles segment X (Fig. 475).

Description. — Antennae brown; antero-mesal faces of scapes darker, glabrous. Vertex of

head dark brown, warts yellowish. Thorax warm yellowish brown. Brush of fore leg of male

slight, on basal third of femur only. Spurs dark brown. Fore wing length of male 15.0 mm;

dark brownish grey, irrorate, with larger patches of solid colour; two major areas of clear

membrane just distad of chord and immediately anterad of Cul+2; veins of chord darker

than others. Venation identical to that of Limnephilus spp.

Male genitalia. (Specimen from George Lake, near Busby, Alberta). Dorsal strap of seg-

ment IX simple, broad; main body of segment almost elipsoidal. Clasper long, thin, triangu-

lar blade with black tip (Fig. 471). Median lobes of segment X short, with denticulate dorsal

edges; distal areas black, each with distinct disto-lateral tooth (Fig. 472). Cercus short, with

cup-like disto-mesal faces, black, toothed posterior edges. Median shaft of aedeagus tapered

slightly distad; directed postero-dorsad from base (Fig. 473); lateral arms curved in semi-

circle, each with lanceolate, spinate distal lobe; two small, spatulate lobes on ventral edge of

each arm; each lobe distally fringed with stout spines.

Female genitalia. (Specimen from Indian Head, Saskatchewan). Vulval scale with short,

conical median lobe (Fig. 475); lateral lobes long, narrow, tapered, directed postero-laterad,

well separated from median lobe. Segment IX massive, of one piece; dorsal portion slightly

narrowed; ventral portion swollen, fused ventrad of segment X (Fig. 474, 475). Supra-genital

plate very wide, short, hyaline. Segment X very short, tubular, not cleft. Cereal lobes not

evident.

Notes on biology. — Adults of this species are found near sedge and cattail sloughs and

lakes. The flight season extends from June 3 to July 23.

Geographical distribution. — The known range of this species extends from British Colum-

bia to Great Slave Lake, Minnesota and Ontario (Fig. 641). In Alberta it is confined to the

plains and eastern edges of the foothills in the lower valleys.

I have examined 1 1 males of this species from the study area and Great Slave Lake, and

a female from Saskatchewan.

The lapponicus group

Two species of this group are known from the study area.

Males of this group are distinguished by large, projected, lanceolate cerci; and by small

median lobes of segment X. Females are distinguished by discrete ventro-lateral lobes of

segment IX; and by large, triangular lateral lobes of vulval scale.

Asynarchus curtus (Banks), 1920

(Fig. 476-481,642)

Anabolia curta Banks, 1920:345. (Type locality: Massachusetts). Betten, 1934:354. Ross,

1938b:28.

Limnephilus curtus\ Ross, 1938b:35. Ross, 1944:298. Ross, 1947:152. Ross, 1950b:428.

429, Fig. 18, 18a. Ross and Merkley, 1952:443. Etnier, 1965:148. Ross, 1966:592.

Limnephilus curtis\ Milne (D. J.), 1943:192, 194. Morse and Blickle, 1953:58.

Asynarchus curtus\ Schmid, 1954b: 60, 81-84, 85, 86. Schmid, 1955:154. Fischer, 1969:

44-45. Flint, 1960:60, 104, 114.

Rhyacophilidae and Limnephilidae

131

Anabolia planifrons; Milne, 1935:43, 44, 49. Schmid, 1954b:60.

Limnephilus batchewana Denning, 1 949b : 90-9 1 . (Type locality: Batchewana Bay, Lake

Superior). Ross, 1950b:426. Ross and Merkley, 1952:443. Ross and Spencer, 1952:48.

Schmid, 1954b:60. Ross, 1965:592.

Limnephilus conerus Ross, 1950b: 426, 428-429. (Type locality: Thunder Bay, Ontario).

Males of this species are distinguished from males of other species of Asynarchus by short,

distally bifid claspers; by massive, disto-ventrally toothed claspers (Fig. 476); and by uncleft

distal edge of claspers. Females are distinguished by partially separate ventro-lateral lobes of

segment IX (Fig. 480); and by laterally cleft segment X, with wide, triangular dorsal cerci.

Females of this species are, however, inseparable from those of A. aldinus, except by associ-

ation with the male in the field.

The males are of the form batchewana Denning (syn., conerus Ross) and differ consistent-

ly from the eastern curtus as illustrated by Ross (1950b). The synonymy followed here is

that of Schmid (1952c), although Ross and Denning still consider batchewana a distinct

species (in lift.).

Description. — Antennae clothed with black hairs, of overall dark brown colour; scapes

clear yellow, glabrous. Vertex of head dark brown except for narrow area mesad of com-

pound eyes. Thorax reddish brown, with intermixed darker areas. Brush of male fore leg

sparse, pale, confined to basal quarter of femur. Spurs reddish yellow. Fore wing length of

male 14.8 mm; red-brown, irrorate, with very few larger areas clear. Costal area also irrorate.

Venation identical to that of Limnephilus spp.

Male genitalia. (Specimen from Gorge Creek beaver pond, 20 miles west of Turner Valley,

Alberta). Dorsal strap of segment IX short, narrow (Fig. 476). Main body of segment mas-

sive, with narrow sternal area. Clasper short, stout, distally bifid, black. Median lobes of

segment X small, distally hooked dorsad, black. Cercus massive, with heavy longitudinal

ridge along mesal face (Fig. 477); distally black. Median shaft of aedeagus long, curved

dorsad from base (Fig. 478); lateral arms shorter, with disto-dorsal lobe long, scythe-like,

curved dorso-anterad; with small ventral lobe setose (Fig. 479).

Female genitalia. (Specimen from Gorge Creek beaver pond, 20 miles west of Turner

Valley, Alberta). Vulval scale large, with narrow median lobe slightly longer than large, tri-

angular, lateral lobes (Fig. 481). Dorsal body of segment IX small, wide, partially separated

from segment X (Fig. 480); ventro-lateral lobes large, rectangular, partially discrete from

dorsal body. Supra-genital plate wide, incised on posterior edge, partly sunken in ventral

depressions of segment IX. Segment X small, shallowly incised laterally, deeply incised

dorsally; tubular. Cerci massive, set well dorsad of segment X, blunt distally.

Notes on biology. — Adults of this species are found in a wide variety of habitats, from

small plains sloughs and lakes, to high mountain pass bogs, and smaller, quieter creeks. The

flight period extends from July 5 to Sept. 25, with peak about the beginning of September.

Geographical distribution. — The known range of this species extends from Alaska to

Colorado and Labrador (Fig. 642). In Alberta it is found throughout the plains, foothills,

and mountains, to altitudes of around 7,350’.

I have examined 246 specimens, 96 males and 150 females, from the study area and Great

Slave Lake.

Asynarchus aldinus (Ross), 1941

(Fig. 482-483, 643)

Limnephilus aldinus Ross, 1941b: 19. (Type locality: Rowe Lakes, Waterton National Park,

Alberta). Ross, 1944:298. Ross and Merkley, 1952:443.

Asynarchus aldinus', Schmid, 1954b:84-86. Schmid, 1955:154. Nimmo, 1965:786-787.

Fischer, 1969:42.

132

Nimmo

Males of this species are distinguishable from males of A. curtus by lateral aspect of cerci,

with meso-distal angle separated from ventral tooth by lunate cleft, in form of distinct,

postero-ventrally hooked tooth (Fig. 482). Females can only be determined in association

with males.

Description. — Antennae light brown, with yellow, glabrous stripe along antero-mesal

faces of scapes. Vertex of head dark brown, with lighter mesal band joining area between

compound eyes with postero-mesal warts. Thorax warm reddish brown; terga lighter dorsal-

ly. Brush of male fore leg of short, black spines in narrow band on basal third of femur.

Spurs yellow. Fore wing length of male 1 1.2 mm; brownish grey, sparsely irrorate. Venation

identical to that of Limnephilus spp.

Male genitalia. (Specimen from Bow Pass, Banff-Jasper Hwy., Alberta). Dorsal strap of

segment IX very narrow, short. Main body of segment large, angular, almost trapezoidal

(Fig. 482). Clasper large, short, stout; distal extremity black, bifid; base long, tapered ven-

trad. Median lobes of segment X small, hooked dorsad distally. Cercus massive, with large

ventro-distal tooth produced basad along mesal face (Fig. 483); in lateral aspect dorso-mesal

extremity separated from ventral tooth as postero-ventrally curved hook (Fig. 482). Aedea-

gus essentially identical to that of A curtus.

Female genitalia. This sex is known (Nimmo, 1965), but is indistinguishable from female

of A. curtus.

Notes on biology. — Adults of this species are found in high alpine meadow and mountain

pass areas, emerging from shallow, quiet pools or streams in sedge meadows. The flight

period extends from July 25 to August 27.

Geographical distribution. — The known range of this species extends from Great Slave

Lake to Idaho (Fig. 643). In Alberta it attains altitudes of up to 7,000’.

I have examined 16 specimens, eight males and eight females, from the study area.

Asynarchus lapponicus Zetterstedt, 1840

This species has been reported recently from the study area (Clifford, 1969), but I have

been unable to confirm the record as the specimens involved are missing. Therefore I do not

include full details on this species, but draw the reader’s attention to the above record, and

refer to Schmid (1954b: 78-81) for figures and full description.

The Genus Philarctus McLachlan

A single species of this genus is known from the study area.

Synopsis of characters. — Head rather elongate, eyes and cephalic warts small. Antennae

thick, shorter than wings. Anterior femora of male very thick; anterior tibiae equally thick,

slightly flattened, each terminated by ridge; both clothed with brushes of stout spines along

opposing faces; apical spur short, flat, triangular. Spur formula 1,3,4. Wings various in size;

some species with Limnephilus- like wings; others quite shortened, apically parabolic. Color-

ation dark brown. Venation of fore wing with chord weakly disrupted, slightly oblique; hind

wing chord strongly disrupted, parallel to body.

Male genitalia with well developed segment IX; dorsal strap short, wide, setose. Cerci very

large, bulky, strongly sclerotized, fused ventrad of anal aperture and located laterad of it.

Intermediate lobes of segment IX triangular or horizontal discs, ventrad of cerci. Claspers

well developed, with long, slender dorsal process. Aedeagus similar to Limnephilus spp.;

lateral arms very slender, expanded distally, finely spinate.

Female genitalia with dorsal body of segment IX short, blunt, projected, expanded laterad

Rhyacophilidae and Limnephilidae

133

at base; ventro-lateral lobes quite small, not contiguous ventrally. Supra-genital plate large,

short. Segment X large, blunt, thick, weakly sclerotized, pilose; deeply cleft tube, more or

less open ventrally. Cerci large, prominent, free. Median lobe of vulval scale long, narrow;

lateral lobes short, thick, sub-quadrangular.

Philarctus quaeris (Milne), 1935

(Fig. 484-489, 644)

Colpotaulius quaeris Milne, 1935:41, 45. (Type locality: Quesnel Lake, British Columbia).

Milne, 1936:60.

Limnephilus quaeris’, Ross, 1944:298. Etnier, 1965:149.

Philarctus quaeris’, Ross and Merkley, 1952:436. Ross and Spencer, 1952:47. Schmid, 1955:

158. Wiggins, 1963: 103-105. Clifford, 1969:583. Fischer, 1969:58.

Description. — Antennae dark brown; antero-mesal faces of scapes yellow-brown, glab-

rous. Vertex of head dark reddish brown in triangular area between three ocelli; lighter

peripherally. Thorax light brown laterally, dorsally. Brush of male fore leg confined to distal

three-quarters of femur; anterior tibia also with brushes on entire length of opposite face.

Spurs reddish brown. Fore wing length of male 8.7 mm; dark chocolate-brown, heavily

irrorate; costal area hyaline throughout. Venation identical to that of Limnephilus spp.

Male genitalia. (Specimen from 17 miles north of Langdon Corner, on Hwy. 9, Alberta).

Dorsal strap high, wide (Fig. 484); heavily setose. Main body of segment irregular in lateral

aspect, with low, triangular postero-dorsal angles. Clasper with parallelogram-like base sur-

mounted by short, thin, acuminate dorsal process. Median lobes of segment X not evident.

Ventrad of dorsally fused cerci are two placoid lobes which appear to be intermediate lobes

(Fig. 484, 485). Cercus large, fused at base to segment IX; distally acuminate plate fused

dorsally as roof to anal passage. Median shaft of aedeagus long, very slender, with distinct,

long distal head (Fig. 486, 487); lateral arms shorter, slightly expanded from narrow bases

to shovel-like distal blades concave on mesal faces.

Female genitalia. (Specimen from 17 miles north of Langdon Corner, on Hwy. 9, Alberta).

Vulval scale with roughly rectangular median lobe; lateral lobes triangular, close to median

(Fig. 489). Dorsal body of segment IX high, narrow, distinct from segment X (Fig. 488);

ventro-lateral lobes small, irregular, flap-like. Supra-genital plate trapezoidal. Segment X

tubular, with lateral bulges; slightly cleft mesally; recessed slightly into segment IX. Cerci

short, narrow, fused at base, projected clear of remainder of segment X.

Notes on biology. — Adults of this species are found near sedge and horse-tail sloughs or

lakes of the plains or low mountain valleys. The flight period extends from June 18 to

September 13, with a peak toward the end of August.