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 Nimmo 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. 44 Nimmo 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 Nimmo 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 Nimmo 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 Nimmo 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 a"d Wnd ™gS °f maleS °f SPedeS °f Lta“>ephilid » »— canadensis (Banks). Rhyacophilidae and Limnephilidae 161 Fig. 138-145. Fore (a) and hind (b) wings of males, and females where indicated, of species of Limnephilidae 138 Limnephilus sublunatus Provancher. 139. L. partimz Walker. 140. L. nigriceps (Zetterstedt). 141. Nemaaulim hostiUs Hagen,. 142^ Anabolu, ozburni (Milne). 143. Philocazca t her Nimmo n. sp. 144. Limnephilidae species 3 (female) 145. Limnephilidae species 4 (female). Fig. 146-169. Dicosmoecus jucundus Banks, 146. Male genitalia, lateral aspect. 147. Male genitalia, posterior aspect (partial). 148. Male genitalia, dorsal aspect (partial). 149. Aedeagus, lateral aspect. 150. Aedeagus, lateral arm, dorsal aspect. 151. Female genitalia, ventral aspect. 152. Female genitalia, lateral aspect. D. atripes (Hagen), 153. Lateral aspect, distal article of clasper, male genitalia. 154. Mesal face of clasper base, posterior aspect. 155. Aedeagus, lateral aspect. 156. Female genitalia, ventral aspect. Onocosmoecus unicolor (Banks), 157. Male genitalia, lateral aspect. 158. Male genitalia, dorsal aspect (partial). 159. Aedeagus, lateral aspect. 160. Aedeagus, lateral arm, dorsal aspect. 161. Aedeagal straps and clasper bases, dorsal aspect. 162. Female genitalia, lateral aspect. 163. Female genitalia, ventral aspect. Amphicosmoecus canax (Ross), 164. Male genitalia, lateral aspect. 165. Male genitalia, dorsal aspect (partial). 166. Aedeagus, lateral aspect. 167. Right aedeagal strap and clasper base, lateral aspect. 168. Female genitalia, lateral aspect. 169. Female genitalia, ventral aspect. Ce. - Cercus. Cl. - Clasper. 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. Nimmo Rhyacophilidae and Limnephilidae 163 Fig. 170-187. Imania tripunctata (Banks), 170. Male genitalia, lateral aspect. 171. Male genitalia, ventral aspect (partial). 172. Aedeagus, lateral aspect. I. cascadis Ross, 173. Male genitalia, lateral aspect. 174. Male genitalia, ventral aspect (partial). 175. Aedeagus, lateral aspect. 176. Female genitalia, ventral aspect. 177. Female genitalia, lateral aspect. I. bifosa Ross, 178. Male genitalia, lateral aspect. 179. Male genitalia, ventral aspect (partial). 180. Aedeagus, lateral aspect. 181. Female genitalia, ventral aspect. 182. Female genitalia, lateral aspect. I. hector Nimmo n. sp., 183. Male genitalia, lateral aspect. 184. Male genitalia, ventral aspect (partial). 185. Aedeagus, lateral aspect. 186. Female genitalia, ventral aspect. 187. Female genitalia, lateral aspect. 164 Nimmo Fig. 188-205. Ecclisomyia maculosa Banks, 188. Male genitalia, lateral aspect. 189. Male genitalia, dorsal aspect. 190. Aedeagus, lateral aspect. 191. Aedeagus, dorsal aspect. 192. Female genitalia, ventral aspect. 193. Female genitalia, lateral aspect. E. conspersa Banks, 194. Male genitalia, lateral aspect. 195. Male genitalia, ventral aspect. 196. Aedeagus, dorsal aspect. 197. Aedeagal strap, ventral aspect. 198. Female genitalia, lateral aspect. 199. Female genitalia, ventral aspect. Ecclisomyia species 1, 200. Female genitalia, ventral aspect. 201. Female genitalia, lateral aspect. Apatania zonella (Zetterstedt), 202. Male genitalia, lateral aspect. 203. Aedeagus, lateral aspect. 204. Female genitalia, lateral aspect 205. Female genitalia, ventral aspect. Rhyacophilidae and Limnephilidae 165 Fig. 206-225. Apatania shoshone Banks, 206. Male genitalia, lateral aspect. 207. Male genitalia, dorsal aspect (partial). 208. Aedeagus, lateral aspect. 209. Female genitalia, ventral aspect. 210. Female genitalia, lateral aspect. A. stigmatella (Zetterstedt), 211. Male genitalia, lateral aspect. 212. Male genitalia, dorsal aspect (partial). 213. Aedeagus, lateral aspect. 214. Female genitalia, lateral aspect. 215. Female genitalia, ventral aspect. A. crymophila McLachlan, 216. Male genitalia, lateral aspect. 217. Male genitalia, dorsal aspect (partial). 218. Aedeagus, lateral aspect. 219. Female genitalia, lateral aspect. 220. Female genitalia, ventral aspect. A. alberta Nimmo n. sp. 221. Male genitalia, lateral aspect. 222. Aedeagal tip, dorsal aspect. 223. Male genitalia, dorsal aspect (partial). 224. Female genitalia, lateral aspect. 225. Female genitalia, ventral aspect. 166 Nimmo Fig. 226-249. Oligophlebodes ruthae Ross, 226. Male genitalia, lateral aspect. 227. Male genitalia, ventral aspect. 228. Aedeagus, lateral aspect. 229. Aedeagus, ventral aspect. 230. Female genitalia, lateral aspect. 231. Female genitalia, ventral aspect. O. sierra Ross, 232. Male genitalia, lateral aspect. 233. Male genitalia, dorsal aspect. 234. Aedeagus, ventral aspect. 235. Aedeagus, lateral aspect. 236. Female genitalia, lateral aspect. 237. Female genitalia, ventral aspect. O. zelti Nimmo n. sp., 238. Male genitalia, lateral aspect. 239. Male genitalia, ventral aspect. 240. Aedeagus, lateral aspect. 241. Aedeagus, ventral aspect. 242. Female genitalia, lateral aspect. 243. Female genitalia, ventral aspect. Neo- thremma alicia Banks, 244. Male genitalia, lateral aspect. 245. Male genitalia, dorsal aspect (partial). 246. Aedeagus, lateral aspect. 247. Female genitalia, lateral aspect. 248. Female genitalia, ventral aspect. 249. Maxillary palpus of male. Rhyacophilidae and Limnephilidae 167 Fig. 250-269. Homophylax crotchi Banks, 250. Male genitalia, lateral aspect. 251. Male genitalia, dorsal aspect. 252. Aedeagus, lateral aspect. 253. Female genitalia, lateral aspect. 254. Female genitalia, ventral aspect. H. acutus Denning, 255. Male genitalia, lateral aspect. 256. Male genitalia, dorsal aspect. 257. Aedeagus, lateral aspect. 258. Female genitalia, ventral aspect. 259. Female genitalia, lateral aspect. H. baldur Nimmo n. sp., 260. Basal flap and scaled pocket at base of wing, male. 261. Male genitalia, lateral aspect. 262. Male genitalia, dorsal aspect. 263. Aedeagus, lateral aspect. LimnephUus sublunatus Provancher, 264. Male genitalia, lateral aspect. 265. Male genitalia, dorsal aspect (partial). 266. Aedeagus, lateral aspect. 267. Right lateral arm, dorsal aspect. 268. Female genitalia, lateral aspect. 269. Female genitalia, ventral aspect. Ce. - Cercus. CL - Clasper. 168 Nimmo Fig. 270a-290. Limnephilus sansoni Banks, 270a. Male genitalia, lateral aspect. 270b. Male genitalia, dorsal aspect (partial). 271. Female genitalia, lateral aspect. 272. Female genitalia, ventral aspect. L. extractus Walker, 273. Male genitalia, lateral aspect. 274. Male genitalia, dorsal aspect. 275. Aedeagus, lateral aspect. 276. Female genitalia, lateral aspect. 277. Female genitalia, ventral aspect. L. hageni Banks, 278. Male genitalia, lateral aspect. 279. Male genitalia, posterior aspect (partial). 280. Aedeagus, lateral aspect. 281. Right lateral arm, dorsal aspect. 282. Female genitalia, lateral aspect. 283. Female genitalia, ventral aspect. L. partitus Walker, 284. Male genitalia, lateral aspect 285. Male genitalia, posterior aspect (partial). 286. Right lateral arm of aedeagus, lateral aspect. 287. Right lateral arm, dorsal aspect (partial). 288. Tip of aedeagus, dorsal aspect 289. Female genitalia, ventral aspect. 290. Female genitalia, lateral aspect. Rhyacophilidae and Limnephilidae 169 Fig. 291-308. Limnephilus susana Nimmo n. sp., 291. Male genitalia, lateral aspect. 292. Male genitalia, dorsal aspect (partial). 293. Aedeagus, lateral aspect. 294. Female genitalia, ventral aspect. 295. Female genitalia, lateral aspect. L. species 1, 296. Female genitalia, lateral aspect. 297. Female genitalia, ventral aspect. No scale given. L. indivisus Walker, 298. Male genitalia, lateral aspect. 299. Aedeagus, lateral aspect. 300. Right lateral arm of aedeagus, dorsal aspect 301. Male genitalia, posterior aspect (partial). 302. Female genitalia, lateral aspect. 303. Female genitalia, ventral aspect L. infemalis (Banks), 304. Male genitalia, lateral aspect. 305. Male genitalia, posterior aspect. 306. Aedeagus, lateral aspect. 307. Female genitalia, lateral aspect 308. Female genitalia, ventral aspect. 170 Nimmo Fig. 309-331. Limnephilus ornatus Banks, 309. Male genitalia, lateral aspect. 310. Male genitalia, posterior aspect (partial). 311. Aedeagus, lateral aspect. 312. Right lateral arm of aedeagus, dorsal aspect. 313. Female genitalia, lateral aspect. 314. Female genitalia, ventral aspect. L. picturatus McLachlan, 315. Male genitalia, lateral aspect. 316. Male genitalia, posterior aspect (partial). 317. Aedeagus, lateral aspect. 318. Female genitalia, lateral aspect. 319. Female genitalia, ventral aspect. L. externus Hagen, 320. Male genitalia, lateral aspect. 321. Male genitalia, posterior aspect (partial). 322. Aedeagus, lateral aspect. 323. Aedeagus, dorsal aspect (partial). 324. Female genitalia, lateral aspect. 325. Female genitalia, ventral aspect. L. sericeus (Say), 326. Male genitalia, lateral aspect. 327. Aedeagus, lateral aspect. 328. Left clasper, posterior aspect. 329. Aedeagal tip, distal aspect. 330. Female genitalia, ventral aspect. 331. Female genitalia, lateral aspect. Rhyacophilidae and Limnephilidae 171 Fig. 332-349. Limnephilus femoralis (Kirby), 332. Male genitalia, lateral aspect. 333. Male genitalia, dorsal aspect (partial). 334. Aedeagus, lateral aspect. 335. Female genitalia, ventral aspect. 336. Female genitalia, lateral aspect. L. nogus Ross, 337. Male genitalia, lateral aspect. 338. Male genitalia, dorsal aspect (partial). 339. Aedeagus, lateral aspect. 340. Female genitalia, lateral aspect. 341. Female genitalia, ventral aspect. L. moestus Banks, 342. Male genitalia, lateral aspect. 343. Male genitalia, dorsal aspect. 344. Aedeagus, lateral aspect. 345. Female genitalia, ventral aspect. 346. Female genitalia, lateral aspect. L. cockerelli Banks, 347. Male genitalia, lateral aspect. 348. Aedeagus, lateral aspect. 349. Male genitalia, dorsal aspect (partial). 172 Nimmo Fig. 350-369. Limnephilus valhalla Nimmo n. sp., 350. Male genitalia, lateral aspect. 351. Male genitalia, dorsal aspect (partial). 352. Aedeagus, lateral aspect. 353. Female genitalia, lateral aspect. 354. Female genitalia, ventral aspect. L. lopho Ross, 355. Male genitalia, lateral aspect 356. Male genitalia, dorsal aspect (partial). 357. Aedeagus, lateral aspect. 358. Female genitalia, ventral aspect. 359. Female genitalia, lateral aspect. L. parvulus (Banks), 360. Male genitalia, lateral aspect. 361. Male genitalia, dorsal aspect (partial). 362. Right lateral arm of aedeagus, lateral aspect. 363. Female genitalia, lateral aspect. 364. Female genitalia, ventral aspect. L. spinatus Banks, 365. Male genitalia, lateral aspect. 366. Male genitalia, posterior aspect (partial). 367. Aedeagus, lateral aspect. 368. Female genitalia, ventral aspect. 369. Female genitalia, lateral aspect. Rhyacophilidae and Limnephilidae 173 Fig. 370-389. Limnephilus hyalinus Hagen, 370. Male genitalia, lateral aspect. 371. Right cercus and median lobe of segment X, posterior aspect. 372. Aedeagus, lateral aspect. 373. Female genitalia, ventral aspect. 374. Female genitalia, lateral aspect. L. secludens Banks, 375. Male genitalia, lateral aspect. 376. Male genitalia, posterior aspect (partial). 377. Aedeagus, lateral aspect. 378. Female genitalia, lateral aspect. 379. Female genitalia, ventral aspect. L. janus Ross, 380. Male genitalia, lateral aspect. 381. Male genitalia, dorsal aspect (partial). 382. Aedeagus, lateral aspect. 383. Female genitalia, lateral aspect. 384. Female genitalia, ventral aspect. L. perpusillus Walker, 385. Male genitalia, lateral aspect. 386. Male genitalia, posterior aspect (partial). 387. Aedeagus, lateral aspect. 388. Female genitalia, lateral aspect. 389. Female genitalia, ventral aspect. 174 Nimmo Fig. 390-413. Limnephilus argenteus Banks, 390. Male genitalia, lateral aspect 391. Male genitalia, dorsal aspect (partial). 392. Aedeagus, lateral aspect. 393. Female genitalia, ventral aspect. 394. Female genitalia, lateral aspect L. alberta Denning, 395. Male genitalia, lateral aspect. 396. Male genitalia, posterior aspect (partial). 397. Male genitalia, dorsal aspect (partial). 398. Aedeagus, lateral aspect 399. Aedeagus, dorsal aspect (partial). 400. Female genitalia, lateral aspect 401. Female genitalia, ventral aspect L. labus Ross, 402. Male genitalia, lateral aspect 403. Male genitalia, posterior aspect (partial). 404. Aedeagus, lateral aspect 405. Right lateral arm, dorsal aspect. 406. Female genitalia, lateral aspect 407. Female genitalia, ventral aspect. L. minusculus (Banks), 408. Male genitalia, lateral aspect 409. Male genitalia, posterior aspect (partial). 410. Aedeagus, lateral aspect 411. Tip of aedeagus, dorsal aspect. 412. Female genitalia, lateral aspect 413. Female genitalia, ventral aspect Rhyacophilidae and Limnephilidae 175 Fig. 414-438. Limnephilus kennicotti Banks, 414. Male genitalia, lateral aspect. 415. Male genitalia, dorsal aspect. 416. Aedeagus, lateral aspect. 417. Tip of aedeagus, dorsal aspect. 418. Clasper of male, posterior aspect. 419. Female genitalia, lateral aspect. 420. Female genitalia, ventral aspect. L. nigriceps (Zetterstedt), 421. Male genitalia, lateral aspect. 422. Male genitalia, posterior aspect (partial). 423. Male genitalia, dorsal aspect (partial). 424. Aedeagus, lateral aspect. 425. Female genitalia, ventral aspect. 426. Female genitalia, lateral aspect. L. rhombicus (L.), 427. Male geni- talia, lateral aspect. 428. Male genitalia, posterior aspect (partial). 429. Aedeagus, lateral aspect. 430. Right lateraL arm of aedeagus, dorsal aspect. 431. Female genitalia, lateral aspect. 432. Female genitalia, ventral aspect. L. canadensis Banks, 433. Male genitalia, lateral aspect. 434. Male genitalia, posterior aspect (partial). 435. Aedeagus, lateral aspect. 436. Aedeagus, dorsal aspect. 437. Female genitalia, ventral aspect. 438. Female genitalia, lateral aspect. 176 Nimmo Fig. 439-460. Clistoronia magnified (Banks), 439. Male genitalia, lateral aspect. 440. Male genitalia, dorsal aspect (partial). 441. Male genitalia, posterior aspect (partial). 442. Aedeagus, lateral aspect. 443. Female genitalia, lateral aspect. 444. Female genitalia, ventral aspect. Grammotaulius interregationis (Zetterstedt), 445. Male genitalia, lateral aspect. 446. Male genitalia, posterior aspect (partial). 447. Aedeagus, lateral aspect. 448. Right lateral arm, dorsal aspect of tip. 449. Female genitalia, lateral aspect. 450. Female genitalia, ventral aspect. Nemotaulius hostilis (Hagen), 451. Male genitalia, lateral aspect. 452. Male genitalia, dorsal aspect (partial). 453. Aedeagus, lateral aspect. 454. Female genitalia, lateral aspect. 455. Female genitalia, ventral aspect. Anabolia consocia (Walker), 456. Male genitalia, lateral aspect. 457. Aedeagus, lateral aspect. 458. Aedeagus, lateral aspect. 459. Female genitalia, lateral aspect. 460. Female genitalia, ventral aspect. Rhyacophilidae and Limnephilidae 177 Fig. 461-481. Anabolia ozburni (Milne), 461. Male genitalia, lateral aspect. 462. Male genitalia, dorsal aspect (partial). 463. Aedeagus, lateral aspect. 464. Female genitalia, lateral aspect. 465. Female genitalia, ventral aspect. A. bimaculata (Walker), 466. Male genitalia, lateral aspect. 467. Male genitalia, dorsal aspect (partial). 468. Aedeagus, lateral aspect. 469. Female genitalia, lateral aspect. 470. Female genitalia, ventral aspect. Asynarchus mutatus (Hagen), 471. Male genitalia, lateral aspect. 472. Male genitalia, dorsal aspect (partial). 473. Aedeagus, lateral aspect. 474. Female geni- talia, lateral aspect. 475. Female genitalia, ventral aspect. A. curtus (Banks), 476. Male genitalia, lateral aspect. 477. Male genitalia, posterior aspect (partial). 478. Aedeagus, lateral aspect. 479. Right lateral arm of aedeagus, dorsal aspect 480. Female genitalia, lateral aspect. 481. Female genitalia, ventral aspect. 178 Nimmo 487 Fig. 482-500. Asynarchus aldinus (Ross), 482. Male genitalia, lateral aspect. 483. Male genitalia, posterior aspect (partial). Philarctus quaeris (Milne), 484. Male genitalia, lateral aspect. 485. Male genitalia, dorsal aspect. 486. Aedeagus, lateral aspect. 487. Aedeagus, dorsal aspect (partial). 488. Female genitalia, lateral aspect. 489. Female genitalia, ventral aspect. Arctopora pulchella (Banks), 490. Male genitalia, lateral aspect. 491. Male genitalia, posterior aspect. 492. Aedeagus, lateral aspect. 493. Female genitalia, lateral aspect. 494. Female genitalia, ventral aspect. Lenarchus crassus (Banks), 495. Male genitalia, lateral aspect. 496. Male genitalia, posterior aspect (partial). 497. Male genitalia, ventral aspect. 498. Aedeagus, lateral aspect. 499. Tip of aedeagus, dorsal aspect. 500. Tip of left lateral arm of aedeagus, dorsal aspect. Rhyacophilidae and Limnephilidae 179 Fig. 501-519. Lenarchus fautini (Denning), 501. Male genitalia, lateral aspect. 502. Male genitalia, dorsal aspect. 503. Aedeagus, lateral aspect. L. vastus (Hagen), 504. Male genitalia, lateral aspect. 505. Male genitalia, dorsal aspect (partial). 506. Male genitalia, dorsal aspect. 507. Aedeagus, lateral aspect. 508. Female genitalia, lateral aspect. 509. Female genitalia, ventral aspect. L. brevipennis Banks, 510. Male genitalia, lateral aspect. 511. Male genitalia, dorsal aspect. 512. Aedeagus, lateral aspect. 513. Female genitalia, lateral aspect. 514. Female genitalia, ventral aspect. Hesperophylax occidentalis (Banks), 515. Male genitalia, lateral aspect 516. Segment X, posterior aspect. 517. Aedeagus, lateral aspect. 518. Female genitalia, lateral aspect. 519. Female genitalia, ventral aspect. 180 Nimmo Fig. 520-539. Hesperophylax incisus (Banks), 520. Male genitalia, lateral aspect. 521. Male genitalia, posterior aspect (partial). 522. Aedeagus, lateral aspect. 523. Female genitalia, ventral aspect. 524. Female genitalia, lateral aspect H. consimilis (Banks), 525. Male genitalia, lateral aspect. 526. Male genitalia, posterior aspect (partial). 527. Aedeagus, lateral aspect. 528. Female genitalia, lateral aspect 529. Female genitalia, ventral aspect. Chyranda centralis (Banks), 530. Male genitalia, lateral aspect. 531. Aedeagus, dorsal aspect. 532. Male genitalia, dorsal aspect. 533. Female genitalia, lateral aspect. 534. Female genitalia, ventral aspect Pycnopsyche subfasciata (Say), 535. Male genitalia, lateral aspect. 536. Aedeagus, lateral aspect 537. Male genitalia, dorsal aspect (partial). 538. Female genitalia, lateral aspect 539. Female genitalia, ventral aspect. Rhyacophilidae and Limnephilidae 181 Fig. 540-561. Pycnopsyche guttifer (Walker), 540. Male genitalia, lateral aspect 541. Male genitalia, posterior aspect (partial). 542. Aedeagus, lateral aspect. 543. Female genitalia, lateral aspect. 544. Female genitalia, ventral aspect. Philocasca thor Nimmo n. sp., 545. Male genitalia, lateral aspect. 546. Male genitalia, dorsal aspect. 547. Aedeagus, lateral aspect. Chilostigmodes areolata (Walker), 548. Male genitalia, lateral aspect. 549. Male genitalia, dorsal aspect (partial). 550. Aedeagus, lateral aspect Glyphopsyche irrorata (Fabricius), 551. Male genitalia, lateral aspect. 552. Male genitalia, posterior aspect (partial). 553. Aedeagus, lateral aspect. 554. Aedeagus, dorsal aspect. 555. Female genitalia, lateral aspect. 556. Female genitalia, ventral aspect Phanocelia canadensis (Banks), 557. Male genitalia, lateral aspect. 558. Male genitalia, dorsal aspect (partial). 559. Aedeagus, lateral aspect. 560. Female genitalia, lateral aspect 561. Female genitalia, ventral aspect 182 Nimmo Fig. 562-581. Psychoglypha prita (Milne), 562. Male genitalia, lateral aspect 563. Male genitalia, dorsal aspect. 564. Aedeagus, lateral aspect. 565. Aedeagus, dorsal aspect (partial). P. schmidi Nimmo, 566. Male genitalia, lateral aspect. 567. Male genitalia, dorsal aspect. 568. Aedeagus, lateral aspect. 569. Aedeagus, dorsal aspect (partial). 570. Female genitalia, lateral aspect. 571. Female genitalia, ventral aspect. P. alaskensis (Banks), 572. Male genitalia, lateral aspect. 573. Male genitalia, dorsal aspect. 574. Aedeagus, lateral aspect. 575. Aedeagus, dorsal aspect. 576. Female genitalia, lateral aspect. 577. Female genitalia, ventral aspect. P. ulla (Milne), 578. Male genitalia, lateral aspect. 579. Male geni- talia, dorsal aspect. 580. Aedeagus, lateral aspect 581. Aedeagus, dorsal aspect (partial). Rhyacophilidae and Limnephilidae 183 Fig. 582-591b. Limnephilidae species 1, 582. Female genitalia, lateral aspect. 583. Female genitalia, ventral aspect. Limnephilidae species 2, 584. Female genitalia, lateral aspect. 585. Female genitalia, ventral aspect. Limnephilidae species 3, 586. Female genitalia, lateral aspect. 587. Female genitalia, ventral aspect. Limnephilidae species 4, 588. Female genitalia, lateral aspect. 589. Female genitalia, ventral aspect. Limnephilidae species 5, 590a. Female genitalia, lateral aspect. 590b. Female genitalia, ventral aspect. No scale given when only females known. Neothremma laloukesi Schmid, 591a. Male genitalia, lateral aspect. 591b. Male genitalia, ventral aspect. (From Schmid, 1968). 184 Nimmo Fig. 592-597. Maps of distribution in Alberta and North America of: 592. Dicosmoecus atripes (Hagen). 593. D. jucundus Banks. 594. Onocosmoecus unicolor (Banks). 595. Amphicosmoecus canax (Ross). 596. Imania tripunctata (Banks). 597. I. bifosa Ross, I. cascadis Ross, and I. hector Nimmo n. sp. Rhyacophilidae and Limnephilidae 185 Fig. 598-603. Maps of distribution in Alberta and North America of: 598. Ecclisomyia maculosa Banks. 599. E. con- spersa Banks, 600. E. species 1, Apatania zonella (Zetterstedt). 601. A. crymophila McLachlan, A. shoshone (Banks). 602. A. stigmatella (Zetterstedt), A. alberta Nimmo n. sp. 603. Oligophlebodes ruthae Ross. 186 Nimmo Fig. 604-609. Maps of distribution in Alberta and North America of: 604. Oligophlebodes sierra Ross, O. zelti Nimmo n. sp. 605. Neothremma alicia Banks and Neothremma laloukesi Schmid. 606. Homophylax crotchi Banks, H. acutus Denning. 607. H. baldur Nimmo n. sp., Limnephilus sublunatus Provancher. 608. L. sansoni Banks. 609. L. extractus Walker. Rhyacophilidae and Limnephilidae 187 Fig. 610-615. Maps of distribution in Alberta and North America of: 610. Limnephilus hageni Banks. 611. L. partitus Walker. 612. L. susana Nimmo n. sp., L. species 1. 613. L. infemalis (Banks). 614. L. indivisus Walker. 615. L. ornatus Banks. 188 Nimmo Fig. 616-621. Maps of distribution in Alberta and North America of: 616. Limnephilus picturatus McLachlan. 617. L. extemus Hagen. 618. L. sericeus (Say). 619. L. nogus Ross. 620. L. femoralis (Kirby). 621. L. moestus Banks. Rhyacophilidae and Limnephilidae 189 Fig. 622-627. Maps of distribution in Alberta and North America of: 622. Limnephilus cockerelli Banks, L. valhalla Nimmo n. sp. 623. L. lopho Ross, L. parvulus (Banks). 624. L. spinatus Banks. 625. L. secludens (Banks). 626. L. janus Ross. 627. L. hyalinus Hagen. 190 Nimmo Fig. 628-633. Maps of distribution in Alberta and North America of: 628. Limnephilus perpusillus Walker. 629. L. alberta Denning, L. argenteus Banks. 630. L. labus Ross. 631. L. minusculus (Banks). 632. L. kennicotti Banks. 633. L. nigriceps (Zetterstedt). Rhyacophilidae and Limnephilidae 191 Fig. 634-639. Maps of distribution in Alberta and North America of: 634. Limnephilus rhombicus (L.). 635. L. cana- densis Banks, Clistoronia magnifica (Banks). 636. Grammotaulius interregationis (Zetterstedt). 637. Nemotaulius hostilis (Hagen). 638. Anabolia consocia (Walker). 639. A. ozbumi (Milne). 192 Nimmo Rhyacophilidae and Limnephilidae 193 Fig. 646-651. Maps of distribution in Alberta and North America of: 646. Lenarchus crassus (Banks), L. vastus (Hagen). 647. L. brevipennis Banks, L. fautini (Denning). 648. Hesperophylax occidentalis (Banks), H. consimilis (Banks). 649. H. incisus (Banks). 650. Chyranda centralis (Banks). 651. Pycnopsyche subfasciata (Say). 194 Nimmo Fig. 652-657. Maps of distribution in Alberta and North America of: 652. Pycnopsyche guttifer (Walker). 653. Philo- casca thor Nimmo n. sp. 654. Glyphopsyche irrorata (Fabricius). 655. Chilostigmodes areolata (Walker). 656 .Phanocelia canadensis (Banks). 657. Psychoglypha prita (Milne). Rhyacophilidae and Limnephilidae 195 Fig. 658-663. Maps of distribution in Alberta and North America of: 658. Psychoglypha schmidi Nimmo. 659. P. alaskensis (Banks). 660. P. ulla (Milne). 661. Limnephilidae species 1, Limnephilidae species 2. 662. Limnephilidae species 3, Limnephilidae species 4. 663. Limnephilidae species 5. 196 Nimmo ORIGINS AND RELATIONSHIPS OF THE FAUNA Aims of the study. — The putative origins and relationships of the fauna of Rhyacophili- dae and Limnephilidae, represented by 22 and 91 species respectively, of Alberta and eastern British Columbia, are described. Dispersal into the study area from unglaciated areas, or.refugia, after the last major glaciation of North America, the Wisconsin, is dis- cussed, and an attempt is made to determine the contributions of the glacial faunas of possible source areas to the present day fauna. At the glacial peak the study area was, to all intents and purposes, a slate wiped clean by total ice cover. There was at least one minor exception to this, mentioned below. Such faunal movements should be amenable to clarification to a greater extent than pre-Wisconsin faunas at the species level and provide a very convenient unit with which to work due to the essentially clean sweep of the study area by the ice. Procedures employed. — I shall attempt to arrive at a reasonable answer to the problem thus stated by an examination of a variety of factors. Firstly, the Wisconsin and post- Wisconsin history of North America is reviewed, especially the full extent of the ice sheets and major outliers, the locations of possible refugia, the subsequent retreat of the ice fronts, and the major post-glacial lakes and river systems and their development to the present. Secondly, a brief review of the major weather systems at the Wisconsin peak and at the present time is presented. Thirdly, an examination is made of the ranges of the Alberta and eastern British Columbia Rhyacophilidae and Limnephilidae, and of Alberta’s position with respect to the range areas, and each species is relegated to the range pattern which it best fits. Also, the species are examined as to groupings by similar habitats, to determine how this may be instructive. Lastly, an examination is made of the range relationships of each species to the remainder of its genus, or species group within a genus. Literature on aquatic orders of North American insects. — While there have been many major faunal works on the wholly aquatic orders of North American insects, such as Betten et al. (1934), Milne (1934, 1935, 1936), Ross (1944), Flint (1960), and Denning (1963) on Trichoptera, Needham and Heywood (1929), Needham and Westfall (1955), and Walker (1953, 1958) on Odonata, Needham and Claassen (1925), and Frison (1935) on Plecoptera, and Burks (1953) on Ephemeroptera, none has attempted to elucidate the zoogeography or origins of the fauna dealt with. Ross (1967) presented a study of evolution and dispersal of the world Trichoptera as a whole. In 1956 he dealt more fully with three families of Trichoptera. He also dealt (1953, 1958, 1965) with the effects of the Pleistocene and subsequent events on various nearctic insect groups, utilising primarily members of the wholly aquatic orders. Various smaller papers dealing with members of discrete groups within wholly aquatic orders in North America also exist, such as those by Ross (1951, 1959) on Trichoptera, Ricker (1963), and Ross, Rotramel, Martin, and McAlpine (1967) on Plecoptera. But there is none, so far as I am aware, which deals with the aquatic fauna, or part of it, of a distinct geographical area such as I deal with here. While reference is made in the following to papers which deal with non-aquatic groups of insects, where appropriate, in the elucidation of the problem before us, I make no reference to any such papers at this point. In dealing with fresh water insects, more especially the wholly aquatic orders, one is dealing with insects of a distinctly circumscribed habitat, at least in the immature stages, which would seem to set distinct Emits to their dispersal patterns and pose problems peculiar to such groups. Ross (1956) mentions this especially with reference to the cool-adapted Trichoptera. Rhyacophilidae and Limnephilidae 197 Synopsis of the glacial history of North America from the peak of the Wisconsin glaciation to the presnt Extent of Wisconsin glaciation. — Figure 664 (from Flint, 1957 and Prest, 1969) outlines the maximum extent of ice cover at the peak of the Wisconsin glaciation in North America. While the continental ice cover originated from several centers to form a series of coalescing ice fields, in effect the land mass of northern North America appears to have been under a single mass of ice which connected, by way of the Arctic archipelago, to the still extant Greenland ice cap. In the west the southern ice margin closely but irregularly paralleled the Canada-United States boundary. South of this margin were located many ice fields (as many as 70 according to Flint, 1957) of varied extents scattered throughout the Cordillera. In the east the ice margin extended further south. Both coasts and their outlying islands were apparently icebound, as was also the Arctic archipelago. Alaska was only partly glaciated, most of the Yukon valley being ice free; this condition extended over into north east Siberia. The line passing north west from southern Alberta to the Arctic Ocean is the loca- tion of the line of coalescence between the Laurentide ice sheet to the east and the Cordil- leran ice sheet to the west, the eastern edge of which flowed east through the Rocky Moun- tains to the Great Plains, there to be forced south east by the Laurentide sheet. The time of initiation of final degradation of the Wisconsin ice is uncertain, and could have varied from place to place, but evidence suggests an average value of about 10,000 B. P., despite at least some readvances in certain areas. Gravenor and Bayrock (1961) esti- mate the final recession, or melting back, of the Wisconsin ice from southern Alberta to have started about 1 1,000 years B. P. In that area they consider the ice to have receded in a northerly or north easterly direction. This refers to the Laurentide ice sheet retreating from the line of coalescence with the Cordilleran ice sheet of the Rocky Mountains. The Laurentide sheet is now extinct but the Cordilleran sheet still exists in the scattered glaciers and ice fields of the Rocky Mountains, and can be assumed to have melted much more slowly than the Laurentide, presumably due to higher altitudes and location in deeper valleys which might be supposed to provide some protection from ablation processes. Pre- sumably, also, the ice would be deeper in the major valleys and simply take longer to melt. It is interesting to speculate on the possibility that, at the time of general recession, while the greater part of Canada may still have been under ice, a corridor of deglaciated territory may have opened between the northwestern United States, western Canada, and the Arctic Ocean, possibly even through to the unglaciated part of Alaska, as this was relatively close to the line of coalescence (Fig. 664). Such a corridor would, in all probability, permit biotic interchange between Alaska and southern North America prior to complete deglaciation, possibly with results different from those that might be expected if the ice were to disappear uniformly, leaving the northern half of North America open to un- impeded colonisation. I have been unable to discover any record of such a corridor in the literature. Locations of possible unglaciated areas. — A.reas which were unglaciated during the Wis- consin, or any other glaciation, are of prime interest to the biogeographer, as they may have acted as refugia or foci of redispersal for plants and animals at a later date. In Fig. 664 roman numerals indicate the locations of known, or postulated, unglaciated areas and, there- fore, possible refugia. The Beringian refugium (I) is known definitely to have existed and includes the unglaci- ated areas of Alaska and northeast Siberia (Ball, 1963; Munroe, 1956). Prest, Grant, and Rampton (1968) in their ‘Glacial map of Canada’, and Prest (1969), show the western areas of the Yukon to have been unglaciated also. As Pewe, Hopkins, and Giddings (1965), 198 Nimmo in their Fig. 2, show the unglaciated area of central Alaska to have extended widely to the Alaska-Yukon boundary, to coincide at least approximately with the unglaciated portion of the western Yukon, this area must have formed part of the Beringian unglaciated area and, therefore, of the Beringian refugium. Flint’s map of Eurasia (1957: Fig. 24-1, and plate 3) shows a large unglaciated area between northeast Siberia and the remainder of Eurasia. From his map, however, it also appears that, at least in the Wisconsin glaciation, corridors may have existed to north or south of the glaciated area of Siberia, by which biotic movements may have occurred. Flint (1957) shows all of northern Canada under ice, but three areas are thought to have been unglaciated and acting as refugia, at least for plants, at the time of the Wisconsin peak (Ball, 1963). There is some doubt as to the existence of the Mackenzie refugium (II) which, if it existed, was small. A refugium has been postulated for the western Arctic islands (III) but Savile (1961) and Munroe (1956) both express doubt on this point, for rather different reasons. Savile believed the islands to have been unglaciated but covered with snow for too long periods to permit survival. However, Leech (1966) does not believe that periods of snow cover, of one or more years, necessarily eliminate life. Munroe states that the islands were under such a load of ice that they were very largely submerged, as evidenced by pro- nounced emergence from the sea in recent times. Prest (1969) shows areas II and III to have been unglaciated. Leech (1966) is of the opinion that a refugium existed in northern Elles- mere Island, having investigated the spider fauna of the Lake Hazen area. The third possible refugium of the high Arctic is Peary Land in northern Greenland (IV), which seems defi- nitely to have been unglaciated and to have acted as a refugium, for plants at least (Savile, 1961). Leech (1966), without limiting his statement to any one part of the biota, upholds the existence of the Peary Land refugium. As he decides that adjacent northern Ellesmere Island was a refugium for spiders, it is logical to assume that Peary Land would have been capable of supporting other animal life also. One or more unglaciated areas have been postulated for eastern North America (V) (Femald, 1925; Munroe, 1956; Lindroth, 1963) but, again it is uncertain whether they existed at all and, if so, where. One other area within the ice sheet is known to have been unglaciated. This is the height of land of the Cypress Hills of southeastern Alberta and southwestern Saskatchewan (VI) (Westgate, 1964; Gravenor and Bayrock, 1961). The remaining unglaciated area, undoubt- edly a refugium, is the entire southern half of North America (VII), except for the highland glaciers of the Cordillera. Ross (1965) surmises that small, very local, refugia may have existed within the Cordil- leran ice mass, on the higher mountain peaks rising above the ice. Ball (1963) states that the distributions of most of the northern biota are explainable by dispersal from the two major unglaciated areas of central Alaska (I) and the area south of the main ice sheet (VII). The distribution of the present day nearctic glaciers is given in Fig. 665. Lakes of Wisconsin and post-Wisconsin time. — As an aid to understanding present distributions and dispersal routes of aquatic insects, the distribution and drainage patterns of the lakes of North America during the Wisconsin glaciation and after are examined. The information presented is derived from Flint (1957), Elson (1967), and Prest, Grant, and Rampton (1968). Figure 666 outlines the major lakes of North America during and after Wisconsin time. Lakes Lahontan and Bonneville lay south of the ice sheet. The re- mainder were covered by ice at the peak of the Wisconsin and formed along the southern edges of the ice as it retreated northward. In most cases each lake is reduced in size at the present time but only Lakes Ojibway-Barlow and Lahontan have essentially disappeared. Rhyacophilidae and Limnephilidae 199 There were multitudes of small lakes, then and now, but they do not concern us here. The outlets of each lake are indicated, each having had several over a period of time as the ice edge retreated or readvanced. Flint (1957) presents a history of the Great Lakes. Briefly, they drained south to the Mississippi River by various routes until eventually they were allowed to take their natural course east to the St. Lawrence River by the final recession of the ice. Lake Ojibway-Barlow was relatively small and drained south and east via the Ottawa valley. It was at one time joined to Lake Agassiz. A history of Lake Agassiz is given by Elson (1967) in some detail. The lake varied greatly in size and position, draining first to the south and the Mississippi, then through several out- lets, to the south as before, to the Great Lakes via Lake Nipigon, and to the north west and the Athabasca River. Finally, as the ice receded, it drained northeast to Hudson Bay and shrank to the present Lake Winnipeg. The Saskatchewan River drained eastward to Lake Agassiz and may at various times, though Elson does not provide information on this, have successively formed part of the Gulf of Mexico, Arctic, and finally the Hudson Bay water- sheds. Prest, Grant, and Rampton (1968), in their ‘Glacial map of Canada’, show all western Canada, virtually to the western mountains, to have been under lake water at one time or the other since Wisconsin time. Presumably, apart from Lake Agassiz, small lakes or sloughs were involved. Lake Lahontan apparently had no outlet. Lake Bonneville at one stage in its history drained north west to the Columbia River and the Pacific Ocean. The study area of Alberta and eastern British Columbia embraces the headwaters of large rivers important to each of the major watersheds of North America except the Atlantic (Fig. 666; the darkened rivers). This curious juxtaposition of major watersheds seems to provide ideal conditions for dispersal of those aquatic organisms which, having their immature stages in aquatic habitats are, nevertheless, able to fly between neighbouring bodies of water as adults. The large post-glacial lake system of North America, temporary though some of its con- stituents may have been, probably provided very suitable pathways for dispersal of aquatic organisms from east to west and vice versa, more particularly for still water forms. The entire system stretching from Alberta to Ontario, Quebec, and the northeastern United States has, at various times since the last glaciation, formed a gigantic, shifting network of waterways. It seems possible that the distribution patterns of Trichoptera may at least partially reflect this, and the point is examined below. Synopsis of climatic history of North America from Wisconsin time to the present Bryson and Wendland (1967) proposed a sequence of weather pattern shifts from the peak of the Wisconsin glaciation to the present. They describe the effects of the shrinking Laurentian ice mass on air masses and the subsequent effects on weather. They consider the effects of climatic changes on the biota, largely the forest and grassland biotas along the southern edges of the ice sheet as it contracted. They do not consider it reasonable to regard vegetation zones to be controlled and located where they are simply by single factors, no matter how closely the boundaries of the zone and any value of the factors may coincide. The control must be the result of a variety of factors acting as annual, or at least seasonal, means. They utilise the mean positions of the various frontal zones and determine the pos- sible past positions by an examination of what is known about former and present vegeta- tion zones. I summarize their plotting of the postulated winter and summer mean positions 200 Nimmo of the Arctic frontal zones for the period about 10,000-13,000 years B. P., and for the present time (Fig. 667). The general trend towards the present was for the frontal zones to migrate north as the ice sheet dwindled. In the period 3,500-5,000 years B. P. they estimate that the fronts were actually further north of their present positions, but have readvanced slightly southward. The Arctic frontal means are used as indicators of cooler areas of the continent, and hence of increasing warmer conditions south of the mean frontal zones as they tended northward. The result, as shown in Bryson and Wendland, has been a shift northward of the North American biotic zones, more particularly, for the central plains of North America, of the boreal forest, and the grasslands. At about the time of retreat of the glaciers from Alberta the area was under boreal forest, which has since moved northward to make way for the present day grasslands of the southern part of the province. A similar effect can be expected in the mountains, except that it would be an altitudinal shift. Bryson and Wend- land indicate that there was essentially no tundra belt between the receding ice front and the encroaching boreal forest belt; that, in fact, the forest may have established itself imme- diately icebound moraine was uncovered. Exactly where tundra may have survived as such is not indicated, but it may reasonably be expected to have existed in the various far north- ern unglaciated areas, or refugia. The overall result was a general shift of the various floras and faunas northward, and upward in the Cordillera, with subsequent isolation of at least some species on scattered mountain ranges. As to the speed with which the biota responded to climatic shifts, Bryson and Wendland indicate that deglaciation is due to climatic changes and lags behind these changes. The result is that the areas beyond, and close to the edges of the ice sheet, were climatically suitable for vegetation prior to removal of the ice. That is, there is no reason to doubt that, for example, there could be forest right up to the edges of the ice (see preceding paragraph). Lindroth (1965), investigating the vegetated area of Skaftafell adjacent to a receding glacier in Iceland, concludes that there is nothing unusual in vegetation occurring in immediate proximity to glacial ice, whether advancing or receding. Presence or absence of vegetation in such a situation is dependent on macro-climate rather than on the presence of the ice itself. A superb example of dense Nothofagus forest in Argentina within feet of an active glacier is given in plate 14 of Tilman (1957). Westgate (1964) presents evidence of ponds with vege- tation and animals not only at the ice edge, but on the ice itself, wherever debris was suffi- ciently accumulated to provide a substrate for the biota. In the west the mean frontal zones clearly trend north west toward Alaska or the Macken- zie delta area (Fig. 667). This is true of both the ancient and modern frontal zones, and for the zones of intermediate periods postulated by Bryson and Wendland. This appears prob- ably to be due to the presence of the western Cordillera, which acts as a barrier, preventing the fronts from extending westward to the coast. Bryson and Wendland show the remaining ice mass (at about 8,000 years B. P.) to be located between Great Slave Lake and Labrador, and between northern Baffin Island and northern Ontario. It is reasonable then to suppose that the ice tended, at least at its southern and western boundaries, to retreat along the frontal zones. Here is some support for the corridor between Alaska and the unglaciated portion of North America, east of the mountains, and right through the study area, as postulated above. Distribution of the Rhyacophilidae and Limnephilidae of Alberta and eastern British Columbia The species of Rhyacophilidae and Limnephilidae known from the study area have been Rhyacophilidae and Limnephilidae 201 found to fall into 12 basic distribution patterns. These are outlined in Fig. 668 and 669. 1 have grouped them into two primary types for convenience. The first type comprises those species which are confined strictly to western North America, tending generally to follow the various ranges of the western Cordillera (Fig. 668). The second type comprises those species which extend beyond, or exist only east of the Cordillera (Fig. 669). The patterns as illustrated outline the general area occupied by the species from which they are drawn. They are not exact delineations of species ranges. The ranges of Trichoptera in North America are, as yet, relatively poorly known, and much intensive collecting is required before accurate statements can be made about range patterns. However, for the present study, current knowledge will have to suffice. The species ranges used in determining these range patterns do not necessarily extend through- out the full extent of the patterns to which they belong. The boundaries are really com- posites of species ranges which appear to follow a common pattern, however incom- pletely. Table 2 lists the range patterns in numerical order, giving the species known from each, and the range map number of each species in the taxonomic portion of this study. Follow- ing is presented a brief outline of each range pattern, in numerical order. Range pattern 1. This pattern extends from central Alaska south, embracing the enitre western Cordillera, as far south as the southwestern United States. The Alberta species included are mostly mountain stream species, but a few are usually found at higher alti- tudes, in the bogs or pools of the high passes and alpine meadows. This pattern is equivalent to Munroe’s (1956) type W5. Range pattern 2. This pattern extends from central Alaska to Colorado, along the eastern ranges of the Cordillera. Only three Alberta species are included, two of which are species of the lower mountain streams; the third, Imania tripunctata, is found at higher altitudes and is rather uncommon. Range pattern 3. This pattern is similar to type 1, but without the extension to the Yukon and Alaska. It contains a large number of species, most of which are inhabitants of mountain streams of varying degrees of rapidity and turbulence. This pattern is equivalent to Munroe’s type W3. Range pattern 4. This pattern extends from the Alberta Cordillera to California, via Idaho, Washington, and the Cascade ranges. It contains a small selection of species which inhabit the lower mountain streams and lakes. Range pattern 5. Species included in this pattern follow the eastern ranges of the Cordil- lera from Alberta to Colorado, and inhabit mountain streams. This pattern is equivalent to Munroe’s type W2. Range pattern 6. This pattern embraces species which are either very restricted in range, largely to the Cordillera of Alberta, or are known only from the type localities; several are described above as new. It is not so much a pattern as a collection of species which cannot yet be assigned to any of the other patterns. Specimens of several of the species may simply be rarely taken, and thus are poorly known. The range pattern embraces the Cordillera of Alberta, British Columbia, Washington, and Idaho. The great majority of included species are found near mountain creeks or ponds ranging to the higher alpine meadows, and largely comprise members of genera which are reputedly rare (Ross, 1950b; Denning, 1964; Wiggins and Anderson, 1968). Range pattern 7. This pattern embraces the greater part of North America, extending from Alaska east to Newfoundland and Greenland, and south east to New Mexico. In the United States it is restricted largely to the eastern ranges of the Cordillera. The species included in this pattern inhabit streams and rivers. 202 Nimmo Fig. 664-669. 664. Maximum distribution of glacial ice in North America at the peak of the Wisconsin glaciation, showing locations of known and suspected unglaciated areas (roman numerals), and line of coalescence between the major ice masses (toothed line). Compiled from Flint (1957), Westgate (1964), and Prest, Grant, and Rampton (1968). 665. Present day distribution of nearctic glaciers. From Flint (1957). 666. Composite map of major periglacial and pluvial lakes of North America (black areas) and present major lakes (hatched areas), showing various drainage patterns which existed post-glacially and which are now extinct (arrows). A - Lake Aggasiz; B - Lake Bonneville; G - Great Lakes; L - Lake Lahontan; O-B - Lake Ojibway-Barlow. Compiled from Flint (1957), and Elson (1967). 667. Map of North America showing the mean positions of the Arctic frontal zone 10,000-13,000 years B. P. in summer (B. P. - S.) and winter (B. P. - W.). Derived from Bryson and Wendland (1967). 668. Range patterns 1-6 exhibited by Alberta species of Rhyacophilidae and Limnephilidae. 669. Range patterns 7-12 as exhibited by Alberta species of Rhyacophilidae and Limnephilidae. Rhyacophilidae and Limnephilidae 203 Table 2. Species of Rhyacophilidae and Limnephilidae from the study area listed under the range pattern to which each belongs. Pattern Map Pattern Map no. Species Fig. no. Species Fig. 1 Rhyacophila vobara Milne 115 Rhyacophila rickeri Ross 1 1 2 Rhyacophila vepulsa Milne 1 1 3 Rhyacophila tucula Ross 105 Rhyacophila vofixa Milne 107 Rhyacophila verrula Milne 1 1 8 Psychoglypha ulla (Milne) 660 Ecclisomyia conspersa Banks 599 Lenarchus brevipennis Banks 647 Lenarchus fautini Denning 647 Lenarchus vastus (Hagen) 646 Asynarchus aldinus (Ross) 643 Hesperophylax incisus (Banks) 649 2 Rhyacophila alberta Banks 104 Limnephilus sansoni Banks 608 Imania tripunctata (Banks) 596 3 Rhyacophila hyalinata Banks 1 1 1 Rhyacophila vaccua Milne 1 1 6 Rhyacophila pellisa Ross 1 1 4 Rhyacophila bifila Banks 109 Rhyacophila coloradensis Banks 1 1 0 Dicosmoecus atripes (Hagen) 592 Oligophlebodes sierra Ross 604 Oligophlebodes ruthae Ross 603 Limnephilus spinatus Banks 624 Neothremma alicia Banks 605 Hesperophylax occidentalis (Banks) 648 Ecclisomyia maculosa Banks 598 Amphicosmoecus canax (Ross) 595 Clistoronia magnifica (Banks) 635 4 Limnephilus nogus Ross 619 Limnephilus lopho Ross 623 Dicosmoecus jucundus Banks 593 Hesperophylax consimilis (Banks) 648 5 Limnephilus cockerelli Banks 622 Homophylax baldur Nimmo 607 Rhyacophila vagrita Milne 119 Apatania shoshone Banks 60 1 6 Rhyacophila vemma Milne 1 20 Rhyacophila belona Ross 1 1 2 Rhyacophila glaciera Denning 1 06 Rhyacophila milnei Ross 1 1 9 Rhyacophila chilsia Denning 1 1 5 Rhyacophila species 1 121 Rhyacophila species 2 121 Homophylax crotchi Banks 606 Homophylax acutus Denning 606 Oligophlebodes zelti Nimmo 604 Neothremma laloukesi Schmid 605 Limnephilus susana Nimmo 6 1 2 Limnephilus valhalla Nimmo 622 Limnephilus alberta Denning 629 Limnephilus species 1 612 Philocasca thor Nimmo 653 Apatania alberta Nimmo 602 Psychoglypha prita (Milne) 657 Psychoglypha schmidi Nimmo 658 Ecclisomyia species 1 600 Imania bifosa Ross 597 Imania cascadis Ross 597 Imania hector Nimmo 597 Limnephilidae species 1 661 Limnephilidae species 2 661 Limnephilidae species 3 662 Limnephilidae species 4 662 Limnephilidae species 5 663 7 Limnephilus infernalis (Banks) 613 Limnephilus ornatus Banks 6 1 5 Limnephilus ex ternus Hagen 617 Limnephilus femoralis (Kirby) 620 Limnephilus minusculus (Banks) 63 1 Limnephilus hyalinus Hagen 627 Limnephilus rhombicus (L.) 634 Limnephilus sublunatus Provancher 607 Limnephilus perpusillus Walker 628 Pycnopsyche guttifer (Walker) 652 Nemotaulius hostilis (Hagen) 637 Asynarchus curtus (Banks) 642 Anabolia bimaculata (Walker) 640 Apatania stigmatella (Zetterstedt) 602 Nimmo 204 Table 2 (continued) Pattern Map Pattern Map no. Species Fig. no. Species Fig. 8 Rhyacophila acropedes Banks 108 Limnephilus janus Ross 626 Rhyacophila angelita Banks 117 Lenarchus crassus (Banks) 646 1 1 Limnephilus indivisus Walker 614 Onocosmoecus unicolor (Banks) 594 Limnephilus argenteus Banks 629 Limnephilus sericeus (Say) 618 Limnephilus extractus Walker 609 Limnephilus moestus Banks 621 Limnephilus canadensis Banks 635 Glyphopsyche irrorata (Fabricius) 654 Limnephilus parvulus (Banks) 623 Arctopora pulchella (Banks) 654 Anabolia consocia (Walker) 638 Chyranda centralis (Banks) 650 Anabolia ozburni (Milne) 639 Phanocelia canadensis (Banks) 656 9 Limnephilus nigriceps (Zetterstedt) 633 Pycnopsyche subfasciata (Say) 651 Limnephilus secludens (Banks) 625 Limnephilus picturatus McLachlan 616 1 2 Limnephilus kenicotti Banks 632 Asynarchus mutatus (Hagen) 641 Limnephilus hageni Banks 610 Philarctus quaeris (Milne) 644 Limnephilus partitus Walker 611 Psychoglypha alaskensis (Banks) 659 Chilostigmodes areolata (Walker) 655 Apatania crymophila McLachlan 601 Grammo taulius in terregationis (Zetterstedt) 636 1 0 Limnephilus labus Ross 630 Apatania zonella (Zetterstedt) 600 Range pattern 8. This pattern comprises an eastern and a western region. The western region extends from Alaska to California and Colorado, and eastward almost to Manitoba. The eastern region extends from Manitoba eastward to Newfoundland. There are no records known, for any of the species which are included within this pattern, for the intermediate area, which variously includes all or parts of at least Saskatchewan and Manitoba. The species included here variously inhabit, in the study area, mountain creeks, the swamps of the lower mountain valleys, and the higher passes. Range pattern 9. This pattern extends from Alaska southeastward, in a widening belt, to Colorado and Ontario. In the United States Cordillera it is confined to the eastern ranges south of Idaho. The included species inhabit primarily lakes, sloughs, or streams in low mountain valleys. Range pattern 10. This pattern embraces the great central plains of North America and includes only two species. The pattern is equivalent to Munroe’s (1956) type C4. Range pattern 1 1 . This pattern extends from the eastern edges of the Cordillera in Alber- ta, to the Great Lakes states of the United States, Labrador, and Newfoundland. Alberta represents the western limits for all the included species of this type. These species occur in lakes, sloughs, and slower streams. This pattern coincides with Munroe’s (1956) type B. Range pattern 12. This pattern embraces virtually all of Canada, with its southern bound- ary extending into the United States south of the Great Lakes. At least one species, Apa- tania zonella, is known to extend northward to the Arctic archipelago. The included species emerge from lakes, sloughs and, perhaps, the slower rivers. Rhyacophilidae and Limnephilidae 205 Distributional relationships of Alberta species of Rhyacophilidae and Limnephilidae to the remainder of their respective genera or species group In this section are examined the spatial relationships of the Alberta Trichoptera dealt with in this study to other species of their respective genera, or species groups within genera. The purpose in so doing is to determine whether it is possible to educe the provenance of the Alberta species after the Wisconsin glaciation. The Rhyacophilidae. - None of the North American species of Rhyacophila is known to be holarctic. With the exception of two transcontinental species, the Alberta species are restricted to the western Cordillera. The alberta group. Of the four species known to belong to this group, kincaidi , alberta, tucula, and glaciera, the last three are represented in the study area; kincaidi is known only from the Cascade ranges of Washington (Ross, \956), glaciera is known only from Montana and Alberta (Fig. 106). The two species alberta and tucula are known from Alaska to the Rocky Mountains of the United States (Fig. 104, 105). These two species are very similar and occupy somewhat mutually exclusive ranges, tucula to the west, alberta to the east; there are no records from the Yukon. The vofixa group. Of the two very similar species of this group only vofixa is known from the study area; its range extends from Alaska and the Yukon to Idaho (Fig. 107). The other species, harmstoni, is known only from Colorado and Utah. The acropedes group. Ross (1956) lists 11 species in this group, three of which are eastern Asiatic. According to Ross (1956) the seven North American species are confined to the western Cordillera, except for acropedes , which is also known from eastern North America. Ross considers that the group originated in eastern Asia, spread to western North America, whence a second, reverse, interchange occurred, back to Asia, to give the present set of species. As this undoubtedly occurred prior to the Wisconsin glaciation it is of little interest at present. According to Ross the North American species of the group are re- stricted to the western coastal ranges, except for acropedes (Fig. 108). Smith (1968) records vao from as far east as Idaho. I record vemna from Alberta; Smith records it from Idaho (Fig. 120). Thus these two species are not as restricted as Ross supposed. Ross derives acropedes from vao as the result of an eastward spread, after which acropedes spread west to coincide with vao. Thus the origin of the North American species of this group is apparently located in the coastal mountain ranges. It may be supposed that the ranges of the species prior to the Wisconsin glaciation were much as they are today, south of the southern limit of ice. The invaria group. This group has an Appalachian branch of six species, and a Cordilleran branch of five species. Two of the western species, bifila and coloradensis , are known from the study area (Fig. 109, 1 10) and are general throughout the Cordillera. Two other species, amabilis and kemada, are known only from southern California, and the fifth, insularis, is known only from Vancouver Island. The hyalinata group. This group, comprising four species, is of peculiar distribution; three species are located in western North America, while the fourth is known from the Caucasus. Ross (1956) cannot be certain that this European species belongs to this group, as he has seen only drawings. Only one North American species, hyalinata , is known from the study area; it is widely distributed in the Cordillera as a whole (Fig. 111). Of the remaining two species, sonoma is known only from California, and vocala from the Cascade ranges from Oregon to British Columbia. The sibirica group. This is a large group of 26 species, four of which are known from the study area. These four species are rather dissimilar belonging to different lineages within the 206 Nimmo group (Ross, 1956:95). One species is known from Europe, several more from eastern Asia, and the greater number from North America, being divided into eastern and western groups. Ross (1956) postulates a western North American origin for the group, with dispersal in time to Asia and eastern North America. There are no holarctic species. Of the Alberta species, two, vepulsa and pellisa, are widespread; vepulsa ranges from Alaska to California (Fig. 113 ), pellisa appears to be limited to Alberta in the north, is not recorded from British Columbia, though it almost certainly occurs there, and extends south by both the coastal and eastern ranges of the Cordillera (Fig. 1 14). Of the other two species, belona is restricted to the Idaho-Alberta area, and rickeri extends from the Alberta-British Columbia Cordillera to Alaska. The betteni group. This group comprises eight species, all confined to the Cordillera of western North America. All but the two species known from the study area are confined to the Cascade ranges of California and Oregon. Of the two known from Alberta one, chilsia, is known only from the Alberta Cordillera (Fig. 115); the second, vaccua, is widespread, but barely extends south along the eastern ranges of the Cordillera (Fig. 1 16). Ross (1956) is of the opinion that the group originated in the coastal ranges and has spread eastward from there. The vobara group. Two species comprise this group, iranda and vobara, of which the latter is known from the study area. The other species is known from the coastal ranges of Oregon. The angelita group. This group contains three species, two of which, perplana and vuzana, are known only from the Cascade ranges of the coast; the third species, angelita, is wide-' spread throughout the Cordillera, from the Yukon to California and Colorado (Fig. 117). It is also recorded from the northern Appalachians, in what must surely be an isolated population; it seems reasonable to suppose that isolation occurred post-glacially. The verrula group. Only one species, verrula, is known in this group. It ranges widely from Alaska to California and Colorado (Fig. 118). The vagrita group. The two very similar species of this group are both known from the study area. These are vagrita and milnei (Fig. 1 19). Only vagrita is widespread, extending to Utah; milnei is known only from the type locality at Banff, Alberta. The Limnephilidae. — The 91 species known from the study area are not distributed so narrowly as the Alberta Rhyacophilidae. Some species are known only from the western Cordillera of North America; others are widespread transcontinental^, and from north to south; others are holarctic, if not circumpolar. The genus Dicosmoecus. Of the 10 species of Dicosmoecus (Schmid, 1955), two are known from the study area. Of these two, atripes is widespread throughout the Cordillera, from Alberta and British Columbia to California and New Mexico (Fig. 592); jucundus (Fig. 593) has a similar distribution except that it is unknown from the eastern ranges. The remainder of the genus is distributed between eastern Asia (two species) and western North America (eight species). Of the North American species, the general distributional pattern is centered on the coastal ranges from California to British Columbia, with one species known only from Alaska. The genus Onocosmoecus. Of the eight species of Onocosmoecus (Schmid, 1955) only one is known from the study area {unicolor). This species is distributed from Alaska to California, New Mexico (Fig. 594), and eastern North America, with a large gap be- tween the eastern and western ranges. As this species is known from streams of the plains regions of Alberta the gap must be viewed with suspicion, as being due to insufficient collecting. One species of Onocosmoecus is known from eastern Asia, the remainder be- ing North American and confined to the Cordillera with the exception of quadrinotatus, Rhyacophilidae and Limnephilidae 207 which ranges from Newfoundland to Michigan and the White Mountains of New Hamp- shire. The genus Imania. Of the 10 known species of Imania, Schmid (1955) lists eight; the ninth is described above as new; the tenth is described by Schmid (1968) as new. Four species are known from the study area: bifosa, cascadis, tripunctata, and hector. Of these, hector is known only from Alberta (Fig. 597); bifosa is known only from Alberta and British Columbia; cascadis from Washington, Oregon, and Alberta (Fig. 597); and tripunc- tata is widespread from Colorado and Washington to Alaska (Fig. 596). Ross (1950b) refers to tripunctata as occurring as isolated, higher altitude, populations, particularly in the southern extremities of its range. Of the genus as a whole one species is known from eastern Asia; the remaining four species are known from isolated localities in Colorado ( gnathos ), Nevada ( renoa ), and Washington (acanthis and cidoibes). The genus Amphicosmoecus . This genus is represented by a single species, canax, which is widespread from Alberta and British Columbia to California and Utah (Fig. 595). The genus Ecclisomyia. Of the six species of this genus (Schmid, 1955), two are known from the study area {maculosa and conspersa)', a third, undescribed species is represented by a single female. Both Alberta species are widespread in the Cordillera. The species conspersa , which ranges from Alaska to California and New Mexico (Fig. 599), has a more extensive known range than maculosa. The latter ranges from Alberta and British Columbia to Colorado (Fig. 598). Of the remaining four species two, digitata and kamtshatica, are known only from eastern Asia, and the other two, scylla and bilera, are known only from British Columbia to Oregon, and California respectively. The genus Apatania. This genus contains 50 species, five of which are known from the study area; one is described above as new. Twelve species are recorded from North America; of these, three are holarctic and known from Alberta or just north of Alberta. The remaining species of the genus are widespread through Eurasia. The species known from the study area, or closely adjacent areas, are zonella, stigmatella, shoshone, crymo- phila, and alberta. Of these species zonella (Fig. 600), stigmatella (Fig. 602), and crymo- phila (Fig. 601) are northern in distribution in North America, and holarctic in total. A. stigmatella has been recorded from Colorado; this is either an error, or represents a pe- culiarly isolated population. Of the remaining two species shoshone is known from Alberta south, along the eastern Cordillera, to Colorado; alberta is known only from around Banff, Alberta. The genus Oligophlebodes. Of the seven species of Oligophlebodes three are known from the study area, one being described above as new. Of the Alberta species zelti is known only from several localities in Alberta (Fig. 604); sierra is widespread from Alberta and British Columbia to Colorado and California (Fig. 604); and ruthae is confined to the eastern ranges of the Cordillera, from Alberta and British Columbia to Oregon and Utah (Fig. 603). Of the four remaining species of the genus one is known only from Colorado ( ardis ), another is known only from New Mexico and Utah (sigma), the third is widespread from New Mexico to Wyoming and South Dakota, and is the most easterly species of Oligophle- bodes ( minuta ), and the fourth (mostbento) is known only from Oregon. The genus Neothremma. Of the four species of Neothremma (Schmid, 1955; 1968) two, alicia and laloukesi, are known from the study area; the total known range of alicia extends from Alberta and British Columbia to Oregon and Colorado (Fig. 605). At present laloukesi is known only from Alberta. The other two species (didactyla and galena) are known only from Washington. The genus Homophylax. Denning (1964) revised this genus, and listed eight species. One further species was described above as new, raising the total to nine. Three species are 208 Nimmo known from the study area ( crotchi , acutus, and baldur ). Of these crotchi is known only from a very small range, from southern Alberta to Vancouver Island and Washington (Fig. 606); acutus is known only from Idaho and Alberta (Fig. 606) but, being only recently described by Denning (1964) will almost certainly be found to extend further; and baldur is known only from Utah and far southwestern Alberta (Fig. 607). Of the remaining six species andax is known from southeast Alaska to Oregon; flavipennis is known from Colo- rado to Montana; insulas is known only from California; rentzi only from California; neva- densis from California and Nevada; and adriana only from New Mexico. More so than other rare genera, Homophylax appears to be divided into distinct eastern and western species groups. Much more almost certainly remains to be known of the range of species of Homo- phylax. The genus is rarely represented in collections and appears to be rare in nature. This apparent rarity may, however, be due to secretive habits and high altitude distribution, thus making collection difficult. The genus Limnephilus. This is one of the larger genera of Trichoptera, and the largest of the Limnephilidae. Schmid (1955) lists 140 species in the genus of which seven are incertae sedis. In this study two new species of Limnephilus are described, raising the total to 142. Of this number of species one is listed from South America, 63 from North America only, and seven are holarctic. Of the 33 species known to occur in the study area one is un- described (the female only is known), and six are holarctic. Of the rhombicus group only one species, rhombicus , is known from the study area. This species is holarctic in distribution with an extensive North American range (Fig. 634); the remaining four species are variously known from Asia and eastern Europe, but one is known from Spain (Schmid, 1955). This group would appear to be decidedly Eurasian in origin. Of the 1 1 species of the subcentralis group six are known from the study area, including one new species. These species are sansoni, extractus, hageni, sublunatus, partitus, and susana. One other species is undescribed as only the female is known. None of the Alberta species are holarctic; sansoni extends from Alaska to Colorado, along the eastern ranges of the Cordillera (Fig. 608); extractus extends from Alberta and Great Slave Lake to New Hampshire (Fig. 609); sublunatus extends from British Columbia to Colorado and Quebec (Fig. 607); partitus extends from British Columbia to the Northwest Territories and New- foundland (Fig. 611); and susana is known only from the type locality in Alberta (Fig. 612). The remaining five species are known variously from Europe to Siberia, with one species, elongatus, from the Northwest Territories. Of the seven species of the stigma group two, infernalis and indivisus, are known from the study area. Both are distributed in a northern transcontinental pattern, infernalis from Alaska to New Hampshire (Fig. 613), and indivisus from British Columbia to Illinois and Nova Scotia (Fig. 614). The remaining five species are variously known from Europe to eastern Siberia. Only one species, ornatus, is known in the ornatus group. Its range in North America is northern transcontinental, from Alaska to Illinois, Newfoundland and Greenland (Fig. 615). Schmid (1955) also reports it from Japan. One of the two species of the picturatus group is known from the study area. This species, picturatus, is holarctic, ranging from Sweden to North America, where it extends from Alaska to Colorado and Hudson Bay (Fig. 616). The second species is known only from British Columbia. Of the two species of the externus group, like the picturatus group, one, thorns, is isolated in North America, in Utah; the second species, externus, is holarctic, ranging from Europe to North America. In North America it ranges from Great Slave Lake to California Rhyacophilidae and Limnephilidae 209 and Newfoundland (Fig. 617). The sericeus group, with two species, is limited to North America. The species known from the study area, sericeus, is northern transcontinental in range, from Alaska to Oregon and Maine (Fig. 618) with an apparent mid-continental gap. The second species, fagus, is known only from Oregon. Of the three species of the morrisoni group one, lopho, is known from the study area. This species is restricted to Oregon, British Columbia, and Alberta (Fig. 623). The re- maining two species, castor and morrisoni, are restricted to the western ranges of the Cordillera. The sitchensis group contains seven species, of which three are known from the study area. These are moestus, cockerelli, and valhalla. The group is restricted to North America. With the exception of moestus, all are restricted to the western Cordillera. Of the Alberta species moestus is most widespread, ranging from British Columbia to Colorado, Newfound- land, and Greenland (Fig. 621); cockerelli is restricted to the eastern ranges of the Cordillera (Fig. 622); and valhalla is known only from the area of the Albertan continental divide (Fig. 622). The single species of the luridus group known from the study area,/ay 1971 CONTENTS 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 Editorial — Publish or Perish? The admonition to publish or perish, with its oral equivalent: present a paper or stay at home, may be of value to the occasional administrator who has no better device for allo- cating travel monies, increments, or other rewards of virtue, but has certainly had some unhappy consequences. All of us, I am sure can recall more than one occasion when, sitting through some usually duller than normal submitted paper, the feeling has come over us that we have heard it all before. Anybody who has ever really tried to keep up with the literature will know of many frustrating examples of related one or two page papers, published in several usually obscure journals, by the same author (or authors in varied sequence) under different titles. When assembled through due process of inter-library loan or microfilm and subjected to critical comparative study, such papers often prove to say exactly the same thing. Often it wasn’t worth saying anyhow. Then there are the papers, often longer, which appear in the proceedings of a meeting, and again in a regular journal, with no great change. There are also coherent theses which are decimated for publication in a diversity of peri- odicals; for this the blame is also diverse: the facile acceptance of long lists of publications as an index of merit in both the papers and their author, editorial boards with a terror of long papers, and authors with an inordinate love for every word they have ever drafted. But all of this is at the personal level; the current Canadian wave of nationalistic or per- haps we should call it tribalistic fervour, since our aboriginal peoples appear to be speci- fically excluded, has expanded the publish or perish dictum to the level of national publish- ing houses. These must be helped, maintained, kept Canadian, cherished, and above all kept uncontaminated by any other funds than Canadian dollars. The first requirement for a Canadian publishing house to flourish, however, is that Cana- dians buy, even if they do not read them, Canadian books, because there are obstacles of various kinds, all unjustifiable, to the passage of books across many international bounda- ries. Canadians do neither of these things; at least not on the scale of most progressive peoples. Some 10 years ago I lived for a while in an African town of some 15,000 souls, more than half of them black or brown, which supported as many shelf-feet of bookshops as a Canadian city of 350,000. 236 Per million of population, Canada publishes one third of the average number of books published by western European countries and only about half the number published in the U.S.S.R. Some 10 years ago we could at least take comfort in the thought that we pub- lished nearly twice as many books per million people as our neighbours to the South, but their production is now well ahead of that of Canada. It is a strange fact that the more books per head a country publishes, the more copies of each it prints. This leaves Canada still further behind in numbers of copies of books printed, so that while bookish Britain prints a book per head in about 16 months, it takes Canada over six years to achieve this. Business men tell me that Canadian publishers lack business ability; this may well be true. At the receiving end 1 know them to be slow, inaccurate, and expensive. It is almost always both quicker and cheaper to buy a British book from a British retailer than from a Canadian distributor, who is usually also a Canadian publisher. Prices may be as much as 65 per cent above British retail prices, and I am told that the average price increase is 30 per cent. But this money buys little service; from the consumer’s viewpoint it would be hard to imagine a commercial group less interested in his needs than Canadian publishers. Telegraphic en- quiries get surface mail replies if any, letters may languish for weeks. Foreign publishers have similar and more serious complaints about Canadian businesses which claim to act as distributors for them. Neither principal nor client is served. If and when books from a foreign publisher eventually arrive through a Canadian distributor they may be the wrong edition or the wrong book or both, they may be so damaged that they must be returned, they will probably be too few and too late for the purpose for which they were ordered, and the price will certainly be an anti-educational shock. Such businesses should be investi- gated, not subsidized. In respect of many one wonders, not that they go out of business, but that they have managed to stay in it for so long. In most countries most of those who deal in books have an interest in education and take pleasure in the contributions they can make to it. In Canada this attitude is rare, though it pays no less than elsewhere; more often, distributors here prove to be inhibitors. The book trade should be cooperating with librarians to, in three words, make Canadians read: surely our climate is favourable. Clearly, until this is done it will remain possible, both at the personal and at the public level, for a nationalistic Canadian to both publish and perish. Brian Hocking CARABIDAE (COLEOPTERA) OF AN ARABLE FIELD IN CENTRAL ALBERTA J. H. FRANK Sugar Manufacturers ’Association Research Department Mandeville P. O. , Jamaica Quaestiones entomologicae 7: 237-252 1971 Sixty-three species of carabid beetles were collected in pitfall traps in an arable field in central Alberta. Data on life histories or biology are given for 26 of the more abundant species and discussed, and population densities are estimated for several of these. Six spe- cies of staphylinid beetles and 12 of spiders also taken in the study are listed. The spider species included Xysticus californicus Keys, a first record in Canada. Dreiundsechzig Arten von Laufkafern (Carabidae) wurden in Fallen auf einem Ackerland in Mittelalberta gesammelt. Angaben iiber die Leben oder Lebensgewohnheiten wurden fur 26 der haufigeren Arten angegeben und erlautert; ausserdem wurde die Populationsdichte fur verschiedene von ihnen geschdtzt. Sechs Arten von Kurzfiuglern und 12 Spinnenarten wurden ebenso in die Studie einbezogen und aufgefuhrt. Die Spinnenarten schlossen Xysti- cus californicus Keys, eine Art, die zum ersten Mai in Kanada entdeckt wurde, ein. Larvae and adults of most species of Carabidae prey upon other insects. As part of a study of the economic importance of Carabidae as predators of cutworms, a study was made of the species inhabiting an arable field in Central Alberta. Mention is made of Staphylinidae and of spiders from the same habitat. This was selected following a report of a cutworm attack on a field of barley near Calahoo, Alberta. The designation of the site is: section SW8, township 55, range 27, meridian W4, which is roughly 32 km (20 miles) northwest of Edmonton. The area is mapped as a mixture of three soils: two formed on fine-textured, stone-free lacustrine sediments (these are Mico silty clay loam, comprising 50% of the area and Maywood clay loam, comprising 20%); the other on a loam-textured, stony glacial till (this is Cooking Lake loam and comprises 30% of the area). The species of cutworm was determined as Euxoa ochrogaster Guenee (Lep; Noctuidae), the red-backed cutworm. A study area of about 0.2 hectare (0.5 acre) was used, near the centre of the field. Partly because of its smallness and partly because of agricultural practices, which inhibit growth of other than the crop plant, the area was very uniform. The duration of the study was the frost-free periods of 1967 and of 1968, from June 1967. In 1968 a crop of oats was grown in the field. Frost occurred from early November 1967. In 1968 thawing was during the first week of May with more or less permanent frost from the first week in October. Frost limited the movement of Carabidae severely, so that the operation of pitfall traps during frost periods THE STUDY AREA THE CLIMATE 238 Frank was unproductive. Meteorological information was supplied by the Weather Office, Inter- national Airport, Edmonton. Maximum temperatures occurred in September 1967 (max. 33.6 C, 92.5 F), but in July 1 968 (max. 3 1 .0 C, 87.8 F). Minimum temperature of the study period was -32.3 C (-26.1 F), in January 1968. This air temperature range of 65 C (over 118 F), although modified by the field crop, must impose severe restrictions on the move- ment of surface active insects. Refuge from temperature extremes is provided by the ability of the adult and larval carabids to burrow into the soil. From the temperature aspect the season was more ‘advanced’ in 1968 than in 1967. The rainfall patterns were similar over the two years, with July and August being the wettest months (total 120-145 mm, 5-6 inches, for the two months together) but September 1967 was an extremely dry month (0.76 mm, 0.03 inch) compared with September 1968 (37.34 mm, 1.47 inches). METHODS The principal means of capturing beetles was a grid of 10 x 10 unbaited pitfall traps spaced at 1 m intervals. This was in place throughout the study period except when agricultural operations (ploughing, seeding, and harvesting) were in progress, at which times it was taken up, to be replaced as soon as possible. The traps were of high impact poly- styrene plastic and were 10.2 cm in height, 8.7 cm (O.D.) at top and 7.2 cm (O.D.) at base. The shape allowed stacking for transportation and the light weight also facili- tated this. Individual traps which had become soiled while in use were replaced by clean traps as necessary, although the plastic resisted adherence of mud. A vinyl square 10.8 x 10.8 cm (quarter of a 9 inches x 9 inches floor tile) was supported on four small wooden stakes of length about 10 cm, at a height of about 2.5 cm above each trap to keep it dry. Insects caught in the pitfall traps were removed from the study area for identification and were not returned unless they were to be used for mark, release, and recapture studies. Those not returned were used to establish and maintain cultures, or were dissected. Population estimates were made in a way similar to that described by Frank (1967). Released beetles had been marked by cutting a minute notch in the right elytron with spring scissors instead of marking with paint. Larvae taken in pitfall traps were reared individually, at first in plaster of Paris blocks, later in 60 x 15 mm plastic petri dishes contained in a glass battery jar to maintain an adequate level of humidity. Entomophagous larvae were fed larvae of Musca or pieces yof Periplaneta, and phytophagous larvae (species of Amara and of Harpalus would not survive for long on a diet of insects) were fed pieces of maize (corn) kernel. An identification label was attached to the inside cover of each petri dish with a smear of vaseline. Food had to be replaced daily to prevent growth of mould. Cultures of adults were maintained on a damp peat substrate in 8 x 17.5 x 4.5 mm transparent plastic boxes and were fed in much the same way as were the larvae. Eggs and larvae, when found, were removed from these cultures and treated as were the pitfall- trapped larvae. THE CARABIDAE Names of the 63 species collected arranged according to Lindroth (1961), with data on those less frequently collected, are given in Table 1. Data on the more abundant species follow. Carabidae of an arable field 239 Table 1. A list of names of carabid species collected in the study area near Calahoo, Alberta, with notes, numbers, and dates for the less frequently encountered species (mature adults except where otherwise noted). Name of species Notes Cicindela limbalis Klug Carabus maeander Fischer von Waldheim Carabus taedatus Fabricius Carabus serratus Say Calosoma calidum Fabricius Notiophilus semistriatus Say Notiophilus aquaticus Linnaeus Loricera pilicornis Fabricius Patrobus lecontei Chaudoir Bembidion nitidum Kirby Bembidion grapei Gyllenhal Bembidion bimaculatum Kirby Bembidion rupicola Kirby Bembidion obscurellum Motschoulsky Bembidion nudipenne Lindroth Bembidion rapidum LeConte Bembidion versicolor LeConte Bembidion quadrimaculatum oppositum Say Bembidion mutatum Gemminger and Harold Bembidion canadianum Casey Pterostichus lucublandus Say Pterostichus corvus LeConte Pterostichus adstrictus Eschscholtz Pterostichus femoralis Kirby Calathus ingratus Dejean Synuchus impunctatus Say Agonum quadripunctatum DeGeer Agonum retractum LeConte Agonum cupripenne Say Agonum cupreum Dejean Agonum placidum Say Amara lacustris LeConte 2 - 31.VII.1967 & 20.V.1968 3 - 30.VI. 1 967 & TT.VI. 1 968 5 - VIII-IX.196Z; 1 1 - V-VIII.1968 few - VII-VIII. 1967 & 1968; 1 instar III - 12.VII.1968 text p. 241 1 - 18.X. 1967 29 - X-XI.1967; 2 - VII. 1968 2 — VII. 196^ infrequently; 4 gravid 99 — 18. VII, 24. VII, & 21 .VIII.L968 text p. 241 3 - between X& XI. 1967 text p. 241 text p. 243 text p. 243 1 - 1967; 12 - VI. 1968; 7 - VII. 1968; gravid 9 — 26.IX.1968 2- VI. 1968 text p. 243 text p. 243 text p. 244 text p. 244 text p. 245 text p. 245 text p. 245 1 - 30.IX.1968 text p. 246 few, all brachypterous; 9 — 17.VII.1967 with 43 well-developed eggs; 9 - 21. VIII. 1967 with 40 eggs; 9 - 21. VIII. 1967 with tachinid larva parasite 1 - 26.X. 1967 1 - 18.X. 1967; 2 — VI. 1968 2- 17.VII. 1967 &5.VII. 1968 text p. 246 text p. 246 few - VII & VIII. 1967 & 1968; gravid 9 - 3.VIII.1968 text p. 246 Amara torrida Panzer 240 Frank Table 1 (continued) Name of species Amara latior Kirby Amara apricaria Paykull Amara avida Say Amara obesa Say Amara quenseli Schonherr Amara sinuosa Casey Amara farcta LeConte Amara patruelis Dejean Amara laevipennis Kirby Amara ellipsis Casey Amara littoralis Mannerheim Amara cupreolata Putzeys Amara convexa LeConte Amara pallipes Kirby Harpalus amputatus Say Harpalus funerarius Csiki Harpalus uteanus Casey Harpalus pleuriticus Kirby Harpalus desertus LeConte Harpalellus basilaris Kirby Trichocellus cognatus Gyllenhal Brady cellus lecontei Csiki Brady cellus congener LeConte Brady cellus species ? Stenolophus comma Fabricius Badister obtusus LeConte Chlaenius alternatus Horn Metabletus americanus Dejean Cymindis planipennis LeConte Cymindis cribricollis Dejean Notes text p. 247 text p. 247 text p. 247 1 gravid 9 - 28.VIII. 1967 30; gravid 99 - 28.VIII.1967 & 24.VII- 30.VIII. 1968; 7 parasitised each by single dipterous larva — V-VIII.1968 4 - 23 .VIII & 27.IX.1968 including 2 gravid 99 31 - V-VIII.1968; gravid 99 - V & 3.VII.1968; immature adults - 30. VIII. 1968; 2 66 each with 1 parasitic dipterous larva - 10&20.V.1968 text p. 248 1 - 1967; 2 - 1968, 1 immature (30.VIII) text p. 248 text p. 248 few - 30. VI & 5. IX. 1967 & VIII. 1968; gravid 9- 12.VII.1968 3 1 - X.1967; 4 - VII-VIII. 1968; gravid 9 - 24.VII.1968; 2 immature 99 - 30.VIII. 1968 text p. 249 few - VI-VIII; immature adult - 9.VIII.1968 few - IX. 1967 & VII. 1968; gravid 9 - 18.VII.1968 text p. 249 d- 18.VII.1968 6 — VI-VII.1968 text p. 249 1 - 2.XI.1967 5 - VI-VII.1968; 2 gravid 99 - 26.VII.1968 2 99, elytra iridescent — 27.X.1967 & 23.IX.1968 1 - 17.VI.1968 1 -VI. 1968 1 - 18.IX.1968 text p. 249 1 - 18.X. 1967 6 - 17.VII. 1967; gravid 9 - 31. VII. 1968 63 species Carabidae of an arable field 241 Calosoma calidum Fabricius No trace of this species was seen at Calahoo in 1967, but several examples were captured in 1968, with immature adults found on 16 and 26 August. Two eggs were laid by a captive 9 on 20 June 1968 and one of these was reared to the third instar. A few living larvae were taken in pitfall traps (between 5 July and 3 August) and reared to the adult state. The aver- age length of each stage was: egg 4 days, instar I 5 days, instar II 8 days, instar III 14 days, pupa 8 days. The life cycle is therefore complete in 5 to 6 weeks. No evidence suggests that there is more than one generation per year or that any larvae overwinter. Bembidion nitidum Kirby Specimens were trapped in June, September, and October 1967, and in May-July and again in late September-October 1968. The population density was estimated for 1968 as <1.0/m2. A final instar larva, found in a culture of adults in the laboratory, on 2 January 1968, pupated on 6 January, but became infected with a fungal growth by the middle of the following month and died. Nineteen other larvae were taken from the same culture which was replenished by the addition of newly-trapped adults, in summer 1968. These appeared between 24 and 28 May and then not again until the end of July. None was reared to the adult; one, taken as an instar I larva on 24 May, underwent ecdysis on 28 May and again on 31 May, but the instar III larva was unable to pupate successfully and died on 10 June. Of the larvae which appeared at the end of July, some had reached instar III by 19 August. An instar II larva was trapped on 23 August. Lindroth (1963) recorded immature beetles from early August and stated that hiberna- tion is probably imaginal i.e. that larval growth takes place in the summer. However, there seem to be two breeding periods indicated first by peaks in number or activity (i.e. pitfall trap catches) in early May and late June (Fig. 1) and second by instar I larvae produced in culture only in late May and in late July. If this interpretation is correct, the immature adults recorded by Lindroth were probably from the second generation. The larva found in culture in January 1968 was perhaps properly from the first annual generation which had been produced early because of warm temperatures in the laboratory. Likewise the number or activity peak of September-October is possibly an early manifestation of that of the following May which is interrupted by freezing temperatures. A situation in some ways the reverse of this is discussed by Mitchell (1963) where a few adults of Trechus quadristriatus (Schrank) (which is stated to breed in September) overwinter and breed in the following spring. The development period of the supposed second annual larval generation is not necessarily of the same length as that of the first and if a third takes place during the winter, this last would obviously be much lengthened. Bembidion bimaculatum Kirby Many more specimens of this species were taken in August than in any other month and it appeared to have a single peak of numbers or activity (Fig. 1). Captured 99 contained eggs between the end of July and early September, when as many as 17 mature eggs were contained in the abdomen of a single 9. Larvae were taken from a laboratory culture on 4 November 1967. Lindroth (1963) considered the larva to hibernate. The population size was estimated at <0.8/m2 in 1968. The related species B. sordidum Kby. was not recognised from Calahoo. 242 Frank Bembidion nitidum A Agonum placidu m 2 n I - O ^ Bembidion bimaculatum 2 -| Amara torrida 3 1 2 H I O Bembidion \ rupicola ~I 2 - I - o-l- Amara latior Amara apricaria Amara avida 2 -i I _ r \ Amara 0J J V_Patruelis. Amara ellipsis 2 - I - o-* Bembidion mutatum 2 -j I _ Amara littoralis Q J 2 n I Bembidion \ canadianum Harpalus amputatus Pterostichus lucublandus Harpalus pleuritic us 2 n 0 - 2 1 - O - ASONDJ FMAMJ J A SON ASONDJ FMAMJ JASON 1967 1968 1967 1968 Fig. 1. Pitfall catches of Carabidae over fortnightly periods (log. scale) x time. Carabidae of an arable field 243 Bembidion rupicola Kirby A common species at Calahoo, with an estimated population size (1968) of 2.6/m2. Adults showed a very similar pattern of numbers or activity to that shown by B. nitidum, with peaks in early May, late June, and October. Several pairs of adults were observed mating in pitfall traps on 10 May 1968. Larvae were taken from a culture of adults in the laboratory at various dates between 24 May and 10 June 1968, but none was reared successfully, so that the duration of the larval instars cannot be given. However, an instar III larva, taken at Worsley, Alberta, on 11 July 1967, pupated on 14 July and the adult emerged on 20 July. Lindroth (1963) recorded immature beetles between 22 July and 3 August and stated that adults hibernate. If an analogy is drawn with B. nitidum and B. quadrimaculatum , below, it would seem likely that the numbers or activity peaks of early May and late June (Fig. 1) correspond to two separate breeding periods. The first generation reaches the adult state in late June. Either these adults and their parents, or both, then breed giving rise to the second genera- tion which reaches the adult state at the end of July and beginning of August. This second generation may normally overwinter and perhaps breed in May of the following year, but a certain number of larvae may be produced in October and may overwinter. A single instar III larva trapped at Calahoo on 11 September 1968, pupated in the laboratory on 16 September and the adult emerged on 28 September. This raises the pos- sibility of a third generation in the year, but because of the absence of a peak in numbers or activity in early August, it seems more probable that the occurrence of this larva was exceptional. Bembidion obscurellum Motschoulsky Far fewer adults of this species were trapped in late 1967 than of B. rupicola, but the former were more numerous at the beginning of 1968. Either B. obscurellum adults were not influenced so greatly to activity by the warm autumn of 1967, or they survived the winter better. Population size between June and August 1968 was estimated at 3.9/m2. The pattern of activity appears to be identical to that of B. rupicola, with three peaks in numbers or activity, one in early May, the second in late June and a smaller third in October (Fig. 1). Like B. rupicola, adults were found mating in pitfall traps on 10 May 1968. Larvae and eggs were taken from a culture of adults on 24, 25 and 26 May and not again until 26-31 July. Lindroth (1963) recorded immature adults in late July in Alberta. Bembidion versicolor LeConte This species had a rather similar pattern of numbers or activity (Fig. 1) to that of B. nitidum. It showed a peak in early May, a second in late June and a third in November. The third peak had its beginning earlier in 1967 than in 1968. Gravid 99 were taken on 19 and 26 July 1968. No larvae were obtained. The species is somewhat hygrophilous. Specimens of the related species B. timidum LeConte were not collected at Calahoo. Bembidion quadrimaculatum oppositum Say By far the most numerous carabid at Calahoo, its population size was estimated to be 20/m2 in 1968 with a pattern of numbers or activity (Fig. 1) similar to that of B. 244 Frank nitidum. Several pairs of a laboratory culture were observed mating in late May 1968. Instar HI larvae were recovered from this culture on 13 June and one of these pupated on 20 June and the adult emerged on 1 July. A 9 taken in a pitfall trap on 26 July was found to be gravid. Larvae were again taken from culture between 26 and 31 July of which one pupated on 5 August but subsequently died. Rivard (1964) has indicated for the subspecies a breeding period extending from May until July in eastern Ontario. This was based on dissection of 72 females captured between May and November 1963, of which 24 were gravid. During most of this May-July period, according to Rivard’s Fig. 1 , no more than 50% of the female beetles dissected were gravid, except in early July when approximately 100% were gravid. In Rivard’s Fig. 1, a line is drawn through 10 points on a graph which relate to the 24 gravid beetles and a similar number of non-gravid ones. Thus a difference of one or two gravid beetles at any date could make a large difference to the shape of the graph line. The numbers of individuals taken in pitfall traps at Calahoo (Fig. 1) are based upon the summation of pitfall trap catches over fortnightly periods. The number taken in early May was 924; in late May 769; in early June 294; in late June 1234; in early July 872 and in late July a mere 66. Unless the substantial drop in numbers in early June and substantial increase in late June are related to breeding activity they are difficult to ac- count for; the explanation is hardly to be sought in direct temperature or rainfall variation effects. If no significant drop in numbers occurs, but merely a period of quiescence, then the second peak must represent a resumption of breeding activity by the same generation of adults. Thus a single generation would have two temporally (at least partially) separated breeding periods. If a real drop in numbers occurs, then the second peak must be associated with an increase in the adult population by the metamorphosis of pupae — but the continuance of breeding activity after this date is not disputed — and this suggests true bivoltinism. On this basis, because adults of this new generation must outnumber adults of the pre- vious generation by at least four to one (Fig. 1) in late June and thereafter, then no more than 20% of trapped females should be gravid from late June onwards, if there is no bivoltinism. Rivard found 100% of females captured to be gravid only in early July. The evidence points to the existence of two breeding periods at Calahoo. Several other species of Bembidion show a very similar pattern of numbers or activity. Bembidion mutatum Gemminger and Harold Less numerous than the similar B. quadrimaculatum . The numbers trapped were small particularly in 1968, but indicate numbers or activity peaks (Fig. 1) similar to those of the above species, perhaps slightly delayed. A gravid 9 was captured on 9 August 1968. Bembidion canadianum Casey As frequently trapped in late 1967 as B. rupicola, this species was only as numerous in 1968 as B. mutatum. The numbers taken in pitfall traps decreased from May 1968 until June, after which there was no activity until October (Fig. 1). The absence of a second summer peak in 1968 may indicate that the species is univoltine, if meaning can be inferred from the small numbers of individuals trapped. Carabidae of an arable field 245 Pterostichus lucublandus Say The population size of this species was estimated at <0.7/m2 in the study area in 1968. The adults were active only during May-July of 1968, but in the milder autumn of 1967 there was another peak of numbers or activity in late September (Fig. 1). Approximately equal numbers were caught during each half-month period between early May and early July and there is no evidence to suggest that there are two activity peaks with the corollary of bivoltinism. Rivard (1964) also arrived at this conclusion of univoltinism for this species and his Fig. 1 indicated that the breeding period extends from May into September in eastern Ontario. Females dissected on 20 May 1968 and 17 July 1967 contained large eggs. Mating of pairs caught in pitfall traps was observed on 21 May 1968 as well as in laboratory cultures during the last week of May. Eggs were produced by these cultures from 23 May until 5 June and from 8-16 July. The egg stage lasted 4-7 days, instar I 6-7 days, instar II 6-1 1 days, instar III 14-21 days and the pupa 8-10 days. Towards the end of July and in early August larvae were taken in pitfall traps frequently and one of these was reared to produce an adult on 23 Au- gust (pupal stage 7 days). Immature adults were taken in pitfall traps on 5 September 1968. Larvae (apparently instar III) were described briefly by Schaupp (1881), one of which pupated on 26 August and the adult emerged on 5 September; another pupated on 18 August and the adult emerged on 29 August. Pterostichus corvus LeConte Not a single example of this species was taken in the study area in 1967, but 16 were taken in May-July 1968. By analogy with the related P. lucublandus , the population size may have been of the order of 0.07/m2 in 1968. Eggs were obtained from a laboratory culture on 30 June, 14, 15 and 26 July and 3 August 1968. The average duration of each stage was as follows: egg 6 days, instar I 6 days, instar II 6 days, instar III 16 days, pupa 7 days. Because of the small numbers trapped it is not possible to specify the breeding period or periods with any certainty. The northernmost locality given by Lindroth (1966) for the occurrence of this species in Alberta is Morin, some 150 miles SSE of Calahoo. Pterostichus ads trie tus Eschscholtz More than twice as numerous at Calahoo as P. lucublandus (estimated population size in 1968 1.8/m2), but apparently with a very similar life cycle. The second numbers or activity peak of late September 1967 shown by P. lucublandus was even more marked in this species and was repeated to some extent the following year (Fig. 1). There was a very slight diminu- tion in numbers trapped from late May until June. Gravid 99 were found as early as 10 May and as late as 17 July, while immature adults were trapped from 21 August until 5 September. Adults in a laboratory culture produced eggs between 23 May and 26 July and the resultant larvae were reared in the same manner as those of P. lucublandus. The egg stage lasted 4-8 days, instar I larva 4-6 days, instar II larva 4 days, instar III 17-18 days and pupa 6-7 days. Adults reared from instar III larvae taken in pitfall traps emerged at the end of July and beginning of August. The related P. pennsylvanicus LeC., although abundant at George Lake, Alberta, a marshy, wooded locality little more than 16 km (10 miles) NNW of Calahoo, was never found in the study area. 246 Frank Calathus ingratus Dejean A few individuals only were taken, in June-August of both 1967 and 1968; the normal habitat is wooded localities. Two 99 (with very reduced wings) were trapped on 17 July 1967, one of which contained eight large eggs, the other five. Another 9, taken on 24 July 1968, was also gravid. A (macropterous) 9, taken on 21 August 1967, contained no eggs, neither did brachypterous 99 taken at Scandia and at Tofield, Alberta in April 1968. Agonum cupreum Dejean Estimated to have a population size of 1.0/m2 in 1968 at Calahoo, this species seemed to be less abundant than during the previous year. All examples from Calahoo, in which the wings were examined, proved to be macropterous. None of the 99 caught at various states between 21 April (1968) and 17 July (1967) contained mature eggs (though one contained the pupa of a parasitic hymenopteron), but a 9 taken on 21 August (1967) contained (only) two eggs, perhaps indicating the approaching end of the breeding season. An instar III larva, trapped on 30 August 1968, pupated on 15 September and the adult emerged on 21 September. An immature adult was taken in a pitfall trap on 16 September 1968. A pair was observed in copula, in the laboratory, on 20 November 1967. Lindroth (1966) recorded immature adults at the end of July and in August. The graph of numbers or activity (Fig. 1) is difficult to interpret and it may be that there is a single breeding period extending from May into July, but numbers trapped are too small for certainty. The September- October peak possibly represents early incidence of the ac- tivity of the following year. If breeding occurs from May to July, immature adults would be expected from late July until late September. The apparent decline in numbers between 1967 and 1968 might be the result of unsuccessful overwintering possibly as a result of the high and perhaps untimely level of activity in late 1967. Agonum placidum Say Less frequently captured in 1967 than was A. cupreum , the situation was reversed in 1968, when A. placidum was more often captured and its population size was estimated at <1.5/m2. Females with eggs were taken on 17 July 1967, the maximum number of eggs recorded per 9 being 20. Of 20 99 captured on 14 April 1968 at Scandia, Alberta, none contained eggs, but one contained two nematode parasites. Eight instar III larvae caught between 23 August and 9 September 1968 produced adults between 4 Sept, and 17 Sept., the average time for the pupal stage being 6 days. A numbers or activity peak in July is well marked in this species (Fig. 1) and there is some evidence of a minor peak in the year (October 1967 and September-October 1968). The breeding period seemed to be restricted to July in 1968 at Calahoo. Rivard (1964) found gravid 99 between June and September in eastern Ontario. Lindroth (1966) quoted records of gravid 99 from June to August in southern Ontario. Amara torrida Panzer Lindroth (1968) wrote that the habitat of this species was similar to that of Pterostichus Carabidae of an arable field 247 adstrictus. Certainly both species were common at Calahoo. The population density of A. torrida was estimated by the mark, release and recapture technique as <0.6/m2 in 1968. In the same year, numbers taken in pitfall traps gradually rose to a single peak in early August and then declined more rapidly (Fig. 1). Gravid 99 were trapped in 1967 on 17 July, 21 August, 24 August and 1 September, while in 1968 gravid 99 were taken on 19 July and 9 August, but 99 trapped on 20 May and 30 September were not gravid. The maximum number of eggs contained by a single 9 was 20 (24 August 1967). Pairs were observed in copula on 6 August 1967 and 16 August 1968. A pitfall trap emptied on 21 August 1968 contained a single adult 9 A torrida together with two eggs (no other beetles present) which, unfortunately, did not prove to be viable. Five eggs were removed from a culture of adults in the laboratory, between 30 July and 2 August 1968, but these too were not viable. An immature adult was trapped on 17 July 1968. This bears out the suggestion by Lindroth (1968) of larval hibernation and indicates a lengthy larval growth period. The single peak in numbers trapped, in early August, may w^ll indicate the coincidence of the peak of the breeding period with the peak period of emer- gence of the new generation of adults. Adults, each with a single parasitic dipterous larva, were captured on 10 May 1968 (1 example); 20 May 1968 (1 example); 9 August 1968 (3 examples). A male captured on 9 August 1968 was the host of a nematode. Amara latior Kirby Only a few individuals of this species were trapped in either year at Calahoo, the greatest number appearing in early July (Fig. 1). A gravid 9 was taken on 5 September 1967. A larva, taken from soil on 1 1 July 1967 at Worsley, Alberta, pupated the same day and the adult emerged on 17 July. Lindroth (1968) recorded gravid 99 in September and October in Ontario, while Rivard (1964) recorded others during August-October in eastern Ontario. Amara apricaria Paykull This species was commoner at Calahoo than species of Amara previously discussed, but numbers trapped were still too low to allow interpretation of the pattern of numbers or activity (Fig. 1). A single gravid 9 was taken on 17 July 1968. The larva was described briefly by Schi^dte (1867) but the date of capture was not given. An individual parasitised by a dipterous larva was captured on 10 May 1968. Amara avida Say This species was apparently commoner at Calahoo in 1968 than was A torrida (<0.9/m2) and like that species with a single peak in numbers or activity (Fig. 1). The peak, however, occurred in early July, a month earlier than the A. torrida peak. Some evidence of a second peak in September-October was seen in 1967 but not in 1968. Gravid 99 were taken on 17 July, 21 August and 24 August 1967 and 9 August 1968. As many as 1 1 eggs were dissected from a single 9. Several eggs were laid singly by a 9 in a laboratory culture on 3 August 1 968 and some of these hatched between 9 and 1 2 August, but none of the resultant larvae survived instar I. A single immature individual was taken among 17 mature adults on 10 July 1968. Lindroth (1968) noticed immature adults in June in British Columbia and Alberta. Two individuals, each parasitised by a dipterous larva, were captured on 9 August 1968. 248 Frank Amara patruelis Dejean Specimens of this species were most frequently taken in late September and early Octo- ber. Two gravid 99 were taken at Calahoo on 17 July 1967 and two others on 21 April 1968 at Mill Creek, Edmonton. A pair was found in copula on 6 June 1968 and an immature adult was taken in a pitfall trap on 16 August 1968. Lindroth (1968) described A. patruelis as “a pronounced spring species” and stated that adults hibernate. It appears as if the larvae have a summer development period in contrast to the foregoing species of the subgenera Curtonotus C Amara aulica group Lth.) e.g. A. torrida , and Brady tus {Amara apricaria group Lth.) e.g. A. avida , but the pattern of numbers or activity (Fig. 1) is difficult to interpret because of the small numbers trapped. Three 66, the first captured on 20 May 1968. the others on 1 1 September 1968, each contained a single parasitic dipterous larva. Amara ellipsis Casey Not uncommon at Calahoo. The greatest number of individuals to be taken in a two- week period was in early October 1967 and a peak at this time of year was repeated in 1968 (Fig. 1). This late-year peak is a possible forerunner of the small peak of early May. There appears to be a second peak in July. There is similarity in this to the condition shown in the Bembidion species and on the basis of this and the following evidence two breeding periods per year are suggested. An overwintering 9, hand collected on 3 April 1968, contained eggs which were not fully developed. Pitfall trapped 99 yielded eggs on 10 May and on 3 and 15 July 1968; none was trapped between the middle of May and the end of June; no 9 trapped later in the year than July was gravid. An adult emerged successfully on 19 July 1967 from a pupa which had been collected as a larva on 22 June, the date of pupation being 7 July. An immature adult was taken at Calahoo on 24 July 1968. Lindroth (1969) found immature adults at the end of July. Three 66 captured on 26 July 1968, 21 August and 30 September each contained a single parasitic dipterous larva. Amara littoralis Mannerheim Present and active in largest numbers in July at Calahoo, the commonest species of sub- genus Amara {Amara lunicollis group Lth.). It seems to have a life cycle similar to that of A. ellipsis , with a peak in numbers or activity (Fig. 1) in early May and a second one in July. A decline in numbers trapped in early August is not accounted for unless the peak of late August is the result of the emergence of adults of the second generation. Two breeding periods per year are suggested. Dissected 99 contained eggs on 10 May and on 20 May 1968; none was trapped thereafter until early July. In the latter month 99 contained eggs on 3, 10, 12, 15, 18, 24, 26 and 31 July, but not thereafter. Lindroth (1968) found immature adults in July and at the begin- ing of August. A 9 taken on 17 July 1967 contained a larva of an unidentified tachinid, which occupied about half the volume of the host’s abdomen. The empty chorion of the tachinid egg lay under the right elytron of the host. Four 66, captured on 24 July 1968, 31 July (2 ex- amples) and 3 August each contained a single parasitic dipterous larva. Carabidae of an arable field 249 Harpalus amputatus Say According to Lindroth (1968) a xerophilous species, the most abundant of the large carabids at Calahoo in 1968, when its population size was estimated at 4.2/m2. The pattern of numbers or activity probably indicates a single peak in early June (1968) and extending from May until July, with slight fluctuation. A slight autumnal peak was seen in 1967 (Fig. 1), but the species was not trapped in that year until early September. Dissected 99 contained eggs on 8 May 1968, 17 July 1967, 9 August 1968 and 21 August 1967. Unfortunately no females captured in June 1968 were put aside for dissection, but all were used to establish cultures or in population estimations. The presence of gravid 99 in June would have established the existence of a single breeding period as indicated by the pitfall trap captures. Harpalus pleuriticus Kirby The population size of this species at Calahoo in 1968 was of the order of 1.0/m2. The species is thus not only of smaller size but was of smaller population size than H. amputatus. It reached the zenith of what may be a single extensive peak in numbers or activity in July (Fig. 1). If this is, in fact, a single peak, then the breeding period may extend from May until August, as seems to be indicated with H. amputatus. An autumnal peak was seen in late September 1967. Dissected 99 contained mature eggs on 17 July 1967, 12, 15, 17, 18, 19 and 24 July and 9 August 1968, but not thereafter and not in May 1968. As with H. amputatus , no 99 captured in June were set aside for dissection. A pair was observed mating in a laboratory culture on 8 July 1968. Immature adults were taken in pitfall traps on 18 and 24 July, 9 and 16 August 1968. Adults captured on the following dates in 1968 contained parasitic dipterous larvae: 20 May (1 example); 9 July (1 example); 24 July (2 examples each with 2 parasites); 9 August (1 example with 2 parasites). Trichocellus cognatus Gyllenhal The adults of this species were taken at Calahoo only in late September-November, when large numbers (387 during October 1967, 465 during October 1968) were caught in pitfall traps. Freezing interfered with assessment of population size. A pair was found mating in a pitfall trap on 2 October 1968. Lindroth (1968) stated that immature beetles are abundant from mid-July to the end of August, but this was not so at Calahoo. Larvae were taken from culture between 1 and 17 October 1968. Oviposition might well take place in October and November when temperatures allow, and larvae may overwinter. Metabletus americanus Dejean Adults of the species were present and active in May to mid-July and late September to November only (Fig. 1). None was captured between mid-July and late September, but a 9 taken on 19 July 1968 was gravid. In the laboratory six instar I larvae were taken from a culture of adults between 10 and 18 July 1968, but none was reared successfully. Several instar I larvae were taken from culture on 20 November 1968, of which two underwent ecdysis to instar II on 5 December. 250 Frank There was a decline in pitfall trap captures in early June, similar to that of some Bem- bidion species, so that two generations per year may be indicated. Some larvae may be pro- duced in May and early June. Other Arthropods The following species of Staphylinidae were captured in the study area: Leptacinus batychrus Gyll., Philonthus occidentalis Horn, P. concinnus Grav., P. furvus Nordm., Que- dius spelaeus Horn, Tacky porus sp. The third, fourth, and fifth of these were trapped only occasionally. Spiders taken in the study area were: Trochosa terricola Thor., Pardosa groenlandica (Thor.), P. mackenziana Keys, P. moesta Banks, Pardosa sp. nr. saxatilis (Htz.), Pardosa sp. of metlakatla complex, Gnaphosa sp. nr. muscorum (Koch), G. parvula Banks, Micaria sp. nr. alberta Gertsch, Xysticus californicus Keys (1 9, 21 June 1968, first Canadian record), Paraphidippus marginatus (Walck.). Pardosa groenlandica was the most frequently trapped of these. DISCUSSION Life cycles Perhaps more ecological studies of Carabidae have been made than of any other family of Coleoptera, but the volume of knowledge of their life cycles is slender. Rivard (1964) concluded that for 13 species studied in eastern Ontario, each had only a single generation per year, based on occurrence of mature eggs in the ovaries of adult 99. He classified them into two major groups, spring breeders and autumn breeders, with a small third group overlapping the two periods. Greenslade (1965) arrived at similar conclusions in a study of 26 species of Carabidae in England, based on observations of activity periods of adults and larvae through numbers of individuals caught in pitfall traps and the association of these activity periods with breeding activity. However, Heydemann (1963) considered a number of species of carabids in maritime Germany to have more than one breeding period per annum. He cited some species of Bembidion and observed that (apparently in respect of insects in general) while carnivorous forms are mostly univoltine, saprophagous forms are frequently bivoltine or polyvoltine. Rivard’s (1964) assumption that the breeding periods are coincident with the presence of mature eggs in [a high percentage of] 99, with perhaps a 2-3 weeks delay between the maturity of the eggs and the onset of oviposition, is doubtless a good approxima- tion for the population as a whole; but no allowance is made for periods of inactivity e.g. if gravid 99 are inactive over the winter before oviposition. Nor is distinction made between different generations present in the population. Thus if the premise be expanded (beyond Rivard’s statements) one is led to infer that adult generation I breeds, producing eggs, larval instars, pupae and thus the resultant adult generation II. The fate of adult generation I is ignored, but possibly a considerable number of adults of generation I may survive and may breed again (at a time not necessarily coincident with the breeding pe- riod of generation II). This possibility is higher in species of lower reproductive potential or lower survival rate. The survival of these adults depends upon the combined effects of parasitism (including disease), predation, inter- and intraspecific competition, climatic factors and ‘natural senescence’. Lack of information on ‘natural senescence’ is a result of the difficulty of observing individuals in the field over long periods of time. Conditions in the laboratory must pro- Carabidae of an arable field 251 duce anomalous results. No individual beetle in the present study survived in the lab- oratory for much more than 1 year. However, an adult 6 and 9 of Omophron americanum Dejean (for which Rivard (1964) gives the breeding period as May to June) were cap- tured at Chappice Lake, Alberta, on 29 May 1967, along with five other adults, and survived in captivity until May 1969. These individuals must have been produced, as eggs, not later than the summer before capture (on Rivard’s evidence) and so their total life span was at least 3 years. The ‘culture’ of seven adults produced eggs in June 1967 (the resultant larvae subsequently died) but none thereafter. This, however, is no proof that the normal life span approaches (or exceeds) 3 years, nor that each generation of adults breeds only once. It merely shows that adults of generation I could still form part of the population at the breeding period of adults of generation II and even of generation III. Breeding periodicity in different parts of the geographical range of a species may differ and may or may not be directly influenced by climatic conditions. Heydemann (1963) (following Lindroth, 1945) remarked upon the different percentages of species with a sum- mer larval growth period or with a winter larval growth period between habitats with a maritime climate and with a continental climate. The implication was that in a harsher (continental) climate, a greater percentage of species has a winter larval growth period. Populations The principal interest of the present study lies in the observed dominance of Carabidae among the larger soil-surface inhabiting insects of the study area and in the large number of species apparently coexisting in an apparently very uniform habitat. On the assumption that there is at least some similarity of requirements between the Bembidion species, then there must be interspecific competition (Odum, 1959). If popula- tion size is taken as a measure of success in this competition, then B. quadrimaculatum, with a population size about double that of all other Bembidion species combined, must be seen as the most successful species. Similarity of requirements may not be unconnected with accepted taxonomic relationship. Nine of the 1 1 species belonging to different species groups of Bembidion (Lindroth, 1963) where there might be expected to be greater differ- ences in requirements, but B. rupicola and B. obscurellum belong to the same species group, as do B. quadrimaculatum and B. mutatum. Bembidion rupicola and B. obscurellum were of comparable abundance in the study area in 1968, not so with B. quadrimaculatum and B. mutatum. Few genera of Carabidae have been recorded in the literature as having restricted food preferences. It is to be expected that those species which were not merely stragglers into the area would have some food-chain relationship to the crop plant. Carabids might eat the crop plant or its seeds or stages of its decomposition, or they might be predators of other animals which do so. ACKNOWLEDGEMENTS Gratitude is due to G. E. Ball, T. L. Erwin, and D. R. Whitehead for assistance in the identification of Carabidae and to R. E. Leech for the identification of spiders. Carl and Henry Bergstreiser of Calahoo kindly allowed the study to be carried out in their field. Technical assistance from A. C. Schaaf, B. M. Rolseth, and Mrs. T. L. Erwin is gratefully acknowledged. The study was financed by a grant to R. H. Gooding from The Alberta Agricultural Research Trust (grant No. A. R. 67-31). 252 Frank REFERENCES Frank, J. H. 1967. The insect predators of the pupal stage of the winter moth, Operophtera brumata (L.) (Lepidoptera: Hydriomenidae). J. Anim. Ecol. 36:375-389. Greenslade, P. J. M. 1965. On the ecology of some British carabid beetles with special reference to life histories. Trans. Soc. Br. Ent. 16:149-179. Heydemann, B. 1963. Die biozonotische Entwicklung vom Vorland zum Koog. Vergleich- end-okologische Untersuchungen an der Nordseekuste. 2. Kafer (Coleoptera). Akad. Wiss. Lit. Mainz; Abh. math, naturwiss. (1962) 1 1 : 173-370 + pis. I-III. Kryger, J. P. and H. P. S. S0nderup. 1945. Biologiske Iagttagelser over 200 Arter af danske Billelarver. Ent. Medd. 24:175-261. Lindroth, C. H. 1945-49. Die fennoskandischen Carabiden. Eine tiergeographische Studie. I-III. Goteborgs k. Vetensk.-o. Vitterh. Samh. Handl., (B)4( 1-3): 1-709, 1-227, 1-911. Lindroth, C. H. 1961. The ground beetles of Canada and Alaska, 2. Opusc. ent., Suppl. 20, 200 pp. Lindroth, C. H. 1963. The ground beetles of Canada and Alaska, 3. Opusc. ent., Suppl. 24, 208 pp. Lindroth, C. H. 1966. The ground beetles of Canada and Alaska, 4. Opusc. ent., Suppl. 29, 240 pp. Lindroth, C. H. 1968. The ground beetles of Canada and Alaska, 5. Opusc. ent., Suppl. 33, 295 pp. Mitchell, B. 1963. Ecology of two carabid beetles, Bembidion lampros (Herbst) and Trechus quadristriatus (Schrank). I. Life cycles and feeding behaviour. J. Anim. Ecol. 32:289-299. Odum, E. P. 1959. Fundamentals of ecology. 2nd ed. Philadelphia, Saunders XVII + 546 pp. Rivard, I. 1964. Observations on the breeding periods of some ground beetles (Coleoptera: Carabidae) in Eastern Canada. Can. J . Zool. 42:1081-1 084. Schaupp, F. G. 1881. Description of the larvae of Pterostichus lucublandus and Pt. mutus. Say. Bull. Brooklyn ent. Soc. 3:88-89 + pi. Schi0dte. J. C. 1867. De metamorphosi eleutheratorum observationes: Bidrag til insekternes Udviklingshistorie (Fortsaettelse). Naturhist. Tiddskr. (3)4:415-552 + pis. 15-22. CORRECTION FOR DRAG OF A FLIGHT MILL, WITH AN EXAMPLE FOR AGROTIS ORTHOGANIA MORR. (LEP. NOCTUIDAE) MARTIN A. CHANCE Department of Entomology University of Alberta Quaestiones entomologicae Edmonton 7, Alberta 7 : 253-254 1971 A method of measuring the drag-speed relationship for an insect flight mill system is reviewed. Compensated free flight speeds and ranges for Agrotis orthogonia Morrison are given for Jacobson's (1965) data. Compte rendu du rapport resistance-vitesse pour insectes montes sur un manege. De plus, je propose pour les donnees de Jacobson (1965) un facteur de correction pour la vitesse de vol en liberte et pour les distances de Agrotis orthogonia Morrison. The use of insect flight mills as described by Hocking (1953) necessitates a consideration of mill drag. The additional drag of the device substantially lowers flight speed, and should be corrected for as follows. The difference between power required to maintain a mill alone, and a mill loaded with a dead insect at a constant speed, is the power required to overcome drag on the insect alone at that speed. Free flight speed would be the speed at which power to overcome drag on the insect alone would equal power to overcome drag on the mill-insect combination at the observed mill speed. Correction for this requires the measurement of drag-speed relation- ships for the mill itself and for the insect. I tested the mill used by Jacobson (1965). It is similar to those described by Hocking (1953), but gives a flight circumference of 0.893 meters. With the mill pivoted on a steel needle, the mill tip sweeps out a horizontal circle. Following Hocking’s (1953) procedure, about the glass stem I wound two natural silk threads, passed these over light weight pulleys on either side of the glass stem, and attached a series of weights to their opposite ends. By measuring the terminal tip speed for each pair of weights the drag at this speed can be obtained from the following relationships: , 2Wrg , „ dc 2Wgrc d = — - — and P = — = — - — Rt where, d = equivalent drag on the system at the arm tip at terminal speed (kg m sec-2 ) W = weight on each thread (kg) r = radius of the glass stem (m) g = acceleration due to gravity (9.8 m sec"2) R = length of the mill arm (m) P = power to maintain a terminal tip speed (Joules sec'1 ) c = circumference swept out by the insect mount on the mill arm (m) t = time for one revolution (sec) I used the means of two readings taken in each direction of rotation at each loading to plot drag-speed relationships (Fig. 1). For speeds too high for counting by eye I used Hocking’s photocell system. I plotted power against terminal speed for the insect and for the mill with a dead insect, without wings or legs, mounted on it in flight attitude. The appropriate drag value, i.e. that on the insect alone or that on the insect plus ‘tip equivalent drag’ for the mill, multiplied by the tip speed gives the power values. As suggested by Hocking (1953) by comparison of the mill mounted insect speed to the free insect speed at equal power, the free flight speed can be estimated. The dotted lines on Fig. 1 illustrate this procedure. Free flight speeds estimated, a proportional correction can also be made to equivalent flight ranges. An ex- ample using Jacobson’s (1965) data follows. 254 Chance LINEAR SPEED m sec'1 Fig. 1. The drag-speed relationships for A. orthogonia and Jacobson’s (1965) flight mill. The maximum mill speed of male Agrotis orthogonia Morrison is given at 4.5 mph or 2.01 m sec'1 . From Fig. 1 the equivalent free speed is found by the projection at equal drag from the insect-mill curve to the free flight curve (dotted lines). The equivalent free flight speed is 2.47 m sec'1 or 5.5 mph. The mean free flight speed of the male is 20% faster than the mill speed, and the equivalent flight range is then 17.6 miles. Equivalent Free Flight Speeds and Ranges of A. orthogonia Mill values Free flight values male female male female Mean speed (mph) 2.5 1.8 3.0 2.1 Max. speed (mph) 4.5 4.8 5.5 5.9 Max. range (miles) 14.7 3.5 17.6 4.1 REFERENCES Hocking, B. 1953. The intrinsic range and speed of flight of insects. Trans. R. ent. Soc. Lond. 104:223-345. Jacobson, L. A. 1965. Mating and oviposition of the pale western cutworm, Agrotis ortho- gonia Morrison (Lepidopera: Noctuidae), in the laboratory. Can. Ent. 97:994-1000. BIOLOGY AND HOST-PLANT RELATIONSHIPS OF AN OLIGOPHAGOUS LEAF MINER PHYTOMYZA MA TRICARIAE HENDEL (DIPTERA:AGROMYZIDAE) VINOD K. SEHGAL Department of Entomology Un i versi ty of Alberta Edmonton 7, Alberta Quaestiones entomologicae 7 : 255-280 1971 The members of this species feed in nature only on representatives of the plant genera Achillea, Chrysanthemum, Matricaria and Tanacetum belonging to the tribe Anthemideae, family Compositae. Natural incidence, mating, adult and larval feeding, oviposition, life cycle and immature stages on Tanacetum vulgare L. were observed. Thirty-eight plant species belonging to 17 families were tested for acceptability to gravid females for feeding and oviposition. Comparison of index of acceptability for feeding and oviposition with an index of plant relationship show that only plants closely related phylogenetically to the natural host-plant were acceptable for feeding and oviposition. This ovipositional preference by adult females is probably due to host-specific substances present only in selected plants. Studies on feeding and ovipositional preference by adult females when offered a choice of six acceptable plants show that maximum numbers of feeding punctures were on the plant from which flies were bred. Other preferred plants also served as hosts in nature. Two genera, Artemisia and Helianthus, though acceptable but not preferred in these studies do not serve as hosts in nature. The larva being a completely internal plant feeder is unable to select a more suitable food plant which might be available in its range. First instar larvae were manually transferred from the natural host-plants in various test plant species. Comparison of the success index for larval development with the index of plant relationship show that some plants widely removed phylogenetically but presumably lacking toxic or inhibitory substances were nutritionally adequate for completion of larval development. Die zu dieser Art gehorenden Insekten ernahren sich unter naturlichen V erhaltnissen nur von Pflanzen der Gattungen Achillea, Chrysanthemum, Matricaria und Tanacetum die zu der Familie Compositae, Tribus Anthemideae gehoren. Vorkommen, Paarung, Nahrung der Imagines und der Larven, Eiablage, und Entwicklungsstadien einschliesslich der Larven und Puppen, wurde unter naturlichen Bedingungen an Tanacetum vulgare L. beobachtet. Bezu- glich ihrer Eignung fur Nahrungsaufnahme und Eiablage wurden 38 Pflanzenarten von 17 Familien fur ihre Anziehungskraft auf das Trachtige Weibchen untersucht. Ein Vergleich des Indexwertes der Anziehungskraft fur Frass und Eiablage mit dem des Verwandtschaftsgrades der Pflanzen zeigt, dass nur solche Pflanzen fur Nahrungsaufnahme und Eiablage angenom- men werden, die phylogenetisch mit der Wirtpflanze nahe verwandt sind. In Hinsicht auf die Eiablage ist diese Bevorzugung durch das Weibchen wahrscheinlich in wirtspezifischen Sub- stanzen zu suchen, die nur in ausgesuchten Pflanzen vorhanden sind. Wenn eine Wahl von 6 bekommlichen Pflanzen fur Nahrungsaufnahme und Eiablage geboten wurde, zeigte es sich in der folgenden Untersuchung, dass die grosste Anzahl von Einstichen in den Pflanzen gefunden wurde, auf denen die Fliegen vorher herangezogen worden waren. Andere bevor- zugte Pflanzen dienten in naturlicher Umgebung auch als Wirte. Zwei Gattungen, Artemisia und Helianthus, zwar geeignet aber nicht bevorzugt in den Versuchen, dienen in der Natur nicht als Wirtpflanzen. Da die Larve ausschleisslich im Inneren der Pflanze Nahrung sucht, kann sie nicht einen anderen, in der nahe vorhandenen, besser geeigneten Wirt aussuchen. Larven die sich im ersten Stadium vefanden, wurden manuell vom naturlichen Wirt auf die Versuchspflanze ubertragen. Ein Vergleich der Erfolgszifferwerte der Larvenentwicklung mit denen des Verwandtschaftsgrades der Pflanze, zeigte, dass einige phylogenetisch nicht sehr nahe Pflanzen nahrungsmassig bekommlich genug waren, um eine voile Entwicklung der 256 Sehgal Larven herbeizufuhren. Es ist anzunehmen, dass in diesem Falle toxische oder hemmende Substanzen nicht vorhanden waren. Most of our knowledge of the biology of agromyzid flies is due to the late Professor E. M. Hering who in 1951 reviewed all existing information and compiled an extensive bibliogra- phy on this subject. Numerous other workers have studied the biology of many leaf mining species in detail (Webster and Parks, 1913; Smulyan, 1914; Cohen, 1 936; Ahmad and Gupta, 1941; Allen, 1956; Oatman and Michelbacher, 1958, 1959; Tauber and Tauber, 1968). Phytomyza matricariae Hendel is an oligophagous species whose members feed in nature around Edmonton, Alberta, Canada, only on the representatives of the tribe Anthemideae, family Compositae. The plant species attacked are Achillea millefolium Michx. (Fig. 1), Achillea sibirica Ledeb. (Fig. 2), Achillea sp. (cultivated variety), Chrysanthemum sp. (culti- vated soft leaf variety), Matricaria matricarioides (Less.) Porter (Fig. 3), and Tanacetum vul- gare L. (Fig. 4). Hering (1957) lists Achillea, Anacyclus, Anthemis, Cotula and Matricaria as European hosts for the members of this species. However, the identity of flies bred from some of these hosts probably needs confirmation by examination of the male genitalia. Spencer (1969) lists Tanacetum also as a European host. Literature on various aspects of host selection, feeding, and host preference in phyto- phagous insects has been reviewed by many authors (Lipke and Fraenkel, 1956; Friend, 1958; Thorsteinson, 1960; Kennedy, 1965; Dethier, 1966, 1970; Schoonhoven, 1968). Verschaffelt (1910) for the first time demonstrated that host selection in Pieris brassicae and P. rapae is determined by mustard oil glucosides in cruciferous and related plant families. Since then the food ranges of many oligophagous insects have been explained by the botanical distribution of secondary plant chemicals (Fraenkel, 1959). Feeding or token stimuli which evoke special feeding responses in phytophagous insects have been investi- gated (Thorsteinson, 1953; Sugiyama and Matsumoto, 1959; Nayar and Fraenkel, 1962, 1963; Harris and Mohyuddin, 1965; Keller and Davich, 1965; Stride, 1965) and together with deterrents play an important role in determining the selection of host-plants (Thor- steinson, 1960; Jermy, 1961, 1964). Many recent studies on the host range of oligophagous species have shown that plants not closely related to natural host-plants may be acceptable for normal growth and development (Jermy, 1961, 1966; Hsiao and Fraenkel, 1968). The association between insects and their host-plants has been shown to be the consequence of the interaction of two independently mutating systems (Dethier, 1970), and of the evolu- tionary coadaptations to chemical interactions between them (Whittaker and Feeny, 1971). Most research in the field of insect host-plant relationships has been on external plant feeders. Agromyzids having evolved as exclusively internal plant feeders are more closely bound to plants than any group of external feeders and are therefore ideal for the study of insect-food plant relationships. The female agromyzid deposits an egg individually inside the tissue of a selected plant. The emerging larva, unlike that of external plant feeders, is unable to select a more suitable food plant which might be available in its ecological range. The larva either feeds on the plant tissue selected for it by its mother or dies. Although an agromyzid larva is not concerned with the selection of a suitable food plant, it is directly involved with its acceptance. These larvae are therefore most suitable for the study of then- potential to use various food plants for their development. Host-plant relationships in Agromyzidae have been discussed by Hering (1951), Nowa- kowski (1962), Spencer (1964) and Sehgal (1971). Among Agromyzidae, monophagy and oligophagy are both of common occurrence. Strict monophagy is rare outside of monotypic genera. Extreme polyphagy is also rare; all known polyphagous species are restricted in then- range of food plants. Most species therefore feed in nature on related plants. Biology of Phytomyza 257 Fig. 1-4. Leaf mines on natural host plants of Phytomyza matricariae. 1. Achillea millefolium Michx. 2. Achillea sibirica Ledeb. 3. Matricaria matricarioides (Less.) Porter. 4. Tanacetum vulgare L. 258 Sehgal MATERIALS AND METHODS Observations on biology, and host-plant relationships of adult females and larvae were made under laboratory conditions of 70 ± IF and 1 2 hours of daily illumination maintained inside a growth chamber. The flies used in these experiments were bred from Tanacetum vulgare L. Both greenhouse and field grown plants were used. Small cuttings from the test plants were kept in Sach’s culture solution for flowering plants. By changing the solution it was possible to keep the cuttings healthy during the test period. Biology of Phytomyza matricariae Hendel Observations on mating, oviposition, feeding habits, and life cycle, were made on Tana- cetum vulgare L. In order to determine the incubation period, individual leaves were caged with a large population of flies. After six hours the leaves were removed, examined for eggs, and the positions of individual eggs marked. The leaves were then observed at 12 hour intervals. The progress of larval mines was marked with different water soluble colors every 12 hours and the mine examined for moulted mouth hooks, which can be seen inside the mine by transmitted light under a binocular microscope. Duration of larval stadia was esti- mated from the positions of moulted mouth hooks. The time of moulting during any 12 hour period was estimated by measuring the relative length of leaf mine before and after the position of moulted mouth hooks. This method of recording larval activity has been used by Allen (1956) and Tauber and Tauber (1968). It was thus possible to estimate the duration of larval stadia, length of leaf mine excavated by different instars, and observe the mining habits of the larva. Leaf mines were fixed in ‘Formal Acetic Alcohol’ (F.A.A.) for micro- tomy. Host-plant relationship in adult females Range of food plants. - Small twigs of various plant species, bearing young leaves, were exposed individually to five gravid females inside a muslin cage, for a period of 24 hours. At the end of the experiment, flies were removed from the cages and the leaves examined for feeding punctures and punctures with eggs. Feeding and oviposition preference by gravid females. — A circular plastic petri dish 5 Vi inches in diameter was used as a choice chamber to test the feeding and oviposition prefer- ence of adult females. The young leaves of six different plants, grown under greenhouse conditions, were placed around the periphery of the dish equidistant from one another. The petioles of leaves were pulled out through small holes in the periphery of the dish and wrapped with cotton kept moist with distilled water. The plants used in this experiment were Tanacetum vulgare L., Achillea sibirica Ledeb ., Matricaria matricarioides (Less.) Porter, Artemisia sp., Chrysanthemum sp., (cultivated variety) and Helianthus annuus L. Five gravid females from a laboratory culture maintained on Tanacetum vulgare L. were used in each test after being isolated from their food plant for one hour. They were anesthetized with C02 and then introduced at the centre of the petri dish. Host-plant relationships of the larva Transfers of larvae from natural host-plants to test plants. - First instar larvae normally less than 24 hours old were used in these experiments. The supply of healthy first instar larvae was from plants in which eggs were laid in the laboratory. Field collected larvae were not used in order to avoid any early parasitization by braconids or chalcidoids. Only young and tender leaves which are easier to handle than the mature leaves were used in these experiments. A small slit was made in the leaf of a test plant, using fine insect Biology of Phytomyza 259 pins under a binocular microscope. It is normally easier to make the slit near the base of the leaf or near the mid rib, more so on one side of the leaf than the other, depending on the test plant. A first instar larva was then removed by opening its mine on the natural host- plant and transferred with a fine tip of a soft brush into the slit made on the leaf of the test plant. The larva was pushed inside the slit so that it was completely surrounded by the tissue of the test plant. The leaf of the test plant along with a small portion of petiole or twig was then enclosed inside a square plastic petri dish containing moist filter paper in order to prevent any sudden drying of the tissue around the slit. Two small holes were cut in upper corners of the petri dishes and covered with thin muslin cloth to permit transpiration and to prevent excessive condensation. The larva inside the test plant can be observed by trans- mitted light, moving its mouth hooks in an attempt to eat the new tissue. Leaves of the test plants were checked within a couple of hours of making the transfers. If the larva was still moving its mouth hooks, the transfer was considered successful; if the larva did not show any movement it was assumed to have been injured and the transfer was rejected. With patience and experience with particular test plant, it was possible to make good transfers of larvae, except to Artemisia because of the very woolly surface of the leaf. The transferred larva usually ended up inside the fibres on the leaf, rather than inside the leaf tissue. Observations were made every 12 hours on larval feeding and pupation, if any, during the previous 12 hour period. The pupae obtained were kept individually in small vials con- taining moist sand, for emergence of adults. The emergence of adults was also checked every 12 hours. BIOLOGY OF PHYTOMYZA MATRICARIAE HENDEL Natural incidence The adults of this species appear around Edmonton, Alberta during the first week of June. The leaf mines and larvae start appearing on various host-plants by the second week of June. There are numerous overlapping generations during July, August, and up to mid- September, when numbers start declining. At this time the host-plants also decline in vigor due to shorter days and lower temperatures. Towards the end of September, puparia go into winter diapause. Mating Mating was observed in the laboratory and it occurs many times in the life of both sexes, usually on the leaves of the food plant. The mating posture in a superimposed position is typical of other agromyzid flies. Its duration as in other agromyzid flies varies greatly, from Vi hour to approximately 2 hours. Adult feeding The flies feed upon plant exudates soon after emergence. The female selects a suitable spot on the leaf tissue, bends the tip of her abdomen vertically downwards, pierces the epidermis and then rotates the tip of her ovipositor within the leaf tissue. She then with- draws her ovipositor, turns around and imbibes the sap exuding from the wound. The puncture thus made is almost conical in shape. This method of feeding among agromyzid flies is of wide occurrence and has been described for many species. The female spends most of her lifetime making punctures in leaf tissue. These punctures are made both on upper and lower surfaces of the leaf, but are usually more numerous on the upper surface. The males, which are incapable of making such punctures, feed on the sap from punctures made by females or on natural plant exudates and probably also on 260 Sehgal nectar of flowers. Pollen grains could not be found in the guts of about 10 field collected males examined for this purpose. Plants normally survive the injury made by feeding punctures on the leaf, but under severe laboratory infestations they become greatly etiolated and sometimes collapse. Oviposition The eggs are laid singly inside the leaf parenchyma in punctures made in a similar way to feeding punctures. The egg punctures, like feeding punctures, were found both on upper and lower surfaces of the leaf, but unlike feeding punctures were usually more common on the lower surface. Feeding punctures always greatly outnumber oviposition punctures. This method of oviposition is general among the leaf mining agromyzid flies and has been de- scribed in many species. Incubation period The incubation period (Table 1) ranged from 90-102 hours, with an average of approxi- mately 91 hours or 3.8 days. The egg, originally translucent, becomes opaque white within the first 24 hours. The cephalopharyngeal skeleton appears as a darkly sclerotized structure at the end of 72 hours. At this time the embryo is almost fully developed and the mouth hooks can sometimes be seen to move horizontally. Table 1 . Duration of life history of Phytomyza matricariae Hendel on its natural host-plant Tanacetum vulgare L. Stage Number of records Duration in hours Total average duration (days) Egg 44 91.0 (90.0-102.0)* 3.8 First instar 20 64.2 (55.0-73.0) Second instar 20 45.0 (40.0-51.5) Third instar 20 56.4 (44.0-65.0) TOTAL larval 20 165.6 (144.0-180.0) 6.9 Pupal stage 15 13.7 (13.0-14.5) (days) 13.7 TOTAL life cycle 24.4 *The values of 40 observations were 90.0 hours and four 102.0 hours. Larval activity The average duration of first, second, and third instars (Table 1) was 64.2, 45.0 and 56.4 hours respectively. The total larval period was 165.6 hours or 6.9 days on an average. From hatching to shortly before pupation, the larva is completely endophagous. This results in the excavation of a linear leaf mine. The mine starts from the upper or lower surface of the leaf depending upon the site of oviposition and may terminate on either surface, when the larva leaves by cutting a small crescent shaped slit. The larva then falls to the ground and pupates. The larva like most other agromyzid larvae (Hering, 1951) Biology of Phytomyza 261 seems unable to re-enter the leaf once it is removed from it. The ability of the larvae of ‘ Liriomyza pusilla Meigen’ (Tilden, 1950) and at least some members of the Agromyza rufipes group (Nowakowski, 1964; Griffiths, verbal communication) to re-enter the leaf seems very unusual. The mouth hooks can be seen cutting through the leaf tissue in a lateral and semicircular motion. The larvae, like those of other agromyzids (Hering, 1951), mine the leaf while lying on their sides and they alternate from one side to the other. The frass is deposited in discrete granules along the sides of the mine alternately as the larva turns from side to side. Many larvae may start mining the same leaf simultaneously resulting in numerous mines crossing each other, however, every mine remains distinct and contains only one miner. The larvae normally feed only on the leaf tissue, but in heavy infestations also migrate to the petiole or mine under the epidermis of the stem. Transverse sections of the mined leaves (Fig. 5, 6) show that larvae feed indiscriminately on the palisade and spongy mesenchymatous tissue between the two epidermal layers of the leaf. The larvae only consume the entire tissue between upper and lower epidermis when this is very heavily infested. Thus the leaf mine is normally more visible from one side of the leaf. The larvae are capable of crossing the leaf veins but the vascular bundles are not consumed, as also reported in some other agromyzid species (Trehan and Sehgal, 1963; Tauber and Tauber, 1968). The lengths of mines excavated by first, second and third instar are given in Table 2. The total length of 142.0 mm is much shorter than 273.0 mm for “ Phytomyza lanati Spencer” (Tauber and Tauber, 1968). Although the duration of the third larval instar is approximately the same as that of the first instar, the major portion of the mining activity was done by the third instar. Table 2. Lengths of leaf mines excavated by different larval instars of Phytomyza matri- cariae Hendel in Tanacetum vulgar e L. Stage of larva Length of leaf mine in mm (each 19 observations) Range Average First instar 8.0- 18.4 13.5 Second instar 13.4- 42.5 29.1 Third instar 71.2-127.0 99.4 TOTAL larval life 110.0-177.0 142.0 Pupation The mature larva when ready to pupate leaves the leaf mine and falls to the ground, where it seeks a suitable site for pupation. Often the larva remains sticking to the exit slit in the mine and there forms the puparium. Duration of the puparium averages 13.7 days but varies considerably. The pupae from the fall generation undergo winter diapause. 262 Sehgal Fig. 5-6. Transverse sections of leaf mines of Phytomyza matricariae. mine on Matricaria matricarioides (Less.) Porter. 5. leaf mine on Achillea sibirica Ledeb. 6. leaf Biology of Phytomyza 263 DESCRIPTION OF THE IMMATURE STAGES Egg The egg (Fig. 7) when freshly laid is translucent white, smooth, elongate, ovo-cylindrical, slightly broader at the posterior end, and with a small, almost indistinguishable micropyle at the anterior end. The eggs of the members of the Phytomyza syngenesiae group have a similar micropyle (see Smulyan, 1914, ‘ Phytomyza chrysanthemi Kowarz’ and Cohen, 1936, ‘ Phytomyza atricornis Meigen’). The egg dimensions are given in Table 3. Table 3. The dimensions in mm of egg, larva, cephalopharyngeal skeleton*, and puparium of Phytomyza matricariae Hendel. All measurements based on 10 observations. Stage Length average (range) Width average (range) Egg 0.293 (0.283-0.316) 0.129 (0.100-0.141) First instar larva 0.412 (0.300-0.550) 0.152 (0.116-0.208) Cephalopharyngeal skeleton Second instar larva 0.121 (0.116-0.125) 0.732 (0.592-0.825) 0.243 (0.208-0.300) Cephalopharyngeal skeleton Third instar larva 0.220 (0.216-0.233) 2.395 (2.125-2.625) 0.667 (0.625-0.750) Cephalopharyngeal skeleton Puparium 0.316 (0.308-0.316) 1.810 (1.675-1.950) 0.852 (0.800-0.925) * includes mouth hooks, labial sclerite and paired paraclypeal phragma. Larva There are three larval instars which can be easily recognised by the sizes of their cephalo- pharyngeal skeletons (Table 3). The first instar larva when freshly hatched is translucent white, but soon becomes greenish due to the ingested leaf tissue. The second and third instar larvae are yellowish green in colour. The first instar larva is metapneustic, while the second and third instar larvae are amphipneustic. The anterior spiracles (Fig. 8) of the mature third instar larva have 7-9 small oval bulbs, while the posterior ones (Fig. 9) have 19-21 small oval bulbs. De Meijere (1926) illustrated the anterior and posterior spiracles in the European members of this species bred from Matricaria chamomilla L. He reported about 12 bulbs on the anterior and 18 bulbs on the posterior spiracles. The head (Fig. 10) bears two small longitudinal sclerites just above the mouth hooks, small but conspicuous maxillary palps, a pair of small antennae and numerous sense papillae. The muscle scars on the intersegmental membrane (Fig. 11) are small, oval and transversely elongated. The tubercle bands (Fig. 11) consist of small conical processes irregularly scattered along the intersegmental membrane. The tubercle bands as in other agromyzid larvae (Allen, 1957) are best developed along lateral portions of intersegmental membrane. The cephalopharyngeal skeletons of the first, second, and third instar are illustrated in Fig. 12-14. They consist of paired mouth hooks or mandibles, labial sclerite, and paired paraclypeal phragma. The mouth hooks in the first instar larva are small, simple and sickle 264 Sehgal 12-14 Fig. 7-14. Phytomyza matricariae. 7. egg. 8. anterior spiracle. 9. posterior spiracles. 10. facial mask of third instar larva. 11. muscle scars and tubercle band from a lateral portion of first abdominal segment of larva. 12. cephalo- pharyngeal skeleton of first instar larva. 13. cephalopharyngeal skeleton of second instar larva. 14. cephalopharyngeal skeleton of third instar larva. 0. 1MM Biology of Phytomyza 265 shaped; while in second and third instars they are well developed with two teeth each, alternating with one another. The right mouth hook is higher than the left and both mouth hooks are joined at the base (Fig. 10). Labial sclerite and paraclypeal phragma are smaller in first and second instars, but are well differentiated in the third instar. The dorsal process of the paraclypeal phragma consists of a long, single, slender and darkly sclerotized arm; the ventral arm is short, lightly sclerotized, and has a conspicuous foramen towards its posterior end. Puparium The puparium dimensions are given in Table 3; it is conspicuously segmented and shining black in colour. The hardening of the third larval skin is due to the deposition of the calcospherites (Frick, 1952; Allen, 1957) so that it can be softened by treatment with dilute hydrochloric acid and can be cut open for detailed examination. Although the puparium preserves the external morphology of the third instar, this is best studied in the larva itself. HOST-PLANT RELATIONSHIPS OF ADULT FEMALES The ovipositing female comes across numerous other plants besides those normally at- tacked in nature. In order to test the specificity of feeding and oviposition a selection of a wide range of plants was exposed to a batch of five gravid females, for a period of 24 hours. Plants used in this study included some common plants, which the female would encounter in the field, as well as some plants which are known for certain secondary substances like alkaloids, glycosides, etc. A total of 38 plant species belonging to 17 families were tested for feeding and oviposition. Feeding and oviposition preference of females when offered a choice of acceptable plants was also studied. Degree of phylogenetic relationship of the test plant to the natural host -plant was compared with the index of acceptability for feeding and oviposition. Index of plant relationship Botanical relationship of the test plant species, used in experiments on feeding and ovi- position by adult females and transfers of larvae, to one of the natural food plants of P. matricariae was examined. The phylogenetic relationships between plant families and orders is still a matter of con- troversy. Most plant classifications fall into two groups depending on the supposed nature of primitive angiosperm flowers (Davis and Heywood, 1965). One system is based on the assumption that the earliest angiosperms were wind-pollinated and that the monocotyledons and dicotyledons have arisen independently from hypothetical gymnosperms. According to the second system dicotyledons and monocotyledons were both derived from primitive an- giosperms which were insect-pollinated. The second system has the support of most recent botanists (Eames, 1961 ; Hutchinson, 1964; Takhtajan, 1969). Hutchinson (1964) has main- tained a basic division of dicotyledons into woody “Lignosae” and herbaceous “Herbaceae”, a system which allegedly leads to the wide separation of certain plant families which other- wise seem closely related in the structure of their flowers. The arrangement of plant families used in this study is after Takhtajan (1969), which is considered to reflect more closely the phylogenetic relationships between plant families. An index of plant relationship from 1 to 10 was used as follows: Relationship of the test plant species to natural host-plant Index of plant relationship 1 . Same species 1 0 266 Sehgal 2 Same genus 9 3. Same tribe (Anthemideae) 8 4. Same family (Compositae) 7 5. Same order (Asterales) 6 6. Same superorder (Asteranae) 5 7. Same subclass (Asteridae) 4 8. Same class (Dicotyledoneae) 3 9. Same division (Angiospermae) 2 10. Same phylum (Tracheophyta) 1 Acceptability index for feeding and oviposition Acceptability index (A. I.) of a test plant for feeding and oviposition relative to that of the natural food plant from which flies were obtained was calculated for comparison with the index of plant relationship. In experiments with feeding and oviposition studies, females used were obtained from the natural host-plant Tanacetum vulgare L. Data given in Tables 4 and 5 were used to calculate the acceptability index. In order to give a theoretical maximum value of 1 to the acceptability index, the sum of the components was divided by 2; thus: A. I. Feeding punctures Feeding punctures in T. vulgare Oviposition punctures Oviposition punctures in T. vulgare Range of food plants Data on the acceptability of plants for feeding and oviposition by gravid females is sum- marized in Table 4. Indices of plant relationship and of acceptability for feeding, and ovipo- sition are given for each plant species. Of the 38 plant species tested only seven belonging to the family Compositae tribes Anthemideae and Heliantheae were acceptable both for feed- ing and oviposition. Among these Artemisia, Helianthus and Zinnia were not found attacked in nature. The acceptability index for feeding and oviposition based on the data in Tables 4 and 5 is plotted against the index of plant relationship in Fig. 15. Indices of acceptability were very low for most species of test plants, but high for indices of plant relationship of 8 or over. Feeding and oviposition preference by gravid females The preference of gravid females for feeding and oviposition when offered a choice of six acceptable plants was examined. The plants used in this study were Achillea sibirica Ledeb., Artemisia sp., Chrysanthemum sp., Helianthus annuus L., Matricaria matricarioides (Less.) Porter and Tanacetum vulgare L. belonging to the family Compositae. Results of this experiment are summarized in Table 5. The number of feeding punctures on Tanacetum was significantly higher than on other plants tested. The numbers of feeding punctures on Chrysanthemum, Achillea, Matricaria and Helianthus were not significantly different from each other, but were significantly lower than on Tanacetum. The numbers of punctures with an egg on Tanacetum, Chrysan- themum, Achillea and Matricaria were not significantly different from each other, but were significantly higher than on Helianthus and Artemisia. No relationship was found between the numbers of oviposition punctures and the numbers of feeding punctures. Feeding preferences by freshly emerged females Feeding preference by freshly emerged females, which had not been exposed to any food plant, was examined in a similar experiment the results of which are summarized in Table 6. The only difference from Table 5 is that the numbers of feeding punctures on Achillea, Matricaria and Chrysanthemum were significantly higher than on Helianthus and Artemisia. Biology of Phytomyza 267 Table 4. Feeding and oviposition by females of Phytomyza matricariae Hendel on various plant species. Test plant Number of feeding punctures Number of punctures with an egg Index of accepta- bility Index of plant relationship PTERIDOPHYTA Polypod iaceae Nephrolepis 2 0 0.002 1 ANGIOSPERMAE-DICOTYLEDONEAE Ranunculaceae Aquilegia sp. (cultivated) 16 0 0.018 3 Clematis verticillaris DC 12 0 0.013 3 Delphinium sp. (cultivated) 0 0 0.000 3 Papaveraceae Papaver sp. (cultivated) 11 0 0.012 3 Chenopodiaceae Chenopodium sp. 5 0 0.005 3 Cucurbitaceae Cucumis sp. 11 0 0.012 3 Cruciferae Brassica khaber (DC.) Wheeler 3 0 0.003 3 Thlaspi arvense L. 22 0 0.024 3 Rosaceae Potentilla sp. 0 0 0.000 3 Leguminosae Caragana arborescens Lam. 3 0 0.003 3 Lathy rus odoratus L. 0 0 0.000 3 Lupinus sp. (cultivated) 0 0 0.000 3 Pisum sativum L. 32 0 0.036 3 Vicia americana Muhl. 0 0 0.000 3 Tropaeolaceae Tropaeolum sp. (cultivated) 3 0 0.003 3 Solanaceae Lycopersicon esculentum L. 3 0 0.003 4 Nicotiana tabacum L. 4 0 0.004 4 Solanum tuberosum L. 3 0 0.003 4 Scrophulariaceae Antirrhinum sp. (cultivated) 0 0 0.000 4 Labiatae Gale op sis tetrahit L. 5 0 0.005 4 Campanulaceae Campanula sp. (cultivated) 0 0 0.000 5 Compositae Achillea sibirica Ledeb. 567 28 1.039 8 Artemisia sp. 227 10 0.398 8 Aster ciliolatus Lindl. 10 0 0.011 7 268 Sehgal Table 4 (continued) Test plant Number of feeding punctures Number of punctures with an egg Index of accepta- bility Index of plant relationship Chrysanthemum sp. (cultivated) 467 34 1.012 8 Helianthus annuus L. 292 19 0.600 7 Matricaria matricarioides (Less.) Porter 441 30 0.926 8 Senecio vulgaris L. 54 0 0.060 7 Solidago sp. 12 0 0.013 7 Sonchus uliginosus Bieb. 4 0 0.004 7 Tanacetum vulgare L. 443 35 1.000 10 Taraxacum officinale Weber 124 4 0.139 7 Zinnia sp. (cultivated) 44 3 0.092 7 ANGIOSPERMAE-MONOCOTYLEDONAE Liliaceae Allium cepa L. 1 0 0.001 2 Smilacina stellata (L.) Desf. Gramineae 0 0 0.000 2 Hordeum vulgare L. 7 0 0.007 2 Typhaceae Typha lati folia L. 0 0 0.000 2 Table 5. Feeding and oviposition preferences of female Phytomyza matricariae Hendel from a culture raised on T. vulgare. Test plant Average* number of feeding punctures Average number of punctures with an egg Index of success Index of plant relation- ship Tanacetum vulgare 519.5 a** 11.8 a’ 1.000 10 Chrysanthemum sp. 200.5 b 13.3 a’ 0.756 8 Achillea sibirica 181.6 be 10.5 a’ 0.614 8 Matricaria matricarioides 161.0 be 15.0 a’ 0.790 8 Helianthus annuus 58.5 b c 2.3 b’ 0.153 7 Artemisia sp. 36.8 c 1.6 b’ 0.102 8 * Averages are based on six replicates. ** Treatments which are not significantly different from each other have the same letter opposite; as calculated by Duncan’s multiple range significance level test. Biology of Phytomyza 269 Fig. 15. Graph showing the index of acceptability of test plant for feeding and oviposition, and the index of plant relationship. Table 6. Feeding preferences of freshly emerged (<24 hours) females of Phytomyza matri- cariae Hendel from T. vulgare. Test plant Average number of feeding punctures* Tanacetum vulgare 250.8 a* ** Achillea sibirica 174.6 b Matricaria matricarioides 139.5 b Chrysanthemum sp. 128.6 b Helianthus annuus 28.3 c Artemisia sp. 14.3 c * Based on six tests. ** Treatments which are not significantly different from each other have the same letter opposite; as calculated by Duncan’s multiple range significance level test. 270 Sehgal HOST-PLANT RELATIONSHIPS OF LARVAE In nature the larvae feed only on the plants accepted for oviposition by the female. Being completely internal plant feeders, they cannot select a more suitable food plant which might be available in its geographical range. In order to test the ability of larvae to use different plants as food which may be available in their geographical range, the first instar larvae obtained from the natural food plant were transferred into the tissue of the test plant. An index of success for larval development based on larval feeding, pupation, and emergence of imago was calculated for comparison with the index of plant relationship. Index of success for larval development An index of success (S. I.) for larval development was calculated from results obtained in the transfers of larvae from natural host-plant into recipient plant species, for comparison with the index of plant relationship. This calculation was based on three components as follows: \. Duration of larval survival in recipient plant. — This was expressed in half days; the 12 hours immediately preceding the finding of a non-feeding larva was included since observations showed that the average survival time after the cessation of feeding was about 12 hours. Studies on the life history as summarized in Table 1 show that the average duration of a larval stage in P. matricariae on T. vulgare is 6.9 days. The first instar larvae used in transfer experiments were about 1 day old. Therefore under normal conditions the remainder of larval feeding time should average 5.9 days. Success in larval feeding was expressed as the ratio of the duration of feeding of larva on test plant to 5.9 days. 2. Pupation. — This is considered as successful termination of larval development. Success was expressed as the proportion of the transferred larvae pupating successfully on the test plant. 3 . Emergence. — In most plants the larvae which pupated also emerged as adult flies. The emergence was given one-fourth as much weight as pupation in calculation of success index, that is, it was expressed as one quarter of the proportion of transferred larvae which yielded adult flies. In order to give a theoretical maximum value of 1 to the Success Index (S. I.) the sum of these components was divided by 2.25; thus: Survival time (days) Pupations Emergences 5.9 Transfers 4 x Transfers Transfers of larvae from natural host-plant to test plant The number of larvae transferred individually from the natural host-plant to inside the tissue of recipient plant species was 20, except in two species where it was 10. Results of the transfers of larvae are summarized in Tables 7 and 8. Values for the success index and the index of plant relationship are also given graphically in Fig. 16. Among Pteridophyta, only the greenhouse fern Nephrolepis sp. tested for transfers of larvae from Tanacetum, could not be used by larvae as food for completing their develop- ment indicating the plant to be toxic or otherwise unacceptable. Among dicotyledons, 28 plants belonging to 10 plant families were tested for transfers of larvae from Tanacetum (Table 7) and Achillea (Table 8). One larva at least completed its development on 16 plant species belonging to 5 families. Leaf mines formed after trans- fer of larvae from Tanacetum into four of the test plants are shown in Fig. 17-20. Biology of Phytomyza 271 Table 7. Results of transfers of larvae of Phytomyza matricariae Hendel from Tanacetum vulgare L. to other plant species. 20 transfers except where indicated. Note that transfers to T. vulgare represent a control. Recipient species Duration of larval feeding in test plant (days) Number of larvae pupated (**emerged) Index of success Index of plant rela- tionship PTERIDOPHYTA Polypodiaceae Nephrolepis sp. (cultivated) *0.25 ±0.0 0 0.018 1 ANGIOSPERMAE-DICOTYLEDONEAE Ranunculaceae Aquilegia sp. (cultivated) (0.5— 0.5) *2.57 ±3.0 3 0.276 3 Caryophyllaceae Silene noctiflora L. (0.5— 9.0) *1.07 ±0.3 0 0.080 3 Chenopodiacea Chenopodium album L. (1. 0-2.0) *0.97 ±0.5 0 0.072 3 Cruciferae Brassica khaber (DC.) (0.5— 2.0) *2.47 ±2.0 2 0.241 3 Thlaspi arvense L. (1.0— 7.5) *1.80 ±0.8 0 0.135 3 Rosaceae Potentilla sp. (1. 0-4.0) *0.67 ±0.4 0 0.050 3 Leguminosae Lathy rus odoratus L. (0.5— 2.0) *1.55 ± 1.0 0 0.116 3 Lupinus sp. (0.5— 4.0) *1.72 ±0.9 0 0.129 3 Melitotus officinalis (L.) Lam. (1.0— 4.5) *3.07 ± 2.6 2(1) 0.280 3 Pisum sativum L. (1.0— 9.0) *4.47 ± 1.3 12 0.669 3 Umbellifereae Apium sp. (2.0— 6.5) *0.35 ±0.2 0 0.026 3 Solanaceae Nicotiana tabacum L. (0.5-1. 5) *0.25 ±0.1 0 0.018 4 (0.2- 1.0) 272 Sehgal Table 7 (continued) Recipient species Duration of larval feeding in test plant (days) Number of larvae pupated (♦♦emerged) Index of success Index of plant rela- tionship Labiatae Galeopsis tetrahit L. *3.65 ± 1.9 (1. 0-7.0) 10 0.552 4 Compositae Achillea sibirica Ledeb. (10 transfers) *4.65 ± 1.4 (1. 0-6.0) 9(8) 0.594 8 Artemisia sp. (10 transfers) *0.25 ±0.0 (0.5— 0.5) 0 0.018 8 Chrysanthemum sp. (cultivated) *4.07 ± 2.8 (1.5-8. 5) 6 0.472 8 Dahlia sp. (cultivated) *2.37 ± 1.8 (1. 0-8.0) 2(1) 0.228 7 Helianthus annuus L. *2.97 ± 2.3 (0.5— 6.0) 10 0.501 7 Matricaria matricarioides (Less.) Porter (10 transfers) *4.40 ± 1.4 (0.5-5. 5) 9 0.580 8 Senecio vulgaris L. *3.85 ± 2.2 (0.5— 7.5) 6(5) 0.450 7 Sonchus uliginosus Bieb. *3.02 ± 2.6 (0.5— 7.0) 9 0.476 7 Tanacetum vulgare L. *5.57 ± 1.0 (2.0— 7.0) 19 0.966 10 Taraxacum officinale Weber *2.25 ± 2.3 (0.5— 6.0) 4(3) 0.274 7 ANGIOSPERMAE-MONOCOTYLEDONEAE Liliaceae Allium cepa L. *3.95 ± 2.4 (1. 0-9.0) 6(5) 0.458 9 Gramineae Hordeum vulgare L. *2.67 ± 1.7 (1 .0—7.0) 3 0.284 9 Typhaceae Typha lati folia L. *0.55 ±0.2 (0.5- 1.0) 0 0.041 2 * mean ± S. D. * ** all emerged unless otherwise indicated, (range) Biology of Phytomyza 273 Table 8. Results of transfers of larvae of Phytomyza matricariae Hendel from Achillea sibirica Ledeb. to test plants. 20 transfers except where indicated. Note that transfers to A. sibirica represent a control. Recipient species Duration of larval feeding in test plant (days) Number of larvae pupated (**emerged) Index of success Index of plant rela- tionship ANGIOSPERMAE-DICOTYLEDONEAE Cruciferae Brass ica oleracea L. Leguminosae * 1 .50 ± 0.9 (0.5— 5.5) 1 0.140 3 Lupinus sp. (cultivated *1.45 ± 1.3 (0.5— 7.0) 0 0.108 3 Pisum sativum L. Solanaceae *4.17 ± 1.2 (2.0— 6.0) 10 0.591 3 Nicotiana tabacum L. *0.80 ±0.4 (0.5 — 2.0) 0 0.060 4 Solarium tuberosum L. (10 transfers) Compositae *0.50 ±0.4 (0.5— 1 .5) 0 0.037 4 Achillea sibirica Ledeb. *4.72 ± 1.6 (1.0— 7.5) 15 0.772 10 Solidago sp. *0.67 ± 0.4 (0.5— 2.0) 0 0.050 7 Sonchus arvensis L. *1.50 ± 1.3 (1 .0—6.0) 0 0.112 7 Tanacetum vulgare L. (10 transfers) *5.05 ± 1.1 (2.5— 6.5) 9 0.880 8 Zinnia sp. (cultivated) *2.72 ± 1.5 (1 .0—5.5) 6(5) 0.365 7 mean ± S. D. (range) ** all emerged unless otherwise indicated. 274 Sehgal Fig. 16. Graph showing the success index of test plant for larval development against index of plant relationship. Transfers from Tamcetum vulgare L. ■ Transfers from Achillea sibirica Ledeb. On the basis of available data, although it is not possible to make statistical comparisons of the relative success of larval development on these plants, nevertheless some of these plants were clearly more suitable for larval development than others. Two species outside the Compositae, Pisum sativum L. and Galeopsis tetrahit L., were at least as good as Tana- cetum, their natural food plant, if not better, in their suitability for larval development. Brassica spp. tested were quite resistant to larval development; besides others, one reason was the formation of callus in the injured area of the young leaf. Among other plants tested, the formation of callus in the punctures made for feeding and oviposition, and in other small injuries, was very frequent in the young leaf of Helianthus. Twelve dicotyledons were not used by larvae as food for complete development, though they did survive for a certain length of time in most of these plants. Among these plants the failure of larvae to mature in Artemisia sp. was probably due to the fact that they would bite their way out into the woolly fibres of the leaf. In all other plants the experimental larva died inside the tissue of the recipient plant. Among three monocotyledons, two species, Allium cepa L. and Hordeum vulgare L. were used by larvae to complete development to adult flies. The failure to use the third species Typha latifolia L. was, besides other factors, probably also due to the presence of large air spaces in the leaf tissue. Biology of Phytomyza 275 Fig. 17 20. Leaf mines formed after transfers of larvae of Phytomyza matricariae from Tanacetum vulgare to various recipient species. 17. leaf mine on Brassica oleracea L. 18. leaf mine on Pi sum sativum L. 19. leaf mine on Sonchus uliginosus Bieb. 20. leaf mine on Zinnia sp. (cultivated). 276 Sehgal DISCUSSION Host-plant relationships of adult female Range of food plants. - Even though females made varying numbers of feeding punctures on many plants, eggs were only deposited in members of the family Compositae belonging to the tribes Anthemideae and Heliantheae. Plants which are not accepted for both feeding and oviposition are considered as non-hosts. The physical characteristics of the plant do not seem to be important, although they may play some role in host preference. The accepta- bility of closely related plants for both feeding and oviposition indicate that chemical factors are involved in the very high degree of host-plant specialization. That many plants are not acceptable as hosts by P. matricariae is indicative of deterrent effects in the plants. Plants acceptable for feeding must therefore have stimulating substances and at the same time lack deterrent substances. Since the females oviposited only on closely related mem- bers of the family Compositae, the plants accepted for oviposition must also have substances which provide adequate stimuli for egg laying and these are not necessarily the same sub- stances as those which stimulate feeding. Hsiao and Fraenkel (1968) working with Colorado potato beetle, Leptinotarsa decemlineata (Say), suggested that host specific stimuli from some solanaceous plants were required in the oviposition behavior of this species. When the index of success in feeding and oviposition was compared with the index of plant relationship (Fig. 15), only plants most closely related to natural food plants were acceptable for feeding and oviposition, thus confirming the observed oligophagy of P. matri- cariae in nature. Hering (1951) pointed out numerous species of mining insects in which the ovipositing females committed errors in their choice of host-plants and concluded that such instances of erroneous oviposition were quite common and yet escaped our attention. Later Nowa- kowski (1962) examined such known instances in the family Agromyzidae and found that these were mainly due to misidentifications of the insects. The results of the experiments on the host range of P. matricariae indicate that the females are unlikely to make the mis- take of ovipositing on the wrong host under normal circumstances. Feeding and oviposition preference of females. — When females were offered a choice of six acceptable plants for feeding and oviposition, Tanacetum, from which the flies used were obtained, was most preferred for feeding (Table 5); however, the numbers of eggs laid were not significantly higher than on some other plants in the test. In another experiment in which freshly emerged females obtained from pupae bred on Tanacetum were used, Tanacetum was still most preferred (Table 6) in number of feeding punctures. The prefer- ence for feeding on Tanacetum may either be explained by the preconditioning of the fe- males in their own life, or by preconditioning in their larval life as defined by Hopkins’ (1917) host selection principle, or by the greater quantity of substances which stimulate feeding, or just by the taste preference of the females. This however, cannot be clarified at present and would need further detailed studies. However, behavior in which insects prefer the plant species previously eaten is in agreement with the observations of Jermy, Hanson and Dethier (1968) on Manduca sexta (Johanssen) and Heliothis zea (Boddie). It may be pointed out that Tanacetum appeared to be more heavily attacked in nature than other host-plants. This, however, may also be due to various other factors like greater abundance of this plant in the habitat. Among other plants used in the study Achillea, Chrysanthemum, and Matricaria were almost equally preferred, while Helianthus and Artemisia were least preferred for both feed- ing and oviposition (Tables 5, 6). The first three plants belong to the same tribe Anthe- mideae as Tanacetum and also serve as host-plants in nature. In Chrysanthemum only the Biology of Phytomyza 277 soft leaf variety was found to be attacked in nature. This suggests some importance of physical characteristics of plants in their selection. Artemisia , although closely related to Tanacetum, was not preferred, probably because the leaves used had a thick covering of woolly fibres on their lower surfaces, which may act as a physical barrier for females of this species. Helianthus which was also not preferred, is not as closely related to Tanacetum. It was also not found to be attacked in nature. Hussey and Gurney (1962) suggested the use of feeding punctures to egg ratio as a method of assessing host preference in agromyzid species. The most ‘preferred’ host plant would have the lowest feeding puncture to egg ratio. They worked with a polyphagous species ‘‘Phytomyza atricornis Meigen’ which was later shown by Griffiths (1967) to consist of two distinct species, Phytomyza syngenesiae (Hardy) feeding predominantly on compo- ses and Phytomyza horticola Goureau feeding on composits and other families so that their results cannot be properly evaluated. In the populations used they found that feeding punc- ture to egg ratio was lower on preferred plants and concluded that preferred plants are nutritionally superior. It appears that the differential feeding and oviposition in their ex- periments with different varieties of Chrysanthemum was due to chemical factors which act as stimulants or deterrents rather than to nutritional differences. Host-plant relationships of larvae The ability of the larva to use 16 plants belonging to five different families for its devel- opment clearly shows that it is far less sensitive to deterrents than the adults which would only feed and oviposit on certain members of the family Compositae. The larvae having evolved a completely internal parasitic mode of life have reduced or poorly developed sense organs, which in turn reduces their ability to discriminate between various plant species. This is further supported by the observation that the larva starts feeding almost as soon as it is transferred to the test plant. The act of feeding was inferred from the move- ment of larval mouth hooks inside the test plant. However, the larva is capable of distin- guishing between various kinds of tissue within the leaf, as only the mesenchymatous tissue is eaten. It is apparent that plants widely separated phylogenetically are nutritionally ade- quate for the completion of larval development, if they lack substances which are toxic or inhibitory. Buhr (1937) was the first to carry out transplantation experiments with agromyzid larvae. He found that among the plants tested, the transferred larvae developed only on plants phylogenetically related to their natural food plant and died on plants not related to the normal host-plant. Liriomyza eupatorii (Kaltenbach) was exceptional, but it was already known to feed on Eupatorium (Compositae) and Galeopsis (Labiatae). He also succeeded in transferring larvae of Liriomyza cannabis Hendel from Cannabis (Urticaceae), to Eupa- torium and Galeopsis. However it is not known whether these two “species”, Liriomyza eupatorii (Kaltenbach) and L. cannabis Hendel, represent host races or sibling species, as was considered by Nowakowski (1962), since there is no clear morphological distinction between them. Admittedly, the possibility that the agromyzid female would oviposit in nature on plants not closely related to the natural host-plant is very small. But, if the female did oviposit on plants outside the normal range, there is a good possibility that the larva would complete development, if the plant is not toxic or inhibitory. This is contrary to Nowakowski’s (1962) suggestion that the probability of larval survival is very small. This ability of the larvae to use successfully certain plants outside the range of normal host-plants of the ovipositing female in P. matricariae , can explain observed patterns in agromyzids and also in external feeders. Liriomyza eupatorii (Kaltenbach) normally feeds 278 Sehgal in nature on Eupatorium, family Compositae and Galeopsis , family Labiatae. Liriomyza brassicae (Riley), an oligophagous species feeding on Cruciferae and related families, has also been reported to feed on Pisum, family Leguminosae (Spencer, 1964; Sehgal, 1965). Gupta and Thorsteinson (1960) showed that the leaves of non-cruciferous plants were normally accepted by the caterpillars of Plutella maculipennis , which normally feed on cruciferous plants. Jermy (1961) showed that Colorado potato beetle, normally a solana- ceous feeder, accepted the leaves of Asclepias syriaca L., family Asclepiadaceae and Allium cepa , family Liliaceae. He later (1966) suggested that in these plants some other substances replace the specific phagostimulants. Hsiao and Fraenkel (1968) working on Colorado potato beetle found the leaves of Asclepias (Asclepiadaceae) and Lactuca (Compositae) to be the most suitable non-solanaceous plants and these plants could support reproduction and continuous culturing. They further reported that these plants were not fed upon in the presence of normal solanaceous hosts. They therefore concluded that host selection in this species was determined not only by the presence of adequate feeding stimuli and nutri- ents, but also by the presence of host specific substances which induce the initial feeding behavior. In P. matricariae such host specific substances could be important in the speci- ficity of oviposition on certain members of the family Compositae, but not in the larval feeding on various test plants. Jermy (1966) suggested that certain plants like Pisum sativum L. and Malva sylvestris L. seem to be in general free of strong feeding inhibitors, while others like Solidago are strongly deterrent. Results of transfers of larvae of P. matricariae support this view, as Pisum sativum was quite suitable for larval development while Solidago was not. Galeopsis tetrahit prob- ably also belongs to a similar category of non-inhibitory plants. The majority of species in the family Agromyzidae are restricted feeders, being mono- phagous or oligophagous (Sehgal, 1971). This study of insect host-plant relationships of adults and of larvae of P. matricariae , as well as recent studies involving other oligophagous species have shown that botanically unrelated plants can also serve as adequate food plants for normal development. However, in nature an oligophagous species normally selects bo- tanically related plants for feeding and oviposition. Restricted feeding in nature on botani- cally related plant species or on unrelated plant species having similar secondary substances is probably the result of numerous evolutionary coadaptations of the phytophagous insect to the allelochemics (Whittaker and Feeny, 1971), allomones and kairomones, of the host- plants). ACKNOWLEDGEMENTS I am grateful to B. Hocking, Chairman, Department of Entomology, University of Al- berta, for providing the opportunity and support for this project and for his supervision, criticism of manuscript, and keen interest throughout this study. I am grateful to G. E. Ball, Department of Entomology, University of Alberta, for advice and valuable criticism of the manuscript. I am also grateful to K. A. Spencer, London, England and G. C. D. Griffiths, Department of Entomology, University of Alberta, for numerous useful discus- sions and valuable suggestions. I would like to thank D. A. Craig and W. G. Evans, Depart- ment of Entomology, University of Alberta and G. Pritchard, Department of Biology, University of Calgary for numerous suggestions and helpful criticism of the results. I thank J. G. Packer, Department of Botany, University of Alberta for help in identification of host-plants, H. S. Welling for help in greenhouse culturing of plants, J. S. Scott for help in preparing photographs and J. Rickert for help in translating the abstract into German. Biology of Phytomyza 279 REFERENCES Ahmad, T. and R. L. Gupta. 1941. The pea leaf-miner, Phytomyza atricornis (Meigen), in India. Indian J. Ent. 3:37-49. Allen, P. 1956. Observations on the biology of some Agromyzidae (Diptera). Proc. R. ent. Soc. Lond. (A) 31:117-131. Allen, P. 1957. The larval morphology of Agromyzidae (Diptera). Proc. R. ent. Soc. Lond. (A) 32:59-66. Buhr, H. 1937. Parasitenbefall und Pflanzenverwandtschaft. Botan. Jahrb., 68:142-198. Cohen, M. 1936. The biology of the Chrysanthemum leaf -miner, Phytomyza atricornis Mg. (Diptera : Agromyzidae). Ann. appl. Biol. 23:612-632. Davis, P. H. and V. H. Hey wood. 1965. Principles of Angiosperm Taxonomy. Oliver and Boyd. 558 pp. Dethier, V. G. 1966. Feeding behavior, In insect behavior. Symp. R. ent. Soc. London. 3:46-58. Dethier, V. G. 1970. Chemical interactions between plants and insects. In Chemical Ecology, E. Sondheimer and J. B. Simeone (eds.). Academic Press, New York. 83-102. Eames, A. J. 1961. Morphology of the Angiosperms. McGraw-Hill, New York. Fraenkel, G. S. 1959. The raison d’etre of secondary plant substances. Science. 129:1466-70. Frick, K. E. 1952. A generic revision of the family Agromyzidae (Diptera) with a catalogue of New World species. Univ. Calif. Pubis ent. 8:339-452. Friend, W. G. 1958. Nutritional requirements of phytophagous insects. Ann. Rev. Ent., 3:57-74. Griffiths, G. C. D. 1967. Revision of the Phytomyza syngenesiae group (Diptera: Agromy- zidae), including species hitherto known as “ Phytomyza atricornis Meigen”. Stuttg. Beitr. Naturk. 177:1-28. Gupta, P. D. and A. J. Thorsteinson. 1960. Food plant relationships of the diamond-back moth [Plutella maculipennis (Curt.)] 1. Gustation and olfaction in relation to botanical specificity of the larva. Ent. exp. & appl. 3:241-250. Harris, P. and A. I. Mohyuddin. 1965. The bioassay of insect feeding tokens. Can. Ent. 97:830-833. Hering, E. M. 1951. Biology of the leaf miners, ’s-Gravenhage: 1-420. Hering, E. M. 1957. Bestimmungstabellen der Blattminen von Europa. ’s-Gravenhage, 3 vol., I: 1-648, II: 649-1185, III: 1-221. Hopkins, A. D. 1917. A discussion of C. G. Hewitt’s paper on “Insect Behavior”. J. econ. Ent. 10:92-93. Hsiao, T. H. and G. Fraenkel. 1968. Selection and specificity of the Colorado potato beetle for solanaceous and nonsolanaceous plants. Ann. ent. Soc. Amer. 61:493-503. Hussey, N. W. and Gurney, B. 1962. Host selection by the polyphagous species Phytomyza atricornis Meigen (Dipt., Agromyzidae). Entomologist’s mon. Mag. 98:42-47. Hutchinson, J. 1964. The families of flowering plants. Oxford University Press. Jermy, T. 1961. On the nature of the oligophagy in Leptinotarsa decemlineata Say (Cole- optera: Chrysomelidae). Acta Zool. Acad. Sci. Hung. 7:119-132. Jermy, T. 1964. The role of rejective stimuli in the host selection of phytophagous insects. Proc. XII Int. Congr. Ent. London. 1964 (1965) p. 547. Jermy, T. 1966. Feeding inhibitors and food preference in chewing phytophagous insects. Ent. exp. & appl. 9:1-12. Jermy, T., F. E. Hanson and V. G. Dethier. 1968. Induction of specific food preference in Lepidopterous larvae. Ent. exp. & appl. 11:21 1-230. 280 Sehgal Keller, J. C. and T. B. Davich. 1965. Response of five species of insects to water extracts of their host plants. J. econ. Ent. 58:164. Kennedy, J. S. 1965. Mechanisms of host plant selection. 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Amer. 51:557-566. Oatman, E. R. and A. E. Michelbacher. 1959. The melon leaf miner, Liriomyza pictella (Thomson) (Diptera: Agromyzidae). Ann. ent. Soc. Amer. 52:83-89. Schoonhoven, L. M. 1968. Chemosensory bases of host plant selection. Ann. Rev. Ent. 13:115-136. Sehgal, V. K. 1965. Studies on Indian Agromyzidae (Diptera). Beitr. Ent. 15:3-10. Sehgal, V. K. 1971. (in press). A taxonomic survey of the Agromyzidae (Diptera) of Alberta, Canada, with observations on host-plant relationships. Quaest. ent. Smulyan, M. T. 1914. The Marguerite fly or Chrysanthemum leaf miner. Bull. Mass. agr. Exp. Sta. No. 157:21-52. Spencer, K. A. 1964. The species-host relationship in the Agromyzidae (Diptera) as an aid to Taxonomy. Proc. 12th int. Congr. Ent. (London 1964) 1:101. Spencer, K. A. 1969. The Agromyzidae of Canada and Alaska. Mem. ent. Soc. Can., 64: 311 pp. Stride, G. O. 1965. Studies on the chemical basis of host-plant selection in the genus Epilachna (Coleoptera, Coccinellidea). J. Insect Physiol., 1965:21-32. Sugiyama, S. and Y. Matsumoto. 1959. Olfactory responses of the vegetable weevil larvae to various mustard oils. Nogaku kenkyu 46:150-157. Takhtajan, A. 1969. Flowering plants, origin and dispersal, Oliver and Boyd, 205-239 pp. Tauber, J. M. and C. A. Tauber. 1968. Biology and leaf-mining behaviour of Phytomyza lanati (Diptera: Agromyzidae). Can. Ent. 100:341-357. Thorsteinson, A. J. 1953. The chemotactic basis of host plant selection in an oligophagous insect. ( Plutella maculipennis Curt.). Can. J. Zool. 3 1 : 52-72. Thorsteinson, A. J. 1960. Host selection in phytophagous insects. Ann. Rev. Ent. 5: 193-218. Tilden, J. S. 1950. Oviposition and behavior of Liriomyza pusilla (Meigen) (Diptera: Agro- myzidae). Pan-Pacif. Ent., 26:1 19-121. Trehan, K. N. and V. K. Sehgal. 1963. Range of host plants and larval feeding in Phytomyza atricornis (Meigen) (Dipt., Agromyzidae). Ent. mon. Mag. 99:1-3. Verschaffelt, E. 1910. The cause determining the selection of food in some herbivorous insects. Proc. Acad. Sci. Amsterdam 13:536-42. Webster, F. M. and T. H. Parks. 1913. The serpentine leaf-miner. J. Agric. Res. 1:59-87. Whittaker, R. H. and P. P. Feeny. 1971. Allelochemics: chemical interactions between species. Science. 171, No. 3973:757-770. NOTES AND CORRECTIONS TO A RECLASSIFICATION OF BOMBARDIER BEETLES (CARABIDAE, BRACHINIDA) TERRY L. ERWIN Division of Cole op ter a Department of Entomology Smithsonian Institution Washington, D. C. 20560 Quaestiones entomologicae 7 : 281 1971 Since publishing my “A reclassification of bombardier beetles and a taxonomic revision of the North and Middle American species (Carabidae:Brachinida)” (Quaest. ent. 6:4-215, 1970) a number of points have come to my notice. These follow seriatim , page numbers and references are to that paper. Page 17, line 29: Read uniperforate for uniporforate. Pages 34-37: Jeannel (1949:1084) designated Brachinus senegalensis Dejean (1825:308) as type of Pheropsophus Solier (1833:463). Unfortunately, I followed his designation and failed to see Hope’s previous designation of Carabus complanatus Fabricius (1775:242) in his Coleopterist’s Manual (1838:99). Since P. complanatus Fab. is a New World species (type from Santo Domingo) Pheropsophus Solier is the generic name for all the New World species and Pheropsophidius Hubenthal (191 1 : 547) is a synonym ( Pheropsophidius Hubenthal, type Cicindela aequinoctialis Linne 1763:395 . . . aequinoctialis L. = compla- natus F. according to Castelnau 1835:51). Thus, Pheropsophus is not available for the Old World species as they represent a genus different from the New World group. The first available name is Stenaptinus Maindron (1906: 15), type species S. krichna Maindron (1906: 15) designated by Jeannel (1949: 1084). The following list summarizes the correct names of the subtribe Pheropsophina: Genus Stenaptinus Maindron Subgenus Stenaptinus s. str. Subgenus Parapheropsophus Hubenthal (= Pheropsophus auct.) Subgenus Aptinomorphus Jeannel Genus Pheropsophus Solier Subgenus Pheropsophus s. str. (= Pheropsophidius auct.) Subgenus Protopheropsophus Hubenthal The name Parapheropsophus Hubenthal must be raised from synonymy (Darlington, 1968:234) to become available for the taxon called Pheropsophus by Jeannel (1949), myself (1970), and others. The above was drawn to my attention by Basilewsky (in litt.) and I thank him kindly. Page 38: The type species of Aptinus was designated by Hope (1838:99) not Jeannel. Page 41: The type species of Aptinoderus Hubenthal (1919:332) is Aptinus cyaneus Mot- schulsky (1864:214) designated by Jeannel (1949:1110). Pages 59, 81, 108, 153, 155: The following species were described by me in Opusc. ent. 34(3):287-288, cf. 1969b in the list of references: Brachinus microamericanus , B. adusti- pennis , B. kavanaughi, B. fulminatus, and B. vulcanoides. Page 164, line 42: Read extant for extent. Page 211, line 30: Read 21(2) for 1(2). 282 Book Review LOVELAND, R. P. 1970. Photomicrography. A Comprehensive Treatise, Vol. I & II. John Wiley & Sons, Inc., New York. Vol. I., vi + 1-526 + xii, Vol. II, vi + 527-1039 + xii. Cloth bound. Can. $34.50. Loveland, a former research associate of the Kodak Research Laboratories, has produced a book that is indeed a comprehensive account of photomicrography. He deals with all the major aspects of the subject without neglecting the more recent developments in this field. In Volume I he gives a general consideration of the compound microscope, photomacro- graphy, the optics of illuminating lamps, the quality of illumination, image contrasts and finally the eyepiece and roll-film cameras. This is followed by the appendices where depth of field, focal length of lenses and illumination for photomacrography are considered fur- ther; by a list of equations used in the text; and by the index which includes the subjects in Volume II. Loveland believes illumination to be very important and he pursues this subject further in Volume II where he considers special methods of illumination, flash photo- micrography and the photographic spectrum. Consideration is then given to the selection of photographic materials, the determination of exposure, and processing and printing. He also deals with the advantages and problems of color photomicrography, with fluorescence micrography and cinemicrography. This is followed by appendices where he deals, with exposure formulas, special immersion fluids and an illuminator for far-ultraviolet photo- micrography, the equations used in the text, and an index as in Volume I. In the foreword, Loveland indicates that the book has been written not only for the professional but also for the “neophyte” and he never loses sight of this. Ample warnings of the difficulties and disadvantages of various techniques are given and in some cases neo- phytes are warned off certain techniques. Each chapter is begun with a general consideration of the topic and then moves rapidly into the theoretical aspects. Loveland never neglects the practical problems and he always gives a good account of the various instruments avail- able from the major manufacturers. He has obviously had much experience for when there is not a commercially manufactured piece of equipment to solve a problem, he gives photo- graphs or line diagrams of the equipment he had built. Loveland’s style of writing is good; even in the more technical portions of the text it is a pleasure to read. The book’s usefulness as a reference text is much enhanced by the printing of important sections of the text in italics or bold face type. These two volumes will be a very useful teaching aid for photomicrography and should be in every laboratory where photomicrography is practised. D. A. Craig Department of Entomology University of Alberta Edmonton 7, Alberta 283 Book Review ARNETT, R. H., JR. 1970. Entomological Information Storage and Retrieval. The Bio-Rand Foundation, Inc., Baltimore, xii + 210 pp., 1 plate, 10 text Fig., c. 237 refs. $7.95. This interesting little book is a guide to sources of entomological information, and an introduction to current information storage and retrieval theories, problems, and accom- plishments as they relate to entomology. The book is intended not only as peripheral reading material, but as a student text. Accordingly, each chapter except the first contains suggested assignments, optional assignments, and a series of review questions. The writing is clear, the printing quality is good, and the illustrations are pertinent and well reproduced. Arnett’s book has a rather complex organization, with 1 1 text chapters, a glossary, four appendices, and subject and author indices, in addition to preface, postface, and colo- phon. Chapter one contains introductory observations, chapters two through eight deal with original information, and the closing three chapters deal with secondary information. Chapter two is of particular importance, since it concerns information storage and retrieval theory, the main subject of the book. In chapters three, four, and five Arnett discusses stored information, and original documents and their preparation for storage. Chapter six concerns the preparation of these documents for information retrieval. The remaining five chapters, which are more general and to some extent superficial, concern search resources, abstracts and indices, synthetic literature, popular literature, and societies, institutions, and personnel. The four appendices are: a bibliography of basic types of entomological litera- ture; a selected list of commercial publishers and dealers in this literature; a list of entomo- logical equipment and supply companies; and a list of major North American entomological libraries. The arrangement of the references, or bibliography, is unfortunate. This bibliography contains 237 entries, at least one being a repetition. As a key to various types of literature, its organization by chapter and subject headings is quite suitable. Perhaps, though, the many additional text references to serials, and to other works such as the Zoological Record, should be included in it. As a key to references cited, however, this bibliography leaves much to be desired. Some citations repeated in two or more text chapters are not listed accordingly in the bibliography, and thus are not readily located (retrieved?). For example, there is in chapter four a reference to the Conference of Biological Editors, but this is listed in the bibliography under the heading of chapter five. I suggest that, if a second edition of this book is published, there should be a separate section for references cited. Several acronyms used repeatedly in the text add to the complexity of the book, and any reader unfamiliar with these acronyms should turn first to the glossary and learn their various meanings. The information boxes used in numerous places are useful to summarize main points and to supply ancillary information, but some seem to me to detract from the book. I wonder, for instance, if the list of purposes of publication on page 66 is really needed; if this list is to be included, then I think each of its elements needs further dis- cussion. These criticisms are minor, but relate to the information storage and retrieval features of the book itself! I strongly recommend that student entomologists, in particular, acquire and use this book as a guide. Many will find that the last several chapters will help reduce the time required for, and increase the efficiency of, any literature searches they may need to do. Regrettably, the book is somewhat regional in scope, and as a guide will be of limited use outside of North America. 284 However, the book is especially timely and pertinent to the information explosion prob- lems of the present decade. Chapters two through six are of especially great importance to all entomologists, since procedures for the preparation and dissemination of entomological information are already beginning to undergo some radical changes and will continue to do so in the near future. Indeed, in terms of its information storage and retrieval concerns, Arnett hopes that his book will introduce entomologists to those changes now in progress, and that by so doing it will help accelerate its own obsolescence. For the professional ento- mologist, therefore, the time to read this book is now. Donald R. Whitehead Department of Entomology University of Alberta Edmonton 7, Alberta Book Review MATSUDA, R. 1970. Morphology and Evolution of the Insect Thorax. Memoirs of the Entomological Society of Canada. Ottawa. No. 76. 431 pp., 172 Fig., 24 tables, 744 references, subject and author indices. (Memoirs are included in the subscription price for the Canadian Entomologist.) This is the second in a series of monographs by Matsuda analyzing structural evolution in insects. He began this project to organize the large quantity of published information which had accumulated since the appearance of Snodgrass’ textbook on the subject in 1935. Perusal of the bibliography supports his rationale: of 744 references only 186 appeared before 1935. There are 424 references in English, 180 in German, 79 in French and 61 in other languages chiefly Russian and Italian. As the German and French contributions have importance out of proportion to their numbers the appearance of a review in English is of great value to English-speaking entomologists. This book deals with the thorax but considers the wings and legs only briefly. Matsuda’s conclusions are based largely on his reading, interpretation, and digestion of published works although he contributes original information where required to fill in gaps. The book is divided into two parts: a discussion of general topics on 87 pages, and a discussion on 314 pages which treats in detail selected representatives of each insect order. For most biologists the first part is of greater use. In it Matsuda establishes the primitive organization of the pterygote thorax. This necessitates a summary of his conclusions from part II. He discusses the neck, tergum, sternum, intersegmental regions, pleuron and aspects of the wings, coxae and spiracles, comments on their embryological and evolutionary ori- gins, analyzes the various theories proposed to explain their evolution, and emphasizes the strengths and weaknesses of each theory. Finally, he presents his own conclusions, synthe- sizing a theory of homology from descriptive and experimental embryology, postembryonic development, genetics, comparative morphology, paleontology and phylogeny. In his discussion of wings Matsuda concentrates on their origin, a subject which has in- trigued many workers as evidenced by the plethora of theories published to explain then- presence. His most interesting discussion here concerns the validity of separating the ptery- gotes into the Paleoptera and Neoptera. Matsuda concludes that this separation is phylo- genetically unsound since the wing mechanism of the Ephemeroptera is very much like that of most neopterous pterygotes, while that of Odonata is not. 285 In the second half of part I, Matsuda considers the thoracic musculature. First, he pre- sents an illustrated and tabulated description of the musculature of the apterygote Lepisma saccharina L. based on the work of Barlet. This serves as a standard of reference for his general discussion of pterygote musculature which follows. Here, he introduces a system of abbreviations used in part II for naming muscles. The final section of part I (Major evolutionary features of thoracic musculature) is the strongest section in the book. Matsuda concludes that the predominant evolutionary trend of the adult thoracic muscles in insects has been a reduction in number. At the same time however, some muscles have been added and in their turn these secondary muscles have decreased in number also in the higher pterygotes. Thus, the thoracic musculature includes two kinds of muscles of different evolutionary origin: (i) those inherited from a wingless antecedant and (ii) those which have arisen de novo in the Pterygota. The former Matsuda calls paleogenetic muscles; the latter neogenetic muscles. A third group, the caenogenetic muscles, are those which have developed exclusively in the immature stages and which have no recognizable homologues in the adults. Matsuda discusses the gross developmental pattern of the thoracic muscles. There are few if any caenogenetic muscles in paurometabolous insects, but in holometabolous insects these tend to increase. Also, the time of appearance of imaginal muscles and degeneration of larval muscles during metamorphosis varies from one order to another (heterochrony). Immature pterygote insects tend to preserve muscles present in the Apterygota whereas the adults of these insects lack them. There is also a tendency for nymphal and larval insects to preserve muscles present in the adults of related but less derived groups. This last observation, as emphasized by Matsuda, accords with the biogenetic law of Muller-Haeckel i.e. that onto- geny recapituales phylogeny. Matsuda closes part I with a discussion of the underlying developmental mechanisms for production of new muscles. Differences in the growth rate of epidermal cells and differences in times of connection between muscles and the epidermal cells in various insects result in the production of homologous muscles with new points of attachment and hence new func- tions. The development of ectodermal parts is often dependent on muscles after the connec- tion of the two is established but not before. Here, Matsuda should have cited the work of Sahota and Beckel (1967. Can. J. Zool. 45:407-434) who showed experimentally that in Galleria mellonella L. the topographic relationships between flight muscle myoblasts and the epidermis is the causative factor in determining the orientation of developing flight muscles. Homologous muscles may be inserted on the cuticle in Pterygota and on the epidermis in Apterygota and larval Pterygota. Some muscles are replaced by ligamentous structures which are ectodermal in origin. The flight muscles have been derived differently in different orders. Fibrillar and close- packed types of flight muscles are specializations of the normal, non-flight muscles which are tubular. Much of this argument is based on Tieg’s (1955) classic study of flight muscles, a paper which, in this book, finally receives the recognition it deserves. In part II, each account of an order is illustrated with fully-labelled drawings. Most of these are modified from other works, but some are based on Matsuda’s own observations. Many of the copied illustrations are not of the quality of the originals but all of them show clearly what they are intended to show. Most of the analyses are accompanied by a table providing a uniform system of designation for the muscles found by different investigators in different species. Each table is followed by a list of remarks muscle by muscle. Where information is available both adult and larval musculature and its metamorphosis are dis- cussed. The structural basis for flight is summarized and alary polymorphism is referred to if it occurs in the group under discussion. 286 Matsuda’s phylogenetic conclusions, which are highly controversial, are derived with an appreciation for Hennig’s (1966) principles and terminology. He suggests that Embioptera and Phasmida are derived sister groups of Plecoptera. His belief that the Blattaria, Isoptera, Mantodea, Grylloblattodea, and Orthoptera belong together is evidenced by his tabular comparisons of the thoracic muscles of individuals in these different orders. He supports Crampton’s (1918) idea that the Dermaptera and Coleoptera are closely related to each other and thus implies that the beetles are only distantly related to the other holometa- bolous orders. Zoraptera are related both to the hemipteroid orders and to the Isoptera and Plecoptera and Matsuda proposes that the hemipteroids were derived from the same protor- thopteroid ancestry as Isoptera. Suprisingly, Matsuda does not discuss the relationships of the panorpoid complex but in grouping most of these orders (Mecoptera, Diptera, Lepidop- tera, Trichoptera) together he implies support for Hinton’s (1958. A Rev. Ent. 3: 181-206) interpretation. Hymenoptera he considers to be a sister group of the Mecoptera and the placing of Strepsiptera, he says, requires further study. In leaving consideration of Odonata to the end of the book, Matsuda underlines his belief that these insects are only distantly related to the other pterygotes. He proposes that this order could have arisen from a machilid- or japygid- like ancestor and intimates that odonate peculiarities have been derived in such a way as to obscure this order’s relationships with the Machilidae and Diplura. I am not convinced by Matsuda that the ectognathous Odonata are closely related to entognathous Diplura. As the structures comprising the thorax of insects form a functional complex, they are liable to change with changes in the mode of life of the organisms. Thus, insects in different orders with similar methods of locomotion could have similar thoraces arising through con- vergence. Adaptive features often are of limited value in phylogenetic hypothesizing if they are not used in conjunction with many other characters of different functional significance. I suggest that phylogenies erected on the basis of thoracic structure alone will not persist for very long. I respect this author for his ability at synthesis. Some workers have criticized Matsuda for relying too heavily on the work of others. Nevertheless Matsuda’s conclusions are his own and could only be arrived at by one who has a comprehensive appreciation of thoracic com- plexity in the whole of the Insecta. To have mastered this amount of information and still found time for original investigation is a remarkable accomplishment. The book reveals that insect morphology is an active field of investigation, points out that a great amount of information remains to be obtained particularly in the develop- mental area, and hopefully, shows non-morphologists that this is not the dull subject that many of them believe it to be. I am looking forward to the appearance of future volumes by Matsuda. Bruce S. Heming Department of Entomology University of Alberta Edmonton 7, Alberta Publication of Quaestiones Entomologicae was started in 1965 as part of a memorial project for Professor E. H. Strickland, the founder of the Department of Entomology at the University of Alberta in Edmonton in 1922. It is intended to provide prompt low-cost publication for accounts of entomological research of greater than average length, with priority given to work in Professor Strickland’s special fields of interest including entomology in Alberta, systematic work, and other papers based on work done at the University of Alberta. Copy should conform to the Style Manual for Biological Journals published by the American Institute of Biological Sciences, Second Edition, 1964, except as regards the abbreviations of titles of periodicals which should be those given in the World List of Scientific Periodicals, 1964 Edition. The appropriate abbreviation for this journal is Quaest. ent. An abstract of not more than 500 words is required. All manuscripts will be reviewed by referees. Illustrations and tables must be suitable for reproduction on a page size of 93/4x63/4 inches, text and tables not more than 73/4X43/4 inches, plates and figures not more than SV2 X5 inches. Reprints must be ordered when proofs are returned, and will be supplied at cost. Subscription rates are the same for institutions, libraries, and individuals, $4.00 per volume of 4 issues, normally appearing at quarterly intervals; single issues $1.00. An abstract edition is available, printed on one or both sides (according to length) of 3X5 inch index cards (at $1.00 per volume) or on 5X8 inch standard single row punched cards ($1.50 per volume) . Communications regarding subscriptions and exchanges should be addressed to the Subscription Manager and regarding manuscripts to: The Editor, Quaestiones Entomologicae, Department of Entomology, University of Alberta, Edmonton, Canada. t D- Q i Quaestiones entomologicae LIBRARY AUG 20 1971 HARVARD UNIVERSITY A periodical record of entomological investigations, published at the Department of Entomology, University of Alberta, Edmonton, Canada. VOLUME VII NUMBER 3 JULY 1971 QUAESTIONES ENTOMOLOGICAE A periodical record of entomological investigation published at the Department of Entomology, University of Alberta, Edmonton, Alberta. Volume 7 Number 3 23 July 1971 CONTENTS 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 (Dipteral Tabanidae) from Moss 407 Announcement - 409 Guest Editorial A committee has been established in the Province of Alberta to enquire into non- Canadian influence in post-secondary education in the province. There is a non-Canadian component in this university and especially in the distribution list of Quaes tiones entomo- logicae. To these people among others both the existence of this committee and the follow- ing statement of the president of the university will be of interest: Excellence has no Nationality Let me begin by making it clear to members of the Committee that I appear before you as one person, not as a representative of The University of Alberta. Although I am aware that other people and other groups from The University of Alberta will present ideas to you, no consensus of opinion has been sought, and there will be no opinion given by anyone or any group which might be interpreted as the official opinion of The University of Alberta. It is also important for me to say that I am aware of the unemployment situation in Canada, and that the possibility of high unemployment at all levels is of major concern to me. The University of Alberta employs several thousands of people, both with and without university degrees. It therefore goes without saying that our University must adopt employ- ment procedures which will help to alleviate the high incidence of unemployment among Canadian citizens, and among those to whom our government has granted the right to live and work in Canada. However, the purpose of your inquiry is not economic in nature, and I mention employ- ment procedures for the sole purpose of saying that universities are aware that their situ- ation in the 1970’s will be far different from what it was in the 1960’s. The question of “who should teach at The University of Alberta” is not new. Let me quote from the minutes of a meeting of a governing body of this University: “Dr. Jenkins expressed his disapproval of the idea of appointing all the professors and assistants outside of the province. He was strongly in favor of utilizing the material available in the province and he felt sure that many men now engaged in educational work were emin- ently qualified to undertake the higher branches of learning and he felt an injustice would be done the whole of the teaching profession of the province if only outsiders were ap- pointed.” 288 The sentiment was expressed at the first meeting of the Senate of The University of Al- berta on March 30, 1908. This was before a single member of the staff, other than the President, had been appointed to The University of Alberta. Fortunately the matter was not taken further. I say “fortunately” because I feel strongly that this University would not enjoy the respect it has today if Dr. Jenkins had had his way sixty-three years ago. Since you are a Committee of Inquiry into non-Canadian Influence in Alberta Post- Secondary Education, sooner or later you will have to define what will be meant by a “non-Canadian influence,” and to propose the procedures by means of which such an in- fluence can be measured. Presumably you will have to be able to identify a good “non- Canadian influence” from one that, by some means or other, will be deemed to be bad. Your task will not be easy because in a rather restricted sense one might say that there is no real Canadian content in the science and mathematics courses taught in Canada today. Let me explain what I mean. Education generally, and higher education particularly, is a world of ideas. If one traces the sources of the big ideas of science and mathematics, they will not be found to be Canadian, nor, in fact, were they discovered in the United States. Up until the middle of the present century, these ideas came from Great Britain, France, Germany, Switzerland, Italy, and even smaller countries like Denmark made major contributions to the world of ideas. During the first fifty years of the present century, it was common practice for the men and women of Canada and the United States to go abroad for post-graduate education. Indeed, these people coupled with the thousands of people who were brought to this continent from abroad developed the educational systems of Canada and the United States. Who were the people who dominated the world of scientific ideas during the first half of the twentieth century? As illustration only, there was Einstein of Germany, Fermi of Italy, Weyl of Switzerland, Dirac of England, Bohr of Denmark, Wigner and Von Neumann of Hungary, and De Broglie of France. Even if the list were complete, no Canadian and few Americans would be contained among those who contributed to the world of big ideas up till 1950. Since that time the United States and Russia have come to the fore. The ideas that led to the development of computer technology, to the exciting ideas being developed in genetics and the life sciences, to laser technology, and to transportation and communications technologies, to name but a few, should make us aware that the time has not yet come when Canadians can arrogantly say that we shall lock out the world of ideas that exists beyond our borders, and go the rest of the way by ourselves. How does one classify the study of arithmetic, dependent as it is on the Arabic number system? Is this a non-Canadian influence? How does one classify the study of geometry, dependent as it is on the ideas of Euclid and Pythagoras? Is this a non-Canadian influence? I do not envy you the task that has been placed into your hands, and do not know how you will accomplish it. It is my fear however that groups are raising the '"non-Canadian influence” issue on a meaningless statistical basis, and that conclusions will be drawn, and, indeed, have already been drawn, which are not warranted by the statistics. If these issues develop into a witch- hunt in Canada, then I want no part of it. I lived through the McCarthy era, and friends of mine were made uncomfortable and unwelcome at their universities. Some were dismissed and some went to jail, not because they were Communists, but because they chose to fight for academic freedom, the right to seek truth as they saw fit and the right to teach and publish the truth as they saw it. During the 1 960’s, Canada sought and brought men and women from the four corners of the world to help us solve what then seemed to be an insoluble problem: the problem of expanding enrolments with too few people qualified to teach. If, after making a major con- 289 tribution to our country, such men and women are made to feel uncomfortable and unwel- come, then this will indeed be a sorry way to show our gratitude. There is no reason to believe that the search for truth will ever end at the borders of Canada. We are a young country and a small nation. There is every reason to believe that the majority of the big ideas of science, by means of which scientific knowledge is acquired, will come from outside our borders for many years to come. But Canada must pursue excel- lence, and it must pursue excellence wherever excellence exists. Therefore, Canadians must continue to go outside of our country for study, and leaders of excellence must be brought to Canada to teach their ideas on Canadian soil. Although I have nothing but respect and admiration for the accomplishments of Canadian universities during recent years, we have no Harvards nor Oxfords in our midst. Banting and Best shared the Nobel Prize for their discovery of insulin about half a century ago. No othei Nobel Prize in science has been awarded to a Canadian nor have we been able to attract such people to work in and for Canada. As I said at the beginning, concern over unemployment in Canada is genuine, and univer- sities must help our people to find interesting and rewarding work. However, we must not adopt a policy which will give a short-term gain but will turn into a long-time evil. It will be a tragedy if ever Canada posts signs, or enacts legislation, which will tell the members of the world of ideas they are not welcome here. Excellence attracts excellence, and such a policy will discourage desirable people from coming to Canada, and may, in the long run, encourage desirable Canadians to leave Canada. Max Wyman University of Alberta A TAXONOMIC SURVEY OF THE AGROMYZIDAE (DIPTERA) OF ALBERTA, CANADA, WITH OBSERVATIONS ON HOST-PLANT RELATIONSHIPS VINOD K. SEHGAL Department of Entomology University of Alberta Quaestiones entomologicae Edmonton 7, Alberta 7 : 291-405 1971 Thirty-one new species are described and eleven further are recorded from Alberta for the first time . This brings the total of Albertan Agromyzidae to 170 and that for Canada and Alaska to 321 species. Genitalia of all males representing new species are illustrated. Keys to Albertan genera and species, diagnostic characters and data on larval host-plants are given. Host-plant relationships are discussed. In der vorliegenden Arbeit einunddreizig neu Agromyzid Arten sind beschrieben und weitere elf von Alberta sind zum erstemal aufgefiihrt. Das bringt die gesamte Zahl von Alberta auf 170 und diese von Kanada und Alaska auf 321 Arten. Die mannliche Genitalien alle neue Arten sind abgebildet. Die Bestimmungsschliissel fur die Gattungen und Arten von Alberta, ihre Bezeichnungen den Merkmale und Einzelhalten iiber Raupenfutter sind angegeben. Die Angaben zur Bionomie sind besprochen. This survey of the family Agromyzidae in Alberta was started in the summer of 1966, when this was one of the most poorly known families of cyclorrhaphous Diptera in Canada. Strickland (1938, 1946) listed 20 species as occurring in Alberta. The onlv comprehensive treatment for North American Agromyzidae was by Frick (1959) who included those known from Canada. Since the publication of Frick’s synopsis, the concept of certain genera of Agromyzidae has been revised by Nowakowski (1962) on the basis of his studies of male genitalia. As new characters of male genitalia were discovered, it became necessary to con- firm all determinations of species, which were previously based on external morphology and sometimes only on female specimens. One of my main objectives was to collect, determine, and describe as many species as possible in order to assess the family Agromyzidae in Alberta and in Canada, and to provide keys for the determination of genera and species. This investigation in Alberta was started parallel to Spencer’s 1969 synopsis of the Agro- myzidae of Canada and Alaska. As a preliminary report on these investigations I described 13 new species from Alberta (Sehgal, 1968). Spencer examined all previously known records of Canadian and Alaskan Agromyzidae and confirmed a total of 290 species. Among the Canadian species Spencer (1969) recorded 128 described species as occurring in Alberta. As a result of this study 3 1 new species are described from Alberta and 1 1 further species have been recorded for the first time as occurring in Alberta. This brings the total of Albertan Agromyzidae to 170 and that of Canadian and Alaskan Agromyzidae to 321 described species (Table 1). The male genitalia of almost all Albertan species have been examined. Any record of Albertan Agromyzidae outside the present work must be con- sidered tentative, until confirmed on the basis of male genitalia. 292 Sehgal Table 1. Distribution among genera of new species, new Alberta records, and all known species of Agromyzidae from Alberta, and Canada and Alaska. Total number of species now known from: New New Alberta Canada and Genus species records Alberta Alaska Agromyza 2 2 17 36 Melanagromyza 3 — 9 20 Hexomyza — — 1 2 Ophiomyia 1 — 17 27 Phytobia Cerodontha 1 3 8 Dizygomyza — 2 7 18 Poemyza — — 5 8 Icteromyza — — 2 6 Cerodontha — — 2 3 Calycomyza — 1 2 13 Amauromyza 2 — 2 5 Nemorimyza — — 1 1 Liriomyza 6 1 27 42 Lemurimyza — — 1 3 Metopomyza 1 — 2 4 Praspedomyza — — 1 1 Haplomyza — — 1 1 Phytoliriomyza — — 1 2 Paraphytomyza 1 — - 5 7 Pseudonapomyza — — 2 2 Napomyza — — 3 9 Phytomyza 15 4 59 98 Other genera — — — 5 TOTAL 31 11 170 321 MATERIALS AND METHODS As a part of this study I made intensive collections of agromyzid specimens from the province of Alberta, Canada, including the Alberta Rockies. Some specimens collected by others, especially G. C. D. Griffiths, K. A. Spencer, the late Professor E. H. Strickland, and B. Hocking, from Alberta localities were also examined and are acknowledged in the list of material. Attempts were made to rear adults from immature stages as far as possible. This per- mitted me to clarify the biology of many species. I studied approximately 1 200 specimens and examined genitalia preparations of 620 specimens. Genitalia of all males representing new species and a few other species where necessary for their specific determination have been illustrated. Reference is made to good earlier illustrations of male genitalia. The holo- types and allotypes of all new species will be deposited in the Canadian National Collection, Ottawa. Agromyzidae of Alberta 293 The terminology used in describing new species is that employed by Spencer ( 1 969) and other workers in Agromyzidae. The frons width and eye width are measured at the level of the median front ocellus from above. The term gena here means the area below the eyes including the lower orbits. The ratio of genal and eye heights are measured mid-way be- tween vibrissal and posterior margins. The terminology of the aedeagus is that used by Frick (1952) and Spencer (1969). Abbreviations acr, arcostichal hair; Adap, aedeagal apodeme; Ar, arista; As3, third antennal article; Bsph, basiphallus; C, costa; dc, dorsocentral bristles; Dph, distiphallus; Ejap, ejaculatory apodeme; Ejb, ejaculatory bulb; Ejd, ejaculatory duct; Hypa, hypandrium; m-m, medial crossvein; M1+ 2 and M3 + 4, median veins; Mph, mesophallus; oc, ocellar bristles; Orijower orbital bristles; Ors, upper orbital bristles; os, orbital setulae; Pgo, postgonites; Phph, phallo- phore; Prgo, pregonites; Pvt, postvertical bristles; Rl5 R2+ 3 and R4+ 5 , radial veins; r-m, radiomedial crossvein; Sc, subcosta; Vi, Vibrissa; Vte, outer vertical bristle; Vti, inner verti- cal bristle. CHARACTERISTICS OF THE FAMILY AGROMYZIDAE The main distinguishing characters of the members of this family are as follows: Head. — Postvertical bristles divergent; distinct orbital bristles present, normally two strong upper orbital bristles, Ors, directed upwards and two lower orbital bristles, Ori, directed inwards and upwards; orbital setulae present; distinct vibrissal hair present, repre- sented by a bunch of fused hairs or ‘vibrissal horn’ in some males of the genus Ophiomyia Braschnikov; centre of frons without bristles or setulae. Mesonotum. — Distinct dorsocentral bristles present, normally 3+1, sometimes a few anterior bristles are reduced or lost; variable number of acrostichals present; scutellum normally with four scutellar bristles, two in subgenus Cerodontha Rondani. Wing. — Costa broken at end of subcosta, extended to apex of vein R4+5 or M1+2; sub- costa weakly developed distally, adjacent to, and either joined to or independent from Rx distally; crossvein r-m present; anal vein shortened, not extended to the wing margin. Male genitalia. — Hypandrium large and well developed; pregonites and postgonites normal- ly well differentiated, former sometimes fused with hypandrium; epandrium large and con- spicuous; surstyli and cerci normally well developed and with characteristic setae or setulae; aedeagus complex; aedeagal apodeme large and darkly sclerotized; aedeagal hood conspicu- ous; aedeagus with distinct basal section consisting of basiphallus and phallophore and distal section consisting of various sclerites forming mesophallus, paraphallus, and disti- phallus; terminal section of ejaculatory duct inside distiphallus normally bifid; ejaculatory apodeme normally well developed, sometimes reduced, bulb conspicuous. Female postabdomen. — Seventh segment completely sclerotized, forming a conical ovi- positor sheath; eighth segment elongate, retractible into seventh, bearing numerous anteri- orly directed denticles; pair of egg guides around gonopore; anus well beyond gonopore; two spermathecae. Larvae. — Anterior spiracles on first abdominal segment approximate; posterior spiracles on last or eighth abdominal segment approximate or widely separated; mouth hooks well developed and almost vertical in relation to labial sclerite; paraclypeal phragma normally with dorsal and ventral arms; muscle scars and tubercle bands strongly developed along lateral portions of abdominal segments. 294 Sehgal Biology. — Larvae feed inside the living tissue of angiosperms (except those of Pterido- myza Nowakowski, Liriomyza felti (Mall.) and Phytomyza scolopenderii R.-D. v/hich feed on ferns and few representatives of the genus Liriomyza Mik feeding on horsetails and liverworts). Larvae show varying host-plant specificity, but normally feed on one plant organ. Most species feed as leaf miners, some feed inside the stems, seeds, and other parts of plants; a few cause galls ( Hexomyza Enderlein). TAXONOMIC TREATMENT The generic concept used in this study is the one which has come to be generally accepted (Spencer, 1969), but is certainly not final in agromyzid classification. As recent studies pro- gress on the male genitalia, the need for defining some of the larger genera on a monophy- letic basis becomes more apparent. No attempt is made in this study to undertake any generic revision of the existing classification. According to the biological species concept (Mayr, 1963), species are defined to be groups of actually or potentially interbreeding natural populations, which are reproductively iso- lated from other such groups. I accept this species definition. The application of this species concept presents many practical difficulties and some of these have been discussed by Mayr (i969). The reproductive isolation of a population is usually inferred indirectly from com- parisons of external morphology. In Agromyzidae, besides the evidence of external mor- phology, various other evidences are helpful. The majority of agromyzid species are restric- ted feeders, either monophagous or oligophagous, confined to botanically related plant species. It seems highly unlikely that significant gene interchange occurs between popula- taions restricted to unrelated host-plants, even if the morphological differences between them are minor. Such populations can be assumed to represent distinct species, so long as the observed differences are shown consistently. Often other evidence such as larval mor- phology’ or shape of mine is available to support this assumption. Spencer (1969) discussed briefly the significance of various taxonomic characters current- ly in use in agromyzid taxonomy; the same criteria have been accepted here. The characters of male genitalia have been used as far as possible in order to provide a basis for more accurate determination of species. Information on Albertan host-plants and biology, wher- ever available, has been included. The following key represents a further development of that originally produced by Hendel (1931) and modified by Frick (1952, 1959) and Spencer (1969). Key to genera of Albertan Agromyzidae 1(0). Subcosta developed throughout its length, coalesced with Rj before contact with costa 2 Subcosia weakly developed distally, joined with costa independent of Rj .... 5 2(1). Prescutellars absent; dorsocentrals, two, three or four pairs, if three or four pairs halteres black 3 Prescutellars present; at least three pairs of dorsocentrals; halteres yellow Agromyza Fallen, p.296 3(2). Mesonotum or abdomen normally with greenish, bluish or coppery metallic sheen; antennae not separated by conspicuous keel; aedeagus with basiphalius U-shaped; larvae normally with conspicuous horn in centre of posterior spiracles; larvae feed inside stems or seeds Melanagromyza Hendel, p.303 Mesonotum and abdomen black; aedeagus with basiphalius with long side arms; posterior spiracles of larva without central horn 4 Agromyzidae of Alberta 295 4(3). 5(1). 6(5). 7(6). 8(7). 9(8). 10(9). 11(10). 12(11). 13(12). 14(7). 15(14). 16(15). 17(6). 18(5). Antennae separated by conspicuous raised facial keel (absent from pulicaria group); male with or without distinct vibrissal horn; larval posterior spiracles with more than three bults; larva feeds below stem epidermis or as leaf miner Ophiomyia Braschnikov, p. 3 10 Antennae approximate; male without vibrissal horn; larval posterior spiracles v/ith three bulbs; larva gall producer Hexomyza Enderlein, p. 309 Orbital setulae erect or reclinate, rarely absent 6 Orbital setulae distinctly proclinate .18 Costa extended to apex of vein M1+2, if only to R4+5 then either crossvein m-m absent ( Phytobia confessa Spencer) or lunule broad, distinctly higher than semi- circle ( Cerodontha (Dizygomyza) frankensis Spencer) 7 Costa extended to apex of vein R4+5 17 Scutellum normally dark, concolourous with mesonotum 8 Scutellum yellow; vein Mj +2 ending nearest wing tip 14 Halteres with knob white or yellow 9 Halteres with knob black or partially darkened, if yellow distiphallus with numer- ous spinules Amauromyza Hendel, p.326 Crossvein m-m absent Haplomyza Hendel, p. 344 Crossvein m-m present 10 Vein R4+s ending nearest wing tip; larvae cambium miners Fhy to bia Lioy , p . 3 1 6 Vein Mj +2 ending nearest wing tip 11 Third antennal article with conspicuous spine anterodorsally, scutellum with only two bristles; or lunule higher than semicircle, narrow or triangular; or lunule broad, antennal bases widely separated, third antennal article in male distinctly enlarged Cerodontha Rondani, p.317 Not so 12 Fore-tibia with lateral bristle; abdomen in male yellowish; larva makes character- istic blotch mine with concentric rings on Aster and Solidago in Nemorimyza posticata (Meigen) Nemorimyza Frey, p. 327 Fore-tibia without lateral bristle; leaf mine not as above 13 Frons dark; orbits raised above plane of frons .... Praspedomyza Hendel, p. 343 Frons yellow, orbits in same plane as frons Calycomyza Hendel, p. 325 Frons yellow; orbits in plane of frons 15 Frons dark; orbits raised above plane of frons Metopomyza Hendel, p. 342 Crossvein m-m absent; one Ors Haplomyza Hendel, p. 344 Crossvein m-m present (absent in Liriomyza singula Spencer); twro Ors 16 Prescutellar area yellow; orbital setulae normally 'erect; aedeagus with sclerotized paired tubules in the distiphallus, epandrium with conspicuous black spines .... Lemurimyza Spencer, p. 341 Prescutellar area normally dark, sometimes yellow; orbital setulae reclinate; aedea- gus variable in form, but not as in the genus Lemurimyza Spencer . Liriomyza Mik, p. 328 Crossvein m-m either absent, or if present well beyond r-m Paraphytomyza Enderlein, p. 345 Crossvein m-m basal to r-m Pseudonapomyza Hendel, p. 345 Crossvein m-m basal to or at same level as r-m Napomyza Westwood, p. 349 Crossvein m-m absent Phytomyza Fallen, p. 350 296 Sehgal Genus Agromyza Fallen Agromyza Fallen 1810:21. The main distinguishing characters of this genus are subcosta developed throughout its length and coalesced with Rj before contact with costa; at least three pairs of dorsocentrals; prescutellars present and halteres yellow. This genus is represented in Alberta by 17 species, of which two are described as new. The species in this genus form a very diverse assemblage of many groups. The species in the ambigua group, albertensis Sehgal, aprilina Malloch and kincaidi Malloch; as well as species in the nigripes group, albipennis Malloch, brevispinata new species, hockingi Spencer and nigripes Meigen; like other members of the ambigua /nigripes groups (Griffiths, 1963) form a single group of grass mining species with similarity in shape of the distiphallus. Members of the ambigua group differ from those of the nigripes group only by a shortened costa, not extended beyond the apex of vein R4+5, and longer distiphallus. The Urtica miner, pseudoreptans Nowakowski, belongs to the reptans group, while the Mertensia miner, cana- densis Malloch, belongs to the rufipes group. The species in the spiraeae group, populoides Spencer, vockerothi Spencer, fragariae Malloch, masculina Sehgal and spiraeae Kaltenbach, are believed to be closely related due to similarities in the male genitalia. This concept was first proposed by Sasakawa (1961). The species in this group have asymmetric sclerotization of the basiphallus and mesophallus. A. aristata Malloch, whose larvae mine the leaves of Ulmus americana L., family Ulma- ceae, is probably close to the rubi /spiraeae group of Sasakawa (1961). Another group of species whose members are characterized by 3+1 strong dorsocentrals and yellow frons is represented in Alberta by nearctica new species. Three further females belonging to this group cannot be identified in the absence of males. The leaf mines on members of Geum allepicum Jacq., Potentilla sp., and Rosa acicularis Lindl., of the family Rosaceae probably represent those of the members of the spiraeae group, but no flies have been bred from these hosts. Key to Alberta species of the genus Agromyza Fallen 1(0). Dorsocentrals 3-6, decreased significantly in size anteriorly; presuturals as well as anterior postsutural dorsocentrals usually not distinguishable from acrostichals 2 Dorsocentrals 3+1 , strong and distinct 12 2( 1 ). Wing tip near apex of vein M x +2 sulfuriceps Strobl, p. 303 Wing tip near apex of vein R4+5 or midway between R4+5 and Mx +2 3 3(2). Legs largely yellowish 4 Legs largely black or brown 5 4(3). Antennae yellow, smaller specimens, wing length 2.2 to 2.7 mm; larvae leaf miner on Ulmus americana L aristata Malloch, p.298 Antennae dark brown; larger specimens, wing length about 3.0 mm canadensis Malloch, p. 299 5(3). Costa extended to apex of vein R4+5 6 Costa extended to apex of vein M j +2 .• 8 6(5). Squamal fringe brown or black; larger specimens, wing length about 2.7 to 3.5 mm kincaidi Malloch, p300 Squamal fringe pale or whitish, smaller specimens 7 7(6). Third antennal article with distinct angle or point anterodorsally; eyes upright, normal albertensis Sehgal, p. 297 Agromyzidae of Alberta 297 8(5). 9(8). 10(9). 11(9). 12(1). 13(12). 14(13). 15(14). 16(15). Third antennal article rounded at end; eye distinctly slanted aprilina Malloch, p. 298 Mesonotum at least weakly shining black; mesophallus and distiphallus single S-shaped sclerotization 9 Mesonotum distinctly mat black; mesophallus and distiphallus separate scleroti- zations pseudoreptans Nowakowski, p. 302 Third antennal article with distinct angle anterodorsally 10 Third antennal article rounded at tip 11 Squamal fringe pale or whitish; ejaculatory bulb very broad albipennis Meigen, p.297 Squamal fringe dark or brown; ejaculatory bulb narrower nigripes Meigen, p. 302 Surstyli with 3-6 distinctly large spines (Fig. 6) hockingi Spencer, p. 300 Surstyli with smaller spines (Fig. 5); aedeagus as in Fig. 3,4 brevispinata n. sp., p. 298 Frons reddish yellow; orbits black; second and third antennal articles black; aedeagus as in Fig. 10 nearctica n. sp.,p. 301 Frons dark, brown or black 13 Squamal fringe pale populoides Spencer, p. 302 Squamal fringe darker, brown or black 14 Mid-tibia with a strong bristle posteriorly vockerothi Spencer, p. 303 Mid-tibia without a distinct bristle . 15 Frons distinctly brown fragariae Malloch, p. 300 Frons distinctly mat black 16 Gena narrow one-eighth to one-tenth eye height; basiphallus and mesophallus with sclerotized strips masculina Sehgal, p301 Gena broader, about one-fifth eye height; distiphallus separated from basiphallus by long membranous section spiraeae Kaltenbach, p.302 Agromyza albertensis Sehgal Agromyza alber tensis Sehgal 1968:57; Spencer, 1969:32. Comparisons and diagnostic characters. — The members of this species differ from those of a similar species, kincaidi Malloch, in smaller size, wing length 2. 0-2. 7 mm and in having pale squamal fringe, and from those of ambigua Fallen in having frons less projected above eyes in profile. The main distinguishing characters are the conspicuous angle on the third antennal article and distinct male genitalia. Sehgal (1968) illustrated the head, wing and male genitalia characteristic of this species. Spencer (1969) also figured the aedeagus. Biology. — Not confirmed, but larvae probably mine leaves of grasses (Gramineae). Geographical distribution. — Known only from Alberta, from the following localities: CANADA. Alberta: Banff, Blairmore, Elkwater. Agromyza albipennis Meigen Agromyza albipennis Meigen 1830: 171 ; Spencer, 1969:32. Diagnostic characters. — The members of this species may be recognised by the combina- tion of characters given in the key. The distinctive aedeagus and ejaculatory bulb are as illustrated (Fig. 1, 2). Sasakawa (1961) and Griffiths (1963) described in detail and illus- trated significant diagnostic characteristics of this species. 298 Sehgal Biology \ — Larvae are known to mine the leaves of grasses (Gramineae) (Sasakawa, 1961 ; Griffiths, 1963). Geographical distribution. — The members of this species are Holarctic in distribution, known from numerous localities in Europe (Griffiths, 1963), Japan (Saskawa, 1961) and Canada (Spencer, 1969). I have examined three specimens from the following localities: CANADA. Alberta: 1 9 Cypress Hills, near Elkwater Lake, 24.vi.1966; 1 6 Devon, University of Alberta botanical garden, 22.vi.1966; 1 6 Edmonton, Rainbow Valley, 14.vi. 1968. Agromyza aprilina Malloch Agromyza aprilina Malloch 191 5b: 35 9; Spencer, 1969:36. Comparison and diagnostic characters. — The members of this species resemble closely those of A. kincaidi Malloch and can be reliably separated only by examination of the characters of male genitalia. Other external differences are pale squamal fringe, smaller size and distinctly slanting eyes. Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — Not confirmed, but the larvae probably mine the leaves of grasses (Gramineae). Geographical distribution. - The members of this species are Nearctic in distribution and are known from United States and Canada. The known Alberta locality is: CANADA. Alberta: Banff (Spencer, 1969). Agromyza aristata Malloch Agromyza aristata Malloch 1915a: 13; Spencer, 1969:38. Agromyza ulmi Frost 1924:54; Frick, 1957:199. Diagnostic characters. — The members of this species may be recognized by the combina- tion of characters given in the key. Spencer (1969) illustrated the distinctive aedeagus. Biology. — Larvae make elongated blotch mines on the upper surface of the leaves of Ulmus americana L., family Ulmaceae. The leaf mines appear in great numbers around Edmonton during the second week of June. The members of this species have only one generation a year in Alberta. Geographical distribution. — The members of this species are widespread in United States (Frick, 1959) and are also known from Canada (Spencer, 1969). I have examined the fol- lowing material from Alberta: CANADA. Alberta: 2 99 Edmonton, University of Alberta campus from leaf mines on Ulmus americana L., coll. 6.vi. 1968, emerged 2-27. v. 1969, coll. G. C. D. Griffiths; 1 9 same locality, swept over same host, 3.vi.l969; Numerous leaf mines on Ulmus americana L., same locality, 9.vi.l966, 5.vi.l967, lQ.vi.1968. Agromyza brevispinata new species Comparison and diagnostic characters. — The male of this species differs from those of the similar species hockingi Spencer and lucida Hendel in having a weakly shining or some- what mat mesonotum and distinct male genitalia. This species is included in Spencer’s (1969) key to Canadian species of the genus Agromyza Fallen by amending and extending the couplet as follows: 26. Third antennal segment distinctly cut away below (Spencer, 1969, Fig. 5) nigripes Meigen Agromyzidae of Alberta 299 Third antennal segment rounded at tip 26a 26a. Mesonotum somewhat mat or weakly shining black; surstyli with 8-10 small spines; aedeagus as in Fig. 3, 4 brevispinata n.sp. Mesonotum entirely shining black 27 Description. — Head. Frons slightly wider than width of eye at level of front ocellus, not projected in front of eye margin in profile; two strong Ors directed upwards; two Ori direc- ted inwards and upwards; orbital setulae reclinate; eyes oval, approximately 1 .3 times higher than their length; gena deepest at rear, approximately one-fifth eye height midway between vibrissal and posterior margins; third antennal article rounded at tip; arista long and pubes- cent. Mesonotum. Two strong dorsocentrals; acr in about nine irregular rows. Wing. Length in male about 2.8 mm; costa extended to apex of vein M1+2; wing tip nearest to vein R4 +5 ; crossvein r-m approximately at centre of discal cell. Male genitalia (Fig. 3-5). Hypandrium with short apodeme and broad pregonites; surstyli (Fig. 5) with 8-10 small spines; aedeagus as illustrated (Fig. 3, 4). Colour. Frons and orbits mat black; ocellar triangle weakly shining black; mesonotum weakly shining mat black; squamae yellow, fringe dark brown. Derivation of the specific name. — The name brevispinata is given in view of the small spines on surstyli. Biology. — Not confirmed, but larvae probably mine leaves of grasses (Gramineae). Geographical distribution — This species is known from a single male collected at the following locality: CANADA. Alberta: Holotype 6 St. Albert near Edmonton, 18.vi.1967. Agromyza canadensis Malloch Agromyza canadensis Malloch 19 13a: 299; Spencer, 1969:39. Comparison and diagnostic characters. - The members of this species are large flies, wing length approximately 3.0 mm and are distinctive in having yellowish brown legs and dark antennae. They have male genitalia which appear indistinguishable from those of A. pseudonifipes Nowakowski. The two previously known Canadian specimens of this species are brownish flies (Shewed, 1953) while the members of European A. pseudonifipes are darker in colour. This was the basis of Spencer’s (1969) accepting them as different species. The bred male from Yukon Territory, Canada, however, is darker in colour. The discovery of this dark specimen casts doubt on the separation of these two species on the basis of colour. The name A. pseudonifipes Nowakowski will probably prove to be a junior synonym of A. cana- densis Malloch. Biology. — Larvae make blotch mines on the leaves of Mertensia paniculata (Ait.) G. Don, family Boraginaceae. Pupation takes place outside the leaf mine. Geographical distribution. — The members of this species are known from Western and Eastern Canada (Frick, 1959; Spencer, 1969). I have examined the following material refer- able to this species: CANADA. Alberta: Numerous leaf mines on Mertensia paniculata (Ait.) G. Don, Edmon- ton, river bed near University of Alberta campus; White Mud Creek park, July to September 1968; Yukon Territory: 1 6 Dawson City, from leaf mines on Mertensia paniculata (Ait.) G. Don, 5 .viii. 1 968, emerged 22.vi.1969, coll. G. C. D. Griffiths. 300 Sehgal Agromyza fragariae Malloch Agromyza fragariae Malloch 19 13a: 307; Spencer, 1969:42. Comparisons and diagnostic characters. — The members of this species resemble closely those of A. spiraeae Kaltenbach and A. masculina Sehgal but differ in having distinctly brownish frons and distinctive aedeagus. The aedeagus has a characteristic sclerotization in the mesophallus as illustrated by Spencer (1969). Biology. — Larvae are known to mine the leaves of Fragaria virginiana Duchesne, family Rosaceae in United States. Geographical distribution. — The members of this species are known from United States and Canada (Spencer, 1969). From Alberta, Canada they are known from the following localities: CANADA. Alberta: Blairmore; Onefour (Spencer, 1969). Agromyza hockingi Spencer Agromyza hockingi Spencer 1969:44. Diagnostic characters. — The members of this species are small shining black flies, having pale or slightly brownish squamal fringe and distinct male genitalia. Spencer (1969) illustra- ted the aedeagus. The number of bristles on surstyli (Fig. 6) is variable, from three to six. One male collected from Edmonton, White Mud Creek park, 1 3.vi. 1 966 is tentatively referred here as it has the aedeagus very similar to that of hockingi Spencer, but has distinc- tive surstyli (Fig. 7) with two very long spines and a small indistinguishable bristle. This probably represents a further species, but more material is necessary to confirm this opinion. Biology. — Larvae probably mine leaves of grasses (Gramineae). Geographical distribution. — Known from Alberta, New Brunswick, Ontario and Quebec (Spencer, 1969). I have examined the following material from Alberta: CANADA. Alberta: 1 6 Edmonton, White Mud Creek park, 6.vii.l966; 1 6 same locality, viii.1968; 2 99 same locality, 29.vi.1966; 1 6 Elk Island park, 2.viii. 1966; 1 9 same locality, 3 1 vii. 1 966; 1 6 George Lake near Busby, 21 .viii.1966. Agromyza kincaidi Malloch Agromyza kincaidi Malloch 1913a:285; Spencer, 1969:45. Comparisons and diagnostic characters. — The members of this species are large shining black flies, wing length about 3.0 mm, with costa extended to vein R4+5 and dark squamal fringe. The aedeagus (Fig. 8, 9) and surstyli are also very distinctive. Hendel (1931) synonymised the name kincaidi Malloch with ambigua Fallen. Frick (1952, 1959) also accepted it synonymous with ambigua Fallen. Spencer (1965d) concluded that specimens of ambigua Fallen sensu Hendel represented nigrella Rondani and not the true ambigua Fallen. He later (1969) rejected Hendel’s synonymy and re-established this species. Biology. — Larvae probably mine leaves of grasses (Gramineae). Geographical distribution. — Known from Alaska and widespread in Canada (Spencer, 1969. I have examined the following material from Alberta: CANADA. Alberta: 1 9 Cypress Hills, Elkwater Lake, 24.vi.1966; 1 6 Edmonton, White Mud Creek park, 10.vi.1966, 1 6 same locality, 6.vii.l966; 1 6 same locality, viii. 1968; 1 6 Edmonton, 1 10 St. 84 Ave., 15.vi.1968; 1 > Agromyzidae of Alberta 333 Liriomyza eboni Spencer Liriomyza eboni Spencer, 1969:173. Comparison and diagnostic characters. — The members of this species differ from those of the similar species, L. baptisiae (Frost), in having first and second antennal article yellow and acrostichals in two rows. The aedeagus has been illustrated by Spencer (1969). Geographical distribution. — The members of this species are known only from Alberta, from the type locality: CANADA. Alberta: Blairmore (Spencer, 1969). Liriomyza edmontonensis Spencer Liriomyza edmontonensis Spencer, 1969: 174. Comparison and diagnostic characters. — The members of this species resemble closely those of L. sylvatica new species in external morphology and can be reliably separated only by comparison of the characters of male genitalia. Spencer (1969) illustrated the aedeagus characteristic of this species. Geographical distribution. — The members of this species are known from Alberta and British Columbia in Canada. Known Alberta locality is as follows: CANADA. Alberta: Edmonton, University of Alberta campus (Spencer, 1969). Liriomyza eupatorii (Kaltenbach) Agromyza eupatorii Kaltenbach, 1874:320. Liriomyza eupatorii (Kaltenbach), Hendel, 1920: 143; Spencer, 1969: 174. Comparison and diagnostic characters. — The members of this species are very close to those of L. montana Sehgal in external characteristics and are reliably separated only by examination of male genitalia. Spencer (1969) figured the distinctive aedeagus of this spe- cies. The aedeagus of a caught specimen from Alberta is illustrated in Fig. 55, 56. The disti- phallus of this species is very close to that of L. pictella (Thompson) and L. munda Frick, from which it differs only in minor details. Spencer (1965c) illustrated the aedeagus charac- teristic of L. pictella (Thompson) and of L. munda Frick. Biology. — Larvae mine the leaves of members of the genera Solidago, Helianthus, Eupa- torium, Aster , and Lampsana, family Compositae, and Galeopsis, family Labiatae, in Europe (Hering, 1957). Geographical distribution. — The members of this species are widespread in Europe and are known from Canada (Spencer, 1969). Frick’s (1953, 1959) description of L. eupatorii (Kaltenbach) refers to L. munda Frick (Stegmaier, 1966, 1968). I examined the following material from Alberta: CANADA. Alberta: 1 6 Edmonton, Aberhart Hospital lawns, 13.vi. 1967. Liriomyza fricki Spencer Liriomyza trifolii Frick, 1959:410 (not Burgess, 1879). Liriomyza fricki Spencer, 1965c:35. Comparison and diagnostic characters. — The members of this species are very close to those of L. balcanicoides new species in external characteristics, but the male genitalia are very different. Spencer (1965c, 1969) illustrated the distinctive aedeagus of this species. Biology. — Larvae mine the leaves of various species of the genera Medicago, Melitotus, 334 Sehgal Trifolium and Vigna , of the family Leguminosae (Stegmaier, 1968). The flies were also bred from two other genera Lathyrus and Vicia of the family Leguminosae. The leaf mine is a small blotch with a short linear beginning. Geographical distribution. - L. fricki Spencer is a Nearctic species whose members are known from Northern United States and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 d Banff, 28.vi.1966; 1 d Edmonton, 12.vi.1937, coll. E. H. Strick- land; 1 d, 1 9 Elk Island park, from leaf mines on Trifolium repens L., coll. 3 1 .vii. 1 966, emerged 1 4-1 5.viii. 1966; 2 66 same locality, from leaf mines on Vicia americana Muhl., 3 1 .vii.-l 1 .viii. 1 966; 1 9 same locality, from leaf mines on Lathyrus ochroleucus Hook., viii. 1 967 ; 5 66 same locality, 31. vii. 1966; 2 66 same locality, 7.vi. and 2.viii. 1 966; 2 66 Jasper, 18.vi. and 23 .vii. 1966. Liriomyza kenti Spencer Liriomyza kenti Spencer, 1969: 176. Comparisons and diagnostic characters. — The members of this species resemble closely those of L. bifurcata new species and are separated reliably only by examination of the characters of male genitalia. The adults differ from those of similar species, L. senecionivora new species and L. nordica Spencer, in having only two rows of acrostichals. Spencer (1969) illustrated the distinctive aedeagus. Geographical distribution. — The members of this species are known only from the locali- ties of its type series (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 6 Blairmore, 26.vi.1966; 1 6 Edmonton, Emily Murphy park, 1 l.vi. 1968; 1 6 Edmonton, Mayfair park, 17.V.1969; Paratype 1 6 Edmonton, White Mud Creek park, 13.vi.1966; 2 66 same locality, 19.V.1968; 1 d same locality; 1 0.vi. 1 968 ; 10 66 same locality, 25.v. 1969; 5 66 Jasper, 19.vi.1966. Liriomyza lathy ri new species Comparisons. - The members of this species resemble closely those of L. veluta Spencer, L. trifolii (Burgess) and L. taraxaci Hering in external morphology and can be separated reliably only by examination of the characters of the male genitalia. This species is included in Spencer’s (1969) key to Canadian species of the genus Liriomyza Mik by amending couplet 43 and adding couplet 44 as below: 43 Aedeagus as in Fig. 57, 58 lathyri n. sp. Aedeagus not so 44 44. Aedeagus as illustrated (Spencer, 1969) trifolii (Burgess) Aedeagus as illustrated (Spencer, 1969) veluta Spencer Description. — Frons approximately 1.8 times width of eye at level of front ocellus, not projected in front of eye margin in profile; eyes oval, approximately 1.25 times higher than their length; gena approximately one-fifth of eye height midway between vibrissal and pos- terior margins; ocellar triangle small; two Ors directed upwards; two Ori directed inwards; orbits narrow; orbital setulae approximately six, reclinate; third antennal article rounded at tip, with normal pubescence; arista pubescent. Mesonotum. Dorsocentrals 3+1; acr in three irregular rows. Wing. Length in male approximately 1.7 mm; costa extended to vein M1+2; costal seg- ments 2-4 in the ratio of 1 : 0.27 : 0.27; crossvein m-m present; last segment of M3+4 approximately 2.5 times the penultimate. Agromyzidae of Alberta 335 Male genitalia (Fig. 57-59). Hypandrium U-shaped with slender side arms; pregonites broad; postgonites elongated; surstylus typical with a conspicuous spine placed anteriorly; small cone-like projection present on epandrium; phallophore and aedeagus (Fig. 57, 58) as illustrated; ejaculatory apodeme (Fig. 59) broad, darkened at its stem, bulb small, sclero- tized along lower margin. Colour. Frons, orbits, lunule, gena and antennae all yellow or reddish; vte and vti on yellow areas; mesonotum mat greyish black; humeral area yellow, with a dark spot anteri- orly; notopleural area yellow; mesopleura essentially yellow, with dark area centrally and along ventral half; sternopleura black, with a narrow yellow band along its dorsal margin, femora essentially yellow; tibiae and tarsi brownish; squamal fringe dark brown; halteres yellow. Derivation of the specific name. — This species is named lathyri after the generic name of its larval food plant. Biology. — Larvae make large blotch mine with a small linear beginning on the leaflets of Lathyrus ochroleucus Hook., family Leguminosae. Pupation occurs outside the mine. Geographical distribution. - The members of this species are known only from the locali- ties of its type species: CANADA. Alberta: Holotype 6 Edmonton, White Mud Creek park, from blotch mines on leaflets of Lathyrus ochroleucus Hook., coll. 4.ix.l968, emerged 7.ii. 1969; Paratypes 13 66 Elk Island park, 31.vii.-2.viii.1966. Liriomyza lima (Melander) Agromyza lima Melander, 1 9 1 3 :265 . Liriomyza lima (Melander), Frick, 1952:404, 1959:406. Diagnostic characters. — The main distinguishing characters are mat black mesonotum, yellow femora and third antennal article. The pale squamal fringe differentiates this from other species in this group. Spencer (1969) illustrated the distinctive aedeagus. Geographical distribution. — Members of this species are known from United States and Canada. The Alberta locality is: CANADA. Alberta: Edmonton (Spencer, 1969). Liriomyza millefolii Hering Liriomyza millefolii Hering, 1927:185; Spencer, 1969: 1 78. Comparison and diagnostic characters. — The members of this species can be easily recog- nised by the presence of conspicuously long whitish pubescence on the third antennal article and the presence of vertical bristles on yellow areas. The adults resemble closely those of L. sinuata new species in external morphology, but the male genitalia are distinct, Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — Larvae mine the leaves of Achillea millefolium L. and A. sibirica Ledeb., family Compositae. Larvae also mine the leaves of Tanacetum vulgare L. in the laboratory. Geographical distribution. — The members of this species are known from Germany (Hen- del, 1931) in Europe, and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 2 66, 3 99 Edmonton, river bed near University of Alberta campus, from leaf mines on Achillea sibirica Ledeb., coll. 26.vii.1966, emerged 1 0-1 3.viii. 1966; 3 66, 1 9 Edmonton, White Mud Creek park, same host, 6-20.X.1968; 6 66, 3 99 Elk Island park, same host, coll. 31.vii. 1966, emerged 1 5-20. viii. 1966; 3 66 same locality, 31.vii. 1966; 1 6 336 Sehgal same locality, viii. 1 967 Liriomyza montana Sehgal Liriomyza montana Sehgal, 1968:67. Comparison and diagnostic characters. — The members of this species resemble closely those of L. eupatorii (Kaltenbach) in external morphology. The position of vertical bristles on yellow areas used to differentiate the adults of this species from those of L. eupatorii (Kaltenbach) is variable as the area of vertical bristles in some specimens is darkened. The male genitalia are, however, quite distinct. Sehgal (1968) illustrated the head, wing and male genitalia characteristic of this species. Spencer ( 1 969) also figured the distinctive aedeagus. Biology. — Larvae probably mine the leaves of grasses (Gramineae). Geographical distribution. — The members of this species are known from various locali- ties in the Rockies in Alberta, Canada (Sehgal, 1968). Liriomyza nordica Spencer Liriomyza nordica Spencer, 1969: 179. Comparisons and diagnostic characters. — The members of this species are very similar to those of L. senecionivora new species and differ only in having femora blackish. The male genitalia are, however, very distinct. The adults differ from those of other similar species, L. bifurcata new species and L. kenti Spencer, in having darker mesopleura and distinct male genitalia. Spencer (1969) illustrated the aedeagus characteristic of this species. Geographical distribution. — The members of this species are known only from the locality of its type series from Canada. I examined the following material from Alberta: CANADA. Alberta: 1 6 Edmonton, Rainbow Valley, 31.V.1969; 2 66 Edmonton, White Mud Creek park, 25.V.1969. Liriomyia pilosa Spencer Liriomyza pilosa Spencer, 1969: 182. Comparison and diagnostic characters. — The members of this species resemble closely those of L. millefolii Hering in having long pubescence on third antennal article and can be reliably separated only by examination of male genitalia. Surstyli in this species are shorter and broader than in millefolii Hering. Spencer (1969) illustrated the aedeagus characteristic of this species. Geographical distribution. — The members of this species are known only from Alberta from the locality of the type specimen as follows: CANADA. Alberta: Edmonton, University of Alberta campus (Spencer, 1969). Liriomyza senecionivora new species Comparisons. - The adults of this species resemble closely those of L. nordica Spencer in external characteristics and can be reliably separated only by the examination of male genitalia. L. senecionivora and L. bifurcata new species described earlier are included in Spencer’s (1969) key to Canadian species of the genus Liriomyza Mik by amending and extending the couplet 38 as below: 38. acr in four rows 38a acr in two rows 38b Agromyzidae of Alberta 337 38a. Aedeagus as illustrated (Spencer, 1969) nordica Spencer Aedeagus as in Fig. 60, 61 senecionivora n. sp. 38b. Aedeagus as illustrated (Spencer, 1969) kenti Spencer Aedeagus as in Fig. 52, 53 bifurcata n. sp. Description. — Head. Frons wide, approximately 1.8 times width of eye at level of front ocellus, slightly projected in front of eye margin in profile; eyes oval, 1.3 times higher than long; gena approximately one-fifth of eye height midway between vibrissal and posterior margins; ocellar triangle small; lunule high, almost flat above; two strong Ors directed up- wards, three Ori directed inwards and upwards; orbital setulae few, approximately seven, reclinate; antennal bases approximate; third antennal article rounded at tip, with normal pubescence; arista long and pubescent. Mesonotum. Dorsocentrals 3+1; acr in four irregular rows. Wing. Length in male 2.0 mm; costa extended to vein M1+2; costal segments 2-4 in the ratio of 1 : 0.23 : 0.26; wing tip at vein Mj +2 ; crossvein m-m present; last segment of M3 +4 approximately three times penultimate. Male genitalia (Fig. 60-63). Hypandrium U-shaped with slender side arms; pregonites broad and membranous; postgonites long and narrow; surstyli (Fig. 63) small, with two spines placed anteriorly, small spine on epandrium anteriorly also present; phallophore small and darkly sclerotized; basiphallus and distiphallus lightly sclerotized; ejaculatory duct swollen between basiphallus; distiphallus small; ejaculatory apodeme (Fig. 62) narrow and darkly sclerotized at base, bulb membranous. Colour. Frons, orbits, lunule, gena and antennae all yellow; ocellar triangle black; vte on black and vti on margin of black and yellow areas; mesonotum mat black; humeral area yellow, with a dark spot anteriorly; notopleural area yellow; scutellum yellow with dark areas at its basal corners; mesopleuron and sternopleuron black with narrow yellow band along upper margins; femora mainly yellow, with slight brownish area towards their base; tibiae and tarsi dark brown; squamal fringe brown, squamae slightly dark; halteres yellow. Derivation of the specific name. — This species is named senecionivora after the name of its food plant. Biology. - The larvae make linear mines on the leaves of Senecio pauciflorus Pursh. Pupa- tion occurs outside the leaf mine. Geographical distribution. - The members of this species are known only from the type localities: CANADA. Alberta: Holotype <5 Jasper National park, near Medicine Lake, from leaf mines on Senecio pauciflorus Pursh, coll. 16.vii.1969, emerged 30.vii.1969, coll. G.C. D. Griffiths. Paratype 1 <3 Blairmore, 26.vi. 1966. Liriomyza septentrionalis Sehgal Liriomyza septentrionalis Sehgal, 1968:70. Comparison and diagnostic characters. — The members of this species resemble closely those of L. cordillerana Sehgal in external morphology, and can be reliably separated only by the examination of the characters of male genitalia. The third antennal article is variable in colour from complete yellow to slightly darkened at the base of arista; orbits are usually yellow. Sehgal (1968) figured the head, wing and the characteristic aedeagus. Biology. - Larvae mine the leaves of grasses (Gramineae). Geographical distribution. — The members of this species are known from various locali- ties in the Rocky Mountains and Cypress Hills in Alberta and from British Columbia (Sehgal, 338 Sehgal 1968). Liriomyza singula Spencer Liriomyza singula Spencer, 1969: 184. Diagnostic characters. — The members of this species are distinct in the absence of cross- vein m-m; the third antennal article is only lightly darkened at the base of arista. Spencer (1969) figured the distinctive aedeagus. Geographical distribution. - The members of this species are known only from its type species in Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: Paratype 1 6 Blairmore, 20.vi. 1966. Liriomyza sinuata new species Comparisons. — The members of this species resemble those of L. millefolii Hering in having long pubescence on third antennal article but differ in having both vertical bristles on dark areas and distinct male genitalia. This species is included in Spencer’s (1969) key to Canadian species of the genus Liriomyza Mik by amending and extending the couplet 26 as below: 26. Orbits shining black; mesopleura black in lower three-quarters; femora distinctly darkened sp. (Constance Bay) Orbits yellow; femora yellow 26a 26a. vte on black and vti on margin of black and yellow ground sinuata n. sp. Both vt on yellow ground 27 Description. — Head. Frons wide, approximately twice width of eye at level of front ocellus, projected in front of eye margin in profile; eyes oval, slightly slanted, 1 .25 times higher than their length; gena deep, approximately one-third of eye height midway between vibrissal and posterior margins; ocellar triangle small; lunule high, narrow at top; two strong Ors directed upwards; two Ori, lower one "directed inwards and upper one directed upwards; orbital setulae one to two, reclinate; antennal bases approximate; third antennal article rounded at tip, with conspicuous pubescence; arista pubescent. Mesonotum. Dorsocentrals 3+1; acr in two rows. Wing. Length in male approximately 1.5 mm; costa extended to vein M1+2; costal seg- ments 2-4 in the ratio of 1 : 0.35 : 0.25; wing tip at M1+2 ; crossvein m-m present; last seg- ment of M3+4 approximately 2.5 times penultimate. Male genitalia (Fig. 64-67). Hypandrium U-shaped with slender side arms; pregonites broad; postgonites elongated; surstylus (Fig. 67) small with short spine anteriorly and small spine dorsally on epandrium; aedeagal apodeme darkly sclerotized; phallophore small; ejacu- latory duct swollen between basiphallus; distiphallus two long tubular S-shaped processes; ejaculatory apodeme (Fig. 66) broad, bulb small and sclerotized along lower margin. Colour. Frons, orbits, lunule, gena and antennae all yellow; vte on black and vti on the margin of dark and yellow areas; mesonotum mat black; humeral area yellow, with a dark spot anteriorly; notopleural area yellow; scutellum yellow, with dark area along its basal corners; mesopleuron yellow with slight dark area anteroventrally; sternopleuron black, with a narrow yellow band dorsally; femora mainly yellow; tibiae and tarsi brown, squamal fringe dark brown; halteres yellow. Derivation of the specific name. — The name sinuata is given in view of the sinuate or wavy distiphallus. Geographical distribution. — The members of this species are known only from the locali- Agromyzidae of Alberta 339 ties of its type specimens: CANADA. Alberta: Holotype 6 Banff, 28.vi.1966; Paratype 1 6 Cypress Hills, Elkwater, 24.vi.1966. Liriomyza smilacinae Spencer Liriomyza smilacinae Spencer, 1969:186. Comparison and diagnostic characters. - The members of this species are close to those of L. undulata Spencer in external morphology and are separated only by examination of the characters of male genitalia. Spencer (1969) illustrated the characteristic aedeagus. Biology. — Larvae form linear leaf mines on the leaves of Smilacina stellata (L.) Desf., family Liliaceae. Spencer (1969) illustrated the leaf mine characteristic of this species. Geographical distribution. — The members of this species are known only from the local- ities of its type series in Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: Paratype 1 6 Edmonton, White Mud Creek park, 1 3 .vi. 1 966; 1 6 same locality, 18.vi.1968, leg. G. C. D. Griffiths; 4 66, 5 99 same locality, from leaf mines on Smilacina stellata (L.) Desf., coll. 10.vi.1968, leg. G. C. D. Griffiths; 1 6 same locality and host, 1 0.vi.l 968-2. ii. 1969; 1 6, 1 9 same locality and host, 28.vi.-14.vii. 1968. Liriomyza socialis Spencer Liriomyza socialis Spencer, 1969: 186. Diagnostic characters. — The main distinguishing characters of the members of this species are mat grey mesonotum and two rows of acrostichals. The colour of third antennal article varies from pale to dark brown (Spencer, 1969). Spencer (1969) illustrated the distinctive aedeagus. Geographical distribution. — The members of this species are known only from Alberta, Canada. The Alberta localities are as follows: CANADA. Alberta: Blairmore, Elk Island park, Jasper (Spencer, 1969). Liriomyza sylvatica new species Comparisons. — A male of this species is very similar to that of L. edmontonensis Spencer in external characteristics and is reliably separated only by examination of male genitalia, which, however, are very distinct. This species is included in Spencer’s (1969) key to Cana- dian Liriomyza species by amending and extending the couplet 37 as below: 37. Femora partially darkened 37a Femora almost entirely bright yellow arcticola Spencer 37a. Aedeagus as illustrated (Spencer, 1969) edmontonensis Spencer Aedeagus as in Fig. 68, 69. sylvatica n. sp. Description. - Head. Frons approximately 1.3 times wider than the width of eye at level of front ocellus; slightly projected in front of eye margin in profile; eyes oval, 1.4 times higher than their length; gena little less than one-fourth of eye height midway between vibrissal and posterior margins; ocellar triangle small; lunule high; two Ors directed upwards; two Ori directed inwards and upwards; orbital setulae three to four, reclinate; antennal bases approximate; third antennal article with a slight angle anterodorsally, with normal pubescence; arista pubescent. Mesonotum. Dorsocentrals 3+1; acr in four irregular rows. 340 Sehgal Wing. Length in male 1.7 mm; costa extended to vein M1+2; costal segments 2-4 in the ratio of 1 : 0.26 : 0.18; wing tip at vein M! +2 ; crossvein m-m present; last segment of M3+4 approximately three and a half times the penultimate. Male genitalia. (Fig. 68, 69). Hypandrium U-shaped with slender side arms; pregonites broad; postgonites elongated; surstylus small and lightly sclerotized; aedeagal apodeme dark- ly sclerotized; phallophore elongate; ejaculatory duct swollen between basiphallus; distiphal- lus as illustrated; ejaculatory apodeme broad, bulb small and sclerotized along lower margin. Colour. Frons, gena, lunule and antennae yellow; upper orbits partially darkened up to lower Ors; vte on black and vti on the margin of dark and yellow areas; mesonotum mat greyish black; humeral area yellow, with a dark spot anteriorly; notopleural area yellow; scutellum yellow, with slight dark at its basal corners; mesopleuron black with a narrow yellow band dorsally; sternopleuron black; femora yellow, darkened towards base, tibiae and tarsi dark brown; squamal fringe brownish; halteres yellow. Derivation of the specific name. — The name sylvatica indicates that the species is woodland-inhabiting. Geographical distribution. — This species is known only from the type locality: CANADA. Alberta: Holotype 6 St. Albert, near Edmonton, 18.vi. 1967. Liriomyza taraxaci Hering Liriomyza taraxaci Hering, 1927: 184; Spencer, 1969: 188. Comparisons and diagnostic characters. — The members of this species resemble closely those of L. veluta Spencer, L. trifolii (Burgess), L. lathyri new species, and differ only in having mesonotum black and not grey. The male genitalia are, however, entirely different. Spencer (1969) illustrated the aedeagus distinctive of this species. Biology. — Larvae form elongate blotch mines on the leaves of Taraxacum officinale Weber, family Compositae. Hering (1927) illustrated the characteristic leaf mine. Geographical distribution. — The members of this species are known from various locali- ties in Europe (Hendel, 1931) and from Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 6 Banff, 28.vi. 1966; 1 6 Blairmore, 26.vi.1966; 1 9 Edmonton, 110 Street, 84 Avenue, from leaf mines on Taraxacum officinale Weber, 1 5-29. vi. 1968; 2 66 Edmonton, University of Alberta campus, same host, 27.vii.-l 1 .viii. 1 966; 1 6 Edmon- ton, White Mud Creek park, 28.V.1967. Liriomyza undulata Spencer Liriomyza undulata Spencer, 1969: 190 Comparison and diagnostic characters. - The members of this species resemble closely those of L. smilacinae Spencer in external morphology, but have distinct male genitalia. Spencer (1969) illustrated the characteristic aedeagus. The distiphallus is distinctive in having a long undulating process distally. Geographical distribution. - The members of this species are known only from the local- ities of its type series from Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 2 66 Blairmore, 26.vi.1966; Paratype 1 6 Edmonton, White Mud Creek park, 23.vi.1966, coll. V. K. Sehgal, 11 66 same locality, 23-29.vi. 1966. Agromyzidae of Alberta 341 Liriomyza veluta Spencer Liriomyza veluta Spencer, 1969: 190. Comparisons and diagnostic characters. — The members of this species resemble closely those of L. lathyri new species and L. trifolii (Burgess) in external morphology and can be separated only by examination of the characters of male genitalia. The adults differ from those of another similar species, L. taraxaci Hering, in having mesonotum grey and not black. Spencer (1969) illustrated the distinctive aedeagus. Geographical distribution. — The members of this species are known from various locali- ties of its type series from Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 6 Blairmore, 26.vi.19 66; 1 6 Edmonton, White Mud Creek park, viii.l968;2 66, 1 9 George Lake, near Busby, 21.vi.1966. Liriomyza viciae Spencer Liriomyza viciae Spencer, 1 969: 191. Comparison and diagnostic characters. — The members of this species resemble closely those of L. melampyga (Loew) in external morphology and differ only in having acrostichals in two rows and distinctive male genitalia. Spencer (1969) illustrated the aedeagus charac- teristic of this species as well as that of L. melampyga (Loew). The adults are small flies, wing length approximately 2.0 mm, with characteristic prescu- tellar yellow and yellow antennae. Biology. — Larvae form blotch mines on the leaflets of Vicia americana Muhl., family Leguminosae. Geographical distribution. — The members of this species are known only from the type series from Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 9 Banff, 28.vi.1966; Paratype 1 6 Blairmore, 27.vi.1966; 1 9 Edmonton, University of Alberta campus, from leaf mines on Vicia americana Muhl., 21.ix. 1 968-13. ii. 1969; 1 6 Elk Island park, 3 l.vii. 1966; 3 99 Jasper, 1 7-23 .vi. 1 966. Genus Lemurimyza Spencer Lemurimyza Spencer, 1965b:26. The main distinguishing characters of the genus Lemurimyza Spencer are: subcosta weak- ly developed distally, joined to costa independent of Rx ; costa extended to vein M1+2; orbital setulae normally erect or slightly proclinate; scutellum yellow; mesonotum with yellow central area adjoining scutellum, epandrium normally with comb-like arrangement of dark spines; aedeagus typical of the genus, with paired sclerotized tubules. The genus Lemurimyza Spencer is represented in Alberta by only one species, L. pallida Sehgal. The members of this genus are extremely similar to those of the genus Liriomyza Mik in external characteristics, but possess very distinct male genitalia. Lemurimyza pallida Sehgal Lemurimyza pallida Sehgal, 1968:72. Comparisons and diagnostic characters. — The members of this species differ from those of two other species, L. dorsata (Siebke) and L. pacifica (Melander), known from Canada (Spencer 1969), in having third antennal article yellow and distinctive male genitalia. Sehgal 342 Sehgal (1968) illustrated the head, wing and male genitalia characteristic of this species. Spencer (1969) also illustrated the aedeagus. Geographical distribution. — The members of this species are known only from the type locality as follows: CANADA. Alberta: Banff (Sehgal, 1968). Genus Metopomyza Enderlein Metopomyza Enderlein, 1936: 180. The main distinguishing characters of this genus are subcosta fold-like distally and joined to costa independent of ; orbital setulae reclinate; costa extended to apex of vein Mj +2 ; scutellum yellow; orbits broad and raised above plane of frons; aedeagus typical of genus. The members of this genus are very similar to those of the genus Liriomyza Mik in ex- ternal morphology, but the male genitalia are very distinct. This genus is represented in Alberta by two species, interfrontalis (Melander) and grif- fithsi new species. Key to Alberta species of the genus Metopomyza Enderlein 1 . Squamal fringe yellow; larger specimens, wing length 2. 0-2-3 mm interfrontalis (Melander), p. 343 Squamal fringe brown, smaller specimens, wing length 1.5 mm in male griffithsi n. sp., p. 342 Metopomyza griffithsi new species Comparisons and diagnostic characters. — A member of this species differs from that of interfrontalis (Melander) in having smaller size and brown squamal fringe. It resembles that of a Palaearctic species, flavonotata (Haliday), but possesses distinct male genitalia. This species is distinguished in Spencer’s (1969) key to Canadian species of the genus Metopo- myza Enderlein by amending and extending the couplet 1 as below: 1. Notopleural area black la Notopleural area yellow 2 la. Larger specimens, wing length 2. 0-2. 3 mm; squamal fringe yellow interfrontalis (Melander) Smaller specimens, wing length 1 .5 mm in male; squamal fringe brown griffithsi n. sp. Description. — Head. Frons approximately 1 .8 times width of eye at level of front ocellus, slightly projected in front of eye margin in profile; orbits broad, slightly raised above the plane of frons; eyes oval, strongly slanted along posteroventral margin, vertical height is almost equal to their length; ocellar triangle small; gena deep, approximately 0.3 times ver- tical height of eye. Two strong Ors directed upwards; two strong Ori directed inwards ;orbital setulae numerous, reclinate; third antennal article slightly angulate antero-dorsally and rounded below, pubescent. Mesonotum. Dorsocentrals 3+1 decreased in length anteriorly, acr in three to four irregu- lar rows. Wing. Length in male 1.5 mm; costa extended to vein M! +2 ; costal segments 2-4 in ratio of 1 : 0.33 : 0.26; vein M1+2 at the wing tip; crossvein m-m present; last segment of M3+4 approximately 0.4 times penultimate. Male genitalia (Fig. 70-72). Hypandrium with narrow side arms; surstyli (Fig. 70) with Agromyzidae Of Alberta 343 two rows of conspicuous spines as illustrated; phallophore long; aedeagus (Fig. 71) typical of genus; basiphallus broad and sclerotized; mesophallus long and slender; distiphallus as small divergent tubules distally; ejaculatory apodeme (Fig. 72) small and narrow, bulb small, membranous. Colour. Frons darker below and yellowish above, orbits black; lunule dark; gena greyish black; antennae black; mesonotum mat black, slightly brownish; scutellum almost entirely yellow; mesopleura, sternopleura and pteropleura all black; femora black, with distal tips yellow; tibiae and tarsi brownish black; squamae pale, fringe brownish; halteres yellow. Derivation of the specific name. — This species is named in honour of G. C. D. Griffiths of the Department of Entomology, University of Alberta, Canada. Geographical distribution. — A member of this species is known only from the type locality: CANADA. Alberta: Holotype 6 Edmonton, White Mud Creek park, 18.vi.1968, coll. G. C. D. Griffiths. Metopomyza interfrontalis (Melander) Agromyza interfrontalis Melander, 1913:263. Liriomyza interfrontalis (Melander), Frick, 1952:403. Metopomyza interfrontalis (Melander), Frick, 1957:204, 1959:41 2; Spencer, 1969:198. Comparison and diagnostic characters. — The members of this species differ from those of griffithsi new species in having larger size and pale squamal fringe. Spencer (1969) illustrated the distinctive aedeagus. Geographical distribution. — The members of this species are known from Canada and United States. The Alberta locality is: CANADA. Alberta: Elkwater (Spencer, 1969). Genus Praspedomyza Hendel Dizygomyza (Praspedomyza) Hendel, 1 93 1 : 77. Phytobia (Praspedomyza) Frick, 1952:395, 1959:394. Praspedomyza Hendel, Spencer, 1966b: 146. Nowakowski (1962) on the basis of his studies on male genitalia proposed that this genus should be merged with the genus Liriomyza Mik. Later Spencer (1966b, 1969) in view of the dark colouration, raised orbits and distinct male genitalia justified the retention of Praspedomyza Hendel as a distinct genus. This genus is represented in Canada by only one species, galiivora Spencer, the common leaf miner on Galium. Praspedomyza galiivora Spencer Praspedomyza galiivora Spencer, 1969:199. Diagnostic characters. — The members of this species are quite distinctive in having yellow third antennal article and distinct male genitalia. Spencer (1969) illustrated the aedeagus characteristic of this species. The colour of third antennal article varies slightly from bright yellow to reddish. Biology. - Larvae mine the leaves of Galium boreale L., family Rubiaceae. Geographical distribution. — The members of this species are known from Europe and Canada (Spencer, 1969). I examined the following material from Alberta: 344 Sehgal CANADA. Alberta: 1 6 Banff, 28.vi. 1966; 1 <3, 3 99 Edmonton, White Mud Creek park, 1 3-23 .vi. 1 966; 3 99 same locality, from leaf mines on Galium boreale L., coll. 28.vi. 1968, emerged 1 l-12.vii.1968; 1 6 same locality and host, coll. 4.ix.l968, emerged 16. ix. 1968, coll. G. C. D. Griffiths; 1 6 Edmonton river bank near University of Alberta campus, 14.vi. 1 969; 4 66, 1 9 Elk Island park, 3 1 .vii.-2.viii. 1 966. Genus Haplomyza Hendel Antineura Melander, 1912:219. Haplomyza Hendel, 1914:73, new name for Antineura Melander, not Osten Sacken 1881. The members of this genus resemble externally those of the large genus Liriomyza, but possess distinct male genitalia. They are represented in Alberta by only one species, togata (Melander). Haplomyza togata (Melander) Antineura togata Melander, 1913:250. Haplomyza togata (Melander); Frick, 1953:73, 1959:413; Spencer, 1969:201. Diagnostic characters. — The main distinguishing characters of the members of this species are wing length 1.75-2.2 mm, costa extended to vein M1+2; crossvein m-m absent; one Ors and three Ori; eyes slanted; frons, gena, face and antennae yellow; mesonotum mat grey, few acrostichals and distinct male genitalia. The ninth sternite is greatly elongate. Biology. — Larvae are known to make irregular blotch mines on the leaves of Amaranthus spp., family Amaranthaceae, in United States (Frick, 1959). Geographical distribution. — The members of this species are known from United States, and Alberta and Saskatchewan in Canada (Spencer, 1969). The Alberta locality is: CANADA. Alberta: Drumheller. Genus Phytoliriomyza Hendel Liriomyza (Phytoliriomyza) Hendel, 1921:203. Phytoliriomyza Hendel; Frey, 1941 : 19: Frick, 1952:410; Spencer. 1964b:662. Xyraeomyia Frick, Spencer, 1964b: 662. The members of this genus differ from those of the genus Liriomyza Mik in having dark scutellum and proclinate orbital setulae. They are represented in Alberta by only one species, arctica (Lundbeck). Phytoliriomyza arctica (Lundbeck) Agromyza arctica Lundbeck, 1900:304. Odinia immaculata Coquillett, 1902:185. Agromyza formosensis Malloch, 191 4b: 3 1 5 . Dizygomyza (Icteromyza) arctica (Lundbeck), Hendel, 1931:57. Phytoliriomyza arctica (Lundbeck); Shewell, 1953:469; Frick, 1959:414. Diagnostic characters. - The main distinguishing characters of the members of this species are: eyes oval, slanted, slightly pilose; acrostichals present; wing length approximately 2.0 mm; costa strongly extended to vein M1+2; crossvein m-m present; and aedeagus with characteristic two long, membranous coiled tubules. Spencer (1963, 1964b, 1969) discussed in detail and illustrated the male genitalia of members of this species. Agromyzidae of Alberta 345 Biology. - Larvae feed as stem miners on Sonchus asper L., family Compositae in Germany (Spencer, 1963). No host plant is yet known in North America. Geographical distribution. — The members of this species are most widely distributed being known from Europe, Formosa, Canada, United States and South America (Spencer, 1963). I examined the following material from Alberta: CANADA. Alberta: 4 66 Cypress Hills, near Elkwater Lake, 24.vi.1966. Genus Pseudonapomyza Hendel Pseudonap omyza Hendel, 1920: 1 15. The members of this genus differ from those in the genus Phytomyza Fallen in having crossvein m-m basal to r-m and reclinate orbital setulae. They are represented in Alberta by two species, atra (Meigen) and lacteipennis (Malloch). Key to Alberta species of the genus Pseudonapomyza Hendel 1 . Mesonotum weakly shining black; tarsi dark brown; wings normal atra (Meigen), p. 345 Mesonotum mat grey; tarsi yellow; wings whitish lacteipennis (Malloch), p. 345 Pseudonapomyza atra (Meigen) Phytomyza atra Meigen, 1830:191. Pseudonapomyza atra (Meigen); Hendel, 1 932:302; Spencer, 1969:209. Comparison and diagnostic characters. — The members of this species are quite distinctive in having angulate third antennal articles. The adults differ from those of similar species, P. lacteipennis (Malloch), in having dark tarsi and weakly shining black mesonotum. Biology. - Larvae mine the leaves of grasses (Gramineae). Geographical distribution. — The members of this species are Holarctic in distribution, known from Europe (Hendel, 1932), United States (Frick, 1959) and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 9 Edmonton, 26.v. 1946, coll. W. R. M. Mason; 1 9 Edmonton, University of Alberta campus from leaf mine on grass; coll. 22.vi.1968, emerged 1 2 .vii. 1 968. Pseudonapomyza lacteipennis (Malloch) Phytomyza lacteipennis Malloch, 1913b: 152. Pseudonapomyza lacteipennis (Malloch); Frick, 1952:419, 1959:419; Spencer, 1969:210. Diagnostic characters. — The main distinguishing characters are mat greyish mesonotum, yellow tarsi and whitish wings. Biology. - Larvae probably mine the leaves of grasses (Gramineae). Geographical distribution. — The members of this species are known from United States and Canada. The Alberta localities are as follows: CANADA. Alberta: Elkwater; Medicine Hat; Orion (Spencer, 1969). Genus Paraphytomyza Enderlein Paraphytomyza Enderlein, 1936: 180; Nowakowski, 1 962: 1 02; Spencer, 1969:203. Rubiomyza Nowakowski, 1962:102. The name Phytagromyza Hendel which has long been used (Hendel, 1920, 1932; Frick, 346 Sehgal 1952, 1959) for members of this genus, cannot be used now as its type, P. flavocingulata (Strobl), is now referred to the genus Cerodontha Rondani (Nowakowski, 1962, 1967). The main distinguishing characters of the genus Paraphytomyza Enderlein are: subcosta weakly developed distally, joined to costa independent of Rj ; orbital setulae erect or re- clinate or absent; costa extended to vein R4+5; crossvein m-m usually absent, if present, always beyond crossvein r-m. This genus is represented in Alberta by five species. All Alberta species discussed here probably form a single group within the genus Paraphytomyza Enderlein, whose members feed on the representatives of the family Caprifoliaceae and other related families of the order Rubiales. Nowakowski (1962) proposed a new genus, Rubiomyza , for this group of flies. The name proved to be synonymous with Paraphytomyza Enderlein. Another group of leaf miners on Salicaceae is probably also represented in Alberta. Linear leaf mines on the under surface of the leaves of Populus tremuloides Michx., quite common around Edmonton, are very similar to those of Paraphytomyza tremulae (Hering) in Europe on Populus tremula L. Since no flies have yet been bred, their identity cannot be confirmed. Key to Alberta species of the genus Paraphytomyza Enderlein 1(0). Crossvein m-m present 2 Crossvein m-m absent 4 2(1). Dorsocentrals two; mouthparts elongate nitida (Malloch), p. 347 Dorsocentrals three or more; mouthparts normal 3 3(2). Notopleural area yellow plagiata (Melander), p. 347 Notopleural area brownish black lonicerae (Robineau-Desvoidy), p. 346 4(1). Small specimens, wing length 1.6-1. 8 mm in males; aedeagus as illustrated (Fig. 76) spenceri n. sp., p. 348 Larger specimens, wing length 2.0 to 2.4 mm orbitalis (Melander), p. 347 Paraphytomyza lonicerae (Robineau-Desvoidy) Phytomyza lonicerae Robineau-Desvoidy, 1851:396. Phytagromyza lonicerae (Robineau-Desvoidy); Hering, 1951:36; Frick, 1953:74. Paraphytomyza lonicerae (Robineau-Desvoidy); Spencer, 1969:205. Comparison and diagnostic characters. — The members of this species are very close to those of P. orbitalis (Melander) in the general shape of aedeagus, but differ in lacking cross- vein m-m. Spencer (1969) illustrated the aedeagus characteristic of this species. The poste- rior spiracles of the puparium are distinctive in having a dark spine in centre. Biology. — Larvae mine the leaves of various members of the genera Lonicera and Sym- phoricarpos, family Caprifoliaceae. Frick (1953) reared this species from Lonicera involu- crata (Richards) Banks and Symphoricarpos albus (L.). I observed the leaf mines of this species in Alberta on Lonicera dioica L., L. tartarica L. and Symphoricarpos albus (L.). The leaf mine is whitish, linear with distinct frass granules disposed alternately along the mine. Hering (1951) illustrated the characteristic leaf mine. This species is the first to appear in early spring and there is only one generation a year. Geographical distribution. — The members of this species are known from Europe, United States and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 2 66 Edmonton, White Mud Creek park, 19.V.1968; 2 66 same locality, 7.V.1969. Agromyzidae of Alberta 347 Paraphytomyza nitida (Malloch) Agromyza nitida Malloch, 191 3a: 288 ; Frick, 1952:373. Phytagromyza nitida (Malloch); Frick, 1953:74, 1959:417. Paraphytomyza nitida (Malloch); Spencer, 1969:207. Diagnostic characters. — The members of this species are distinctive in having elongate mouthparts and absence of crossvein m-m. Spencer (1969) illustrated the distinctive aedea- gus. Biology. — Not confirmed. Spencer (1969) noted the similarity between this species and P. orphana (Hendel), a stem miner on Galium in Europe, and has suggested as host one of the Galium species occurring in Alberta. Geographical distribution. — The members of this species are known from United States (Frick, 1953, 1959) and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 9 Elk Island park, 7.vi.l966. Paraphytomyza orbitalis (Melander) Phytomyza orbitalis Melander, 1913:271. Phytagromyza orbitalis (Melander); Frick, 1952:416, 1959:417. Paraphytomyza orbitalis (Melander); Spencer, 1969:207. Comparison and diagnostic characters. — The members of this species resemble externally those of a sympatric species, P. spenceri new species, but differ in having a distinct aedeagus and larval leaf mine (Fig. 73). The aedeagus has been illustrated by Spencer (1969). The females unless represented by bred series cannot be determined definitely. Biology. — Larvae mine the leaves of Lonicera dioica L. and Symphoricarpos albus (L.), family Caprifoliaceae. The leaf mine (Fig. 73) is broad, linear in shape. Pupation occurs outside the leaf mine. Geographical distribution. — P. orbitalis (Melander) is a Nearctic species, whose members are known from United States (Frick, 1952, 1959) and Canada (Spencer, 1969). I examined 10 males and one female from Alberta: CANADA. Alberta: 1 d Blairmore, 27.vi.1966; 1 d Edmonton, 24.V.1946, coll. E. H. Strickland; 1 d Edmonton, University of Alberta campus, from leaf mines on Lonicera dioica L., 29.v.-22.vi.l966; 1 9 Edmonton, White Mud Creek park from leaf mines on Symphoricarpos albus (L.), coll. 10.vi.1966; 3 dd same locality, 1 2-1 9.vi. 1966; 2 66 same locality, 15.vii. 1966; 1 d Edmonton, Mayfair park, 4.v. 1969; 1 o Elk Island park, 4.vi. 1967; 1 6 St. Albert, near Edmonton, 14.vi. 1966. Paraphytomyza plagiata (Melander) Napomyza plagiata Melander, 1913:273. Agromyza plagiata (Melander); Malloch, 1918: 130. Phytagromyza plagiata (Melander); Frick, 1952:416, 1959:417. Paraphytomyza plagiata (Melander); Spencer, 1969:208. Diagnostic characters. — The members of this species can be easily recognised by the characters given in the key. Biology. — Larvae mine the leaves of Lonicera involucrata (Richards) Banks, family Caprifoliaceae. The leaf mine (Fig. 74) is linear and light greenish in colour. Geographical distribution. — The members of this species are known from United States 348 Seghal (Frick, 1952) and Car.ada (Spencer. 1969). I examined the following material from Alberta: CANADA. Alberta: 1 6 George Lake, near Busby, from leaf mines on Lonicera involu- crata (Richards) Banks, 7.vi. 1968 emerged 30.iv.1969, coll. G. C. D. Griffiths; 1 9 St. Albert, near Edmonton, same host, 14.vi. 1966-5. iii. 1967; 1 9 same locality, 14.vi. 1966. Parapliytomyza spenceri new species Comparisons and diagnostic characters. — The members of this species resemble exter- nally those of a sympatric species, P. orbitalis (Melander), and can only be reliably differ- entiated by the examination of the male genitalia. The females unless from bred series are very difficult to determine definitely. The linear leaf mine of this species (Fig. 79) is similar to that of P. luteoscutellata (de Meijere) illustrated by Spencer (1969), but the adults are distinct in having completely black scutellum. This species is distinguished in Spencer’s (1969) key to Canadian species of the genus Parapliytomyza Enderlein by extending the couplet 5 as below: 5. Scuteilum yellow, at least between basal scutellar bristles luteoscutellata (de Meijere) Scutellum entirely dark 5a 5a. Wing length up to 2.0 mm; aedeagus as in Fig. 76 spenceri n. sp. Larger specimens 6 Description. - Head (Fig. 75). Frons almost equal to width of eye at level of front ocel- lus; upper orbits slightly projected in front of eye margin in profile; eyes oval, 1.3 times higher than their width, bare; ocellar triangle small; gena deepest posteriorly, approximately one-sixth of eye height mid-way between vibrissal and posterior margins; two strong Ors directed upwards; two Ori directed inwards; orbital setulae 4-6, reclinate; third antennal article rounded at tip. arista long and pubescent. Mesonotum. Dorsocentrals 3+1; acr numerous in approximately four rows. Leg. Mid-tibia without a differentiated bristle medially. Wing. Length in 66 1.6- 1.8 mm. in 99 approximately 2.0 mm; costa extended to vein R4+s: wing tip between R4+5 and M1+2; crossvein m-m absent; costal segments 2-4 in the ratio of 1.0 : 0.23 : 0.28. Male genitalia (Fig. 76). Hypandrium U-shaped with slender side arms and no apodeme; surstyli broad and rounded, without spines; pregonites broad; postgonites elongate; phallo- phore short and darkly sclerotized; basiphallus a pair of broad arms; distiphallus removed from basiphallus by a short membranous gap, of two distinctive curved tubes; aedeagal apodeme weakly sclerotized; ejaculatory apodeme small and fan-shaped, bulb small and membranous. Colour. Frons darker above the lunule; orbits yellow; ocellar triangle black; antennae, gena and lunule yellowish brown; mesonotum and scutellum mat grayish black; humeral and notopleural areas yellow: femora dark brown; tibiae and tarsi mostly yellowish or slightly brownish: squamae yellow, fringe slightly brownish; halteres yellow. Description of immature stages. — Puparium brownish yellow, oval and deeply segmented, measures approximately 1.5 mm x 0.8 mm. Larval mouth parts obtained from puparium are illustrated (Fig. 77). Right mandible larger than left, each with two distinct teeth alternate with one another; labial sclerite short and darkly sclerotized: paraclypeal phragmata with darkly sclerotized dorsal and weakly sclerotized ventral arms. Muscle scars on abdominal segments small and oval; tubercles numerous in approximately six to eight rows. Anterior spiracles small, each with about six to eight bulbs; posterior spiracles (Fig. 78) small and rounded, each with 11-12 bulbs. Agromyzidae of Alberta 349 Derivation of the specific name. — This species is named in honour of Dr. K. A. Spencer, who has contributed greatly to the knowledge of world Agromyzidae. Biology. — Larvae mine the leaves of Lonicera dioica L. and Symphoricarpos occidentalis Hook., family Caprifoliaceae. The leaf mine (Fig. 79) is linear, greenish black, without dis- crete frass granules. Pupation occurs outside the mine. Geographical distribution. — I examined the members of this species only from the province of Alberta. CANADA. Alberta: Holotype 6 Edmonton, White Mud Creek park, from leaf mine on Lonicera dioica L., 21.ix.-10.x.l968; allotype 9 same locality and host, 21 ,ix.-22.x. 1968; paratypes 2 99 same locality and host, 21.ix.-10.x.l968; 1 6 same locality and host, 6.ix. 1 968-7. ii. 1969; 1 9 same locality and host, 6.ix.-22.x.l968; 1 6, 1 9 same locality and host, coll. 6.ix.l968, emerged 2.vi.l969; 2 66 same locality, from leaf mines on Symphoricarpos occidentalis Hook., coll. 10. ix. 1966, emerged 25.iii.1967 and 2.vi.l967;4 66 same locality, 6-23.vi. 1966; 1 6 same locality, 8.vi.l967; 2 66 Edmonton, Mayfair park, 17.V.1969; 1 6 Drumheller, 14.vi.1946, coll. W. R. M. Mason. Genus Napomyza Westwood Napomyza Westwood, 1840:152 The members of this genus differ from those in the large genus Phytomyza Fallen in the presence of crossvein m-m. Male genitalia are, however, distinct. This genus is represented in Alberta by three species. Key to Alberta species of the genus Napomyza Westwood 1(0). Third antennal article with conspicuous pubescence plumea Spencer, p. 350 Third antennal article almost bare 2 2(1). Smaller specimens, wing length about 2.5-3. 1 mm; distiphallus paler nugax Spencer, p. 349 Larger specimens, wing length about 3. 5-4. 5 mm; distiphallus darkly sclerotized immanis Spencer, p. 349 Napomyza immanis Spencer Napomyza immanis Spencer, 1969:215. Comparison and diagnostic characters. — The members of this species differ from those of a similar species, nugax Spencer, in larger size, wing length 3. 5-4. 5 mm; third antennal article slightly less quadrate and aedeagus with darker distiphallus. Spencer (1969) illus- trated the distinctive aedeagus. Geographical distribution. — Known from Alaska, Alberta, Northwest Territories and Yukon Territory. The Alberta locality is as follows: CANADA. Alberta: Edmonton, White Mud Creek park (Spencer, 1969). Napomyza nugax Spencer Napomyza nugax Spencer, 1969:215. Comparison and diagnostic characters. — The members of this species differ from those of a similar species, immanis Spencer, in having smaller size, wing length 2.5-3. 1 mm; quadrate third antennal article and paler distal process on the distiphallus. Spencer (1969) illustrated the distinctive aedeagus. They also differ from lateralis (Fallen) in having distinct aedeagus. Geographical distribution. - Known from Alberta, British Columbia, Ontario and Quebec in Canada (Spencer, 1969). I examined the following material from Alberta: 350 Sehgal CANADA. Alberta: 1 d, 1 9 Blairmore, 26.vi. 1966. Napomyza plumea Spencer Napomyza plumea Spencer, 1969:217. Diagnostic characters. — The main distinguishing characters are the pubescent third antennal article and distinct male genitalia. The aedeagus has been illustrated by Spencer (1969). Geographical distribution. - Known from Alaska, Alberta, British Columbia, Manitoba and Quebec. The Alberta locality is as follows: CANADA. Alberta: Banff, Mt. Eisenhower (Spencer, 1969). Genus Phytomyza Fallen Phytomyza Fallen, 1810:21. The main distinguishing characters of this genus are subcosta weakly developed distally and joined to costa independent of Rj ; orbital setulae proclinate; costa extended to vein R4+s and crossvein m-m normally absent. The members of this genus as defined presently on the basis of the direction of orbital setulae and shortened costa, form a very diverse assemblage of many groups. The discovery and use of the characters of male genitalia in agromyzid taxonomy have proved beyond doubt that species extremely similar in external characteristics may have very conspicuous differences in genitalic structures. A close look at any of the recent keys shows that many species and even sometimes genera can be distinguished by examination of characters of male genitalia only. Attempts to divide this genus into various groups have not been success- ful as characters of male genitalia were not taken into consideration. It is not possible to undertake the full scale revision of this genus as at present the phallic structures of numer- ous species have not been illustrated. This is the largest genus of agromyzid flies with about 400 described species in the world. Spencer (1969) reported 83 species for Canada, of which he recorded 41 as occurring in Alberta. Fifteen new species are described in this genus here and four additional species are recorded as new to Alberta. P. flavicornis Fallen which has been reported as occurring in Alberta (Spencer, 1969) is not considered here as the Alberta specimens collected from the same locality as those of Spencer’s proved to be a new species luteiceps described here, distinguishable from flavicornis Fallen in the characters of the male genitalia. This genus is now represented in Alberta by 59 described species and in Canada by 98 species. Necessary amendments to include the further new species in Spencer’s (1969) key to Canadian species are given. Key to Alberta species of the genus Phytomyza Fallen 1(0). Frons basically pale, yellow, orange or reddish 2 Frons basically dark, brown or black 41 2( 1 ). Scutellum all or partially yellow 3 Scutellum dark, grey or black 5 3(2). Third antennal article black or dark brown 4 Third antennal article yellow major Malloch, p. 368 4(3). Upper Ors shorter than lower; aedeagus with up to eight coils ranunculi (Schrank), p. 375 clematiphaga Spencer, p. 361 Two Ors equal Agromyzidae of Alberta 351 5(2). 6(5). 7(5). 8(7). 9(8). 10(9). 11(10). 12(7). 13(12). 14(13). 15(14). 16(15). 17(14). 18(17). 19(12). 20(19). 21(20). 22(21). Femora mostly yellow 6 Femora mostly dark, at most with yellow distal tips 7 Third antennal article black miranda Spencer, p. 370 Third antennal article yellow; aedeagus as in Fig. 106 .... luteiceps n. sp., p. 368 Sides of thorax including humeral and notopleural areas yellow 8 Sides of thorax dark, at most upper margins of mesopleura with narrow yellow band 12 Third antennal article with normal pubescence 9 Third antennal artcle with conspicuously long pubescence riparia n. sp., p. 376 Upper Ors shorter than lower or lacking 10 Two Ors equal; hypopleuron and sternopleuron largely yellow petasiti Spencer, p. 373 Second costal segment two and a half to three times length of fourth 11 Second costal segment longer, approximately four times length of fourth spondylii R.-D., p. 379 Second antennal article black; hind margins of eyes black solidaginivora Spencer, p. 378 Second antennal article yellow; hind margins of eyes yellow matricariae Hendel, p. 369 Upper Ors shorter than lower or lacking 13 Two Ors equal 19 Frons partly darkened prava Spencer, p. 374 Frons almost entirely yellow. 14 Second costal segment more than three and a half times length of fourth .... 15 Second costal segment less than three and a half times length of fourth 17 Upper Ors present; aedeagus as in Fig. 81 , 82 aquilegioides n. sp., p. 355 Upper Ors invariably lacking; larvae leaf miner on Heracleum 16 Second costal segment three and a half times length of fourth lanati Spencer, p. 367 Second costal segment four to four and a half times length of fourth spondylii R.-D., p. 379 Upper Ors normally lacking 18 Upper Ors present; aedeagus as in Fig. 93, 94 columbinae n. sp., p. 362 Second costal segment more than three times length of fourth, approximately 3.3 times; notopleural areas dark; larvae make linear leaf mines on Aster con- spicuus Lindl asterophaga Spencer, p. 358 Second costal segment less than three times length of fourth; notopleural areas yellowish; larvae make linear leaf mines on Aster ciliolatus Lindl ciliolati Spencer, p. 360 Third antennal article with conspicuously long pubescence; aedeagus as in Fig. 104 lactuca Frost, p. 366 Third antennal article with normal pubescence 20 Broad epistoma present; gena deeply extended 21 Mouth margin normal 24 Second antennal article black 22 Second antennal article yellowish 23 Larger specimens, wing length 3. 0-3. 4 mm illustris Spencer, p. 365 Smaller specimens, wing length about 2.4 mm in male; aedeagus as in Fig. 88, 89 352 Sehgal 23(21). 24(20). 25(24). 26(25). 27(26). 28(27). 29(26). 30(29). 31(24). 32(31). 33(32). 34(32). 35(34). 36(31). 37(36). 38(37). 39(36). 40(39). blairmorensis n. sp., p. 358 Mesonotum light grey lupini Sehgal, p. 367 Mesonotum darker, blackish grey aquilegiophaga Spencer, p. 356 Acrostichals in three to six rows 25 Acrostichals at most in two rows 31 Frons slightly darkened above lunule; aedeagus as in Fig. 126 solidaginophaga n. sp., p. 378 Frons entirely pale 26 Gena deep, approximately one-third to one-half eye height 27 Gena narrower, one-sixth to one-fifth vertical eye height 29 Orbits yellow; third antennal article distinctly elongate banffensis Spencer, p. 358 Orbits black; third antennal article not so elongate 28 Mesonotum paler grey; frons entirely yellow urbana Spencer, p. 381 Mesonotum darker grey; frons slightly brownish yellow subtilis Spencer, p. 380 Fore-tibia yellowish; gena approximately one-fifth eye height; aedeagus as in Fig. 133 timida Spencer, p. 381 Fore-tibia dark; gena approximately one-sixth eye height; larvae blotch-miners on leaves of Aquilegia and Thalictrum 30 Aedeagus as in Fig. 81, 82 aquilegioides n. sp., p. 355 Aedeagus as figured by Spencer (1969) aquilegiana Frost, p. 354 Acrostichals lacking or at most three to four isolated hairs present 32 Acrostichals in two rows 36 Squamal fringe dark; second costal segment about twice length of fourth .... 33 Squamal fringe pale; second costal segment one and a half times length of fourth 34 Aedeagus with distiphallus distinctly curved (Fig. 123) senecionella n. sp., p. 377 Aedeagus with distiphallus paler and not so curved, as figured by Spencer (1969) syngenesiae (Hardy), p. 380 Second antennal article black; larva leaf miner on Penstemon penstemonis Spencer, p. 373 Second antennal article yellow or slightly brownish 35 Fore-coxae bright yellow; second antennal article yellow plantaginis R.-D., p. 374 Fore-coxae dark; second antennal article brownish; aedeagus as in Fig. 91 colemanensis n. sp., p. 361 Fore-coxae dark 37 Fore-coxae yellow. 39 Squamal fringe dark 38 Squamal fringe pale; aedeagus as in Fig. 130, 131 subalpina n. sp., p. 379 Aedeagus with distiphallus membranous, as figured by Spencer (1969) fuscula Zetterstedt, p. 364 Aedeagus with distiphallus darkly sclerotized as in Fig. 84; larva leaf miner on Arnica .’ amicivora n. sp., p. 357 Frons distinctly projected above eyes in profile 40 Frons not so projected; aedeagus as in Fig. 114 . misella Spencer, p. 371 Gena deep, about two-third eye height; aedeagus as illustrated (Spencer, 1969) subtenella- Frost, p. 380 Agromyzidae of Alberta 353 41(1). 42(41). 43(42). 44(43). 45(43). 46(45). 47(46). 48(42). 49(48). 50(41). 51(50). 52(51). 53(51). 54(53). 55(53). 56(55). 57(56). Gena narrower, about one-third eye height; aedeagus as in Fig. 102 jasper ensis n. sp., p. 365* Upper Ors shorter than lower or absent 42 Two Ors equal 50 Upper Ors present 43 Upper Ors absent 48 Second costal segment more than three times length of fourth. 44 Second costal segment less than three times length of fourth 45 Larva leaf miner on Aralia aralivora Spencer, p. 357 Larva leaf miner on Angelica sp. indet. (Angelica), p. 382 Acrostichals in two rows; larvae leaf miner on Delphinium delphinivora Spencer, p. 363 Acrostichals in approximately four irregular rows 46 Frons partly yellowish 47 Frons darker; aedeagus as in Fig. 110, 111; larvae leaf miner on Mertensia mertensiae n. sp., p. 369 Third antennal article small; acrostichals strong sehgali Spencer, p. 376 Third antennal article larger, oval; acrostichals normal; larvae blotch-miners on leaves of Anemone canadensis L prava Spencer, p. 374 Second costal segment more than three times length of fourth; larger specimens, wing length about 2.4 mm; tibiae and tarsi yellowish brown; aedeagus as in Fig. 97 edmontonensis n. sp., p. 363 Second costal segment less than three times length of fourth; smaller specimens, wing length about 1 .6-1.9 mm; tibiae and tarsi dark 49 Frons slightly paler; acrostichals absent; aedeagus with distiphallus straight .... aquilegivora Spencer, p. 356 Frons darker; acrostichals present; aedeagus with distiphallus wavy thalictrivora Spencer, p. 38 1 Tarsi yellowish brown; larva leaf miner on Cornus .... agromyzina Meigen, p. 354 Tarsi dark brown 51 Second costal segment at least three times length of fourth 52 Second costal segment less than three times length of fourth 53 Larger specimens, wing length 2. 8-3. 3 mm; mesonotum greyish; third antennal article elongate involucratae Spencer, p. 365 Smaller specimens, wing length about 2.7 mm; mesonotum blackish; third anten- nal article rounded milii Kaltenbach, p. 370 Mesonotum brilliantly shining black 54 Mesonotum distinctly mat, greyish or black 55 Orbits normal in width ; only fore-femur with yellow distal tip canadensis Spencer, p.359 Orbits broad; distal tips of femora variable from yellow to almost black; wing base yellow; aedeagus as in Fig. 116 multifidae n. sp., p.371 Third antennal article elongate; frons distinctly projecting above eyes cineracea Hendel, p. 360 Third antennal article normal, rounded at tip 56 Acrostichals in two rows 57 Acrostichals in approximately four irregular rows 58 Second costal segment about one and a quarter times length of fourth; orbits dark lupinivora Sehgal, p.367 Second costal segment almost equal to iourth; aedeagus as in Fig. 119 354 Sehgal oxytropodis n. sp., p.372 58(56). Mesonotum black 59 Mesonotum paler, greyish 60 59(58). Gena deep, about one-half of eye height; broad rings below eyes formed by orbits merula Spencer, p. 370 Gena narrower at most one-fourth eye height; aedeagus as in Fig. 99 gregaria Frick, p. 364 60(58). Frons distinctly projected; orbits well differentiated evanescens Hendel, p. 364 Frons not projected; orbits normal 61 61(60). Frons slightly pale above; distiphallus with distinctly curved distal processes, as figured by Spencer (1969) queribunda Spencer, p. 375 Frons entirely black 62 62(61). Smaller specimens, wing length about 1.75-2.0 mm; aedeagus as figured by Spen- cer ( 1 969) caprifoliae Spencer, p. 360 Larger specimens, wing length 2. 0-2. 3 mm; aedeagus with smaller hypophallus and stouter distiphallus, as figured by Spencer (1969) periclymeni de Meijere, p. 373 Phytomyza agromyzina Meigen Phytomyza agromyzina Meigen, 1 830: 191. Comparison and diagnostic characters. - The members of this species belong to the group having dark frons and two Ors equal. The adults are quite distinctive in having brownish yellow tibiae and tarsi. They resemble those of P. notopleuralis Spencer from which they may be separated by having predominantly dark pleura and distinct male genitalia. Spencer ( 1 969) illustrated the distinctive aedeagus. Other distinguishing characters of the adults are: wing length approximately 2.0 mm; mesonotum with slight yellow on humeral and noto- pleural areas; antennae dark; third antennal article rounded apically, with normal pubes- cence; and dark femora. Biology. - Larvae make linear mines in the leaves of Cornus stolonifera Michx. and C. canadensis L., family Cornaceae. Geographical distribution. — Known from Europe (Hendel, 1935) and in the United States from California and Washington (Frick, 1959) and Canada (Spencer, 1969). I exam- ined the following material from Alberta: CANADA. Alberta: 1 6 Edmonton, White Mud Creek park, 19.v. 1968; 2 66 same locality, lO.vi. 1968, coll. G. C. D. Griffiths; 2 66 same locality, 4.v. and 8.vi. 1969. Numerous leaf mines around Edmonton on Cornus stolonifera Michx. Phytomyza aquilegiana Frost Phytomyza aquilegiana Frost, 1930:459. Comparison and diagnostic characters. — The members of this species belong to the group having yellow frons; dark scutellum and pleura; third antennal article black, with normal pubescence; two Ors equal. The adults resemble those of a sympatric species, P. aquilegi- oides new species, in external characteristics and can be reliably separated only by examina- tion of male genitalia. Spencer (1969) illustrated the distinctive aedeagus. Biology. - Larvae make blotch mines in the leaves of Aquilegia spp. and Thalictrum spp., family Ranunculaceae. Agromyzidae of Alberta 355 Geographical distribution. - Known from United States (Frick, 1959) and Canada (Spen- cer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 9 Edmonton, University of Alberta campus, from blotch leaf mines on Aquilegia sp. (cultivated), coll. 16-18.vii.1966, emerged 1 4.iii. 1 967 ; numerous leaf mines around Edmonton on Aquilegia sp.; Yukon Territory: 1 6 Teslin Lake, from leaf mines on Aquilegia brevistyla Hook., coll. 1 1 .viii. 1968, emerged, lO.v.1969, coll. G. C. D. Griffiths. Phytomyza aquilegioides new species Comparisons and diagnostic characters. - The members of this species belong to the group characterized by yellow frons; normal mouth margin; dark scutellum; mostly dark femora and pleura; and three to six rows of acrostichals. The upper orbital bristles vary in length from almost equal to two-thirds the length of lower. Therefore, this species.has been included in two couplets in Spencer’s (1969) key to Canadian species of the genus Phyto- myza Fallen as amended below: 19. Only 1 Ors present; second costal section at most three and one-third times length of fourth asterophaga Spencer Both Ors present; second costal section three and one-half to four times lensth of fourth 19a 1 9a. Lower Ors only slightly weaker than upper; aedeagus as in Fig. 81,82 aquilegioides n. sp. Invariably small upper Ors present; aedeagus not so 20 36. Jowls deep, almost one-half eye height; third antennal segment distinctly elongate; aedeagus as illustrated (Spencer, 1969) banffensis Spencer Jowls narrower, about one-sixth eye height; third antennal segment rounded 36a 36a. Aedeagus as illustrated (Spencer, 1969) aquilegiana Frost Aedeagus as in Fig. 81,82 aquilegioides n. sp. Description. — Head. Frons approximately 1.5 times width of eye at level of front ocellus, not projected in front of eye margin in profile. Mouth margin normal; lunule high. Two Ors, directed upwards, length of upper Ors varies from equal to two-thirds length of lower; two Ori, directed inwards and upwards, almost equal in size; orbital setulae six to seven, procli- nate. Eyes oval, approximately 1.35 times higher than their length, bare; ocellar triangle small. Gena approximately one-sixth vertical eye height. Third antennal article rounded at tip, with normal pubescence; arista normal, with long pubescence. Mesonotum. Dorsocentrals 3+1 strong bristles; acr four to five irregular rows. Wing. Length approximately 2. 1-2.5 mm; costa extended to vein R4+5; costal segments 2-4 in the ratio of 1 : 0.28 : 0.27; crossvein m-m absent. Male genitalia (Fig. 80-82). Hypandrium (Fig. 80) V-shaped, with narrow side arms and short, broad apodeme; pregonites broad, postgonites elongate; surstyli normal; aedeagus (Fig. 81, 82) as illustrated; ejaculatory apodeme small, fan-shaped, with small bulb. Colour. Frons varies from bright yellow to orange; orbits and gena yellow; ocellar triangle weakly shining black; both vertical bristles on dark ground; antennae black; mesonotum, scutellum and pleura mat greyish black; humeral areas with slight yellow; coxae black; femora dark, with yellow oh distal tips; tibiae and tarsi yellowish brown; squamae and fringe pale; halteres yellow. Derivation of the specific name. — The name aquilegioides indicates that the members of this species have similar biology to those of P. aquilegiae Hardy. Biology. — Larvae make blotch mines on the leaves of Aquilegia formosa Fisch. and 356 Sehgal Thalictrum venulosum Trel., family Ranunculaceae. Pupation takes place outside the mine. The dark brown puparia measure approximately 1.75 mm x 0.8 mm, and are covered all over with conspicuous tubercles and spines, as in the palaearctic species P. thalictricola Hendel. Geographical distribution. — The members of this species are known only from the local- ities of their type specimens as below: CANADA. Alberta: Holotype <5 Edmonton, White Mud Creek park, from leaf mines on Thalictrum venulosum Trek, coll. 5.ix.l968. em. 5.xii.l968; paratypes 1 6 same data, emerged 29.xi.1968. ALASKA. Paratypes 1 <3, 2 99 Chilkat, near Haines, from leaf mines on Aquilegia formosa Fisch., coll. 29.vi.1968, emerged 22.vii.1968, 12.x. 1968 and 21.V.1968, coll. G. C. D. Griffiths. Phytomyza aquilegiophaga Spencer Phytomyza aquilegiophaga Spencer, 1969:227. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum; mostly dark pleura and femora; third antennal article black, with normal pubescence and broad epistoma. The adults resemble those of P. lupini Sehgal but differ in having darker or blackish grey mesonotum and distinct male genitalia. They differ from other related species, P. affinalis Frost, P. blairmorensis new species and P. illustris Spencer, in having second antennal article yellowish brown. Spencer (1969) illustrated the distinctive aedeagus. Biology. — Larvae bore inside the stems of Aquilegia sp. (cultivated), family Ranuncula- ceae, and pupate at the stem base. The dark brown pupae can be found during late summer. Geographical distribution. - The members of this species are known from Canada (Spen- cer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 <3 Edmonton, Aberhart Hospital, University of Alberta, from stem- base of Aquilegia sp. (cultivated), coll. 4.ix.l968, emerged 3.x. 1968; 1 <3, 3 99 same locality, swept over Aquilegia sp., l.vi.1967; 2 66, 1 9 same locality, from stem-base of Aquilegia sp., coll. 3.ix.l968, emerged 26.ix.-26.x.l968, coll. G. C. D. Griffiths. Phytomyza aquilegivora Spencer Phytomyza aquilegiv ora Spencer, 1969:229. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by dark frons, mesonotum and scutellum; upper Ors lacking; essentially dark femora and pleura and second costal segment less than three times length of fourth. The adults resemble those of P. thalictrivora Spencer but differ in having yellowish frons and lacking acrostichals. They also differ from the similar species, P. minuscula Goureau, in having very different male genitalia. Spencer (1969) illustrated the aedeagus characteristic of the species. Biology. — Larvae make linear mines on the leaves of Aquilegia sp. (cultivated), family Ranunculaceae. Pupation takes place outside the mine. Geographical distribution. — The members of this species are known only from the local- ity of its type series from Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 5 66 Edmonton, Aberhart Hospital, University of Alberta campus, swept over Aquilegia sp. (cultivated), 25.vi.1969; 1 <3 same locality, l.vi.1967; 3 66 same Agromyzidae of Alberta 357 locality, from leaf mines on Aquilegia sp., coll. 4.ix.l968, emerged 16-1 9.ix. 1 968, 30.x. 1968; 2 66 Edmonton, Garneau, from same host, coll. 3.ix.l968, emerged 1 5-1 7.ix. 1 968. Phytomyza aralivora Spencer Phytomyza aralivora Spencer, 1969:230. Comparison and diagnostic characters. - The members of this species belong to the group characterized by dark frons, mesonotum and scutellum; upper Ors shorter than lower; and second costal segment more than three times fourth. The adults resemble those of P. osmor- hizae Spencer but differ in having yellow tarsi and distinct male genitalia. Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — Larvae make linear mines on the leaves of Aralia nudicaulis L., family Aralia- ceae. Pupation takes place outside the mine. Geographical distribution. - The members of this species are known only from Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 6 Edmonton, river bed near University of Alberta campus, 14.vi. 1969; Numerous leaf mines around Edmonton and in Elk Island park on Aralia nudicaulis L. Phytomyza arnicivora new species Comparison and diagnostic characters. — The main distinguishing characters of the mem- bers of this species are pale frons; two equal Ors; third antennal article normal; mouth mar- gin normal; mesonotum, pleura and scutellum all mat greyish black; femora black, with slight yellow on distal tips and squamal fringe dark. The adults resemble those of P. fuscula Zetterstedt and can be reliably distinguished only by examination of male genitalia. The members of this species are included in Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen by extending the couplet 50 as below: 50. Mesopleura entirely grey 50a Mesopleura with upper margins narrowly yellow 51 50a. Aedeagus with distiphallus membranous, as illustrated (Spencer, 1969) fuscula Zetterstedt Aedeagus with distiphallus sclerotized, as in Fig. 84 arnicivora n. sp. Description. — Head. Frons approximately twice width of eye at level of front ocellus, not projected in front of eye margin in profile. Mouth margin normal. Two Ors, equal in size, directed upwards; one large Ori and a small hair below, directed inwards and upwards; orbital setulae few, approximately eight to nine, proclinate. Eyes oval, almost equal in height to their length; ocellar triangle small. Gena approximately one-fourth vertical eye height. Third antennal article rounded at tip, with normal pubescence, arista normal, pubes- cent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr in two rows. Wing. Length in male 2.4 mm; costa extended to vein R4+5 ; costal segments 2-4 in the ratio of 1 : 0.3 : 0.5; crossvein m-m absent. Male genitalia (Fig. 83-85). Hypandrium (Fig. 83) small, side arms broad, no apodeme; pregonites broad; postgonites elongate; surstyli normal; aedeagus complex as illustrated in Fig. 84; ejaculatory apodeme (Fig. 85) small, bulb small and membranous. Colour. Frons pale whitish, slightly darkened at centre; orbits pale;gena yellowish; ocel- lar triangle weakly shining black; vte on black and vti on margin of dark and yellow grounds; antennae black; mesonotum, scutellum and pleura mat greyish black; coxae black; femora black, with slight yellow on distal tips; tibiae and tarsi black; squamae pale, fringe dark; 358 Sehgal halteres pale. Derivation of the specific name. — This species is named after the generic name of its food plant. Biology. — Larvae make linear leaf mines on Arnica cordifolia Hook., family Compositae. Pupation occurs inside the leaf mine and the whitish puparia can be collected during July and August. Geographical distribution. - The members of this species are known only lrom the type locality: CANADA. Alberta: Holotype 6 Jasper National Park, near Medicine Lake; from leaf mines on Arnica cordifolia Hook., coll. 16.vii.1969, emerged 24.vii.1969, coll. G. C.D. Griffiths; Numerous leaf mines on the same host around Jasper. Phytomyza asterophaga Spencer Phytomyza aster ophaga Spencer, 1969:230. Comparison and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum; essentially dark femora and pleura and upper Ors lacking. The adults resemble those of P. ciliolati Spencer and differ in having a darker notopleural area and different biology. Spencer (1969) illustrated the aedeagus and leaf mine distinctive of this species. Biology. - Larvae make linear mines on the leaves of Aster conspicuus Lindl., family Compositae. The leaf mines are distinctive in having frass disposed in the form of discrete granules alternately in the mine. Pupation occurs outside the mine. Geographical distribution. — The members of this species were previously known only from the locality of type series from Western Canada (Spencer, 1969). I examined the fol- lowing material from Alberta: CANADA. Alberta: 1 6 Blairmore, 26.vi.1966; 1 <5 Edmonton, White Mud Creek park, from leaf mines on Aster conspicuus Lindl., coll. 10.ix.1966, emerged 8.iii. 1 967 ; 2 66 Elk Island park, same host, emerged 4.vi. 1967. Phytomyza banffensis Spencer Phytomyza banffensis Spencer, 1969:231. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; essentially dark scutellum, pleura and femora; acrosti- chals in three to six rows and third antennal article with normal pubescence. The adults resemble those of P. aquilegiana Frost and P. aquilegioides new species but differ in having deeper gena, approximately one-half of vertical eye height, elongate third antennal article and distinct male genitalia. Spencer (1969) illustrated the distinctive aedeagus. Geographical distribution. — The members of this species were previously known only from the locality of its type series from western Canada (Spencer, 1969). 1 examined the following material from Alberta: CANADA. Alberta: 2 66 Jasper, 17.vi.1966. Phytomyza blairmorensis new species Comparison and diagnostic characters. — A member of this species belongs to the group characterized by yellow frons; two equal Ors; broad epistoma; normal third antennal seg- ment; dark scutellum and mostly dark femora and pleura. The adult resembles those of Agromyzidae of Alberta 359 P. lupini Sehgal and P. aquilegiophaga Spencer but differs in having second antennal article black and distinct male genitalia. It also resembles other similar species, P. illustris Spencer and P. affinalis Frost, and may be separated from them as shown below in extension to Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen. 29a. Large specimens, wing length 3. 0-3. 4 mm; normally one Ors . . . .illustris Spencer Smaller specimens, wing length at most 2.4 mm in male; two Ors 29b 29b. Frons entirely yellow; aedeagus as illustrated (Spencer, 1969) affinalis Frost Frons slightly darkened; aedeagus as in Fig. 88, 89 blairmorensis n. sp. Description. - Head. Frons approximately 2.5 times width of eye at level of front ocellus; orbits broad, distinctly projected in front of eye margin in profile; broad epistoma. Two equal Ors, directed upwards; two Ori, directed inwards and upwards, lower one weaker than upper; orbital setulae few, six to seven, proclinate. Eyes oval, slightly slanted, their vertical height being approximately 1 .25 times their length; ocellar triangle small. Gena approxi- mately one-third vertical eye height. Third antennal article large, circular, with normal pubescence; arista normal, pubescent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr approximately eight to nine, in two rows. Wing. Length in male approximately 1.75 mm; costa extended to vein R4+5; costal seg- ments 2-4 in ratio of 1 : 0.35 : 0.65; crossvein m-m absent. Male genitalia (Fig. 86-90). Hypandrium (Fig. 86) V-shaped, narrow side arms, no distinct apodeme; pregonites broad; postgonites (Fig. 87) long, with hook-like process anteriorly; aedeagus (Fig. 88, 89) as illustrated; ejaculatory apodeme (Fig. 90) small, well sclerotized, bulb small, membranous. Colour. Frons yellow, very slightly darkened above; orbits yellow, darkened slightly near upper Ors; gena and lunule yellow; ocellar triangle weakly shining black; both Vt’s on dark ground; antennae black; mesonotum, scutellum and pleura mat grey; legs black, only distal tips of femora with slight yellow; squamae pale, fringe brown; halteres yellow. Derivation of the specific name. — This species is named after the locality of its type specimen. Geographical distribution. — A member of this species is known only from the type locality: CANADA. Alberta: Holotype 6 Blairmore, 26.vi.1966. Phytomyza canadensis Spencer Phytomyza canadensis Spencer, 1 969: 23 1 . Comparison and diagnostic characters. — The main distinguishing characters of the mem- bers of this species are: dark frons; two equal Ors; brilliantly shining black mesonotum and scutellum; acrostichals in approximately two rows; dark tarsi and second costal segment less than three times length of fourth. The adults resemble those of the very similar species, P. multifidae new species, but differ in having narrower orbits and different puparia. Biology. — The larvae were stated by Spencer (1969) to make linear mines in the leaves of Anemone canadensis L., family Ranunculaceae. The characteristic leaf mines have been illustrated by Spencer (1969). However, a confusion seems to have arisen, since the leaf figured by Spencer is clearly not of this species. Mr. Griffiths and I have found similar leaf mines only on Anemone riparia Fern., never on A. canadensis L. The mined leaf figured by Spencer was probably also of A. riparia Fern. Geographical distribution. — The members of this species are known only from Canada 360 Sehgal from the type locality (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: Numerous empty leaf mines on Anemone riparia Fern., Edmonton, White Mud Creek park, 5-6.ix.1968, and in Elk Island park. Phytomyza capri foliae Spencer Phytomyza caprifoliae Spencer, 1969:233. Comparison and diagnostic characters. — The main distinguishing characters of the mem- bers of this species are: frons black, not projected; two equal Ors; normal third antennal article; mat grey mesonotum, scutellum and pleura; approximately four rows of acr; black tarsi and second costal segment less than three times length of fourth. The adults resemble those of P. periclymeni de Meijere and can be reliably separated only by examination of male genitalia. Spencer (1969) illustrated the distinctive aedeagus. Biology. - Larvae mine the leaves of Symphoricarpos sp., family Caprifoliceae. Geographical distribution. - The members of this species are known only from the local- ity of its type series from Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 2 66, 6 99 Edmonton, White Mud Creek park, from leaf mines on Symphoricarpos sp., coll. 10. ix. 1966, emerged 9.i.-7.iii.l967; 2 66 same locality, 14.V.1968 and 8.vi. 1967. Phytomyza ciliolati Spencer Phytomyza ciliolati Spencer, 1969:234. Comparison and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum; essentially dark femora and pleura and upper Ors absent. The adults resemble those of P. asterophaga Spencer but differ in having yellow on notopleural areas and different biology. Biology. — Larvae make linear mines on the leaves of Aster ciliolatus Lindl., family Com- positae. The leaf mines are distinctive in having frass disposed in continuous streaks. Pupa- tion occurs outside the mine. Spencer (1969) illustrated the characteristic leaf mine. Geographical distribution. - The members of this species are known only from the type locality (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: Numerous leaf mines on Aster ciliolatus Lindl. around Edmonton during July and August. Phytomyza cineracea Hendel Phytomyza cineracea Hendel, 1920: 160. Comparison and diagnostic characters. — The main diagnostic characters of the members of this species are: yellowish brown frons; mat grey mesonotum and scutellum; black tarsi and second costal segment approximately two times the length of the fourth. The adults are distinctive in having an elongate third antennal article. They differ from those of the similar species, P. erigerontophaga Spencer, in having frons distinctly projected above eyes and distinct male genitalia. Griffiths (1968) and Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — Larvae of this species feed inside the stems of Ranunculus spp., Ranuncula- ceae (Griffiths, 1968). Geographical distribution. - The members of this species are known from Europe, Ice- Agromyzidae of Alberta 361 land (Griffiths, 1968) and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 8 66 Blairmore, swept over Ranunculus acris L., family Ranuncula- ceae, 26.vi.1966, 3 66 Jasper, 16-1 9.vi. 1966. Phytomyza clematiphaga Spencer Phytomyza clematiphaga Spencer, 1969:236. Comparison and diagnostic characters. — The members of this species belong to the group characterized by yellow frons and partially yellow scutellum. The adults differ from those of P. major Malloch in having their body darker and third antennal article black. They differ from those of P. ranunculi (Schrank) in having both Ors of equal length and distinct male genitalia. Spencer ( 1 969) illustrated the distinctive aedeagus. Biology. — Larvae make linear mines on the leaves of Clematis verticillaris DC. family Ranunculaceae. Pupation occurs inside the leaf mine. Geographical distribution. — The members of this species are known from Canada only from the type locality. I examined the following material from Alberta: CANADA. Alberta: Holotype 6 (in K. A. Spencer’s collection) Edmonton, river bed near University of Alberta campus, from leaf mines on Clematis verticillaris DC, coll. 26.vii. 1966, emerged 7.viii.l966, coll. B. Hocking; paratype 1 9 (in K. A. Spencer’s collection) Edmon- ton, University of Alberta campus, from same host, coll. 24.ix.1966, emerged 9.x. 1966; 2 66, 4 99 Edmonton, river bed near University of Alberta campus, same host, coll. 26.vii. 1966, emerged 6-12.viii. 1966, coll. B. Hocking; 4 99 Edmonton, University of Alberta campus, same host, coll. 23.ix.1966, emerged 1.x. 1966, 4.ii. 1967 and ll.iii.1967. Phytomyza colemanensis new species Comparison and diagnostic characters. — A member of this species belongs to the group characterized by yellow frons; two equal Ors; mouth margin normal; third antennal article with normal pubescence; dark scutellum; mostly dark femora and pleura; and acrostichals approximately three to four scattered hairs. The adult resembles those of P. penstemonis Spencer and P. plantaginis R.-D. from which it may be separated as shown below in the extension to Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen: 41. Fore-coxae essentially dark 41a. Fore-coxae conspicuously yellow plantaginis R.-D. 41a. First and second antennal segment black; aedeagus as illustrated (Spencer, 1969) penstemonis Spencer First and second antennal segment yellowish brown; aedeagus as in Fig. 91 colemanensis n. sp. Description. — Head. Frons almost twice width of eye at level of front ocellus, slightly projected in front of eye margin in profile. Mouth margin normal. Two equal Ors directed upwards; one strong Ori incurved, one small hair present below Ori; orbital setulae only two, proclinate. Eyes slightly slanted, their vertical height 1.2 times their length, bare; ocellar triangle small. Gena approximately one-third vertical eye height. Third antennal article with slight angle anterodorsally, with normal pubescence; arista normal, pubescent. Mesonotum. Dorsocentral 3+1 strong bristles; acr few, three to four scattered hairs. Wing. Length in male 1.6 mm; costa extended to vein R4+5 ; costal segments 2-4 in the ratio of 1 : 0.3 : 0.7; crossvein m-m absent. Male genitalia (Fig. 91). Hypandrium U-shaped, with broad side arms; pregonites broad; 362 Sehgal postgonites elongate, with curved process anteriorly; surstyli normal; aedeagus (Fig. 91) with distinctive hypophallus; ejaculatory apodeme broad, bulb small and membranous. Colour. Frons, orbits and gena yellow; ocellar triangle weakly shining black; Vte on black and Vti on margin of dark and yellow ground; first and second antennal articles yellowish brown; third antennal article black; legs black; mesonotum, scutellum and pleura mat grey- ish; squamae and fringe pale. Derivation of the specific name. — This species is named colemanensis after the name of the type locality. Geographical distribution. — A member of this species is known only from the type locality: CANADA. Alberta: Holotype 6 Coleman, 27.vi.1966. Phytomyza columbinae new species Comparison and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; upper Ors shorter than lower; dark scutellum; essentially dark femora and pleura. The adults differ from those of the similar species, P. timida Spencer, and may be separated as shown below in extension to Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen: 24. Jowls narrow, one-fifth to one-fourth vertical eye height 24a Jowls deeper, one-third to one-half vertical eye height 25 24a. Fore-tibia yellowish; aedeagus as illustrated (Spencer, 1969) .... timida Spencer Fore-tibia dark; aedeagus as in Fig. 93, 94 columbinae n. sp. Description. — Head. Frons approximately twice width of eye at level of front ocellus, not projected in front of eye margin in profile. Mouth margin normal; lunule high. Two Ors, directed upwards, upper one shorter than lower; two Ori, directed inwards and upwards; orbital setulae few five to six, proclinate. Eyes oval, approximately 1.2 times higher than their length, bare; ocellar triangle small. Gena approximately 0.22 times vertical eye height. Third antennal article rounded at tip, with normal pubescence; arista normal, pubescent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr in four to five irregular rows. Wing. Length 1.5- 1.8 mm; costa extended to vein R4+5 ; costal segments 2-4 in the ratio of 1 : 0.33 : 0.4; crossvein m-m absent; wing tip at M3+4. Male genitalia (Fig. 92-95). Hypandrium (Fig. 92) V-shaped, with small apodeme; prego- nites broad; postgonites elongate; surstyli normal; aedeagus (Fig. 93, 94) as illustrated; ejaculatory apodeme (Fig. 95) small, fan-shaped, bulb small, membranous. Colour. Frons and gena pale; orbits slightly darkened, lunule dark; ocellar triangle shining black; antennae black; both Vt’s on dark ground; mesonotum, scutellum and pleura mat greyish black; legs black; squamae pale, fringe brown; halteres yellow. Derivation of the specific name. — This species is named columbinae after the common name of its food plant, columbine ( Aquilegia ). Biology. — Larvae make blotch mines in the leaves of Aquilegia sp. (cultivated) and Thalictrum venulosum Trel., family Ranunculaceae. Pupation occurs outside the mine. The dark brown puparium measures approximately 1.5 mm x 0.75 mm and is densely covered with small spinules. Geographical distribution. — The members of this species are known only from the type locality: CANADA: Alberta: Holotype 6 Edmonton, White Mud Creek park, from blotch mines on Thalictrum venulosum Trel., coll. 6.ix.l968, emerged 19.x. 1968; paratypes 2 66 Edmon- ton, Aberhart Hospital, University of Alberta campus, from leaf mines on Aquilegia sp. Agromyzidae of Alberta 363 (cultivated), coll. 4.ix.l968, emerged 1 8.ix. 1 968 and 2.ii. 1 969 ; 2 66 same locality, swept over Aquilegia sp., 25.vi.1969; 1 6 Edmonton, University of Alberta campus, from leaf mines on Aquilegia sp., coll. 13.vi. 1969, emerged 17.vi.1969; 2 66 Edmonton, Rainbow Valley, from leaf mines on Thalictrum venulosum Trel., coll. 14.vi. 1968. Phytomyza delphinivora Spencer Phytomyza delphinivora Spencer, 1969:238. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by dark frons, mesonotum and scutellum; essentially dark femora and pleura; upper Ors shorter than lower and second costal segment less than three times length of fourth. The adults differ from those of the similar species, P. mertensiae new species and P. prava Spencer, in having only two rows of acrostichals and distinct male genitalia.. Spencer ( 1 969) illustrated the aedeagus characteristic of this species. Biology. - Larvae mine the leaves of Delphinium sp. (cultivated), family Ranunculaceae. Spencer (1969) illustrated the characteristic linear mine. Pupation occurs outside the mine. Geographical distribution. — The members of this species are known only from the type locality (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 2 66 Edmonton, White Mud Creek park, 28.V.1967; Numerous leaf mines around Edmonton on Delphinium sp. (cultivated). Phytomyza edmontonensis new species Comparisons and diagnostic characters. — A member of this species belongs to the group characterized by dark frons; upper Ors absent; essentially dark femora and pleura; mat grey- ish black mesonotum and scutellum; and second costal segment more than three times length of fourth. The adult resembles that of P. modica Spencer from which it may be separated as shown below in an extension to Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen: 78. Second costal section long, five times length of fourth pallipes Spencer Second costal section shorter, up to 3.5 times length of fourth 78a 78a. Second costal section slightly over three times length of fourth; veins pale; aedea- gus as illustrated (Spencer, 1969) modica Spencer Second costal section 3.5 times length of fourth; veins brownish; aedeagus as in Fig. 97 edmontonensis n. sp. Description. — Head. Frons approximately twice width of eye at level of front ocellus, not projected in front of eye margin in profile. One Ors, curved upwards; three strong Ori directed inwards; orbital setulae numerous, proclinate. Eyes almost circular, their vertical height being 1 . 1 times their length; ocellar triangle small. Gena narrow, approximately 0.3 times eye height midway between vibrissal and posterior margins. Third antennal article slightly enlarged, rounded at tip, with normal pubescence; arista normal, pubescent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr in three irregular rows. Wing. Length in male 2.4 mm; costa extended to vein R4+5 ; costal segments 2-4 in the ratio of 1 : 0.26 : 0.3; crossvein m-m absent. Male genitalia (Fig. 96-98). Hypandrium (Fig. 96) U-shaped, with broad side arms, pre- gonites elongate; postgonites broad anteriorly and with a small hook; surstyli small; aedea- gus (Fig. 97) darkly sclerotized and as illustrated; ejaculatory apodeme (Fig. 99) very broad, bulb small and membranous, latter with darkly sclerotized areas. Colour. Frons, orbits and gena brown; ocellar triangle black; mesonotum, scutellum and 364 Sehgal pleura mat black; femora black, tibiae and tarsi yellowish brown; wing veins brownish; squamae and fringe pale; halteres yellow. Derivation of the specific name. — This species is named after the type locality. Geographical distribution. — A member of this species is known only from the type locality: CANADA. Alberta: Holotype 6 Edmonton, White Mud Creek park, 18.vi. 1968, coll. G. C. D. Griffiths. Phytomyza evanescens Hendel Phytomyza evanescens Hendel, 1920: 167. Comparisons and diagnostic characters. — The diagnostic characters of the members of this species are: dark frons; two equal Ors; normal third antennal article; mat grey mesono- tum, scutellum and pleura; dark tarsi; second costal segment less than three times length of fourth and acrostichals in approximately four rows. The adults differ from those of the similar species, P. caprifoliae Spencer, P. periclymeni de Meijere and P. queribunda Spencer in having frons distinctly projected and characteristic male genitalia. The surstyli have long wing-like processes. Griffiths (1964) illustrated the male genitalia characteristic of this species. Spencer (1969) also figured the aedeagus. Biology. — Larvae feed inside the stems of Ranunculus spp., family Ranunculaceae (Griffiths, 1 969). Geographical distribution. — The members of this species are known from Europe, Ice- land, Faroes (Griffiths, 1968) and Canada (Spencer, 1969). I examined the following materi- al from Alberta: CANADA. Alberta: 1 6 Blairmore, 26.vi.1966; 2 66 Jasper, 16-19.vi. 1966. Phytomyza fuscula Zetterstedt Phytomyza fuscula Zetterstedt, 1848:2831; Spencer, 1969:242. Comparison and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum; essentially dark femora and pleura; mouth margin normal; third antennal article black, with normal pubescence; two Ors equal; acr in two well-defined rows and dark fore-coxae. Spencer (1969) groups this species both under yellow frons and dark frons. The Alberta specimens correspond to the colour form having yellow frons. The aedeagus of Alberta specimens corresponds exactly to that figured by Spencer (1969) including the weakly sclerotized membranous processes in the distiphallus. Griffiths’ (1969) description of fusc- ula Zett. from Greenland refers to puccinelliae Spencer (see Spencer, 1969). Biology. — Larvae of this species mine the leaves of grasses (Gramineae) in Canada (Spen- cer, 1969). Geographical distribution. — The members of this species are known from Europe (Hen- del, 1935) and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 6 Edmonton, University of Alberta campus, 6.vi. 1968; 1 6 Elk Island park, 31.vii.1966. Phytomyza gregaria Frick Phy tomyza gregaria F rick , 1954:371. Comparison and diagnostic characters. — The distinguishing characters of the members of Agromyzidae of Alberta 365 this species are: dark frons; two equal Ors; normal third antennal article; mat black meso- notum, scutellum and pleura; acrostichals in approximately four rows; dark tarsi and second costal segment less than three times length of fourth. The adults resemble those of the similar species, P. periclymeni de Meijere, but differ in having darker mesonotum; narrower gena, approximately one-fourth eye height and distinct male genitalia. The aedeagus of an Alberta specimen is illustrated in Fig. 99. Spencer (1969) also illustrated the aedeagus. Biology. — The larvae mine the leaves of Lonicera involucrata (Richards) Banks, family Caprifoliaceae. Geographical distribution. — The members of this species are known from United States (Frick, 1959) and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 d St. Albert, near Edmonton, 1 8. vi. 1967; Numerous leaf mines on Lonicera involucrata (Richards) Banks around Edmonton Phytomyza illustris Spencer Phytomyza illustris Spencer, 1969:247. Comparison and diagnostic characters. — The members of this species differ from those of similar species, blairmorensis new species and affinalis Frost, in larger size, wing length 3. 0-3.4 mm and normally only one Ors. Spencer (1969) illustrated the aedeagus character- istic of this species. Geographical distribution. — Known from Alberta, British Columbia and Yukon Territory. The Alberta locality is: CANADA. Alberta: Blairmore (Spencer, 1969). Phytomyza involucratae Spencer Phytomyza involucratae Spencer, 1969:249. Comparison and diagnostic characters. — The members of this species differ from those of a similar species, milii Kaltenbach, in having larger size, wing length 2. 8-3. 3 mm, grey mesonotum and third antennal article large and elongate. Spencer (1969) illustrated the distinctive aedeagus. Geographical distribution. — Known from Alberta and British Columbia. The Alberta locality is: CANADA. Alberta: Frank; St. Albert. The following type specimen was examined: CANADA. British Columbia: Paratype 1 6 Prince George, 17.vi.1966, coll. K. A. Spencer. Phytomyza jasperensis new species Comparisons and diagnostic characters. - A member of this species belongs to the group characterized by yellow frons; normal mouth margin and third antennal article; two equal Ors; dark scutellum; mostly dark femora and pleura and acrostichals in two rows. The adult resembles that of P. pedicularicaulis Spencer and can be reliably separated only by examina- tion of male genitalia. This species may be included in Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen by amending couplet 43 and extending 44 as below: 43. Frons strongly projected above eyes; jowls conspicuously deep, two-thirds eye height subtenella Frost Frons less projected; jowls at most one-half eye height 44 44. Arista conspicuously thickened basally 44a 366 Sehgal Arista more slender, normal 45 44a. Aedeagus as illustrated (Spencer, 1969) pedicularicaulis Spencer Aedeagus as in Fig. 102 jasperensis n. sp. Description. — Head. Frons approximately 2.5 times width of eye at level of front ocellus, projected in front of eye margin in profile. Mouth margin normal. Two equal Ors directed upwards; two Ori, lower one smaller than upper, incurved; orbital setulae few, approximate- ly eight, proclinate. Eyes oval, their vertical height being 1 .3 times their length, bare; ocellar triangle small. Gena approximately one-third eye height. Third antennal article rounded at tip, with normal pubescence; arista slightly thickened at base, pubescent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr in two rows. Wing. Length in male 2.6 mm; costa extended to vein R4+5 ; costal segments 2-4 in ratio of 1 : 0.27 : 0.45; crossvein m-m absent. Male genitalia (Fig. 100-103). Hypandrium (Fig. 100) V-shaped, with narrow side arms; pregonites broad; postgonites (Fig. 101) long, with hook-like process anteriorly, surstyli normal; aedeagal apodeme exceptionally long; aedeagus (Fig. 102) relatively short and as illustrated; ejaculatory apodeme (Fig. 103) broad, bulb small and membranous. Colour. Frons, orbits and gena bright yellow; maxillary palpi black; ocellar triangle weak- ly shining black; lunule yellow; both Vt’s on dark ground; first antennal article yellowish, second and third articles black; mesonotum, scutellum and plerua mat grey; only mesopleura with narrow yellow band along upper margin; legs with fore-coxae yellowish, femora dark with yellow on distal tips, tibiae and tarsi black; squamae yellow, fringe brownish; halteres yellow. Derivation of the specific name. — The species is named jasperensis after the name of the type locality. Geographical distribution. — This species is known only from the following locality: CANADA. Alberta: Holotype 6 Jasper, 17.vi.1966. Phytomyza lactuca Frost Phytomyza lactuca Frost, 1924:85. Comparison and diagnostic characters. — The members oi this species belong to the group characterized by yellow frons; dark scutellum; essentially dark pleura and femora; and two Ors equal. The adults differ from all other species in this group by having conspicuously long pubescence on the third antennal article and distinct male genitalia. The aedeagus of an Alberta specimen bred from Crepis tectorum L. is illustrated in Fig. 104. Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — Larvae make long linear mines, usually on under surface of the leaves of Taraxacum officinale Weber, Crepis tectorum L. and Sonchus uliginosus Bieb., family Com- positae. Larvae are also known to mine the leaves of Lactuca scariola var. integrifolia (Bogenh.) G. Beck in Pennsylvania, U. S. A. (Frost, 1924). Geographical distribution. — The members of this species are known from United States (Frick, 1959) and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 6 Blairmore, 28.vi.1966; 1 6 Edmonton, river bed near University of Alberta campus, from leaf mines on Crepis tectorum L., coll. 15.vi.1969 emerged 12.vi. 1969; 1 9 same locality, from leaf mines on Sonchus uliginosus Bieb., coll. 15.vi.1969, emerged 6.vii.l969; 3 99 Edmonton, University of Alberta campus, from leaf mines on Taraxacum officinale Weber, coll. 7.x. 1966, emerged 19.xii.1966 and 5-6. iii. 1967; 2 66, 2 99 Waterton National park, same host, coll. 6.ix.l966, emerged 10-22.ix.1966 and 25.ii. 1967. Agromyzidae of Alberta 367 Phytomyza lanati Spencer Phytomyza lanati Spencer, 1969:250. Comparison and diagnostic characters. — The members of this species differ from those of a similar species, spondylii R.-D., in having second costal segment shorter, about 3.5 times length of fourth. These specimens cannot be satisfactorily separated on the basis of external characteristics alone; however, the male genitalia are distinct. Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. - Larvae mine leaves of Heracleum, family Umbellifereae. Details of leaf mine not known (Spencer, 1969). Geographical distribution. — The members of this species are known from California in United States and Alberta in Canada. The Alberta locality is: CANADA. Alberta: Jasper (Spencer, 1969). Phytomyza lupini Sehgal Phytomyza lupini Sehgal, 1968:73. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum; mostly dark pleura and femora; two Ors equal; third antennal article black, with normal pubescence and broad epistoma. The adults resemble those of P. aquilegiophaga Spencer and differ in having slightly paler grey mesonotum and distinct male genitalia. They differ from another similar species, P.blair- morensis new species, in having the second antennal article yellowish brown and distinct male genitalia. Sehgal (1968) illustrated the head, wing and male genitalia characteristic of this species. Spencer (1969) also illustrated the aedeagus. Biology. — Larvae bore inside the stems of Lupinus sericeus Pursh, family Leguminosae. The pale whitish puparia are found inside the stems. The puparia are characteristic in having a small horn in the posterior spiracles. Geographical distribution. — The members of this species are known only from western Canada: Alberta and British Columbia (Sehgal, 1968). The material examined remain the same as reported earlier (Sehgal, 1968). Phytomyza lupinivora Sehgal Phytomyza lupinivora Sehgal, 1968:74. Comparison and diagnostic characters. - The main distinguishing characters of the mem- ber of this species are: dark frons; distinctly mat greyish mesonotum and scutellum; dark tarsi; normal third antennal article and acrostichals in two rows. The adult resembles that of P. oxytropidis new species from which it is separated by having slightly longer second costal segment, approximately one and a quarter times the length of the fourth, and darker orbits. Sehgal (1968) illustrated the head and wing characteristic of this species. Biology. — Larvae make linear mines on the leaves of Lupinus sericeus Pursh, family Leguminosae. Pupation occurs outside the mine. Geographical distribution. — A member of this species is known only from the type locality: CANADA. Alberta: Blairmore (Sehgal, 1968). 368 Sehgal Phytomyza luteiceps new species Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum and yellow femora. The adults resemble those of P. flavicornis Fallen and can be separated reliably only by examination of charac- ters of the male genitalia. Spencer (1965d, 1969) illustrated the aedeagus of P. flavicornis Fallen. It is doubtful at present if the true P. flavicornis Fallen occurs in Alberta. This spe- cies is included in Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen, by amending and extending the couplet 10 as below: 10. All coxae yellow; jowls exceptionally deep at rear, at least two-thirds of eye height 10a Mid- and hind-coxae black rufipes Meigen 10a. Mesonotum black, weakly shining; aedeagus as illustrated (Spencer, 1969) flavicornis Fallen Mesonotum mat greyish black; aedeagus as in Fig. 106 luteiceps n. sp. Description. - Head. Frons wide, little more than three times width of eye at level of front ocellus, conspicuously projected in front of eye margin in profile. Broad epistoma present; lunule low. One strong Ors and three strong Ori; orbital setulae 10-1 1, proclinate. Eyes oval and slanting; their vertical height being almost equal to their length, bare; ocellar triangle small. Gena deep, approximately 0.7 times vertical eye height. Third antennal article rounded at tip, with short upcruved pubescence; arista normal and pubescent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr in approximately two rows. Wing. Length 2.5 to 2.8 mm; costa extended to vein R4+5; costal segments 2-4 in the ratio of 1 : 0.3 : 0.4; crossvein m-m absent. Male genitalia (Fig. 105-107). Hypandrium (Fig. 105) almost circular below, with broad arms as illustrated; pregonites broad; postgonites long with small process anteriorly; aedea- gus (Fig. 106) as illustrated; ejaculatory apodeme (Fig. 107) short, bulb small. Colour. Frons bright yellow; orbits and gena yellow; Vte on black and Vti on yellow ground; ocellar triangle shining black; lunule yellow; antennae completely yellow; arista brown; mesonotum and scutellum mat grey; humeral and notopleural areas yellow; stemo- pleura slightly brownish at base; meso- and pteropleura yellow; legs with coxae, femora and tibiae yellow, tarsi slightly brownish; squamae yellow, fringe brown; halteres yellow. Derivation of the specific name. — This species is named luteiceps because of the mostly yellow head. Biology. — Not confirmed, but the larvae will probably prove to feed in stems of Urtica. Geographical distribution. — The members of this species are known only from the local- ities of its type series as below: CANADA. Alberta: Holotype 6 St. Albert, near Edmonton, 14.vi.1966; allotype 9 same data; paratypes 3 66, 3 99 same data; 2 66 Jasper, 16.vi.1966. Phytomyza major Malloch Phytomyza major Malloch, 1 9 1 3b: 1 50. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons and yellow scutellum. The adults differ from those of other species in this group. P. clematiphaga Spencer and P. ranunculi (Schrank), in having a mostly yellow body, yellow third antennal article and distinct male genitalia. The adults are largely yellow flies, wing length approximately 4.0 mm. Spencer (1969) illustrated the distinctive aedeagus. The aedeagus of an Alberta specimen is as in the Fig. 108. Agromyzidae of Alberta 369 Geographical distribution. - The members of this species are known from Labrador (Frick, 1959) and western Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 6, 1 9 Banff, 1 4.vii. 1 949, coll. E. H. Strickland; 1 9 George Lake, near Busby, University of Alberta field station, Malaise trap collection, 1 7-2 l.vi. 1966, coll. P. Graham. Phytomyza matricariae Hendel Phytomyza matricariae Hendel, 1920: 161 ; Spencer, 1969:254. Comparison and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum; essentially dark femora; upper Ors shorter than lower; and with slight yellow on upper parts of mesopleura, humeral and notopleural areas. The adults resemble those of P. spondylii R.-D. but differ in having the second costal segment shorter, approximately three times the length of the fourth and distinct male geni- talia. Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — In Alberta the larvae make linear mines in the leaves of Achillea millefolium L., A. sibirica Ledeb., Chrysanthemum sp. (cultivated), Matricaria matricarioides (Less.) Porter, and Tanacetum vulgare L., belonging to the family Compositae. Pupation occurs outside the mine. Detailed biology and host-plant relationships are discussed in a separate paper (Sehgal, 1971). Geographical distribution. — The members of this species are known from Europe (Hen- del, 1935) and Canada (Spencer, 1969). I examined numerous specimens bred from all hosts listed above from various localities around Edmonton. Phytomyza mertensiae new species Comparisons and diagnostic characters. — The members of this species belong to the group characterized by dark frons; upper Ors shorter than lower; dark mesonotum and scutellum; essentially dark femora and pleura; and second costal segment less than three times length of fourth. The adults differ from those of the similar species, P. prava Spencer and P. sehgali Spencer, in having darker frons and distinct male genitalia. This species is included in Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen by amending and extending couplet 88 as below: 88. Third antennal article distinctly enlarged nepetae Hendel Third antennal article small 88a 88a. Frons paler, brownish above; acr strong sehgali Spencer Frons dark brown; acr normal; aedeagus as in Fig. 1 10, 1 1 1. . . . mertensiae n. sp. Description. — Head. Frons approximately two and a half times width of eye level of front ocellus, slightly projected in front of eye margin in profile. Mouth margin normal; lunule high. Two Ors, directed upwards, upper smaller than lower; two Ori, directed inwards and upwards, lower one weaker than upper; orbital setulae few, approximately six to seven, proclinate. Eyes oval, approximately 1.17 times higher than their length, bare; ocellar tri- angle small. Gena approximately 0.22 times vertical eye height. Third antennal article normal, rounded at tip; arista normal, pubescent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr in four irregular rows. Wing. Length approximately 2.0 mm; costa extended to vein R4+5; costal segments 2-4 in the ratio of 1 : 0.32 : 0.35; crossvein m-m absent. Male genitalia (Fig. 109-112). Hypandrium (Fig. 109) small, V-shaped, with broad side 370 Sehgal arms; pregonites broad; postgonites elongate; surstyli normal; aedeagus (Fig. 110, 111) com- plex as illustrated; ejaculatory apodeme (Fig. 112) broad, bulb small and membranous. Colour. Frons, orbits, gena and lunule dark brown; ocellar triangle weakly shining black; antennae black; mesonotum, scutellum and pleura mat greyish black; coxae black; femora black, with yellow on distal tips; tibiae and tarsi dark brown; squamae yellow, fringe brown; halteres pale. Derivation of the specific name. — This species is named after the generic name of its food plant. Biology. — Larvae make linear mines in the leaves of Mertensia paniculata (Ait.) G. Don, family Boraginaceae. Pupation occurs outside the mine. Geographical distribution. — The members of this species are known only from the type locality: CANADA. Alberta: Holotype d Edmonton, White Mud Creek park, from leaf mines on Mertensia paniculata (Ait.) G. Don, coll. 10.ix.1966, emerged 1 0.iii. 1 967 ; paratypes 1 9 same data; 1 6 same locality, 8.vi.l967. Phytomyza merula Spencer Phytomyza merula Spencer, 1969:254. Comparison and diagnostic characters. — The members of this species differ from those of a very similar species, gregaria Frick, in having deeper gena, about one-half eye height and orbits in form of a broad ring below eyes. Spencer (1969) illustrated the distinctive aedeagus. Geographical distribution. — Known only from Alberta, Canada from the following local- ity: CANADA. Alberta: Jasper (Spencer, 1969). Phytomyza milii Kaltenbach Phytomyza milii Kaltenbach, 1864:248; Spencer, 1969:255. Phytomyza intermedia Spencer; Griffiths, 1964:405. Comparison and diagnostic characters. — The members of this species belong to the group characterized by dark frons; mat black mesonotum and scutellum; two equal Ors, black tarsi and second costal segment at least three times the length of the fourth. The adults resemble those of P. involucratae Spencer and can be reliably separated only by examination of male genitalia. The sclerotization of distiphallus varies in this species (Griffiths, 1 964). The aedeagus of an Alberta specimen is as illustrated in Fig. 1 13. Griffiths (1964) illustrated the aedeagus of European and Faeroese specimens. Spencer (1969) also illustrated the aedeagus. Biology. — Larvae probably mine the leaves of grasses (Gramineae) in Alberta. Geographical distribution. — The members of this species are known from Europe, Ice- land, Faeroes (Griffiths, 1964) and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 6 Banff, 3.ix.l966; 2 66 Jasper, l-2.ix.1966; 3 66 Jasper, Mt. Edith Cavell, l.ix.1966. Phytomyza miranda Spencer Phytomyza miranda Spencer, 1969:255. Comparison and diagnostic characters. — The members of this species differ from those of Agromyzidae of Alberta 371 a similar species, luteiceps new species, in having black third antennal article. The elongate surstyli and aedeagus as figured by Spencer (1969) are quite distinct. Geographical distribution. - The members of this species are known only from Alberta, Canada from the following locality: CANADA. Alberta: Blairmore (Spencer, 1969). Phytomyza misella Spencer Phytomyza misella Spencer, 1969:256. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; normal mouth margin and third antennal article; two equal Ors; essentially dark femora and pleura; acrostichals in two rows; and yellow fore- coxae. The adults differ from those of the similar species, P. subtenella Frost, by having frons less projected, narrower gena and distinct aedeagus. They also resemble those of P. pedicularicaulis Spencer and P. jasperensis new species but have entirely different male geni- talia. The aedeagus of an Alberta specimen is illustrated (Fig. 114). Spencer (1969) also illustrated the aedeagus. Geographical distribution. — The members of this species are known only from western Canada from the type locality (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 4 66 Jasper, 17.vi.1966. Phytomyza multifidae new species Comparisons and diagnostic characters. — The members of the species belong to the group characterized by dark frons; two equal Ors; brilliantly shining black mesonotum, scu- tellum and pleura; and second costal segment less than three times length of fourth. The members of this species were included in Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen at couplet 61 as Phytomyza sp. (Sehgal). This couplet is amended as below: 61. Orbits normal in width; only fore knees yellowish canadensis Spencer Orbits broad; knees variable from yellow to almost dark; wing base yellow; aedea- gus as in Fig. 116 multifidae n. sp. Description. — Flead. Frons approximately 1.6 times width of eye at level of front ocellus, not projected in front of eye margin in profile. Mouth margin normal; lunule low. Two Ors, directed upwards, equal in size; two Ori, directed inwards, the lower one smaller than upper; orbital setulae few, approximately seven to eight, proclinate. Eyes almost circular, approxi- mately 1.1 times higher than their length, bare; ocellar triangle small. Gena approximately one-third vertical eye height. Third antennal article rounded at tip, with normal pubescence; arista normal, pubescent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr in three to four irregular rows. Wing. Length 1.5- 1.6 mm; costa extended to vein R4+5; costal segments 2-4 in the ratio of 1 : 0.33 : 0.66; crossvein m-m absent; M3 +4 at wing tip. Male genitalia (Fig. 115-117). Hypandrium (Fig. 115) small, V-shaped; pregonites broad; postgonites elongate; surstyli normal; aedeagus (Fig. 116) complex, as illustrated; ejacula- tory apodeme (Fig. 117) small, fan-shaped, bulb small, membranous. Colour. Frons, orbits, gena and lunule dark; ocellar triangle shining black; antennae black; mesonotum, scutellum and pleura shining black; legs black; distal tips of femora in females bright yellow, but in male dark; wing base yellow; squamae and fringe pale; halteres bright 372 Sehgal yellow. Derivation of the specific name. — This species is named after the specific epithet of its food plant Anemone multifida Poir. Biology. — Larvae make linear mines in the leaves of Anemone multifida Poir., family Ranunculaceae. Pupation occurs outside the mine. Geographical distribution. - The members of this species are known only from the type locality: CANADA. Alberta: Holotype 6 Tolman bridge, Red Deer valley (badlands), from leaf mines on Anemone multifida Poir., coll. 14.vi. 1969, emerged 8.vii. 1969; paratypes 2 66 same locality and host, emerged 5.vii. 1969; coll. G. C. D. Griffiths. Phytomyza oxytropidis new species Comparison and diagnostic characters. — The members of this species belong to the group characterized by dark frons; two equal Ors; normal third antennal article; mat greyish black mesonotum and scutellum; two rows of acrostichals; and dark tarsi. The adults resemble the member of P. lupinivora Sehgal from which they may be separated as shown below in exten- sion to Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen. 65. Second costal section short; less than 1 .25 times length of fourth 65a Second costal section longer, at least 1.5 times length of fourth h6 65a. Second costal section approximately 1.25 times length of fourth; orbits dark lupinivora Sehgal Second costal section almost equal to fourth; orbits yellowish oxytropidis n. sp. Description. — Head. Frons approximately twice width of eye at level of front ocellus, slightly projected in front of eye margin in profile. Two Ors equal in size, directed upwards; two Ori, lower one smaller than upper, directed inwards; orbital setulae few, six to seven, proclinate. Eyes oval, approximately 1.2 times higher than their length; ocellar triangle small. Third antennal article rounded at tip, with normal pubescence; arista normal, pubes- cent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr few, three to six scattered hairs. Wing. Length in male 1.6 mm; costa extended to vein R4+5; costal segments 2-4 in the ratio of 1 : 0.5 : 0.93; crossvein m-m absent; M3+4 at wing tip. Male genitalia (Fig. 1 18-120). Hypandrium (Fig. 1 18) with broad side arms and conspicu- ously long apodeme; pregonites broad; postgonites elongate and broad anteriorly; surstyli small, without any big spines; aedeagus (Fig. 1 19) with characteristic long spines between two long, darkly sclerotized arms of basiphallus; distiphallus separated by a small mem- branous section; ejaculatory apodeme (Fig. 120) broad, bulb small. Colour. Frons, gena, lunule and antennae all black; orbits slightly yellowish in most speci- mens; ocellar triangle weakly shining black; legs black; mesonotum, scutellum and pleura mat greyish black. Derivation of the specific name. — This species is named oxytropidis after the generic name of its food plant. Biology. — Larvae make linear mines on the leaflets of Oxytropis splendens Dougl. and O. campestris gracilis (A. Nels), family Leguminosae. Pupation occurs inside the leaf mine. Geographical distribution. - The members of this species are known only from the local- ities of its type specimens as below: CANADA. Alberta: Holotype 6 Jasper, 5 miles south of Athabasca Falls, from leaf mines on Oxytropis splendens Dougl., coll. 15.x. 1967, emerged iv. 1968; paratypes Id, 2 99 same Agromyzidae of Alberta 373 data. Yukon Territory: 1 6 Lake Laberge, from leaf mines on Oxytropis camp estris gracilis (A. Nels.), coll. 9.viii.l968, emerged 20.V.1969, coll. G. C. D. Griffiths. Phytomyza penstemonis Spencer Phytomyza penstemonis Spencer, 1969:265. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum; essentially dark femora and pleura; two equal Ors; acrostichals approximately three to four scattered hairs; and pale squamal fringe. The adults resemble those of P. plantaginis R.-D. and P. colemanensis new species but differ in having second antennal article black and distinct male genitalia. Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — Larvae make linear mines on the leaves of Penstemon confer tus Dougl. and P. procerus Dougl., family Scrophulariaceae. Larvae pupate inside the leaf mine. Geographical distribution. - The members of this species were previously known only from the locality of its type series from western Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 9 Blairmore, from leaf mines on Penstemon confertus Dougl., coll. 26.vi. 1966, emerged 2.vii.l966; 1 6 Coleman, 27.vi. 1966, 2 66, 3 99 Nevis, from leaf mines on Penstemon procerus Dougl., coll. 14.vi.1969, emerged 23-26.vi.1969, coll. G. C. D. Griffiths. Phytomyza periclymeni de Meijere Phytomyza periclymeni de Meijere, 1924: 145. Comparison and diagnostic characters. — The main distinguishing characters of the mem- bers of this species are: dark frons; normal third antennal article; two equal Ors; mat grey mesonotum, scutellum and pleura; dark tarsi; second costal segment less than three times the length of the fourth; and acrostichals in approximately four rows. The adults resemble those of P. caprifoliae Spencer and can be reliably separated only by examination of male genitalia. Spencer ( 1 969) illustrated the aedeagus characteristic of this species. Biology. — In Alberta the larvae mine the leaves of Lonicera involucrata (Richards) Banks, family Caprifoliaceae. Geographical distribution. — The members of this species are known from Europe (Hen- del, 1935) and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 4 66, 2 99 Edmonton, river bank, near University of Alberta campus, from leaf mines on Lonicera involucrata (Richards) Banks, coll. 26.vii.1966, emerged 15-16. viii.1966; 1 6 Elk Island park, 4.vi. 1967; 2 66 St. Albert, near Edmonton, 14.vi. 1966. Phytomyza petasiti Spencer Phytomyza petasiti Spencer, 1969:266. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum; essentially dark femora; and mostly yellow pleura. The adults' resemble those of P. spondylii R.-D. and P. matricariae Hendel and differ in having both Ors equal and distinct male genitalia. The aedeagus of this species has been illustrated by Spencer, Biology. — Larvae make linear mines on the leaves of Petasites sagittatus (Pursh) A. Gray, family Compositae. Pupation occurs outside the mine. 374 Sehgal Geographical distribution. — The members of this species are known only from Canada (Spencer, 1969). 1 examined the following material from Alberta: CANADA. Alberta: 1 9 Devon botanical garden, University of Alberta, from linear mines on leaves of Petasites sagittatus (Pursh) A. Gray, coll. 28.vii.1966, emerged 1 3.iii. 1 967 ; 1 6 Elk Island park, same host, coll. 13.vii. 1968. emerged viii.1968. Phytomyza plantaginis R.-D. Phytomyza plantaginis Robineau-Desvoidy, 185 1 :404. Comparisons and diagnostic characters. - The members of this species belong to the group characterized by yellow frons; two equal Ors; third antennal article elongate and with normal pubescence; essentially dark femora and pleura; and acrostichals usually few isolated hairs. The adults differ from those of the similar species, P. syngenesiae (Hardy), by having pale squamal fringe; shorter second costal segment, approximately 1.5 times length of fourth, and conspicuously yellow fore-coxae. They differ from those of other similar spe- cies, P. penstemonis Spencer and P. colemanensis new species, by having yellow second antennal article and distinct male genitalia. Spencer (1969) illustrated the aedeagus charac- teristic of this species. Biology. — In Alberta the larvae make linear mines in the leaves of Plantago major L., family Plantaginaceae. Pupation occurs inside the leaf mine. Geographical distribution. — The members of this species are Holarctic in distribution, known from Europe, Australia, Japan, U. S. A. and Canada. I examined the following mater- ial from Alberta: CANADA. Alberta: 1 9 Edmonton, river bed near University of Alberta campus, from leaf mines on Plantago major L., 26.vii-5.viii. 1966; 4 99 George Lake near Busby, Univer- sity of Alberta field station, coll. 21.viii. 1966, emerged 25-3 l.viii. 1966. Phytomyza prava Spencer Phytomyza prava Spencer, 1969:269. Comparison and diagnostic characters. — The members of this species are distinctive in having frons which is basically yellow, but is conspicuously darkened. This species has therefore been included in both parts of the key having yellow frons and dark frons. Other diagnostic characters of the members of this species are: dark scutellum; dark femora and pleura and only one Ors. The adults resemble those of P. mertensiae new species, but differ in having yellowish frons and gena, and distinct male genitalia. Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — Larvae make dark blotch mines on the leaves of Anemone canadensis L., family Ranunculaceae. The leaf mine of this species was illustrated among undetermined mines in Fig. 531 (Spencer, 1969). Pupation occurs outside the mine. Geographical distribution. - The members of this species were previously known from Canada only from the locality of its type series (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 2 66, 1 9 Edmonton, White Mud Creek park, from leaf mines on Anemone canadensis L. (Ranunculaceae), coll. 4.ix.l968, emerged 20.ix.1968; 8 66, 9 99 same locality and host, coll. 3.ix.l968, emerged 19-25.ix.1968, 26.V.1969 and 4.vi.l969, coll. G. C. D. Griffiths; 1 6 George Lake near Busby, University of Alberta field station, 21. vi. 1966. Agromyzidae of Alberta 375 Phytomyza queribunda Spencer Phytomyza queribunda Spencer, 1969:271. Comparison and diagnostic characters. — The members of this species belong to the group characterized by dark frons; mat greyish mesonotum and scutellum; two Ors equal; dark tarsi; second costal segment less than three times length of fourth; and acrostichals in four rows. The adults resemble those of P. caprifoliae Spencer and P. periclymeni de Meijere but differ in having frons slightly paler above and entirely different male genitalia. Spencer (1969) illustrated the characteristic aedeagus of this species. Geographical distribution. - The members of this species are known only from the type locality in Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 6 George Lake near Busby, University of Alberta field station, 7.vi. 1968, coll. G. C. D. Griffiths. Phytomyza ranunculi (Schrank) Musca ranunculi Schrank, 1803:140. Phytomyza flavoscutellata Fallen, 1823b:4. Phytomyza albipes Meigen, 1830:195. Phytomyza ranunculi (Schrank); Hendel, 1920:153, 1935:463; Frick, 1952:428, 1959:434; Spencer, 1969:271. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark third antennal article and yellow scutellum. They differ from another species in this group, P. clematiphaga Spencer, by having the upper orbital bristle shorter than lower and distinct male genitalia. The Alberta specimens from Elk Island park, bred from leaf mines on Ranunculus abor- tivus L., correspond in colour to the form albipes Meigen having yellow on mesonotum. One specimen from George Lake corresponds in colour to the form flavoscutellata Fallen having darker mesonotum. The number of coils in the distiphallus vary from one coil in a Faeroes specimen (Grif- fiths, 1964) to eight coils in Alberta specimens. There are five coils in the European speci- men illustrated by Nowakowski (1962). Besides, there is variation in the direction of coils, which in some specimens are coiled upwards, while in others downwards. The number of coils and their direction does not seem to be related to their external colour variations in this species as was pointed out by Griffiths (1964). However, it is possible that more than one species is involved in its entire range. The male genitalia of this species are also very close to that of P. vibeana Griffiths, but the latter differs in having 1 1 coils in the distiphallus and dark mesonotum and scutellum. Biology. — In Alberta the larvae make linear mines in the leaves of Ranunculus abortivus L., family Ranunculaceae. Pupation occurs outside the mine. Geographical distribution. — The members of this species are Holarctic in distribution, known from Europe (Hendel, 1935), United States (Frick, 1959), Faeroes, Iceland and Greenland (Griffiths, 1966), Japan (Sasakaw, 1961) and Canada (Spencer, 1969). I exam- ined the following material from Alberta: CANADA. Alberta: 10 66, 6 99 Elk Island park, from linear mines in the leaves of Ranunculus abortivus L., coll. 22.v. 1969, emerged 5-8.vi. 1969, coll. G. C. D. Griffiths; 1 6 George Lake near Busby, University of Alberta field station, 21.vi.1966. 376 Sehgal Phytomyza riparia new species Comparisons and diagnostic characters. - A member of this species belongs to the group characterized by yellow frons; upper Ors shorter than lower; mouth margin normal; mat greyish mesonotum and scutellum; upper margins of mesopleura, humeral and notopleural areas yellow; essentially dark femora; and second costal segment approximately three and a half times length of fourth. The adult resembles that of P. spondylii R.-D., from which it may be separated as shown below in extension to Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen: 13. Two Ors equal 14 Upper Ors shorter than lower or lacking 13a 13a. Third antennal segment with normal pubescence; upper Ors sometimes lacking; aedeagus as illustrated (Spencer, 1969) spondylii R.-D. Third antennal segment with conspicuously long pubescence; upper Ors present; aedeagus as in Fig. 121 riparia n. sp. Description. — Head. Frons almost twice width of eye at level of front ocellus, only slightly projected in front of eye margin in profile. Mouth margin normal. Two Ors, upper shorter than lower, one side of holotype has only one Ors, directed upwards; two strong Ori, directed inwards; orbital setulae few, six to seven, proclinate. Eyes oval, approximately 1.2 times higher than their length, bare; ocellar triangle small. Gena approximately one-fourth of vertical eye height. Third antennal article rounded at tip, with conspicuously long pubes- cence; arista normal, pubescent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr in almost five irregular rows. Wing. Length in male 2.0 mm; costa extended to vein R4+5;costal segments 2-4 in the ratio of 1 : 0.3 : 0.3; crossvein m-m absent. Male genitalia (Fig. 121-122). Hypandrium with broad side arms and inconspicuous or small apodeme; pregonites broad; postgonites elongate; surstyli normal; aedeagus (Fig. 121) as illustrated with two long characteristic processes in the distiphallus; ejaculatory apodeme (Fig. 122) broad, bulb small. Colour. Frons, orbits and gena yellow; both Vt’s on dark ground; third antennal article dark brown; upper parts of mesopleura, humeral and notopleural areas yellow; mesonotum and scutellum mat greyish, slightly paler; coxae black; femora dark brown, with yellow on distal tips; tibiae and tarsi yellowish brown; squamae yellow, fringe brown; halteres yellow. Derivation of the specific name. — This species is named riparia as its holotype was col- lected along the Saskatchewan River bank. Geographical distribution. — The members of this species are known only from the type locality: CANADA. Alberta: Holotype 6 Edmonton, Saskatchewan River bank near University of Alberta campus, 20.vii.1966. Phytomyza sehgali Spencer Phytomyza sehgali Spencer, 1969:274. Comparisons and diagnostic characters. — The members of this species differ from those of mertensiae new species in having paler frons and from prava Spencer in having strong acrostichals, small third antennal article and distinct male genitalia. Spencer (1969) illustra- ted the aedeagus characteristic of this species. Geographical distribution. — Known only from Alberta, Canada from the following local- ity: Agromyzidae of Alberta 377 CANADA. Alberta: Edmonton, White Mud Creek (Spencer, 1969). Phytomyza senecionella new species Comparison and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; two Ors equal; dark scutellum; essentially dark femora and pleura; normal mouth margin and third antennal article; acrostichals only two to three scat- tered hairs; dark squamal fringe and fore-coxae. The adults resemble those of the similar species, P. syngenesiae (Hardy), and males of the two can be separated reliably only by examination of the genitalia. This species is included in Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen as below: 40. Second costal section short; 1.5 times length of fourth; third antennal segment enlarged; squamal fringe pale 41 Second costal section longer, approximately twice length of fourth; third antennal segment normal; squamal fringe dark 40a 40a. Aedeagus as illustrated (Spencer, 1969) syngenesiae (Hardy) Aedeagus as in Fig. 123. senecionella n. sp. Description. — Head. Frons wider than width of eye at level of front ocellus (1 : 0.55), very slightly projected in front of eye margin in profile. Two Ors equal, directed upwards; one strong Ori and one small hair present below directed inwards; orbital setulae few, six to seven, proclinate. Eyes almost circular, their vertical height being almost equal to then- length, bare; ocellar triangle small. Gena deep, approximately two-fifths of eye height. Antennal bases approximate; third antennal article rounded at tip, with normal pubescence; arista normal and pubescent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr two to three scattered hairs. Wing. Length in male 2.75 mm; costa extended to vein K4+5 ; costal segments 2-4 in ratio of 1 : 0.3 : 0.56; crossvein m-m absent; vein M3+4 at wing tip. Male genitalia. (Fig. 123-124). Hypandrium with side arms broad and no conspicuous apodeme; pregonites broad; postgonite with small hook anteriorly; surstyli small and nor- mal; aedeagus (Fig. 123) as illustrated; ejaculatory apodeme (Fig. 124) slightly broad, bulb small and membranous. Colour. Frons, orbits and gena yellow; both Vt’s on dark grounds; all antennal articles black; ocellar triangle shining black; mesonotum, sctuellum and pleura mat grey; legs black, only tips of femora with slight yellow, coxae black; squamae yellow, fringe dark; halteres yellow. Derivation of the specific name. — This species is named after the generic name of its food plant. Biology. — Larvae make broad linear mines on the leaves of Senecio conges tus var. palustris (L.), family Compositae. The leaf mines were more or less communal with more than one larva feeding in them. Pupation occurs usually at the leaf bases or sometimes on the stem. Geographical distribution. — The members of this species are known only from the type locality: CANADA. Alberta: Holotype 6 Elk Island park, from leaf mines on Senecio congestus palustris (L.), coll. 2.vii.l969, emerged 6.vii.l969; paratypes 2 66 same locality and host, emerged 6-1 2.vii. 1969, coll. G. C. D. Griffiths. 378 Sehgal Phytomyza solidaginivora Spencer Phytomyza solidaginivora Spencer, 1969:274. Comparison and diagnostic characters. - The members of this species differ from those of similar species, matricariae Hendel, in having dark second antennal article and distinct male genitalia. Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — Larvae make linear leaf mines on Solidago, family Compositae. Geographical distribution. — Known only from Alberta, Canada from the following local- ity: CANADA. Alberta: Edmonton, University of Alberta campus (Spencer, 1969). Phytomyza solidaginophaga new species Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; two equal Ors; normal third antennal article; dark scutellum; mostly dark femora and pleura; and three to six rows of acrostichals. The adults resemble those of P. aquilegiana Frost and P. aquilegioides new species but differ in having the frons slightly darkened below and distinct male genitalia. They also resemble those of another similar species, P. ilicis Curtis, and may be separated as shown below in extension to Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen: 32. Frons distinctly darkened, either above or below 32a Frons entirely pale, yellow or orange, at most orbits dark 33 32a. Aedeagus as illustrated (Spencer, 1969); larva leaf-miner in Ilex ilicis Curtis Aedeagus as in Fig. 126; larva leaf-miner in Solidago solidaginophaga n. sp. Description. — Head. Frons approximately twice eye width at level of front ocellus, slightly projected in front of eye margin in profile. Mouth margin normal; lunule low. Two equal Ors, directed upwards (one specimen has only one Ors, but has two bristles in the same socket as upper Ori; two Ors have therefore been considered as normal for the mem- bers of this species). Two Ori, directed inwards, lower one weaker than upper; orbital setu- lae few, approximately seven, proclinate. Eyes oval, approximately 1.2 times higher than their length, bare; ocellar triangle small. Gena approximately 0.28 times vertical height of eye. Third antennal article rounded at tip, with normal pubescence; arista normal, pubescent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr in approximately four irregular rows. Wing. Length in male approximately 2.1 mm; costa extended to vein R4+5; costal seg- ments 2-4 in ratio of 1 : 0.26 : 0.3; crossvein m-m absent. Male genitalia (Fig. 125-127). Hypandrium (Fig. 125) V-shaped, with broad side arms; pregonites broad; postgonites elongate; surstyli normal; aedeagus (Fig. 126) with charac- teristic row of small spines between two long arms of basiphallus, as illustrated; ejaculatory apodeme (Fig. 127) small, bulb small and membranous. Colour. Frons yellow, slightly darkened just above lunule; orbits slightly darkened along eye margins; lunule and gena darkened; ocellar triangle weakly shining black; both Vt’s on dark ground; antennae black; mesonotum, scutellum and pleura mat greyish black; femora black, with yellow distal tips; tibiae and tarsi dark brown; squamae yellow, fringe brown; halteres yellow. Derivation of the specific name. — This species has been named after the generic name of its food plant. Biology. — Larvae make linear mines in the leaves of Solidago lepida DC, family Com- positae. Pupation occurs outside the mine. Geographical distribution. — The members of this species are only known from the type Agromyzidae of Alberta 379 locality: CANADA. Alberta: Holotype 6 George Lake near Busby, University of Alberta field station, from mines on the leaves of Solidago lepida DC, coll. 7.vi. 1968, emerged 30.iv.1969, coll. G. C. D. Griffiths; paratype 1 6 same data. Phytomyza spondylii R.-D. Phytomyza spondylii Robineau-Desvoidy, 1851: 147. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum; and essentially dark femora. The colour of the upper margins of the mesopleura, humeral and notopleural areas is variable from yellow to almost dark. A small upper Ors is usually present. The adults having yellow on the sides resemble those of P. matricariae Hendel and differ in having the second costal section longer, approximately 3.5 times the fourth and dark second antennal article. The darker forms resemble those of P. asterophaga Spencer but differ in having entirely differ- ent male genitalia. Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — Larvae make linear mines in the leaves of Heracleum lanatum Michx., family Umbelliferae. Pupation occurs outside the mine. Geographical distribution. — The members of this species are known from Europe (Hen- del, 1935) and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 2 <5d, 2 99 Edmonton, White Mud Creek park, from leaf mines on Heracleum lanatum Michx., emerged 18-19.vii.1966. Phytomyza subalpina new species Comparisons and diagnostic characters. — A member of this species belongs to the group characterize^ by yellow frons; normal mouth margin and third antennal article; two Ors equal; dark scutellum; essentially dark femora and pleura; acrostichals in two rows; and dark fore-coxae. The adult resembles that of P. fuscula Zetterstedt but differs in having a pale squamal fringe and distinct male genitalia. It differs from another similar species, P. atripal- pis Aldrich, as shown below in extension to Spencer’s (1969) key to Canadian species of the genus Phytomyza Fallen: 53. Second costal section short, less than twice length of fourth 53a Second costal section longer, twice length of fourth. gelida Spencer 53a. Gena two-fifths eye height atripalpis Aldrich Gena one-fifth eye height subalpina n. sp. Description. — Head. Frons approximately twice width of eye at level of front ocellus; mouth margin normal; lunule low. Two equal Ors directed upwards; two Ori, lower one weaker, both directed inwards; orbital setulae four to five, proclinate. Eyes oval, their verti- cal height being approximately 1.3 times their length, bare; ocellar triangle small. Gena approximately one-fifth vertical eye height. Third antennal article rounded at tip, with normal pubescence; arista normal, pubescent. Mesonotum. Dorsocentrals 3+1 strong bristles; acr approximately nine hairs, in two rows. Wing. Length in male approximately 2.1 mm; costa extended to vein R4+5; costal seg- ments 2-4 in ratio of 1 : 0.35 : 0.65; crossvein m-m absent. Male genitalia (Fig. 128-132). Hypandrium (Fig. 128) U-shaped with broad side arms; pregonites broad; postgonites (Fig. 129) elongate, with hook-like process anteriorly; surstyli normal; aedeagus (Fig. 130, 131) as illustrated; ejaculatory apodeme (Fig. 132) small, bulb membranous. 380 Sehgal Colour. Frons, gena and lunule yellow; orbits yellow, slightly darkened near upper Ors; ocellar triangle weakly shining black; both Vt’s on dark ground; antennae black; mesono- tum, scutellum and pleura mat grey; femora, tibiae and tarsi black; squamal fringe dirty pale, squamae pale; halteres yellow. Derivation of the specific name. - This species is named subalpina as its holotype was collected in the subalpine zone of foothills of Rocky Mountains. Geographical distribution. - This species is known only from the type locality: CANADA. Alberta: Holotype 6 Coleman, 27.vi.1966. Phytomyza subtenella Frost Phytomyza subtenella Frost, 1924:89. Comparison and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; normal mouth margin and third antennal article; usually two equal Ors (but one specimen has three equal Ors); essentially dark femora and pleura; acros- tichals in two well-defined rows and yellow fore-coxae. The adults, wing length approximately 2.5 mm, differ from those of the similar species, P. jasperensis, in having the frons strongly projected in front of eye margin; gena deeper, approximately two-thirds of eye height and distinct male genitalia. The paraphalli in Alberta specimen were independent of the basiphallus and not joined as it appears from Spencer’s (1969) illustration in which they are overlapping. Geographical distribution. — The members of this species are known from the United States (Frick, 1959) and Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 1 <5 Banff, 28.vi.1966, 5 66 Hinton, 23.vii.1967; 1 6 Jasper, 17.vi. 1966. Phytomyza subtilis Spencer Phytomyza sub tilis Spencer, 1969:276. Comparison and diagnostic characters. — The members of this species differ from those of a very similar species, urbana Spencer, in having darker grey mesonotum and slightly brown- ish frons. The male genitalia as figured by Spencer (1969) are, however, very distinct. Biology. - Larvae make blotch mines on the leaves of Lathy rus ochroleucus Hook., family Leguminosae. Geographical distribution. - Known from Alaska and Alberta. The Alberta locality is as follows: CANADA. Alberta: Wabamun Lake (Spencer, 1969). Phytomyza syngenesiae (Hardy) Chromatomyia syngenesiae Hardy, 1849:391. Phytomyza chrysanthemi Kowarz, 1891:243; Smulyan, 1914:21. Phytomyza atricornis Meigen sensu Hendel,1920: 162 (in part); Frost, 1924:68; Frick, 1952: 424, 1959:425 ( nomen dubium). Phytomyza syngenesiae (Hardy); Griffiths, 1967:7; Spencer, 1969:278. Comparisons and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum; mostly dark femora and pleura; two Ors equal; normal third antennal article; and acrostichals normally lacking or at most three Agromyzidae of Alberta 381 to four isolated hairs present. The adults resemble closely those of an Old World species, P. horticola Goureau, and can be separated reliably only by examination of the male genitalia. Griffiths (1967) and Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — Larvae make linear mines in the leaves of numerous Compositae and rarely on non-Compositae hosts (Griffiths, 1967). In Alberta the flies have been bred from only two host-plants of the family Compositae, Senecio sp. (Spencer, 1969) and Crepis gracilis (D.C. Eat.) Rydb. Geographical distribution. — The members of this species are widespread in Europe, Australia, New Zealand, U. S. A. and Canada (Griffiths, 1967). I examined the following material from Alberta: CANADA. Alberta: 1 <3, 2 99 Edmonton, University of Alberta campus, from leaf mines on Crepis gracilis (D.C. Eat.) Rydb., coll. 4.vi. 1966. Phytomyza thalictrivora Spencer Phytomyza thalictrivora Spencer, 1969:279. Comparison and diagnostic characters. — The members of this species belong to the group characterized by dark frons, mesonotum and scutellum; essentially dark femora and pleura; upper Ors lacking; and second costal segment less than three times length of fourth. The adults resemble those of P. aquilegivora Spencer but differ in having darker frons and few acrostichals. They also resemble those of P. minuscula Goureau, but possess distinct male genitalia. Spencer (1969) illustrated the aedeagus characteristic of this species. Biology. — Larvae make linear mines on the leaves of Thalictrum venulosum Trek, family Ranunculaceae. Pupation occurs outside the mine. Geographical distribution. — The members of this species are known only from Canada (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta 1 6 Edmonton, White Mud Creek park, from leaf mines on Thalictrum venulosum Trek, coll. 12.vi.1966, emerged 17.vi. 1966; 1 6 Edmonton, Rainbow Valley, 31.V.1969. Phytomyza timida Spencer Phytomyza timida Spencer, 1969:279. Comparison and diagnostic characters. — The members of this species belong to the group characterized by yellow frons; dark scutellum; and essentially dark femora and pleura. Spencer (1969) includes this species among those having upper orbital bristle shorter than lower. This character is probably variable as the Alberta specimens examined had both upper orbital bristles almost equal. The aedeagus of a specimen from Banff, Alberta is illus- trated (Fig. 133) and agrees with that figured by Spencer (1969). Geographical distribution. — The members of this species are known from Canada only from the locality of its type series (Spencer, 1969). I examined the following material from Alberta: CANADA. Alberta: 2 66 Banff, 18.vi.1966; 1 6 Jasper, 16.vi.1966. Phytomyza urbana Spencer Phytomyza urbana Spencer, 1969:281. Comparison and diagnostic characters. — The members of this species differ from those of a similar species, subtilis Spencer, in having paler greyish mesonotum and paler frons and 382 Sehgal distinct male genitalia. Spencer (1969) illustrated the aedeagus characteristic of this species. Geographical distribution. — Known only from Alberta, Canada from the following locality: CANADA. Alberta: Blairmore (Spencer, 1969). Phytomyza sp. {Angelica arguta Nutt.) Comparison and diagnostic characters. — The distinguishing characters of the female of this species are: yellowish brown frons; wing length 2.5 mm; second costal segment three times length of fourth; dark squamal fringe; dark mesonotum, scutellum, pleura and femora and acrostichals in two to three rows. It resembles the adults of P. aralivora Spencer but has entirely different biology. This species cannot be definitely determined at present as no males are available for examination. Biology. — Larvae make linear mines in the leaves of Angelica arguta Nutt., family Umbelliferae. Geographical distribution. — The members of this species were examined only from southwestern Alberta as below: CANADA. Alberta: 1 9 Blairmore, from leaf mines on Angelica arguta Nutt., coll. 5.ix. 1966, emerged 1 0.iii. 1 967 ; numerous leaf mines, same locality. INSECT HOST-PLANT RELATIONSHIP IN THE FAMILY AGROMYZIDAE The members of the family Agromyzidae are exclusively internal plant feeders during their larval stage. Larval feeding results in a definite pattern called the mine, and the study of the mining habits is called minology or hyponomology (Hering, 1951). The agromyzid mines fall into two general categories. First, the epidermal leaf mines, in which the mining larva feeds only inside the epidermal layer of the leaf. These are restricted mainly to the old world tropics. The second, the parenchymal mine, in which the larva feeds on the parenchymatous tissue inside the leaf or other part of the plant. The majority of mines belong in this category. Leaf mines are usually seen externally and generally are more visible from one side than the other. Surface mines on other parts of a plant can also be detected as whitish or greenish channels with faecal granules distributed in definite tracks. Mines inside the stem or root are not easily detected, however, the injury caused by the mining larva can be seen by breaking the injured plant part. The shape of the larval mine is usually constant within the species, but varies between species. This helps greatly in the separation of closely related species which cannot otherwise be reliably identified by adult morphology. Hering (1951) dealt with shapes of mines made by mining insects and later, in 1957, illustrated the mines of European species in detail. The relationship between the endophagous larvae of mining insects and their food plants is typically parasitic in nature. However, modern parasitology as a science does not concern itself with the study of such relationships. Nowakowski (1962) commenting on this situa- tion proposed the term “zoophytoparasitology” for the study of animals as parasites and plants as hosts. The most important features of this relationship in the family Agromyzidae are the active choice of host-plant by the mining insect, the varying degree of host-plant specificity, and the adaptations of the maggot for an endoparasitic life in the semi-liquid environment of the leaf parenchyma. The understanding of this relationship is of great im- portance in dealing with the systematics of this group as it permits the use of the host- parasite discrimination method. It provides valuable information for identification of similar species, which cannot be separated on adult morphological characteristics alone. Agromyzidae of Alberta 383 Hering (1951) discussed the distribution of leaf mining species on plants of various families and examined the established phylogenetic relationships between the plant families in a system based on serum diagnosis alone as proposed by Mez (1926). Mez’s system of classification has now been criticized by modern botanists because of similar serum reac- tions obtained for certain plant families which clearly are not closely related. Table 2 lists the known local host-plants for the Albertan species. It is realized that the information on host-plants is not adequate for all species, but some useful observations can still be made. The arrangement of plant families is after the supposedly phylogenetic system of Takhtajan (1969). The phylogenetic relationships between plant families and orders is still a matter of controversy. Most plant classifications fall into two groups depending on the supposed nature of the primitive angiosperm flower (Davis and Heywood, 1965). One group is based on the assumption that the earliest angiosperms were wind pollinated and that the monocotyledons and dicotyledons have arisen independently from unknown gymnosperms. According to the second group of systems, dicotyledons and monocotyledons were both derived from primitive angiosperms which were insect-pollinated. Such a view is supported by many recent workers (Eames, 1961: Hutchinson, 1964; Takhtajan, 1969). Hutchinson (1964) in his revised edition of classification of angiosperms maintained a basic division of dicotyledons into woody “Lignosae” and herbaceous “Herbaceae,” a system which allegedly leads to wide separation of certain plant families which are markedly similar in the structure of their flowers. In the absence of established phylogenies of both angiosperms and agro- myzid parasites, it is difficult to study the trends in their coevolution. It is generally accepted that the larvae of Cyclorrhapha were primitively saprophagous from which various specialized feeding habits like phytophagy, carnivory and parasitism have been derived (Hennig, 1952). The dominance of agromyzids on angiosperm hosts suggest that angiosperms were well established as flowering plants before the agromyzids made their appearance. Although opinions differ as to precisely when angiosperms first appeared on the evolutionary scene, there is a general agreement that they came into prominence suddenly during the late Cretaceous (Eames, 1961). Hennig (1965) reviewed all supposed records of agromyzid fossils and concluded that the family Agromyzidae is not yet known to be represented in the Tertiary baltic amber. In fact there are so far no fossils which can be definitely referred to the family. The occurrence of a large number of closely related and poorly differentiated species and their abundance on hosts belonging to highly evolved plant families suggests that much of the diversification of the family Agromyzidae is relatively recent. However, the possibility that the group goes back to late Cretaceous as suggested by Nowakowski (1962) must be admitted. The recent use of male genitalia in agromyzid taxonomy has split many groups originally supposed to be polyphagous or oligophagous species, into species with much narrower host- plant specificity. It is becoming increasingly apparent that the majority of agromyzid species are restricted feeders, being monophagous or oligophagous. Strict monophagy also appears to be rare unless it results from a plant genus being monotypic. Nowakowski (1962) dis- cussed the subject of host-plant specificity among the European species and revealed many examples where the original wide host range was found to be the result of misidentifications or assemblage of many species under the same name. The most polyphagous species, Phyto- myza syngenesiae (Hardy), appears to be a restricted feeder in Alberta and has been bred from only two plant genera Crepis and Senecio of the family Compositae. This species is known from many Compositae and rarely from other host-plants (see Frick, 1959 asPhyto- myza atricornis Meigen; Griffiths, 1967). Most agromyzid genera occurring on species of monocot families are also represented on 384 Sehgal various dicotyledons. The few exceptions are the species of the genus Cerodontha Rondani occurring only on monocot families Graminiae, Cyperaceae, and Juncaceae; and the mem- bers of the ambigua/nigripes groups of the genus Agromyza Fallen which feed only on grasses. The species in these groups are uniform in external morphology of adults and in the general shape of the male genitalia, and probably represent early specialization of their feeding habit. Various species of the agromyzid genera Liriomyza Mik and Phytomyza Fallen feeding on grasses will probably prove to be oligophagous. The oligophagy of various grass-mining species has not been investigated because of the problems of identification of grasses at the time the mining larvae are collected. The family Ranunculaceae is selected by members of the agromyzid genera Melanagro- myza Hendel, Ophiomyia Braxchnikov and Phytomyza Fallen. Melanagromyza actaeae n. sp. feeding inside the stems o iActaea rubra (Ait.) Willd and an Ophiomyis sp. making surface mines below the stem epidermis of Thalictrum venulosum Trel. appear to be specialized monophagous species. There are many closely related and poorly differentiated Phytomyza species feeding on the plant genera Aquilegia and Thalictrum, some of which are oligopha- gous species feeding on both. The species of the genus Phytomyza Fallen feeding on the plant genera Clematis, Delphinium and Ranunculus are specialized monophagous species. Three local species of the plant genus Anemone support three different leaf miners of the genus Phytomyza Fallen. The members of the agromyzid genera Agromyza Fallen, Melana- gromyza Hendel and Hexomyza Enderlein feeding on Ulmaceae, Urticaceae and Salicaceae are all specific feeders. Two species known to have rosaceous host-plants in Europe and the United States have not yet been discovered on Alberta hosts. One is a specific cambium miner, Phytobia amelanchieris (Greene), feeding on Amelanchier canadensis (L.) (Frick, 1959), and the other Agromyza spiraceae Kaltenbach, an oligophagous species feeding on various genera of the subfamily Rosoideae in Europe. Members of the family Leguminosae are fed on by the representatives of the agromyzid genera Agromyza Fallen, Liriomyza Mik and Phytomyza Fallen. Most of these species are monophagous in Alberta, with the excep- tion of Liriomyza fricki Spencer which is oligophagous. The plant families Cornaceae, Arali- aceae, Umbellifereae and Elaeagnaceae are fed on by specific feeders of the agromyzid genera Melanagromyza Hendel, Phytomyza Fallen and Amauromyza Hendel. Members of the plant family Caprifoliaceae suppdft oligophagous species belonging to the agromyzid genera Paraphytomyza Enderlein and Phytomyza Fallen. The oligophagous species feed on the plant genera Lonicera and Symphoricarpos . The plant families Boraginaceae, Scrophula- riaceae; Plantaginaceae and Labiatae have specialized specific feeders. The family Composi- tae supports a highly specialized agromyzid fauna belonging to the genera Melanagromyza Hendel, Ophiomyia Braschnikov, Liriomyza Mik, Caly corny za Hendel, Nemorimyza Frey and Phytomyza Fallen. Most of these species are specific monophagous feeders. However, some oligophagous species feed on the plant genera Crepis, Taraxacum, and Sonchus', others feed upon members of tribe Anthemideae of the family Compositae, as shown by the host range of Phytomyza matricariae Hendel and Liriomyza millefolii Hering. Agromyzidae of Alberta 385 Table 2. Albertan host-plant records of Albertan agromyzid species. DICOTYLEDONS Family Ranunculaceae Actaea rubra (Ait.) Willd Anemone canadensis L. Anemone multifida Poir. Anemone riparia Fern. Aquilegia sp. (cultivated var.) Clematis verticillaris DC Delphinium sp. (cultivated) Ranunculus abortivus L. Thalictrum venulosum Trel. Thalictrum sp. Melanagromyza actaeae n. sp. Phytomyza prava Spencer Phytomyza multi fidae n. sp. Phytomyza canadensis Spencer Phytomyza aquilegiana Frost, P. aquilegio- phaga Spencer, P. aquilegivora Spencer, P. columbinae n. sp. Phytomyza clematiphaga Spencer Phytomyza delphinivora Spencer Phytomyza ranunculi (Schrank) Phytomyza aquilegioides n. sp .,P. columbinae n. sp.,P. thalictrivora Spencer, Ophiomyia sp. Phytomyza aquilegiana Frost Family Ulmaceae Ulmus americana L. Agromyza aristata Malloch Family Urticaceae Urtica gracilis Ait. Melanagromyza martini Spencer, Phytomyza sp., luteiceps n. sp. Family Salicaceae Populus tremuloides Michx. Agromyza populoides Spencer, Hexomyza schineri (Giraud), Paraphytomyza sp. Family Rosaceae Potentilla sp. Agromyza sp. Family Leguminosae Lathyrus ochroleucus Hook. Lupinus sericeus Pursh Oxytropis splendens Dougl. Tri folium repens L. Vicia americana Muhl. Liriomyza fricki Spencer, Liriomyza lathyri n. sp., Phytomyza subtilis Spencer Phytomyza lupini Sehgal, P. lupinivora Sehgal Phytomyza oxytropidis n. sp. Liriomyza fricki Spencer Liriomyza fricki Spencer, L. viciae Spencer Family Cornaceae Cornus canadensis L. Cornus stolonifera Michx. Phytomyza agromyzina Meigen Phytomyza agromyzina Meigen Family Araliaceae Aralia nudicaulis L. Phytomyza aralivora Spencer 386 Sehgal Family Umbellifereae Angelica arguta Nutt. Heracleum lanatum Michx. Phytomyza sp. Phytomyza spondylii R.-D. Family Elaeagnaceae Shepherdia canadensis (L.) Amauromyza shepherdiae n. sp. Family Caprifoliaceae Lonicera dioica L. Lonicera involucrata (Richards) Lonicera tartarica L. Symphoricarpos albus (L.) Symphoricarpos occidentalis Hook. Symphoricarpos sp. Paraphytomyza lonicerae (R.-D.), P. orbitalis (Melander), P. spenceri n. sp. Paraphytomyza plagiata (Melander), Phytomyza gregaria Frick, P. periclymeni de Meijere Paraphytomyza lonicerae (R.-D.) Paraphytomyza lonicerae (R.-D.), P. orbitalis (Melander) Paraphytomyza spenceri n. sp. Phytomyza capri foliae Spencer Family Rubiaceae Galium boreale L. Praspedomyza galiivora Spencer Family Boraginaceae Mertensia paniculata (Ait.) Agromyza canadensis Malloch, Phytomyza mertensiae n. sp. Family Scrophulariaceae Penstemon confer tus Dougl. Penstemon procerus Dougl. Veronica sp. (cultivated) Phytomyza penstemonis Spencer Phytomyza penstemonis Spencer Phytomyza crassiseta Zetterstedt Family Plantaginaceae Plant ago major L. Phytomyza plantaginis R.-D. Family Labiatae Mentha arvensis L. Calycomyza menthae Spencer Family Compositae Achillea milli folium L. Achillea sibirica Ledeb. Arnica cordifolia Hook Aster ciliolatus Lindl. Aster conspicuus Lindl. Bidens cernua L. Chrysanthemum sp. (cult.) Crepis gracilis (D.C. Eat.) Rydb. Crepis tectorum L. Phytomyza matricariae Hendel Melanagromyza achilleana n. sp., Liriomyza millefolii Hering, Phytomyza maticariae Hendel Phytomyza amicivora n. sp. Phytomyza ciliolati Spencer Phytomyza asterophaga Spencer Melanagromyza bidenticola n. sp. Phytomyza matricariae Hendel Phytomyza syngenesiae (Hardy) Phytomyza lactuca Frost Agromyzidae of Alberta 387 Matricaria matricarioides (Less.) Porter Petasites sagittatus (Pursh) Senecio conges tus palustris (L.) Senecio pauciflorus Pursh Senecio sp. Solidago lepida DC Solidago sp. Sonchus uliginosus Bieb. Sonchus sp. Tanacetum vulgare L. Taraxacum sp. MONOCOTYLEDONS Family Liliaceae Smilacina stellata (L.) Maianthemum canadense Desf. Family Cyperaceae Scirpus sp. Family Gramineae Agropyron repens (L.) Beauv. Agropyron smithii Rydb. Deschampsia caespitosa (L.) Phalaris arundinacea L. Triticum aestivum L. Phytomyza matricarae Hendel Phytomyza petasiti Spencer Phytomyza senecionella n. sp. Liriomyza senecionivora n. sp. Phytomyza syngenesiae (Hardy) Phytomyza solidaginophaga n. sp. Ophiomyia maura (Meigen), Calycomyza ? solidaginis (Kaltenbach), Nemorimyza posticata (Meigen) Phytomyza lactuca Frost Calycomyza sonchi Spencer Liriomyza millefolii Hering, Phytomyza lactuca Frost Calycomyza sonchi Spencer Liriomyza smilacinae Spencer Liriomyza sp. Cerodontha (Dizygomyza) ? scirpi (Karl) Cerodontha (Poemyza) incisa (Meigen) Cerodontha (Poemyza) incisa (Meigen) Liriomyza cordillerana Sehgal Cerodontha (Poemyza) incisa (Meigen) Cerodontha (Poemyza) superciliosa (Zetterstedt) ACKNOWLEDGEMENTS I am grateful to B. Hocking, Department of Entomology, University of Alberta, for providing the opportunity and support for this project in Alberta, Canada. I also express my most sincere thanks to him for his criticism of the manuscript and keen interest throughout this study. I am grateful to G. E. Ball, Department of Entomology, University of Alberta, for his ever available help, supervision and valuable criticism of the manuscript. I am also grateful to K. A. Spencer, London, England and G. C. D. Griffiths, Department of Ento- mology, University of Alberta, for numerous useful discussions and valuable suggestions during this study. I would like to thank J. G. Packer, Department of Botany, University of Alberta, for help in identification of host-plants; and J. Belicek for help in translating the abstract into German. 388 Sehgal REFERENCES Braschnikov, W. C. 1897. Zur Biologie und Systematik eininger Arten minierender Dipteren. Izv. mosk. sel’.-khoz Inst. 3: 19-43. Coquillett, D. W. 1902. New acalyptrate Diptera from North America. J. N. Y. ent. Soc. 10:177-191. Davis, P. H. and V. H. Hey wood. 1965. Principles of Angiosperm Taxonomy. Oliver and Boyd. 558 pp. Eames, A. J. 1961. Morphology of the Angiosperms. McGraw-Hill, New York. Enderlein, G. 1936. Zweiflugler, Diptera. In Brohmer et. al., Tierwelt Mitteleur, 6 (Lief. 2, Insekten, 3), 179-183. Fallen, C. F. 1810. Specimen entomologicum novam Diptera disponendi methodum exhi- bens. 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Notes on the Oriental Agromyzidae - 2. Agromyzidae from the Philippines. Ent. Meddr 34:3-9. Spencer, K. A. 1965b. Agromyzidae, in Diptera from Nepal. Bull. Br. Mus. nat. Hist. Ent. 1 6( 1 ): 25-3 1 . Agromyzidae of Alberta 391 Spencer, K. A. 1965c. A clarification of the status of Liriomyza trifolii (Burgess) and some related species (Diptera: Agromyzidae). Proc. ent. Soc. Wash. 67(1): 32-40. Spencer, K. A. 1966a. A revision of European species of the genera Melanagromyza Hendel and Hexomyza Enderlein, with a supplement on the genus Ophiomyia Braschnikov. Beitr. Ent. 16:3-60. Spencer, K. A. 1966b. Notes on the Neotropical Agromyzidae (Diptera). -1. Papeis Dep. Zool. S. Paulo 19:141-150. Spencer, K. A. 1966c. Notes on European Agromyzidae — 1. Beitr. Ent. 16:285-309. Spencer, K. A. 1969. The Agromyzidae of Canada and Alaska. Mem. ent. Soc. Can., 64: 311 pp. Stegmaier, C. E. 1966. Host plants and parasites of Liriomyza rnunda in Florida (Diptera: Agromyzidae) Fla Entomol. 49:81-86. Stegmaier, C. E. 1968. A review of recent literature on the host plant range of the genus Liriomyza Mik (Diptera: Agromyzidae) in the continental United States and Hawaii, excluding Alaska. Fla Entomol. 51:167-182. Strickland, E. H. 1938. An annotated list of the Diptera (flies) of Alberta, Canada. J. Res., D. 16:175-219. Strickland, E. H. 1946. An annotated list of the Diptera (flies) of Alberta. Additions and corrections. Canad. J. Res. D. 24: 157-173. Strobl, G. 1898. Die Dipteren von Steiermark IV. Theil. Nachtrage. Mitt, naturw. Ver. Steierm. 34:192-298. Takhtajan, A. 1969. Flowering plants, origin and dispersal. Oliver and Boyd. Edinburgh 205:239 pp. Westwood, J. O. 1840. An Introduction to the modern Classification of Insects. II. Synopsis of the genera of British Insects. London, 587 pp. Zetterstedt, J. W. 1848. Diptera Scandinaviae, Lund. 7:2728-2844. Zetterstedt, J. W. 1860. Diptera Scandinaviae, Lund. 14:6449-6471. 392 Sehgal Fig. 1-2. Agromyza albipennis. 1. aedeagus, ventral view. 2. ejaculatory apodeme. Fig. 3-5. A. brevispinata. 3. aedeagus, ventral view. 4. distiphallus, lateral view. 5. surstylus. Fig. 6. A. hockingi, surstylus. Fig. 7. A. ? hockingi, surstylus. Fig. 8-9. A. kincaidi. 8. aedeagus, ventral view. 9. aedeagus, lateral view. Fig. 10-12. A. nearctica. 10. aedeagus, lateral view. 11. ejaculatory apodeme. 12. hypandrium. Agromyzidae of Alberta 393 F* 13-20. Melamgr°myza achilleana. 13. head, lateral view. 14. aedeagus, lateral view. 15. ejaculatory apodeme pt \ SUIS£ u i8' Cephal0pharyngeal skeleton of larva- 19- anterior spiracle. 20 posterior spiracles' g. 22. M. actaeae. 21. head, lateral view. 22. cephalopharyngeal skeleton of larva. 394 Sehgal Fig. 23-25. Melanagromyza actaeae. 23. muscle scars and tubercle band from lateral portion of first abdominal segment of larva. 24. anterior spiracle. 25. posterior spiracles. Fig. 26-31. M. bidenticola. 26. head, lateral view. 27. aedeagus, lateral view. 28. hypandrium. 29. cephalopharyngeal skeleton of larva. 30. anterior spiracle. 31. posterior spiracles. Fig. 32. Melanagromyza sp. ?, aedeagus, lateral view. Agromyzidae of Alberta 395 Fig. 33. Ophiomyia labiatarium, aedeagus, lateral view. Fig. 34. O. maura, aedeagus, lateral view. Fig. 35. O. pulicaria, aedeagus, lateral view. Fig. 36—39. O. stncklandi. 36. head, lateral view. 37. aedeagus, lateral view. 38. ejaculatory apodeme. 39. hypandrium. Fig. 40. Cerodontha ? occidentalis, aedeagus, lateral view. 396 Sehgal Fig. 41-42. Amauromyza riparia. 41. aedeagus, lateral view. 42. ejaculatory apodeme. Fig. 43-46. A. shepherdiae. 43. aedeagus, lateral view. 44. distiphallus, ventral view. 45. ejaculatory apodeme. 46. leaf mine on Shepherdia canadensis OL.) Nutt. Fig. 47-50. Liriomyza balcanicoides. 47. aedeagus, lateral view. 48. aedeagus, ventral view. 49. ejaculatory apodeme. 50. surstylus. Agromyzidae of Alberta 397 ig. 51 54. Linomyza bifurcata. 51. head, lateral view. 52. aedeagus, lateral view. 53. aedeagus, ventral view 54 ejaculatory apodeme. Fig. 55-56. L. eupatorii. 55. aedeagus, lateral view. 56. aedeagus, ventral view. Fig. 57-59. L. latnyn. 57. aedeagus, lateral view. 58. aedeagus, ventral view. 59. ejaculatory apodeme. 0. 25 MM 398 Sehgal Fig. 60-63. Liriomyza senecionivora. 60. aedeagus, lateral view. 61. aedeagus, ventral view. 62. ejaculatory apodeme. 63. surstylus. Fig. 64-67. L. sinuata. 64. aedeagus, lateral view. 65. aedeagus, ventral view. 66. ejaculatory apodeme. 67. surstylus. Fig. 68-69. L. sylvatica. 68. aedeagus, lateral view. 69. aedeagus, ventral view. Agromyzidae of Alberta 399 larva. 78. posterior apir ^ Ce"“^ ***■> * 400 Sehgal Fig. 80— 82. Phytomyza aquilegioides. 80. hypandrium. 81. aedeagus, lateral view. 82. distiphallus, ventral view. Fig. 83-85. P. amicivora. 83. hypandrium. 84. aedeagus, lateral view. 85. ejaculatory apodeme. Fig. 86-90. P. blairmorensis. 86. hypandrium. 87. postgonite. 88. aedeagus, lateral view. 89. aedeagus, ventral view. 90. ejaculatory apodeme. Agromyzidae of Alberta 401 Fig. n. Phytomyza colemanensis, aedeagus, lateral view. Fig. 92-95. P. columbinae. 92. hypandrium. 93 aedeagus ateral view. 94 distiphallus ventral view. 95. ejaculatory apodeme. Fig. 96-98. P. edmontonensis. 96. hypandri^i’ 97. aedeagus, lateral view. 98. ejaculatory apodeme. Fig. 99. P. gregaria, aedeagus, lateral view. 402 Sehgal Fig. 100-103. Phytomyza jasperensis. 100. hypandrium. 101. postgonite. 102. aedeagus, lateral view. 103. aedeagus, ventral view. Fig. 104. P. lactuca, aedeagus, lateral view. Fig. 105—107. P. luteiceps. 105. hypandrium. 106. aedeagus, lateral view. 107. ejaculatory apodeme. Agromyzidae of Alberta 403 Fig. 108. Phytomyza major, aedeagus, lateral view. Fig. 109-112. P. mertensiae. 109. hypandrium. 110. aedeagus, lateral view. 111. distiphallus, ventral view. 112. ejaculatory apodeme. Fig. 113. P. milti, aedeagus, lateral view. 404 Sehgal Fig. 114. Phytomyza misella, aedeagus, lateral view. Fig. 115-117. P. multifidae. 115. hypandrium. 116. aedeagus, lateral view. 117. ejaculatory apodeme. Fig. 118-120. P. oxytropidis. 118. hypandrium. 119. aedeagus, lateral view. 120. ejaculatory apodeme. Fig. 121—122. P. riparia. 121. aedeagus, lateral view. 122. ejaculatory apodeme. Agromyzidae of Alberta 405 Fig. 123-124. Phytomyza senecionella. 123. aedeagus, lateral view. 124. ejaculatory apodeme. Fig. 125-127. P. solida- ginophaga. 125. hypandrium. 126. aedeagus, lateral view. 127. ejaculatory apodeme. Fig. 128-132. P. subalpina. 128. hypandrium. 129. postgonite. 130. aedeagus, lateral view. 131. aedeagus, ventral view. 132. ejaculatory apodeme. Fig. 133. P. timida, aedeagus, lateral view. 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 : 406 1971 Corrigenda. A. Nimmo 1971, Quaest. ent. 7:3-234. p. 16 Line 2. ‘Rhyacophilidae Ulmer’ should read ‘Rhyacophilidae Stephens’. p. 49 Line 44. For: ‘Tergum VIII of male unmodified of variously . . . ’ read ‘Tergum VIII of male unmodified or variously . . . ’. p. 68 Line 14. ‘ Apatania shoshone ; Betten, . . . ’ should read ‘ Apatania shoshove; Betten, . . . ’. p. 75 Line 27. For: ‘The holotype, allotype, and three male and 36 paratypes . . . * read ‘The holotype, allotype, and three male and 36 female paratypes . . . ’. p. 77 Line 7. Omit reference to Fig. 605 (N. laloukesi not shown). See Fig. lb. right side, for Lake Louise locality. p. 104 Above ‘ Limnophilus hingstoni Mosely, . . . ’ insert ‘ Limnophilus moestus; Schmid, 1955:137’. p. 134 Line 19. "Limnephilus (Goniotaulius) Pulchellus; . . . ’ should read ‘ Limnephilus (Goniotaulius) pulchellus ; . . . ’. p. 152 Line 26. Second line of diagnosis for/! prita (Milne), for realtively, read relatively. p. 178 Fig. 497. Male genitalia, dorsal aspect (partial). p. 203 In range pattern no. 6, the first species, for: ‘ Rhyacophila vemma’’ read: ‘ Rhyaco - phila vemna\ p. 203 In range pattern no. 6, N. laloukesi. Omit Fig. 605. See note to p. 77, line 7 (above). p. 222 Line 24. For: ‘ . . . possible 6%, . . . ’ read ‘ . . . possibly 6%, . . . ’. p. 234 Line 5. For: Homosphylax ’ read Homophylax ’. AN APPARATUS AND METHOD FOR THE FIELD SEPARATION OF TABANID LARVAE (DIPTERA: TABANIDAE) FROM MOSS ANTHONY W. THOMAS Department of Entomology University of Alberta Quaes tiones entomologicae Edmonton 7, Alberta 7 ; 407-408 1971 A portable apparatus and its use for the separation of tabanid larvae from moss in the field is described. Thirty-seven hours work yielded 463 larvae in 16 species (Hybomitra 10, Chrysops 3, Atylotus 2, Haematopota 1). Compared with a Berlese funnel drying unit, this apparatus was 80% efficient. Ce texte donne la description et Tutilisation dun appareil facilement transportable, con- struit pour separer, au champ, les larves de tabanide de la mousse. Trente-sept heures de travail ont permi de separer de la mousse 463 larves appartenant a 16 especes fHybomitra 10, Chrysops 3, Atylotus 2, Haematopota 1). Si on compare cet appareil a celui de Berlese, soit des entonnoirs sechant, son efficacite est de 80%. The major habitat of Hybomitra and Atylotus larvae in northern North America is moss (Teskey, 1969). The separation of larvae from moss is tedious and has only been accom- plished with any efficiency by drying the moss (Teskey, 1962). Miller (1951) transported moss back to the laboratory and hand sorted it on a table. He considered a yield of 1 0 to 15 larvae per man per day unusually high. Teskey’s (1962) apparatus is efficient but is depen- dent upon a power supply. It also necessitates the transport of moss from the field to the laboratory and is thus of no use on extended collecting trips. The following apparatus was developed for collecting tabanid larvae from moss when transfer back to the laboratory was not practical. CONSTRUCTION OF THE PORTABLE SEPARATOR The frame was built of Vi inch O. D. aluminum alloy tubing having a 1/16 inch wall. It consisted of two six feet long side pieces, two two-feet-nine-inch pieces for the width and four four feet long legs. In use, the legs were pushed one foot down into the moss as an aid to frame stability. The frame was held together by four copper comer pieces, each made of a standard plumbers’ tee and 90° elbow and three two-inch long copper pipes. This frame supported two nets. The upper one was four mesh/inch, made of string and manufactured as a base for carpets, and received the moss. The lower one was 20 mesh/inch, made of fiberglass and manufactured as window screening, and was to collect larvae. METHOD OF USE The separator is easily portable, either dismantled or assembled when it can be carried upside down on one’s back. When an area was to be searched for larvae it was far easier to take the apparatus to the area than transport the moss to the separator. Excessive water was 408 Thomas removed by hand squeezing and the moss then placed on the top net. Enough moss was collected to cover this to a depth of Vz inch; about % of a cubic foot of loosely packed moss. Collection of a sample took less than five minutes. The moss was then shredded by hand, the aim being to separate individual moss plants. This shredding process took between 15 and 20 minutes. During shredding the larvae leave the moss and crawl or fall through the mesh and become stranded on the lower net. This lower net was examined about every two minutes and the larvae retrieved. When the moss was thoroughly shredded the upper net was hit from beneath with the hands. This tossed the moss into the air causing any remaining larvae to separate out. The moss was then discarded and another sample was worked. It is important to shred the moss thoroughly and not place too much on the net at a time. The above method separates pupae as well as larvae but such pupae are almost always crushed. Precautions are necessary if intact pupae are wanted. The sample must be collected with care and without squeezing. The shredding of such saturated moss is difficult. This apparatus was used in muskegs where the substratum was all moss and in sloughs where there was a layer of moss and dead horsetails ( Equisetum ) on a clay substratum. RESULTS During May and June 1970, 273 larvae of 15 species (2 Atylotus, 1 Haematopota, 9 Hybomitra, 3 Chrysops ) were collected during 25 hr sampling in three localities in Alberta. The smallest return was 27 larvae for five hours work and the maximum yield was 42 larvae for two hours work. On five other occasions the moss, after being subjected to field sorting, was brought back to the laboratory and placed in extracting units (Teskey, 1962) until dry. In 12 hr of field work 190 larvae of nine species were obtained, 45 others were obtained from the drying units. Assuming the drying units to be 100% efficient at extracting larvae the efficiency of the field separator ranged from 70% to 89% (average, 80%). Eighty-nine small larvae (< 1 cm) were obtained with the drying units. No attempt to identify these beyond the family level was made. No small larvae were seen during field separations. DISCUSSION When an absolute quantitative result is required this portable separator is of no use. However, when a power supply is unavailable, or it is not practical to transport moss to the laboratory, it provides an efficient way of sampling moss for tabanid larvae. REFERENCES Miller, L. A. 1951. Observations on the bionomics of some northern species of Tabanidae (Diptera). Can. J. Zool. 29:240-263. Teskey, H. J. 1962. A method and apparatus for collecting larvae of Tabanidae (Diptera) and other invertebrate inhabitants of wetlands. Proc. ent. Soc. Ont. 92:204-206. Teskey, H. J. 1969. Larvae and pupae of some eastern North American Tabanidae (Diptera). Mem. ent. Soc. Canada 63:147 pp. ACKNOWLEDGEMENTS I wish to thank W. G. Evans for financing the drying units and portable separator. 409 Announcement — First International Congress of Systematic and Evolutionary Biology The Society of Systematic Zoology and the International Association for Plant Taxon- omy have joined forces to develop this first opportunity for botanical/zoological interaction at the international level. The University of Colorado (Boulder, Colorado) has extended a gracious invitation to meet on that campus August 4-1 1, 1973. The diversity of ecological situations in the surrounding countryside makes this one of the most attractive sites in North America, both aesthetically and scientifically. The presence of experienced, enthusi- astic biologists on that campus also provides an indispensable ingredient for the success of this Congress. To begin the planning phase, two committees have been appointed by the sponsoring organizations, a Steering Committee and an International Advisory Committee. The follow- ing have been asked to serve on these bodies: Steering Committee F. A. Stafleu (Chairman) J. O. Corliss (Convenor) J. L. Reveal (Secretary) R. S. Cowan J. A. Peters R. W. Pennak W. A. Weber G. S. Daniels (Finance Committee) P. D. Hurd, Jr. (Co-Chairman of Program Committee) B. L. Turner (Co-Chairman of Program Committee) International Committee Botanists and Bacteriologists: H. Banks (U. S. A.) S. T. Blake (Australia) * R. S. Cowan (U. S. A.) J. De Ley (Belgium) M. A. Donk (Netherlands) Th. Eckardt (Germany) K. Faigri (Norway) H. Hara (Japan) A. T. Hunziker (Argentina) R. McVaugh (U. S. A.) Tweede Transitorium, Uithof, Utrecht, Netherlands. Department of Zoology, University of Maryland, College Park, Maryland, U. S. A. Department of Botany, University of Maryland, College Park, Maryland, U. S. A. National Museum of Natural History, Smithsonian Institution, Washington, D. C., U. S. A. Department of Vertebrate Zoology, Smithsonian Institution, Washington, D.C., U. S. A. Biology Department, University of Colorado, Boulder, Colorado, U. S. A. Natural History Museum, University of Colorado, Boulder, Colorado, U. S. A. Hunt Botanical Library, Carnegie-Mellon University, Pittsburgh, Pennsylvania, U.S.A. Department of Entomology, Smithsonian Institution, Washington, D.C., U. S. A. Department of Botany, University of Texas, Austin, Texas, U. S. A. Zoologists: J. G. Baer (Switzerland) E. Beltran (Mexico) B. E. Bychowsky (U. S. S. R.) * J. O. Corliss (U. S. A.) R. B. Freeman (U. K.) W. Hennig (Germany) L. B. Holthuis (Netherlands) D. L. Hull (U. S. A.) * P. D. Hurd, Jr. (U. S. A.) M. A. Klappenbach (Uraguay) 410 * F. A. Stafleu (Netherlands) A. Takhtajan (U. S. S. R.) Sir George Taylor (U. K.) * W. A. Weber (U. S. A.) R. C. Rollins (U. S. A.) P. Sneath (U. K.) E. Mayr (U. S. A.) R. V. Melville (U. K.) C. D. Michener (U. S. A.) E. C. Olson (U. S. A.) * R. W. Pennak (U. S. A.) * J. A. Peters (U. S. A.) R. A. Ringuelet (Argentina) C. W. Sabrosky (U. S. A.) (*Also member of Steering Committee.) The Steering Committee will be the principal organizing group. The International Com- mittee will provide valuable advice and guidance in the development of the Congress and it is recognized by the International Union of Biological Sciences as the special working group responsible for this event. Program plans at this point encompass interdisciplinary symposia and contributed paper sessions. The botanists will not convene a nomenclatural section but a zoological one on this subject is anticipated. In the next few months the outline of the program and other activities will begin to take form. All suggestions will be gratefully received, carefully con- sidered, and as many adopted as practical or feasible. Correspondence may be addressed to any member of the Steering Committee but preferably to the Secretary: Dr. James L. Reveal, Department of Botany, University of Maryland, College Park, Maryland 20740. Publication of Quaestiones Entomologicae was started in 1965 as part of a memorial project for Professor E. H. Strickland, the founder of the Department of Entomology at the University of Alberta in Edmonton in 1922. It is intended to provide prompt low-cost publication for accounts of entomological research of greater than average length, with priority given to work in Professor Strickland’s special fields of interest including entomology in Alberta, systematic work, and other papers based on work done at the University of Alberta. Copy should conform to the Style Manual for Biological Journals published by the American Institute of Biological Sciences, Second Edition, 1964, except as regards the abbreviations of titles of periodicals which should be those given in the World List of Scientific Periodicals, 1964 Edition. The appropriate abbreviation for this journal is Quaest. ent. An abstract of not more than 500 words is required. All manuscripts will be reviewed by referees. Illustrations and tables must be suitable for reproduction on a page size of 9%X63/4 inches, text and tables not more than 73/4x43/4 inches, plates and figures not more than 8V2X5 inches. Reprints must be ordered when proofs are returned, and will be supplied at cost. Subscription rates are the same for institutions, libraries, and individuals, $4.00 per volume of 4 issues, normally appearing at quarterly intervals; single issues $1.00. An abstract edition is available, printed on one or both sides (according to length) of 3X5 inch index cards (at $1.00 per volume) or on 5X8 inch standard single row punched cards ($1.50 per volume) . Communications regarding subscriptions and exchanges should be addressed to the Subscription Manager and regarding manuscripts to: The Editor, Quaestiones Entomologicae, Department of Entomology, University of Alberta, Edmonton, Canada. £ ID.- Qw Quaestiones entomologicae fv'iUS. COMP, 2.00 LIBRARY OCT 29 1371 HARVARD UNIVERSITY A periodical record of entomological investigations, published at the Department of Entomology, University of Alberta, Edmonton, Canada. VOLUME VII NUMBER 4 OCTOBER 1971 QUAESTIONES ENTOMOLOGICAE A periodical record of entomological investigation published at the Department of Entomology, University of Alberta, Edmonton, Alberta. Volume 7 Number 4 1 October 1971 CONTENTS 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 Editorial — Four Men and a Moth Of the 4000 or so species of Noctuidae occurring in North America, few are of any great economic importance. Most are unknown to the man in the street and even to the farmers they are most likely to influence directly. Yet we know a lot today in comparison with what we knew in 1911 when it first became apparent that the interests of the white man in North America were also the interests of one of these species, the pale western cutworm, then known as Porosagrotis orthogonia (Morrison). Most of what we know was discovered by as grand a quartet of entomologists as a single insect species could ever hope to attract: E. H. Strickland, William C. Cook, H. L. Seamans, and Larry A. Jacobson: the sequence is chronological; to distinguish between these men on any other basis would be improper. How little we knew of the pale western cutworm was exemplified by the disastrous recom- mendation to farmers by the Canadian government, to cultivate out all weeds before moth flight. This cutworm lays its eggs in loose soil, not, like some other noctuids, on the leaves of plants. But all this, and more, comes out in Jacobson’s able, though modest review. All these four men were primarily interested in the whole life-cycle of the whole insect; in its relationship to its environment, and in the problems it set for the farmer. In pursuit of these interests they, like the pale western cutworm, ignored the forty-ninth parallel. All four were men of strong views, given to speaking their minds. None who knew them would expect them all to agree on anything, let alone on such complex problems as this insect presented. Yet this very diligence in disagreement was the source of their strength as a group, for each sometimes proved another right; sometimes one suspects, by trying to prove him wrong. Strickland, working from a Canada Department of Agriculture Field Station at Leth- bridge, Alberta, when the trouble first started was quick to provide interim recommenda- tions to farmers for control with insecticides. At the same time he was busy accumulating data on predators and parasites with a view to a more basic solution to the problem. Cook, assistant state entomologist for Minnesota, worked from there and at the Montana Agricul- tural Experiment Station. His major interest in the effects of weather and climate on insects conferred upon us the ability to predict outbreaks from weather data. Back in Lethbridge, Seamans applied Cook’s findings in forecasting, developed practical methods for cultural control, and laid the foundations of our knowledge of feeding habits, rearing methods and nutrition. Jacobson, following Seamans at Lethbridge, took Cook’s interest in the influence of weather into the laboratory and quantified the influence of several factors to yield a more definitive life history. He also developed the first effective chemical control, followed up on the work of Seamans on rearing, and contributed much on the behaviour of adults in the field and the laboratory. Jacobson, in this review, assesses the considerable contribu- tions of others. There have been times when superficial consideration of problems like that presented by the pale western cutworm has seemed to suggest that studies of the kind conducted by these four men are outmoded and redundant. The development of new and superior insecticides and technology for their application, of radio-sterilisation and plant breeding techniques seem to encourage such thoughts. But time has shown and will continue to show that there is no substitute for a knowledge of the biology of the insect and of the plant or animal on which it feeds. There will always be a place in applied entomology for the naturalist, for the man with a flair for revealing those features of the life of an insect which permit some finesse in our attempts to control it. Indeed I see this as the central function of an applied entomologist; as that which distinguishes him from a chemist and an engineer. Despite what has been accomplished, it cannot be said, sixty years later in 1971, that the pale western cutworm problem has been solved. Though the value of crops saved ex- ceeds by many times the cost of research done (none of these men made fortunes or even drew large salaries), the problem is still with us. And so it must be, for just as we select strains of insects resistant to our insecticides merely by using them, we also select strains of insects which damage our crops merely by growing them. And insects evolve so much faster than we do. No better example of the dependence of applied entomology on basic entomology could be found than the pale western cutworm. The mistakes of 1911-1912 could be repeated at any time should our use of land change in favour of any of the other 3999 species of noctuid moths. We are very nearly as ignorant about the lives of most of them today as we were about the pale western cutworm in 1911; it would pay us to study them now. It is a pleasure and a priviledge to publish, on the eve of the 50th anniversary of the Department of Entomology at the University of Alberta, a paper which reflects so much credit on the founder of the department and on one of his students. Brian Hocking Frontispiece: Life stages of Agrotis orthogonia Morrison. (A and B) <5 and 9 moth; (C and D) ventral views of <5 and 9 pupa; (E) larva; (F) egg with fully developed embryo (25x). THE PALE WESTERN CUTWORM, A GROTIS ORTHOGONIA MORRISON (LEPIDOPTERA: NOCTUIDAE): A REVIEW OF RESEARCH L. A. JACOBSON Canada Department of Agriculture Research Station Quaestiones entomologicae Lethbridge, Alberta 7 : 414-436 1971 The pale western cutworm, Agrotis orthogonia Morrison, has been one of the major insect pests of grain production in the open plains of western Canada and the United States at fre- quent intervals since 1911. The history of occurrence and known distribution is described. The generic combinations and synonymy are brought up to date. Life history data in the various areas of occurrence are compared. Research on biology of the various stages is re- viewed. Known parasites and predators are recorded and their role in natural control is dis- cussed. The relationships of infestations and weather and the factors used in forecasting outbreaks are reviewed. Various methods of control in the field in the past, and their present status, are included with suggestions of possible methods for the future. Le ver gris orthogonal, Agrotis orthogonia Morrison, est Tune des principals pestes du grain dans les plaines de Touest canadien et americain. En effet, les attaques de cet insect ont ete observees a plusieurs reprises depuis 1911. L’historique des ravages et la distribution presente sont decrites. Les combinations generiques et la synonymie sont mises a jour. Les donnees d’histoire naturelles sont comparees pour les differentes regions attaquees. On y fait le point des recherches sur la biologie de chaque stage. Les parasites et les predateurs connues sont notes, et leur role est discute en fonction du control naturel. Les relations entre les ravages et la temperature, et les facteurs utilises pour la prediction des attaques sont passees en revue. Plusieurs methodes de control utilisees sur le terrain dans le passe, et leur etat present sont discutees, et des suggestions sont faites en fonction de methodes possibles a employer dans Tavenir. The pale western cutworm, Agrotis orthogonia Morrison, has been one of the major eco- nomic pests of grain production in the open plains of western Canada and the United States for many years. This publication is a review of research on this insect and deals with biology, ecology, and control. References do not include publications on local distribution and control unless they include research data not available elsewhere. History The history of the pale western cutworm as a pest in western Canada has been described (Gibson, 1912, 1914, 1915; Seamans, 1926, 1931, 1952). In the United States the first occurrence and subsequent infestations and outbreaks are recorded: Montana (Parker, Strand, and Seamans, 1921; Cook, 1930); North Dakota (Webster, 1924); Oklahoma (Esh- baugh, 1933); Utah (Sorenson and Thomley, 1941); New Mexico (Eyer, 1957); Nebraska (Pruess and Roselle, 1969). The pale western cutworm was virtually unknown before 1911 and apparently did not become a pest until after the cultivation of range lands and the growing of grain became widespread in the prairies of the United States and Canada. Seamans (1934a, 1934b) de- scribed how numbers increased because of the change from native prairie, where grasses 416 Jacobson predominated, to extensive areas of cultivated land with susceptible crops and with cultural procedures favorable to cutworms. Some research on the biology and control of the pale western cutworm was conducted in most of the areas where the cutworms occurred, particularly when it was first found and during subsequent outbreaks. Because of the importance of the insect as a destructive pest and the devastating losses that occurred, research in western Canada has continued since 1913. Most of the Canadian investigations were centered at Lethbridge, Alberta, and in- volved personnel throughout the Prairie Provinces. Synonymy The generic combinations and synonymy of the pale western cutworm follow: Agrotis orthogonia Morrison 1876. Agrotis orthogonia Morrison, Proc. Boston Soc. Nat. Hist. 18:237; described from Glencoe, Nebr. 1890. Porosagrotis orthogonia ; Smith, Bull. U. S. Nat. Mus. 38:129; generic transfer. 1928. Agrotis orthogonia ; McDunnough, Bull. Nat. Mus. Canada 55:34; generic transfer. 1908. Porosagrotis delorata Smith, J. N.Y. ent. Soc. 16:87; described from High River, Alberta. 1926. Porosagrotis orthogonia duae Barnes and Benjamin, Canad. Ent. 58:303; described from Inyo Co., Calif. DISTRIBUTION Walkden (1950) describes the pale western cutworm as a typical dryland cutworm in semi-arid areas. He states that outbreaks have occurred in western Kansas, northeastern New Mexico, the Panhandle sections of Texas and Oklahoma, eastern Colorado, western South Dakota and North Dakota, Montana, Utah, and Wyoming in the United States. In Canada the pale western cutworm is generally confined to the prairie region (Bowman, 1951). Under severe drought conditions it may be found in the outer fringes of the park belt or Savanna region. Seamans (1938) outlined the outbreak area in Canada as extending from Cowley in the foothills of Alberta to Broadview in eastern Saskatchewan and from the International Boundary northward to Turtleford and Lloydminster, Saskatchewan. The areas where infestations have occurred in Canada and the United States are shown in Figure 1 . The map is based on information obtained from various publications and from the USDA Co-operative Economic Insect Reports for the period 1952-1970. There are localities within the area, such as local mountain ranges or deserts, where the pale western cutworm has not been found. LIFE HISTORY The pale western cutworm has only a single generation annually throughout its area of occurrence. The eggs are laid in the soil in early autumn and hatch the following spring. The larvae first feed on early growth such as volunteer cereals or weeds; later they also feed on seeded crops until early summer. As they mature they become less active and form an earthen cell in the soil 2 to 3 inches below the surface. Here they remain dormant until they pupate. The moths emerge from the cell during the late summer, mate, and lay eggs soon after. Pale western cutworm 417 Figure 1. Areas in Canada and the United States where infestations have occurred since 1911. The life history of this cutworm varies considerably from one geographical area to an- other. Crumb (1929) points out that single-brooded species of the Noctuidae tend to be of northern distribution and that in the southern part of the range, where hatching is earlier, the quiescent periods of prepupae and pupae are longer so that a single generation occupies a full year. Table 1 shows the variation in life cycle between northern, southern, and intermediate areas of occurrence in selected areas of the United 'States and Canada. 418 Jacobson Table 1. Dates of various periods in the life history* of A. orthogonia at various locations of the United States and Canada. Lethbridge, Alberta 49°43’N - 1 1 2°48’W Cedar City, Utah 37°40’N - 1 13°4’W Clovis, New Mexico 34° 14’N - 103° 13’W First hatching April 1 March 1 February 1 Larval period to June 20 to June 10 to June 1 Moth period Aug. 12 - Sept. 15 Sept. 5 - Oct. 10 Sept. 15 - Oct. 25 *Mean times after Seamans (1931), Sorenson and Thornley (1941), and Eyer (1957). Egg Description. — The egg was first described by Parker et al. (1921). When laid, the egg is a glistening milk-white, which later becomes dull grey; it is spheroidal, flattened dorso- ventrally, 1 mm in diameter, and 0.8 mm in height. When an egg is completely incubated the fully formed embryo can be seen under magni- fication through the chorion, imparting to the egg a bluish color (frontis. F). Eggs of noctuid species commonly occurring in Alberta, including A. orthogonia , were identified by differences in the pattern of reticulations or ridges on the chorion in the micropyle area (Seamans, 1933). Incubation and hatching, — The eggs are laid in the early fall and hatch the following spring. Incubation in the field, determined by placing eggs after oviposition in simulated sites in soil, required from 30 to 50 days (Jacobson and Blakeley, 1958a). Lindsay (1954) showed that development rate of the embryo varied directly with temperature. Develop- ment took 11, 14, 21, and 33 days at 30°, 25°, 20°, and 15°C, respectively; no develop- ment was apparent at 5°C. In the field the eggs do not usually hatch until the soil is warmed above freezing. In western Canada hatching usually occurs about April 1, although occasionally when the ground is clear of snow earlier than this date some hatching may occur. Instances have been recorded of hatching in the fall (Cook, 1930) but such occurrences are rare. Table 1 shows the approximate date of hatch of the pale western cutworm in various areas. Contact moisture or high relative humidity is required for hatching. In the laboratory this requirement is provided by adding water directly to the eggs or to the substrate. Under field conditions soil moisture is usually adequate. Laboratory studies showed that the rate of hatching of fully developed eggs increases with temperature and relative humidity, that prolonged exposure to temperatures from -5° to -15°C does not affect ultimate hatch, and that desiccation, particularly in the range 20° to 30°C, may cause considerable mortality. Findings in the laboratory, corroborated by studies outdoors, show that eggs are admirably adapted to develop, withstand climatic factors, and hatch at a time when their survival is ensured (Jacobson and Blakeley, 1958a). Diapause. — Early authors noted that development of the embryo was completed during the fall but the eggs would not hatch readily until a further treatment near 0°C intervened (Cook, 1930). They apparently presumed that only the advent of cold weather prevented the eggs from hatching. Andrewartha and Birch (1954) postulated that a weak diapause was involved. Pale western cutworm 419 Jacobson (1962a, 1962b) showed that fully developed embryos immediately begin to feed and continue development when dissected from eggs but when left in the egg hatch slowly and unevenly. This is considered as a weak diapause, the intensity of which is re- flected by the rate at which they hatch. Investigations showed the rate of hatching is influ- enced by the temperature of incubation and by the duration and temperature of the post- incubation treatment. When diapause is firmly established, temperatures above 15°C are required to break diapause and to produce a complete hatch whereas at lower temperatures only partial hatching occurs. As diapause is eliminated hatching occurs at progressively lower temperatures. Intensity of diapause varies in eggs laid at the same time. Some eggs will hatch at suitable temperatures with moisture when embryonic development is complete. Others will require exposure to temperatures from 0° to 10°C before diapause is eliminated. The weak dia- pause in eggs of the pale western cutworm in its natural habitat is sufficient to prevent hatching in the fall and to ensure that most eggs will hatch in the spring when food is available for larvae. Larvae Description. — The larvae of A. orthogonia were described by Parker et al. (1921). Walkden (1950) described some of the morphological characters that identify cutworms, including the pale western cutworm, that attack cereal and forage crops in the central great plains. The internal morphology of larvae was described by Hocking and Depner (1961). When fully grown the larvae are from 30 to 40 mm long and 5 to 7 mm thick and the general color is usually grey with no definite stripes or markings. The only readily distinguishable characteristic is in the head; the capsule is yellow-brown with two distinct vertical black dashes that form an imperfect H or inverted V (frontis. E). Larval feeding. — On hatching the larvae are small, about 2 to 3 mm long, and difficult to find. During early instars the larvae feed on available young seedlings, such as weeds and volunteer grains. All instars of larvae are subterranean in habit, attacking the plant below the surface of the ground. Occasionally they are forced to the surface during heavy rains and sometimes at night after extremely warm days. Early workers believed that early instars fed above ground since the leaves of grain showed notches or holes. It was later demonstrated that these punctures were made by larvae feeding beneath the soil on the coleoptile and the furled leaves of the wheat plant (Jacobson, Farstad, and Blakeley, 1950). When the larvae become older and larger, and as the host plant grows, the cutworm continues to feed below the surface either cutting off the plant and leaving it to wither and die on the surface or, sometimes, pulling it into the soil there to consume it. A key to the insects, including cutworms, damaging grain plants, based on damage observed was prepared by Strickland (1948). More recently another type of feeding was observed. Plants of almost mature winter wheat were found that had been cut near the soil surface. Fully grown larvae, apparently to satisfy a requirement for fibrous material when green food is unavailable, girdle the stem by peripheral feeding. When the stem becomes brittle it falls over. It was estimated that such damage could exceed 10 per cent (Jacobson, 1967). Number of instars. — The number of instars of A. orthogonia Morrison is usually six, although in some situations may be more (Parker et al., 1921). Hardwick (1965), com- menting on supernumerary molts in another noctuid, noted that the number may vary from species to species and from individual to individual depending on the nutritive value of the food and varying with temperature. In the laboratory, pale western cutworm larvae starved during instars III or IV more often had seven instars than those that were not 420 Jacobson starved (Jacobson and Blakeley, 1960). In an insectary Parker et al. (1921) found that most larvae of this species had seven instars, some had eight, a few individuals passed through nine, and one was noted with 10. No instances are known of the pale western cutworm having fewer than six instars. The exact number of instars that occur in the field is not known. Undoubtedly, it varies from six to eight, or even more, depending upon the temperature, quality and quantity of available food, and the extent of starvation that may occur when a crop is destroyed and food is unavailable. Duration of larval period. — The duration of the entire larval period can also vary even under controlled conditions of food and temperature. Developmental times of larvae reared in the laboratory and hatched on the same day ranged from 40 to 76 days. The duration of development of larvae varies inversely with temperature (Table 2). Table 2. Durations (days) of development of larvae, prepupae, and pupae of A. orthogonia at various constant temperatures. 30°C 25°C 20°C Instar I 3 4 8 II 3 3 6 III 3 4 7 IV 3 5 7 V 5 5 8 VI 7 8 14 Larvae total 24 29 50 Prepupae 32 19 13 Pupae 25 26 28 Hatch to adults, total 81 74 91 When the larvae were fed Thatcher wheat, Marquis wheat, Compana barley, and Exeter oats, the mean durations from hatching to pupation at 25°C were 45.6, 45.7, 51.8, and 52.7 days, respectively, the latter two periods being significantly different at the 1 per cent level from the first two (Jacobson and Blakeley, 1958b). Seamans and McMillan (1935) noted differences in development and other effects when larvae were fed various foods, and Hocking (1953) found differences when different parts of the wheat plant were used. Development in the field obviously must vary greatly since larvae probably hatch at various times and are subjected to varying degrees of food deprivation and have access to various kinds of food plants. Damage to crops. — In the field, feeding by early instars is not readily discernible. As the larvae become larger they are able to cut off and consume more plants. Damage becomes evident when the larvae are in instar III or IV. It is characterized by the appearance of bare Pale western cutworm 421 areas, usually at first in sandy areas or on knolls, later these areas may enlarge encompassing many acres; sometimes entire fields may become bare (Parker et al., 1921; Cook, 1930; Seamans, 1938; Walkden, 1940). Moths apparently select hilly areas because they are more favorable for oviposition. Descriptions of the extent of areas infested and the occurrence severe crop losses have been noted (Gibson, 1914;Cook, 1930; Seamans, 1926, 1938, 1952; Sorenson and Thomley, 1941;Eyer, 1957). The density and distribution of larvae in the field as well as such other factors as weather and stage and condition of host plants may influence the damage that may occur. Seamans (1938) found that 15 or more larvae per square yard (18/m2) will destroy a crop. Previously Seamans (1931) had stated that in some seasons an infestation of about 1/ft2 (ll/m2) is sufficient to completely destroy a crop whereas the next year the same infestation will not be noticed. In 1965 the mean numbers of larvae/ft2 (m2) from 25 samples in various por- tions of an infested field of wheat were: Range Mean Damaged area (100% loss) 0-7 3.04 (33) Margin (50% loss) 0-5 2.44 (26) Part crop (25% loss) 0-3 0.84 ( 9) These counts indicate that variation in population can occur in a relatively short dis- tance as these counts were made in three lines parallel to each other and about 6 feet apart. The counts also indicate relationships between density and the degree of crop dam- age (Jacobson, unpublished). The interrelations between cutworms and the host plant have been investigated (Jacobson and Peterson, 1965). Wheat can withstand feeding by instar I and instar II since the larvae cannot cut off wheat plants. The larvae damage the plants by cutting holes in the leaves as they elongate through the coleoptile. The number of plants affected increases with increased larval population (Jacobson et al., 1950). Other experi- ments in the laboratory, in the greenhouse, and on field plots with pale western cutworm at various stages of larval development and wheat at various stages of growth showed that larvae did not completely sever the plants until instar III; the rate and amount of damage varied directly with size and densities of larvae, temperatures above 15°C, and soil mois- ture; and damage varied inversely with the age and size of the wheat plant (Jacobson and Peterson, 1965). Damage to wheat by the pale western cutworm is indicative of a dynamic situation where both the insect and the host plant are actively developing and the advantage to one or the other is constantly changing. Effects of starvation. — Experiments showed that, if weeds in fields containing first-instar larvae were allowed to grow for a short period and then were destroyed by cultivation, the young larvae would starve in 1 0 days to 2 weeks, after which it would be safe to seed the intended crop. Surveys of infested fields along with history of timing of pre-crop cultivation and seeding showed that delayed seeding resulted in less damage than in those fields where the cultivation and seeding were done simultaneously (Seamans, 1937). Greenhouse studies showed that larvae that had fed were more susceptible to starvation than larvae that had not fed (Seamans and Rock, 1945). In the laboratory this was reconfirmed along with the observation that a digestive disturbance occurred in larvae that were fed after starvation (Salt and Seamans, 1945). Other effects of starvation were studied with all instars. Larvae survived starvation longer when they were larger or when temperatures were lower. When food was made available to larvae that had been starved for some time some of them were unable to resume feeding (Jacobson, 1952). Desiccation is another factor involved in mortality from starvation. Instar IV larvae were more resistant to desiccation than starved instar II larvae. The rate of mortal- ity was lowest at RH50. At RHO and RH100 the rate was almost the same, indicating that 422 Jacobson desiccation and excessive moisture were equally harmful to larvae (Jacobson and Blakeley, 1957b). The kind of food affected mortality of larvae when they were starved. Larvae were fed Marquis wheat, Thatcher wheat, Compana barley, or Exeter oats until instar V and then starved. Larvae that were fed the oats were least resistant to starvation whereas those fed the wheat varieties showed the greatest resistance to starvation (Jacobson and Blakeley, 1958b). The rate of mortality was associated in each case with the amount of weight gain and hence was a reflection of better utilization of food. Larvae at all stages were fed on wheat and starved. When larvae were starved for various periods before instar V and then fed, the larval period was prolonged, supernumerary moults occurred, and the pupae weighed less. When larvae were starved in their ultimate instar their development accelerated, the pupae weighed less, and fecundity was reduced (Jacobson and Blakeley, 1960) . Host plants of larvae. - Lists of plants attacked by larvae of the pale western cutworm have been published (Sorenson and Thornley, 1941; Cook, 1930; Webster and Ainslie, 1924). These include a variety of crops grown in the areas where this cutworm has been known to occur. Generally, however, the larvae prefer cereals and the greatest losses have occurred to crops of wheat, oats, and barley. In gardens and in areas where vegetable crops are grown, the pale western cutworm often appears along with other cutworm species. Nutrition of larvae. — Before 1950, studies on the pale western cutworm primarily in- volved field ecology and the information obtained was directly concerned with life history and field control. Many questions about the biology and behaviour of the insect went un- answered. Over a period of about 15 years McGinnis and Kasting carried out a series of nutritional and biochemical investigations aimed at providing some of the answers. Then- first reports showed that the rate of growth and development depended on both quality and quantity of the food consumed. They found that larvae with free access to Thatcher wheat sprouts were larger and developed more rapidly than larvae allowed the same food for only 2 hours each day (McGinnis and Kasting, 1959). They also found that the variety of wheat sprouts upon which the larvae fed affected growth (Kasting and McGinnis, 1959). The amino acids essential in the diet of this insect were determined using a radioactive tracer technique (Kasting and McGinnis, 1966). Results indicated that it had the same general requirements as other animals; no abnormal amino acid requirements were evident (Kasting and McGinnis, 1962). The amino acids in the normal diet of these larvae are largely bound in the protein form. It was necessary, therefore, to determine whether the larvae could utilize dietary protein. Results of a study with protein-U-C14 showed that the larvae readily digested the protein (McGinnis and Kasting, 1962). The presence of proteolytic enzymes was confirmed in a subsequent study with gut homogenates (Khan and Kasting, 1961) . Other enzymes, including various carbohydrases, peptidases, and lipases, were also shown to be present. A synthetic diet for the pale western cutworm was described in 1967 and growth factor requirements were determined by the classical deletion procedure (Kasting and McGinnis, 1967). Results indicated that niacinamide, choline, pantothenic acid, pyridoxine, riboflavin, folic acid, and thiamin are essential. No requirement for biotin, inositol, or vitamin B12 was demonstrated. Because this insect grows fast and is large in the later instars, it has been used effectively as a laboratory animal. Several techniques connected with the nutrition of the species have been developed and tested, as for example, the use of lyophilized plant tissue in diets (McGinnis and Kasting, 1960) and measurement of consumption and digestibility of food (McGinnis and Kasting, 1964a, 1964b, 1969; Kasting and McGinnis, 1965). Pale western cutworm 423 Prepupae and pupae Prepupae. — The prepupal stage has been described as a quiescent or non-feeding period between the end of the larval period and the pupal period. When feeding ceases, the body becomes shrivelled and assumes a yellowish color. In laboratory rearing the exact date on which larvae become prepupae is difficult to establish. Sometimes the larvae may stop feed- ing for several days only to resume. A more accurate measure of the prepupal period was obtained by daily weighings after the fifth moult until feeding ceased and until weights had decreased about 25 per cent. The duration of the prepupal period is considered to be the time from the date of maximum weight to the date of pupation (Blakeley and Jacobson, 1960). In the field the larva, after completing its feeding, burrows 2 to 6 inches (5 to 15 cm) into the soil and constructs an earthen cell enclosing itself. Each larva forms its cell by injecting fluid through its mouthparts into the surrounding earth and compacting this with head and body movements until a smooth lining is formed. In the laboratory where this was observed in glass tubes the process was completed in a few days (Blakeley, 1954). The mean durations of the prepupal stage of insects that had been reared through the larval stage under similar conditions of food and temperature and assigned to 20°, 25°, and 30°C at the beginning of the prepupal stage were 12.5, 19.8, and 31.8 days, respec- tively. The longer prepupal interval at 30°C was considered to be a form of diapause that enabled the pale western cutworm to survive as a single-brooded species in the wide varia- tion of climate from the prairies of Canada to Texas in the United States (Blakeley and Jacobson, 1960). This relationship with temperature will also allow for a compensatory effect to ensure eclosion at the normal time of year if larval development is accelerated because of increased temperatures. Pupae. — In the field, the change from the prepupal stage to pupae occurs inside the earthen cell. In the laboratory where soil is not used this change occurs normally without a cell. The pupae has been described (Parker et al., 1921). In the field it is difficult to determine when prepupae change to pupae except by periodic collections. The progress of development in 1965 is shown in Table 3. No feeding larvae were found after July 5. These data, which are for one year only, show that pupation was complete about August 1 but experience has shown considerable variation from year to year related to weather and moisture conditions. Density decreased as the season progressed as a result of natural mortality factors such as parasitism, predation, and inclement weather. Table 3. Numbers of larvae, prepupae, and pupae of A. orthogonia collected from 25 square-foot samples from a field near Lethbridge, Alberta, in 1965. Date Larvae Prepupae Pupae Total Number/ft2 June 18 76 0 0 76 3.0 23 66 2 0 68 2.7 29 52 3 0 55 2.2 July 2 29 13 0 42 1.8 5 3 33 0 36 1.4 16 0 19 1 20 0.8 27 0 8 2 10 0.3 August 4 0 1 9 10 0.3 424 Jacobson In the laboratory the duration of the pupal stage varied inversely with temperature and directly with weight. An increase of 10 mg in the pupae was associated with an increase of 0.12 days in the duration of the pupal stage (Blakeley and Jacobson, 1960). Since larvae may vary in vigour and not all may have an equal opportunity to feed, they vary in weight. This, and other causes of individual variation, accounts for the variation in time of change to prepupae, to pupae, and finally to emergence of the moths. At 25°C duration of the pupal stage of 48 pupae ranged from 21 to 46 days and averaged 27 days (Blakeley and Jacobson, 1960). Toward the end of the pupal period the adult can be seen inside the pupal case. Eclosion from the pupal case, the subsequent escape from the earthen cell, and the ascent through the soil using spines on the middle and hind legs has been described (Blakeley, 1954). Sex of the pupae is easily determined (Butt and Cantu, 1962) and the method used is invaluable in laboratory investigations to determine sex ratio before adults emerge. The pupae of each sex are shown in frontis. C and D. Mortality of prepupae and pupae. — The pupal cell and surrounding soil protect the pre- pupa and pupa from desiccation. However, the insect is not completely safe as the popula- tion density decreases during these stages (Table 3). Of 50 non-feeding larvae placed out- doors in early July and examined at regular intervals until September 13, eight died of undetermined causes, eight were killed by parasites and four by predators, and 12 were recorded as missing. Only eight of the original 50 emerged as moths. The category missing is used to describe prepupae and pupae that died from predation by insects or animals or other causes and soon disintegrated (Jacobson, unpublished). Adult stage Moth. — The description of the adult male by Morrison (1876) has been repeated in the literature (Parker et al., 1921; Blakeley, 1954). Both sexes are easily identifiable in the field and usually are readily distinguished from other species that appear coincidently. Moths are readily sexed. The antennae of the males, described originally as strongly serrate, are pectinate; those of the females are filiform (frontis. E and F). Eclosion of moths appears to conform to a diurnal rhythm; 90.7 per cent emerged in the laboratory between noon and midnight and 49.5 per cent between 1400 and 1600 hours (Jacobson, 1965). In the field and laboratory, male moths emerge earlier in the flight period than females (Cook, 1930). Flight. — In western Canada, the moths first appear on flowers or around lights dur- ing the first two weeks in August. Their numbers increase until the end of the month and gradually decline in September. The flight period varies from one area to another and re- flects differences in latitude (Table 1). At Saskatoon, Saskatchewan, moths appear and the flight terminates earlier than at Lethbridge. The duration of the flight varies from year to year according to the weather. In 1950, in the immediate Lethbridge area the flight was terminated in early September by unseasonably high maximum temperatures that com- pletely dried the flowers upon which the moths were feeding. Flights usually terminate dur- ing periods of cool weather, marked by snow and freezing temperatures. Cooley (1922) recorded the end of the flight in 1921 after a severe snowstorm in early September. In years when conditions remain favourable moths can be found in low numbers until late September. Duration of flight undoubtedly influences the numbers of eggs that are laid. Moths of the pale western cutworm are crepuscular and exhibit a distinctive diurnal pat- tern of activity. Only rarely during the day are moths found before noon except when they are unusually abundant and the weather is warm. During the early afternoon males and a few females begin to appear on flowers and numbers of both sexes increase gradually Pale western cutworm 425 until sunset. In the late afternoon females become more abundant. Counts of moths on flowers show that the preponderance of one sex over the other changes. Until 1600 hours males outnumber the females about eight to one; after that time until sunset the propor- tion is reversed. The change in ratio is probably due to differences in feeding habits be- tween the sexes and to the oviposition behaviour of the females. Activity is related to temperature. At 22°C and above, moths are very active and diffi- cult to capture, but as the temperature drops below 22°C, the moths become more passive; below 10°C the moths stop flying and seek shelter at ground level, usually around patches of flowers. As the season progresses the daily interval of activity lessens. The moths feed on flowers and are readily attracted to light as evidenced by capture in light traps. The numbers of moths caught each hour in a light trap designed to capture moths for hourly periods (Seamans and Gray, 1934) were similar until midnight but be- came less after that time until about 0300 hours. The decline in numbers is apparently associated with decrease in temperature during the night. Similar data were recorded by Cook (1930) over a four-year period when numbers were counted on flowers during the dark period commencing at 2000 hours. A study of activity of both sexes was made using electrophysiological apparatus de- scribed by Edwards (1964) wherein the flying activity of males and walking of females were recorded electronically. Peaks of female activity occurred after sunset, near midnight, and at sunrise. Male activity was centered mainly at midnight. It was assumed that activity at that time was associated with mating since it coincided with a similar activity time of females. The crepuscular times were interpreted as times of feeding or oviposition. Laboratory studies showed that the maximum distance flown on a flight mill was 14.7 miles (23.7 km) for males and 3.5 miles (5.6 km) for females, that speed varied from less than 1 to 3 m.p.h. (1.6 to 4.8 km/hr), that total flight times were usually marked by a series of repeated flights, and that flying ceased with exhaustion but would resume after rest or feeding (Jacobson, 1965). Chance (1971) has measured the drag-speed relationship of the flight mill I used and calculated the compensation for this which gives values for the insects in free flight about 20 per cent higher than these. Preliminary investigations in the field using marked-capture techniques appeared to indi- cate that movement of moths was local, confined to adjacent fields and patches of flowers. Other observations showed that during the daily flight period movement in and out of the flowers was almost constant except that the proportions of males to females were reversed between one part of the day and another. No definite records of mass migrations of moths of the pale western cutworm have been observed or recorded. Known data indicate that the moth is not as strong a flier as other noctuid species that are known to be capable of migrating considerable distances. The duration of the flight period and daily activity is associated with availability of suitable flowers from which the moths obtain nectar. In western Canada the favoured flowers are sunflowers ( Helianthus sp.) and goldenrods {Solidago spp.), both of which occur throughout the prairie region along roads and field margins. In some areas of Mon- tana and Utah the moths are attracted to rabbitbrush ( Chrysothamnus sp.) (Cook, 1930; Sorenson and Thomley, 1941), a plant that occurs on the Canadian prairies in eroded and arid areas but is not utilized by the moths of the pale western cutworm since this cutworm does not usually occur in these areas. Moths have been observed feeding on the blossoms of broomweed ( Gutierrezia sp.), Russian thistle ( Salsola pestifer A. Nels.), Canada thistle (Cirsium arvense (L.) Scop.), perennial sow-thistle ( Sonchus spp.), gumweed ( Grindelia squarrose (Pursh) Dunal), and several species of fleabane ( Erigeron spp.). Moths are ener- getic feeders. They crawl about on the flowers, quickly uncoil their long sucking tubes, 426 Jacobson and insert them into blossom after blossom. The length of life of moths varies inversely with temperature, at constant temperatures in the laboratory ranging from a mean of 38 days at 0°C to 7.4 days at 35°C. At room tem- perature, 22°C, there was no difference between sexes, the mean life span being 12 days. Longevity of females at room temperatures was influenced by food; those fed honey and water had a mean length of life of 14 days; those fed with water, 8 days; and those not fed, 5 days. The length of life in the field cannot be determined but must be very variable as it is dependent on temperatures, available food, prevailing weather, and presence of predators. Females collected from flowers in the field usually die in 1 to 5 days after capture. Mating and oviposition. — Mating is rarely observed in the field. A few instances are recorded of copulating pairs being found on flowers during the early evening or at night (Cook, 1930; Seamans, 1931). In the laboratory copulation usually occurs during the night as attested by the numbers of pairs attached together in the morning. Vigils at night showed that most matings occurred from 0100 hours to daylight (Jacobson, unpublished). Evidence of mating determined by dissecting females for the presence of spermatophores showed that mating usually occurred during the first 2 nights after emergence. Most of the females mated only once; multiple matings occurred in less than 20 per cent of the females examined. The most spermatophores found in one female was three. In several instances a male mated with two or three females. Moths mated at all experimental temperatures between 5° and 35°C. High temperatures appeared detrimental for mating as only one out of five females mated at 30° and 35°C, whereas in the range from 5 to 25°C, three or more out of five females usually mated. Continuous lighting deterred mating; at 25°C only one female out of eight was mated whereas in continuous darkness the proportion was 1:2 (Jacobson, 1965). The duration of copulation of the pale western cutworm is not known with any degree of certainty. Data on other noctuid species (Hardwick, 1965) indicate that the interval could extend from 45 minutes to 3 hours. Dissections of females collected from flowers in the field and capture in light traps dis- closed that 96 per cent had mated (Jacobson, 1965). Since mating occurs early in the life of a female and over a wide range of environmental conditions, it is concluded that the opportunity for mating is not a factor in oviposition by the pale western cutworm. Oviposition by the pale western cutworm in the field and laboratory has been studied for some time (Parker et al., 1921; Cook, 1930; Sorenson and Thornley, 1941). These authors found that oviposition occurred in the late afternoon and early evening, that moths laid the eggs in loose dusty soil, and that each female was capable of laying about 100 eggs. In the field eggs are difficult to find unless the exact site of oviposition is observed. Further knowledge was obtained from laboratory studies using moths that had been reared (Jacob- son, 1965). Eggs developed during the pupal stage and were ready for fertilization when the moths emerged. Oviposition began on the first or second day after mating, reached a peak soon after, and continued until just before the females died. Maximum oviposition by one female was 564 eggs. In 19 cages, each containing 4 to 18 females and as many or more males, mean numbers per female ranged from 161 to 488. Oviposition occurred at all tem- peratures from 5 to 35°C, the optimum temperature being between 10 and 25°C. More eggs were laid in the dark or in subdued light than in continuous bright light. Females when unfed could oviposit but when fed water or sugar solutions they laid more eggs. A diurnal rhythm of oviposition occurred with distinct peaks; 80 per cent or more of the eggs were laid between noon and early evening. Rhythm of oviposition was apparently initiated by light and could be reversed by reciprocal light regimens but was maintained in continuous dark or light (Jacobson, 1965). Sex ratio. - References to sex ratio (Cook, 1930; Eyer, 1957) indicate a preponderance Pale western cutworm 427 of males. Cook (1930), who counted the numbers of each sex on flowers over a period of several years determined that the ratio of males to females was three to two. Apparently he was not aware that the ratio may change with the time of day or other factors. Eyer (1957) used the numbers of each sex that were captured in light traps and found the ratio was six or seven males to each female. Light trap records over a number of years has shown that males are more readily attracted to lights, possibly because males are stronger fliers and greater numbers of them may be in flight during hours of darkness. Strickland (1922) found that when pans were baited in the field with a fermented molasses solution nearly 50 per cent of the captured moths were females. Counts of moths reared in the laboratory, both from larvae collected in the field and those reared from eggs in the laboratory, show that both sexes occur in almost equal numbers. REARING METHODS Procedures for conducting research on the pale western cutworm in the laboratory have undergone many changes during the course of investigations. Rearing methods were devel- oped that permitted the conduct of research on biology, physiology, nutrition, or insecti- cide testing with all stages at all times of the year. General methods for all stages have been described (Parker et al., 1921; Cook, 1930; Jacobson and Blakeley, 1957a). Methods for obtaining eggs and procedures for their use in investigations under various environmental conditions have been outlined (Lindsay, 1954; Jacobson and Blakeley, 1957b; Jacobson, 1962a, 1962b). The rearing of larvae for various purposes has been described by various authors; on various food plants (Seamans and Mc- Millan, 1935; Jacobson and Blakeley, 1958b; Hocking, 1953), on artificial diets (McGinnis and Kasting, 1959, 1960; Kasting and McGinnis, 1967), and for insecticide testing (Mc- Donald, 1969). One of the difficulties of mass rearing is that the larvae are cannibalistic (Dethier, 1939) and must be contained separately in suitable containers. Hence, the number that can be reared at a time is limited. The methods of containing pupae until emergence have been described (Blakeley, 1954; Blakeley and Jacobson, 1960). Moths have been used for various purposes: for oviposition (Jacobson and Blakeley, 1957b), for flight mill studies (Jacobson, 1965), and for light trap captures (Cook, 1930; Seamans and Gray, 1934). Many species of Noctuidae cannot be reared successfully in the laboratory beyond three generations (Hardwick, 1965) as vigour and viability progressively decline and susceptibility to laboratory disease increases. Similar results have been obtained when successive genera- tions of the pale western cutworm have been reared in the laboratory. It is, therefore, de- sirable to replace laboratory cultures with eggs obtained from females collected in the field each year if possible. NATURAL ENEMIES AND DISEASE Parasites Internal parasites of the larvae of the pale western cutworm have been recorded in the various areas of occurrence (Strickland, 1921; Parker et al., 1921; Cook, 1930; Seamans, 1931; Sorenson and Thornley, 1941; Walkden, 1950; Brooks, 1952). The biology of para- sitism was admirably described by Strickland (1923). Schaaf (1971) studied the parasitoid complex of Euxoa ochrogaster (Guenee) with emphasis on the identification of immature stages and review of biology. Many of the species noted also parasitize larvae of A. ortho- gonia. Dipterous parasites include the families of Ichneumonidae, Braconidae, and Chalci- didae. Table 4 shows the parasites recorded by various authors in their respective areas. 428 Jacobson Table 4. Recorded parasites of A. orthogonia. Parasite Record by Present status, notes Diptera: Tachinidae Bonne tia compta Fall. a,b,c,d,e Ernestia radicum Fab. b,d Pseudomeriania nigrocornea Tot. Gonia capitata DeG. b,d Gonia aldrichi Tot.; perhaps also G. longiforceps Tot. c,e longiforceps, breviforceps, and longipulvilli (D. M. Wood, correspondence) G. brevipulli Tot. g G. aldrichi Tot. e Metaphyto genalis Coq. a Peleteria robusta Wied. a,b,d Peleteria sp., perhaps anaxias Wagneria rohweri Tnsd. c (Wlk) or haemorrhoa (Wulp) (D. M. Wood, correspondence) Periscepia rohweri Tnsd. Diptera: Bombyliidae Anthrax molitor Loew. c Villa altemata (Say) h Villa willis toni (Coq.) g Poecilanthrax willistoni (Coq.) Poecilanthrax sackenii (Coq.) h Hymenoptera: Braconidae Meteorus vulgaris Cress. c,d,f Meteorus leviventris Wesmael Chelonus sp. d Zele sp. d Hymenoptera: Ichneumonidae Paniscus sp. b Apan teles griffini Vier. f Hymenoptera: Chalcididae Berecyntus bakeri (Howard) c Copidosoma bakeri (Howard) a — Parker et al., 1921; b — Strickland, 1923; c — Cook, 1930; d — Seamans, 1931; e — Sorenson and Thomley, 1941; f - Walkden, 1950; g - unpublished records; h — Brooks, 1952. Pale western cutworm 429 Predators Many species of insect predators have been observed attacking larvae and adults of the pale western cutworm. Species of predaceous wasps and larvae of ground beetles prey on cutworms in the field (Cook, 1930; Seamans, 1931). When moths are feeding on flowers they are often attacked by arthropod predators, which include various species of Arachnida, Mantidae, Coreidae, Reduviidae, and Phymatidae (Sorenson and Thornley, 1941 ; Seamans, 1931). Many native birds including larks, sparrows, buntings, crows, curlews, and Franklin’s gulls have been observed digging cutworms from the soil or carrying them to their nests; some have been seen capturing moths on flowers (Cook, 1930; Seamans, 1931; Sorenson and Thornley, 1941. Value of natural enemies The early workers with the exception of Strickland (1923) doubted that parasites and predators were important in the control of the pale western cutworm (Parker et al., 1921; Cook, 1930). Subsequent investigations showed that the incidence of parasites varied from year to year and was influenced by weather (Seamans, 1923, 1935). Rearing records show that parasitism can vary from 20 to 70 per cent. This indicates that when climatic condi- tions favour their increase, parasites, along with predators, can play an important role in quickly reducing populations of cutworms after years of severe outbreaks. Disease The role of disease in reducing populations of the pale western cutworm in the field has not been adequately assessed. Crumb (1929) listed several diseases that caused reductions in populations of tobacco cutworms, but as these are surface-feeding species the diseases that affect them may not affect the pale western cutworm, which spends most of its time in the soil. Steinhaus and March (1962) identified bacteria found in field-collected larvae. Cook (1930) postulated that disease may be a controlling factor. McMillan (unpublished report) recorded a disease found in one field that accounted for 47 per cent mortality, but this may have been the effect of climatic stress during years of a severe outbreak. Out- breaks of disease frequently occur when, larvae are reared in the laboratory but some doubt exists that similar outbreaks occur in the field. Walkden (1950) listed several disease organ- isms and noted that many larvae collected in the field for rearing died in the laboratory of a disease, which he named “the rearing disease”, that turned larvae black similar to dead larvae found in the field after a heavy rain. A disease of pale western cutworms reared in the laboratory was described by Kasting, McGinnis, and Hawn (1971). They called it “black disease” and identified the causative bacterium as Pseudomonas aeruginosa. RELATIONSHIP OF INFESTATION TO WEATHER The numbers of many animals are largely determined by weather. Andrewartha and Birch ( 1 954), after reviewing earlier publications (Parker et al., 1921; Seamans, 1 923 ; Cook, 1 924, 1926, 1930), used the pale western cutworm as an example of numbers of animals in natural populations. The various effects of precipitation on this insect and other crop insects were reviewed by Beirne (1970). Weather, distribution, and outbreaks Confinement of distribution of the pale western cutworm to the semiarid region of the plains in Canada and the United States indicates a relationship to the weather that prevails in these regions. Cook (1923, 1926, 1927a, 1927b, 1928, 1929, 1930) studied the physical 430 Jacobson ecology of Noctuidae of the American plains with special reference to A. orthogonia. He related distribution to weather by means of climographs for various regions and predicted areas in the United States where the pale western cutworm could be found occasionally and in outbreak numbers. His prediction of economic distribution conforms almost identically to that shown in Figure 1, which includes records to 1970, almost 50 years later. He found a high negative correlation between abundance of the pale western cutworm and years when rainfall in May, June, and July exceeded 5 inches (12.7 cm). Dry weather was favorable to increase. He further indicated that one favorable year may increase the number of cutworms to cause slight damage and local outbreaks but two successive favorable years were necessary to produce a severe and widespread outbreak. Forecasting The relationship of seasonal rainfall and possible outbreaks of the pale western cutworm led to the development of a method of forecasting. Seamans (1923, 1935) based the fore- cast on the assumption, confirmed by rearing records, that increased rainfall forced the larvae to feed and move about above ground where they were exposed to attack by para- sites. The basis of the prediction was the number of “wet days” — days on which 0.25 inches (0.64 cm) or more of rain fell — in May and June. More than 10 “wet days” resulted in a decrease in the numbers of cutworms, less than 10 were followed by an increase. In western Canada a forecast was prepared each year, delineating areas where a hazard was anticipated. This information was made available through the daily and weekly newspapers, radio and television, and extension agencies of government and industry. Accepted methods of preventing infestations by cultural means were given with the forecast. In Montana, Wall (1932) showed that the method of predicting infestations was accurate. Seamans (1923, 1935) considered parasites as the main factor in reducing populations during outbreaks when rainfall was above normal, whereas Cook (1930) indicated that fungous and bacterial diseases were equally or more important. In western Canada from 1945 to 1953 counts of moths were made on flowers during the peak of the flight in three selected locations. The mean number of moths per minute decreased in each location with an increase in the number of wet days during the previous May and June (r = -0.878). CONTROL Cultural Early attempts at cultural control involved the use of ploughs, packers, discers, and other available implements. It was found that packing and the use of a press drill limited the movement of larvae in the soil but did not achieve satisfactory results (Strickland, 1915); Parker et at. , 1921 ; Cook, 1930. The observation that moths could not deposit eggs in fields that were crusted because they had not been cultivated for some time before oviposition (Parker, Strand, and Seamans, 1920) led to a recommendation that infestation could be prevented by allowing a crust to form on the surface before the oviposition period. This method of prevention was the only control measure available for many years and was widely followed throughout the areas in the United States and Canada where the pale western cutworm was an economic pest. Surveys during outbreaks showed that damage was less, or none at all, in fields where the method was practised (McMillan, 1935; Seamans, 1952). This method was later augmented by cultural methods to starve early instars in the spring (Seamans, 1937, 1952) which represented the first control measure that could be used after infestations were found in the field. Since the method involved a delay of 10 days to 2 weeks after cultivation of new growth before seeding and an additional operation it was Pale western cutworm 431 not readily adopted by farm operators. Poisoned baits Another of the early attempts at control in the field made use of poisoned baits (Strick- land, 1915). Many of the available poisonous materials were added to various carriers, mainly bran, and spread on the soil (Parker et al. , 1921; Cook, 1930; Seamans, 1931). The consensus was that the baits were not effective (Cook, 1930; Seamans, 1931) mainly because of the subterranean feeding habits of the pale western cutworm. Chemical control When DDT and related organochlorine compounds came into general use for control of agricultural pests it was felt that sprays and dusts would not be effective, as the pale western cutworm fed almost exclusively below the soil surface. The first successful demon- stration of control in the field (Jacobson et al., 1952) showed that chlordane, dieldrin, aldrin, and, to a lesser extent, toxaphene, when applied as sprays to the soil surface could protect crops from damage. These materials were selected from results of laboratory assess- ment (Brown et al., 1947). Faulkner (1954) found, in laboratory tests, that some of the insecticides killed the embryos in unhatched eggs. Further testing in other areas confirmed these results and led to the general recommendation that dieldrin was the most effective material (Hoerner, 1953; Pfadt, 1956; Eyer, 1957; DePew and Harvey, 1957). Dieldrin was later replaced by endrin because of its effectiveness at low rates. When organochlorine com- pounds were found to present a residue hazard to livestock and humans other less persistent materials were sought. Field testing showed that several organophosphorus insecticides, including AC-47031 [cyclic ethylene (diethoxyphosphinyl) dithioimidocarbonate] and fen- sulfothion (dasanit) were as effective as endrin (Jacobson and McDonald, 1966). Additional tests in the field showed that AC-4703 1 and monocrotophos (azodrin) were slightly superior to endrin (DePew, 1970). Laboratory testing further confirmed the effectiveness of AC- 47031 and indicated that other organophosphorus insecticides could be used to control the pale western cutworm (McDonald, 1969). The use of insecticides provides a further advantage to the farmer. When these materials were not used, fields that had been destroyed could not be reseeded until the larvae had ceased feeding. The plantings, delayed until late in June in western Canada, were often damaged by frost or snow before they were ready for harvesting. Damaged fields can be reseeded immediately after treatment with an insecticide. Future control methods The impetus in the search for alternative methods for controlling insects is toward the use of methods other than the application of insecticides because of the problems that result from the accumulation of residues, the possibility that insects have become resistant to insecticides, and that insecticides contaminate the environment. Some of the alternatives might include microbial pesticides, natural or synthetic pheromones to attract adults to be killed with poisons or other means, the use of antifeeding compounds, the breeding of resistant crops, chemosterilants, and integrated control embodying several methods. ACKNOWLEDGEMENTS I express my gratitude to colleagues, present and past, at the entomological laboratories at Lethbridge, Alberta; Saskatoon, Saskatchewan; and Brandon, Manitoba, who have assis- ted in the furtherance of knowledge of the pale western cutworm. I especially acknowledge 432 Jacobson the early entomologists, E. H. Strickland (deceased) and H. L. Seamans (retired). Their notes and unpublished reports were constantly referred to. I also acknowledge with thanks the assistance of the following in the preparation of this publication: D. M. Wood, W. R. Mason, C. M. Yoshimota, G. E. Shewed, and D. F. Hard- wick of the Entomology Research Institute, Ottawa, for advice in systematics of Agrotis orthogonia Morrison and its parasites; D. K. Edwards, Forest Research Laboratory, Victoria, British Columbia, for his assistance in the activity studies of the adults; A. J. McGinnis and R. Kasting for the summary of their nutrition studies on A. orthogonia at this laboratory; and finally, N. D. Holmes, head of the Entomology Section of this Station for his invaluable assistance at various stages of preparation of the manuscript. REFERENCES Andrewartha, H. G. and L. C. Birch. 1954. The distribution and abundance of animals. Univ. Chicago Press, pp. 594-600. Barnes, W. and F. H. Benjamin. 1926. Notes and new species (Lepid., Phalaenidae). Can. Ent. 58:303. Beirne, B. F. 1970. Effects of precipitation on crop insects. Can. Ent. 102:1360-1373. Blakeley, P. E. 1954. Certain aspects of the biology and behaviour of prepupae, pupae, and adults of the pale western cutworm, Agrotis orthogonia Morr. M. 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Seamans, H. L. 1923. Forecasting outbreaks of the pale western cutworm in Alberta. Can. Ent. 55:51-53. Seamans, H. L. 1926. The pale western cutworm. Can. Dep. Agric. Pam. 71. Seamans, H. L. 1931. The pale western cutworm. Unpublished manuscript. On file, Regional Library, Can. Dep. Agric. Res. Sta., Lethbridge. Seamans, H. L. 1933. A preliminary study of the chorion design in the micropyle area of noctuid eggs. M. Sc. thesis. Univ. Saskatchewan. Seamans, H. L. 1934a. The influence of cultural practices on field crop insects. 65 Ann. Rep. ent. Soc. Ont. pp. 22-28. Seamans, H. L. 1934b. Entomological problems of wheat growing in Canada. Proc. fifth Pacif. Sci. Congr. 1933. pp. 3473-3481. Seamans, H. L. 1935. Forecasting outbreaks of the pale western cutworm ( Agrotis ortho- gonia). J. econ. Ent. 28:425-428. Seamans, H. L. 1937. A probable cultural control for the pale western cutworm, Agrotis orthogonia Morr. Sci. Agric. 17:612-615. Seamans, H. L. 1938. The pale western cutworm and its control. Can. Dep. Agric. Pub. 615 (Farm Bull. 59). Seamans, H. L. 1952. The control of the pale western cutworm. In Agricultural Research in Canada, Pub. by Agric. Inst, of Canada. R. Newton (ed.). pp. 138-142. Seamans, H. L. and H. E. Gray. 1934. Design of a new type of light-trap to operate at controlled intervals. 25th-26th Ann. Rep. Queb. Soc. Prot. PI. (1932-1934), pp. 39-46. Seamans, H. L. and Ellis McMillan. 1935. The effect of food plants on the development of the pale western cutworm ( Agrotis orthogonia Morr.). J. econ. Ent. 28:421-425. 436 Jacobson Seamans, H. L. and P. J. G. Rock. 1945. Starvation of the early instars of the pale western cutworm, Agrotis orthogonia Morr., and its use in the control of this pest. Can. Ent. 77:57-61. Smith, J. B. 1890. Revision of the species of genus Agrotis. U. S. nat. Mus. Bull. 38. p. 129. Smith, J. B. 1908. New species of Noctuidae for 1908. J. N.Y. ent. Soc. 16:87. Sorenson, C. J. and H. F. Thornley. 1941. Pale western cutworm. Utah Agric. Exp. Sta. Bull. 297. Steinhaus, E. A. and R. B. March. 1962. Report of diagnosis of diseased insects 1951-1961. Hilgardia 33:349-490. Strickland, E. H. 1915. Poisoned bait for cutworms. Can. Ent. 47:201-204. Strickland, E. H. 1921. Parasites of the pale western cutworm. Can. Ent. 53:97-100. Strickland, E. H. 1922. Poisoned molasses for the destruction of Noctuid moths. J. econ. Ent. 15:214-220. Strickland, E. H. 1923. Biological notes on parasites of prairie cutworms. Can. Dep. Agric. Bull. 26. 40 pp. Strickland, E. H. 1948. Insect pests of grain in Alberta. Univ. Alberta Fac. Agric. Bull. 24. (Revised). 68 pp. Various authors. 1952-1970. U. S. Dep. Agric. A.R.S. Co-operative Economic Insect reports. Walkden, H. H. 1940. Control of the pale western cutworm in the southern great plains region. U. S. Dep. Agric. E-519. 6 pp. Walkden, H. H. 1950. Cutworms, armyworms, and related species attacking cereal and forage crops in the central great plains. U. S. Dep. Agric. Circ. 849. Wall, R. E. 1932. Prediction of pale western cutworm increase in Montana is fulfilled. J. econ. Ent. 25:1043-1048. Webster, R. L. 1924. Pale western cutworm, pp. 45-48. In Report of Director 1921-1923. N. D. Agric. Coll. Agric. Exp. Sta. Bull. 174. Webster, R. L. and C. N. Ainslie. 1924. Pale western cutworm in North Dakota. N.D. Agric. Coll. Agric. Exp. Sta. Bull. 179. PURIFICATION AND PROPERTIES OF ARGININE PHOSPHOKINASE FROM HONEYBEES APIS MELLIFERA L. (HYMENOPTERA, APIDAE) A. C. CHEUNG Department of Entomology University of Alberta Quaestiones entomologicae Edmonton 7, Alberta 7 : 437-446 1971 Arginine phospho kinase was purified from honeybee thoraces. Its molecular weight was estimated by Sephadex gel chromatography at about 36,000. In the direction of arginine phosphate synthesis, the enzyme had a pH optimum around 8.3. The energy of activation for the reaction from 22-45 C was 7,000 cal/mole. Optimum molar ratio ofMg++:ATP appeared to be 1:1. Besides Mg++, the enzyme was activated to various extents by Mn++, Ca++, Co++, and Cu++. UTP, GTP, CTP, and ADP could not substitute for ATP as substrate. The enzyme phosphorylated L-arginine methyl ester and to a much less extent D-arginine, but did not phosphorylate creatine, guanidoacetic acid, nor hippuryl-L-arginine. L ’arginine phosphokinase a ete purifiee a partir de thoraxes dabeilles. Son poids mole- culaire a ete estime par chromatographie sur gele Sephadex a environ 36,000. Dans la reaction donnant la synthese de Targinine phosphate, Venzime a un pH optimum aux environs de 8.3. L’energie dactivation pour la reaction de 22-45 C etait 7,000 cal./mole. Le rapport molaire optimum de Mg++. ATP apparait etre 1:1. En plus de Mg++, Venzime a ete activee a differents degres par Mn++, Ca++, Co++, et Cu++. UTP, GTP, CTP, et ADP ne peuvent pas remplacer VATP comme substrat. L’enzime phosphorilate L-arginine methyl ester et a un degre moins eleve, la D-arginine, mais ne phosphorilate pas la creatine, Vacide puanido-acetique, et la hippuryl-L-arginine. Using crab-muscle extracts, Lohmann (1935) first discovered the enzyme arginine phos- phokinase (EC 2.1 33) which catalyzed the reaction: ATP + arginine ^ ADP + arginine phosphate + H+ Since then other workers have described the enzyme from various invertebrates. Arginine phosphokinase has been purified or identified from extracts of the following species: fresh- water crayfish Potamobius astacus, and P. leptodactylus (Elodi and Szorenyi, 1956), sea crayfish Jasus verreauxi (Morrison et al., 1957; Uhr et al. , 1966), shrimps Palaemon serratus, P. elegans (Virden and Watts, 1964), crabs Pagurus bernhardus, Callinectus sapidus ( Blethan and Daplan, 1968 ), Atelecyclus septemdentatus, Cancer pagurus, Por tunas depurator, Carci- nus maemas, Maia squinado (Virden and Watts, 1964), lobsters Homarus vulgaris (Pradel et al . , 1964; Virden et al., 1965), H. americanus (Blethan and Kaplan, 1967; Regnouf et al., 1969), and Nephrops norwegicus (Virden and Watts, 1964), molluscs Pecten maximus, and Chlamys opercularis (Virden and Watts, 1964), the horseshoe crab Limulus polyphemus (Blethan and Kaplan, 1968), echinoderms Centrostephanus rodgersii, Heliocidaris erythro- gramma (Griffiths et al., 1957a), and Echinus esculentus, Holothuria forskali, and Asterias rubens (Virden and Watts, 1964), the tunicate Styella mammiculata (Virden and Watts, 1964), the cephalochordate Amphioxus lanceolatus (Virden and Watts, 1964), arachnids Pholcus phalangioides , and Dugesiella hentzi (Blethan and Kaplan, 1968), the annelid Sipun- culus nudus (Regnouf et al., 1969), insects Melanoplus bruneri, Apis mellifera, Porthetria dispar, Sympetrum rubicundulum , (Blethan and Kaplan, 1968), and Calliphora erythroce- phala (Lewis and Fowler, 1962), protozoans Tetrahymena pyriformis (Robin and Viala, 1966; Watts and Bannister, 1970), and Stentor coeruleus (Watts et al. , 1968), and from the bacteria Escherichia coli (Di Jeso, 1967). In the study of the properties of this enzyme most workers used enzymes extracted from 438 Cheung crustaceans. Little work has been done on the enzyme from insect sources. Recently Carlson et al. (1971) reported the crystallization of arginine kinase from honeybee thoraces. These workers indicated that the physical, chemical, and catalytic properties of the enzyme were being studied. So far there has been no report on the properties of an insect APK. I report a method of purifying arginine phosphokinase from honeybee thoraces and the investiga- tions on some properties of the enzyme in the direction of arginine phosphate synthesis. The following abbreviations are used: arginine phosphokinase, APK; arginine phosphate, AP; adenosine triphosphate, ATP; uridine triphosphate, UTP; guanosine triphosphate, GTP; cytidine triphosphate, CTP; adenosine diphosphate, ADP; diethyl aminoethyl cellulose, DEAE-cellulose; disodium ethylenediamine-tetraacetate, EDTA; and Tris (Hydroxymethyl) aminomethane, Tris. MATERIALS AND METHODS Experimental animals Honeybee workers ( Apis mellifera L.) were obtained from a local apiary and frozen until use. Sources of chemicals ATP and L-arginine came from both Sigma Chemical and Calbiochem. UTP, CTP, GTP, ADP, L-arginine methyl ester, hippuryl-L-arginine, guanidoacetic acid, creatine, cytochrome c, Tris, and DEAE-cellulose came from Sigma Chemical. D-arginine, myoglobin, haemo- globin, and bovine albumin came from Nutritional Biochemicals. L-cysteine HC1, 2-mercap- toethanol, reduced glutathion, 7-globulin, and Aquacide I came from Calbiochem. MgS04, MnS04, CuS04, CoCl2, CaCl2, ammonium molybdate, l-amino-2-naphthol-4-sulfonic acid, and EDTA came from Fisher Scientific. Assay procedure The activity of the enzyme was estimated by measuring the inorganic phosphate released after acid hydrolysis of arginine phosphate. The assay procedure was modified from that of Morrison et al. (1957). The reaction mixture contained a final concentration of 50 mM Tris, 1 mM 2-mercaptoethanol, 5 mM ATP, and 10 mM each of arginine and MgS04, pH 8.3. In a typical assay, 0.9 ml of the stock solution was incubated at 30 C for 5 min, the reaction was started by adding 0.1 ml APK solution. The reaction was stopped after 5 min by adding 0.5 ml 30% acetic acid. The solution was placed in boiling water for exactly 1 min, after which it was immersed in an ice-bath. The colorimetric determination of inorganic phos- phate was started by adding 2.0 ml 5% w/v ammonium molybdate in 15% v/v H2S04 followed by 0.5 ml 0.25% aminonapthol sulfonic acid half a minute later. The mixture was diluted with 5 ml glass distilled water. The absorbance of the resulting blue solution was read at 540 nm in a Beckman DU-2 spectrophotometer after 20 min. Controls were run in the same way except that acetic acid was added before adding the enzyme. Enzyme activity was measured as the difference in absorbance between the 5 min assay and the control. When working with the crude extracts and various (NH4)2S04 fractions, correction for ATPase activity was made by subtracting the change in absorbance without arginine in the stock solution. By using a standard curve prepared with various amounts of inorganic phos- phate, enzyme velocity was converted to jumoles arginine phosphate synthesized per min. For higher temperatures or lower substrate concentrations, the duration of the assay was reduced to ensure that only the linear portion of the reaction velocity was measured. Protein concentrations were determined spectrophotometrically according to the method of Layne (1957). Honeybee arginine phosphokinase 439 RESULTS Purification of arginine phosphokinase Purification procedure was carried out at 0-4 C. The buffer used was 10 mM Tris, 5 mM EDTA, pH 7.0, unless stated otherwise. Extraction. — Bee thoraces weighing 50 g were homogenized in 150 ml Tris buffer with an omni-mixer for 3 min and centrifuged at 10,000 g for 30 min. The supernatant was saved. The above procedure was repeated on the precipitate with 100 ml buffer. The supernatants were combined. Ammonium sulfate fractionation. — Granular (NH4)2S04 was added to the combined supernatant until 60% saturated. The precipitate formed after centrifugation at 10,000 g for 30 min was discarded. More (NH4)2S04 was added to the supernatant until 80% satu- rated and the precipitate collected after centrifugation at 10,000 g for 30 min was saved. Precipitate from this (NH4)2S04 fraction gave the highest specific activity reading. The reading was much lower than expected, probably due to the high concentration of S04 = ion which was a potent inhibitor of creatine phosphokinase (Noda et al., 1960). Chloride, nitrate, and acetate as the sodium and potassium salts inhibited arginine phophokinase from lobster (Virden et al. , 1 965). Sephadex G-100 chromatography. — The precipitate from 80% (NH4)2S04 fraction was dissolved in Tris buffer and put through a Sephadex G-100 column (1.4 x 102 cm) equili- brated with 50 mM Tris, 5 mM EDTA, 100 mM KC1, pH 7.0 buffer. The column was eluted with the same buffer. Protein concentration of the eluent was estimated by measuring the absorbance at 280 nm and APK activity was assayed in the direction of AP synthesis as described under assay method. DEAE-cellulose chromatography. - The solution from Sephadex chromatography was dialyzed against two changes of Tris buffer overnight. The dialyzed solution was pumped through a DEAE-cellulose column (2.5 x 24 cm) equilibrated with Tris buffer. The column was eluted by 0.1, 0.2, 0.4 M NaCl, and 30% (NH4)2S04 in Tris buffer. The enzyme appeared shortly after 0.1 M NaCl in Tris buffer was pumped into the column. The enzyme solution obtained was free of ATPase activity. A final concentration of 1 mM 2-mercapto- ethanol was added to the enzyme solution. A summary of the data is listed in Table 1. Table 1. Purification of arginine phosphokinase from honeybee thoraces. Weight of tho- races, 50 g. Step Vol. (ml) Protein (mg) Sp. Act. (/rnioles/ min/mg protein) Purifi- cation Total Act. (jumoles / min) Yield (%) Crude extract 222 910 0.29 1 264 100 (NH4)2S04 fractionation 11.4 340 0.08* - - - Sephadex G-100 47.5 95 1.83 6.3 174 66 DEAE-cellulose 125 5 33.7 116 167 63 * See text for the explanation of the exceptionally low specific activity. 440 Cheung Estimation of molecular weight by Sephadex G- 100 gel chromatography A Sephadex G-100 column (1.4 x 102 cm) was calibrated according to the method of Andrews (1964). The proteins used were 7-globulin, bovine albumin, haemoglobin, myo- globin, and cytochrome c. The molecular weight of the enzyme estimated by this method is 36,000 ± 3,000 (Fig. 1). Electrophoresis of APK The APK solution was concentrated by Aquacide I to a concentration of 9.6 mg/ml. About 5 /d was applied to each Sepraphore III cellulose polyacetate strip (1” x 6%”) and electrophoresed at a constant current of 1 ma/strip for 65 min. The buffer used was 10 mM Tris, 1 mM EDTA, 1 mM 2-mercaptoethanol, at pH’s 9.0, 7.0, and 4.0. At these three pH’s the enzyme migrated as a single protein band, APK activity coincided with the band. Protein was stained with Ponceau S. Effect of pH on APK activity The assays were done in 50 mM Tris, 100 mM bicarbonate buffer. Maximum activity oc- curred around pH 8.3. At pH’s higher than 9.0, the enzyme activity declined rapidly (Fig. 2). Effect of enzyme concentration on velocity The reaction velocity was directly proportional to enzyme concentration over the con- centration range studied (Fig. 3). The reaction time used was 5 min for APK concentrations up to 4 jug/ nil; above this the reaction time was 2.5 min. This eliminated the chance of measuring the non-linear portion of the reaction velocity. Temperature stability of the enzyme Aliquots of the enzyme solution were incubated at various temperatures for 1 5 min and then assayed at 30 C for 15 min. There was no loss of activity up to 40 C, from 40-45 C activity declined slightly, and beyond 45 C a sharp decline in activity was observed (Fig. 4). Effect of temperature on velocity To ensure only the initial velocities were measured, the assays were shortened as the temperature was increased. The durations of the assays for 22, 29, 34, 39, 45, and 50 C were 5', 5', 3', 2 ', 1', and 1', respectively. Velocity increased from 22-45 C, beyond this temperature range velocity declined (Fig. 5). When the reciprocals of absolute temperatures for the range of 22-45 C were plotted against the logarithm of velocity, a linear Arrhenius plot was obtained (Fig. 6). The activation energy calculated from the slope of the graph was 7,000 cal/mole. Effect of sulfhydryl compounds Addition of sulfhydryl compounds to the assay solution increased the enzyme activity by as much as 33%. Cysteine, 2-mercaptoethanol, and reduced glutathion all had similar effects. The activation by 2-mercaptoethanol is shown in Fig. 7. Activation by various bivalent cations The metal salts used were either sulfates or chlorides. No activity was detected without adding bivalent cations. Addition of 10 mM of Mg++, Mn++, Cu++, Ca++, and Co++ activated the enzyme to various extents. The enzyme was more active with Mn++ than with Mg++; Cu++, Ca++, Co++ activated the enzyme to a much less extent. These results are listed in Table 2. Honeybee arginine phosphokinase 441 © 8 pH 10 Fig. 1. Estimation of the molecular weight of honeybee APK by Sephadex G-100 gel chromatography. The proteins used and their molecular weights were: 1. 7-globulin, 160,000; 2. bovine albumin (dimer), 134,000; 3. bovine albumin (monomer), 67,000; 4. haemoglobin, 64,500; 5. myoglobin, 17,800; 6. cytochrome c, 12,400. The molecular weight of APK estimated by this method is 36,000 ± 3,000. Fig. 2. Effect of pH on reaction velocity of honeybee APK. The buffer used was 50 mM Tris, 100 mM bicar- bonate at various pH’s. Optimum activity occurred around pH 8.3. Te mp. Fig. 3. Effect of enzyme concentration on the reaction velocity of honeybee APK. Reaction time was 5 min up to 4 ng APK added, above 4 pg, reaction time was 2.5 min. Fig. 4. Temperature stability of honeybee APK. Aliquots of the enzyme were incubated at various temperatures for 15 min and then assayed at 30 C for 15 min. 442 Cheung Fig. 5. Effect of temperature on reaction velocity of honeybee APK. Duration of the assays were progressively decreased as temperature was increased so that only the linear portion of the reaction velocity was measured. APK: 1.3 Mg/assay. Fig. 6. The Arrhenius plot of the data from Fig. 5. T = absolute temperature; v = /imoles/min. APK: 1.3 /ig/assay. Fig. 7. Effect of 2-mercaptoethanol on reaction velocity of honeybee APK. With sulfhydryl-free APK and reaction solution, addition of 2-mercaptoethanol to the reaction solution increased the enzyme activity up to 33% of its original activity. APK: 1.3 /zg/assay. Honeybee arginine phosphokinase 443 Table 2. Activation of APK by metal ions. Conditions as described in assay procedure except that 10 mM of the following ions was used in place of Mg++. APK: 1.3 jug/assay. Metal ion Enzyme activity (jumoles/min) Percentage activity Mn++ 0.082 111 Mg++ 0.074 (100) Ca++ 0.017 23 Co++ 0.010 13.5 Cu++ 0.006 8.1 None 0 0 Effect of varying magnesium concentration on velocity The effect of increasing magnesium concentration on velocity was studied with three levels of ATP concentration (Fig. 8). The maximum velocity was reached when the molar ratio of Mg++ : ATP was one. Increase in the ratio caused a slight decline in velocity. Similar results were obtained when Mg++ was replaced by Mn++. The Lineweaver-Burk plot of the reaction velocities against ATP concentrations at 5 mM and 10 mM Mg++ indicates that the inhibition by Mg++ was competitive. Specificity of arginine phosphokinase When ATP was replaced by the same concentration of UTP, CTP, GTP, or ADP, no activity was observed. The ability of the enzyme to phosphorylate several guanidino compounds was deter- mined. The final concentration of the guanidines in the reaction mixture was 10 mM. The results of the experiment are shown in Table 3. Both L-arginine methyl ester and D-arginine served as substrates to a limited extent. Fig. 8. Effect of magnesium concentration on reaction velocity of hoenybee APK at three ATP concentrations. Reaction time, 5 min; APK: 1.3 g/assay. 444 Cheung Table 3. Specificity of APK: phosphorylation of guanidines. Conditions as described in assay procedure except that 10 mM of the following guanidines was used in place of L-arginine. APK: 2.6 jig/assay. Guanidines Enzyme activity (pimoles/min) Percentage activity L-arginine 0.13 (100) L-arginine methyl ester 0.056 43 D-arginine 0.016 12.3 Guanidoacetic acid 0 0 Hippuryl-L-arginine 0 0 Creatine 0 0 DISCUSSION Using sedimentation and diffusion experiments, Elodi and Szor6nyi (1956) estimated the molecular weight of APK from Potamobius astacus to be 43,000. Virden et al. (1966) concluded from estimates obtained with ultracentrifuge analysis, gel filtration, and density- gradient centrifugation, that the molecular weight of the enzyme from Homarus vulgaris was 37,000. Blethan and Kaplan (1968) estimated the molecular weights of APK from sev- eral arthropods by gel chromatography to range from 35,000 ± 2,000 to 38,000 ± 2,000. Moreland and Watts (1967) discovered the existence of two forms of APK in some molluscs; one with a molecular weight of 40,000 and the other 80,000. From the distribution of the isoenzymes in different muscle tissues, they suggested that the different forms of enzyme were associated with different muscle functions and structures. Regnouf et al. (1969) had shown that APK from Homarus vulgaris had a molecular weight of 43,000 and consisted of a single polypeptide chain, whereas APK from the annelid Sipunculus nudus with molecular weight of 86,000 was a dimer. Oriol et al. (1970) again showed that APK’s from lobster and crab with molecular weights of about 40,000 were monomers. Robin et al (1969) identified an APK from the polychaetes Sabella pavonina and Spirographis spallanzanii with a molecular weight of 160,000. Thus, various workers have shown that there are at least three forms of APK in invertebrates, a monomer with a molecular weight of about 40,000, a dimer, and a tetramer, with corresponding molecular weights. In the present study, both gel chromatography and electrophoresis indicated that only one form of APK was present in honeybee thoraces, and from gel chromatography, the molecular weight of the enzyme was estimated to be 36,000 ± 3,000. The honeybee enzyme has a pH optimum around 8.3. The activity declines sharply at pH’s higher than the optimum and becomes insignificant beyond pH 9.5. Virden et al. (1965) had similar results with APK from Homarus vulgaris. However, Morrison et al. (1957) reported a pH optimum of 8. 4-8. 5 with APK from Jasus verreauxi with a much broader pH tolerance. Whether this difference in pH tolerance is due to differences between the enzymes or to experimental conditions is still to be investigated. As in Homarus vulgaris (Virden et al., 1965), APK from honeybees is activated by Ca++ and Co++. The enzyme from sea crayfish was not activated by either Ca++ or Co++ (Morrison et al., 1957). Honeybee arginine phosphokinase 445 The enzyme is quite specific with respect to the nucleotide substrate. ATP cannot be substituted by UTP, CTP, GTP, or ADP. It is less specific with the guanidino substrate. The enzyme is able to phosphorylate L-arginine methyl ester and to a much less extent, D-arginine. APK from Sabella pavonina was reported to show significant activity with D- arginine (28% of the activity with L-arginine) and those from Maia squinado, Eupagurus bernhardus, Pecten maximum, Polycelis cornuta, Myxicola infundibulum, and Holothuria forskali were also reported to have some activity with D-arginine (1-7% of the activity with L-arginine) (Virden and Watts, 1964). Without added metal ions, no activity was observed. A similar finding was reported by Virden et al. (1965) with lobster enzyme, and a trace of activity was reported by Morrison et al. (1957) with crayfish enzyme. Optimum molar ratio of Mg++ : ATP appeared to be 1:1, similar to the findings of Griffiths et al. (1957b). An increase in the ratio led to a slight decline in activity. Preliminary studies of initial velocity and product inhibition indicated that the reaction mechanism is random sequential. ACKNOWLEDGEMENTS I thank R. H. Gooding for his excellent advice on the research. I also thank B. B. Chiolino and R. H. Gooding for their comments on the manuscript. The research for this paper was supported (in part) by the Defence Research Board of Canada, Grant 6801-41. REFERENCES Andrews, P. 1964. Estimation of the molecular weights of proteins by Sephadex gel-filtra- Biochem. J. 91:222-233. Blethan, S. L. and N. O. Kaplan. 1967. Purification of arginine kinase from lobster and a study of some factors affecting its reactivation. Biochemistry 6:1413-21. Blethan, S. L. and N. O. Kaplan. 1968. Characteristics of arthropod arginine kinases. Bio- chemistry 7:2123-35. Carlson, C. W., S. C. Fink and R. W. Brosemer. 1971. 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C. Watts and E. Baldwin. 1965. ATP : arginine phosphotransferase from lobster muscle; purification and properties. Biochem. J. 94:536-544. Virden, R., D. C. Watts, R. C. Watts, D. B. Gammack and J. H. Raper. 1966. ATP-arginine phosphotransferase from lobster muscle. Molecular weight. Biochem. J. 99:155-158. Watts, D. C. and L. H. Bannister. 1970. Location of arginine kinase in the cilia of Tetra- hymena pyriformis. Nature 226:450-451. Watts, D. C., B. Moreland, E. C. Tatchell and L. H. Bannister. 1968. A particulate ATP- arginine phosphotransferase in the protozoan Stentor coeruleus. Comp. Biochem. Physiol. 25:553-558. Publication of Quaestiones Entomologicae was started in 1965 as part of a memorial project for Professor E. H. Strickland, the founder of the Department of Entomology at the University of Alberta in Edmonton in 1922. 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