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Quaestiones

Entomologicae

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

VOLUME XIII

1977

11

CONTENTS

Editorial — Comments About a Few Words in the Biological Sciences 1

Rempel, Heming and Church — The Embryology of Lytta viridana LeConte

(Coleoptera: Meloidae). IX. The Central Nervous System, Stomatogastric

Nervous System, and Endocrine System 5

Nimmo — The Adult Trichoptera (Insecta) of Alberta and Eastern British Columbia,

And Their Post-Glacial Origins. I. The Families Rhyacophilidae and

Limnephilidae. Supplement 1 25

Nimmo — The Adult Trichoptera (Insecta) of Alberta and Eastern British Columbia,

And Their Post-Glacial Origins. II. The Families Glossosomatidae and

Philopotamidae. Supplement 1 69

Belicek — Corrigenda on Coccinellidae of Western Canada and Alaska with Analyses

of the Transmontane Zoogeographic Relationships Between the Fauna of British

Columbia and Alberta (Insecta: Coleoptera: Coccinellidae) 73

Book Review — Matsuda, R. 1976. Morphology and Evolution of the Insect

Abdomen 75

Evans — Geographic Variation, Distribution and Taxonomic Status of the Intertidal

Insect Thalassotrechus barbarae (Horn) (Coleoptera: Carabidae) 83

Mendez — Mammalian-Siphonapteran Associations, The Environment, and Biogeography

of Mammals of Southwestern Colombia 91

Sengupta — Changes in Acetycholinesterases and Cholinesterases During Development

of Aedes aegypti (L.) (Diptera, Culicidae) 183

Book Review — Horn, D.J. 1976. Biology of Insects 191

Craig — Mouthparts and Feeding Behaviour of Tahitian Larval Simuliidae (Diptera:

Nematocera) 195

Fredeen — A Review of the Economic Importance of Black Flies (Simuliidae) in

Canada 219

Halffter — Evolution of Nidification in the Scarabaeinae (Coleoptera, Scarabaeidae) ... 231

Picchi — A Systematic Review of the Genus Aneurus of North and Middle America

and the West Indies (Hemiptera: Aradidae) 255

Evans & Baldwin — Larval Exuviae of Attagenus bicolor von Harold (Coleoptera:

Dermestidae) from an Archeological Site at Mesa Verde, Colorado 309

Book Review — Wiggins, G.B. 1977. Larvae of the North American Caddisfly genera

(Trichoptera) 311

Steiner — Observations on Overnight Perch Constancy by a Female Digger Wasp,

Ammophila azteca Cameron (Hymenoptera: Specidae), In Captivity 315

Fredeen — Black Fly Control and Environmental Quality with Reference to Chemical

Larviciding in Western Canada 321

Griffiths — Studies on Boreal Agromyzidae (Diptera). XIII. Some Phytomyza and

Chromatomyia Miners on Cichorieae (Compositae) 327

Reichardt — A Synopsis of the Genera of Neotropical Carabidae (Insecta:

Coleoptera) 346

Editor’s Acknowledgements 495

Ill

CORRIGENDA: Quaestiones Entomologicae, Volume 13

Fredeen, F.J.H. A Review of the Economic Importance

308/ 1 change “hispaniolensis” to “hispaniolensis’

Griffiths, G.C.D. Studies on Boreal Agromyzidae

IV

INDEX

Abbott, D.P. ,84, 85,87

Acacia melanoceras, 104

Acarina, 182

Acer rubrum, 282

acety cholinesterases, 75, 183—189

A chat o carp us n igri ca ns, 102

Acrididae, 77

acropedes group, Rhyacophila, 27, 28

acropedes, Rhyacophila, 28

Adoratopsylla, 173

Adoratopsylla intermedia copha, 1 16,

117, 122, 166

Adoratopsylla (T.) intermedia, 166, 172,

174, 176

Adoratopsyllini, 117

aduncun, Piper, 104

Aedes aegypti, 75, 183-189, 210

aegypti, Aedes, 75, 183—189. 210

Aepopsis robinii, 84

Aepus marinus, 84

aequalis, Dolophilodes, 69

Agonini, 87

Agouti paca, 112, 142, 158, 166

Agouti paca quanta, 110

Agromyzidae, 327, 338, 339, 340, 341

aibonitensis, Aneurus, 255, 256, 263, 266,

268, 279, 289, 290, 291, 292, 293, 295

308

Akodon, 117, 140, 141, 176

Akodon pulcherrimus inambari, 141

Akodont, 132

alascense, Chilostigma, 46

alascensis, Chilostigma, 46

alascensis, Halesus, 46

alascensis, Platyphylax, 46

alascensis, Psychoglypha, 25, 65, 46, 47,

49

alba, Philo casca, 25, 45, 49, 50, 65, 67

albicans, Miconia, 105

albiceps, Phytomyza, 321 , 328, 329, 332,

334,337,338

albiforum, Hieracium, 333

albigularis, Oryzomys, 91, 110, 117, 124,

127, 131, 132, 139, 140, 141, 166

alfaroi, Oryzomys, 117, 132, 142, 166

allamandi, Galictus, 142

Allen, J.A., 113, 178

Allium cepa, 105

almifolia, Turnera, 102

Alouatta, 243

alpicola, Ixeris, 336

alpina, Cicerbita, 330, 331

alvatus, Limnephilus, 25, 27, 39, 40, 48, 50

51,63,67

Alysiinae, 339

americana, Mazama, 112, 113

americana, Periplaneta, 1 5

americana, Persea, 104

ammonites, 97

Ammophila azteca, 315, 316, 318, 320

breviceps, 319

pubescens, 318

amphibians, 1 1 1

amplexicaule, Hieracium, 331

Anabolia fusorius, 42

modesta, 42

Anabolia ( Asynarchus) fusorius, 42

Anacardium excelsum, 104

anatis, Leucocytozoon, 226, 229

Ancistropsyllidae, 180

Anderson, J.R., 226, 228

Anderson, N.H., 28, 34, 37, 53

Anderson, R.C., 226, 228

andinus, Eptesicus, 111

andinus, Sylvilagus, 112,

Andosols, 100

Andres, V., Jr., (See Ellman, G.L.), 184, 186

Andropogon sp., 104

Anduaga, S. (see Halffter, V.), 238, 253

Aneuridae, Aneurinae, 288

Aneurosoma, 255, 270

Aneurus aibonitensis, 255, 256, 263, 266, 268,

277, 289, 290, 291, 292, 293, 295, 308

arizonensis n. sp., 255, 263, 266, 269, 275,

279. 280. 289. 290. 291. 292. 293. 295,

298,301,306

barberi, 263, 268, 289, 290, 291, 292, 298,

301, 308

borealis, n. sp., 255, 263, 266, 270, 282, 289,

290. 291. 292. 293. 295, 298, 301, 308

championi, 262, 266, 268, 278, 289, 290,

291, 292, 293, 296, 298, 301, 307

deborahae, n. sp., 255, 263, 266, 269, 274,

275, 289, 290, 291, 292, 298, 301 306

V

Aneurus dissimilis, 255, 262, 263, 265, 266,

267, 270, 276, 289, 290, 291, 292, 293,

296,299,301,307

fiskei, 262, 266, 267, 269, 280, 281, 282,

289. 290. 291. 292, 293, 296, 299,

301. 306

froeschneri, 263, 265, 266, 269, 280, 289,

290. 291. 292, 293, 296, 299, 301, 307

gallicus, 288

haitiensis, 255, 256, 263, 266, 268, 273,

274, 289, 290, 291, 293, 296, 299,

302, 308

hispaniolensis n. sp., 255, 256, 263, 265,

266, 268, 273, 274, 289, 290, 291,

292, 299, 302, 308

hrdyi, 257, 263, 267, 288

inconstans, 259, 260, 263 266, 269, 271,

272, 280, 281, 282, 283, 289, 290,

292, 293, 296, 299, 302, 305

leptocerus, 262, 265, 266, 270, 272, 289,

290, 291, 292, 294, 296, 299, 302, 307

maryae, n. sp., 255, 263, 266, 270, 283,

289. 290, 291, 292, 295, 296, 299,

302. 307

minutus, 255, 263, 266, 269, 274, 275,

276, 277 , 278, 279, 280, 289, 290,

291, 292, 294, 296, 299, 302, 307

montanus, 263, 266, 270, 285, 289, 290,

291. 292. 294. 296. 299. 302, 308

nasutus, 263, 268, 286, 289, 290, 291,

292, 294, 296, 308

neojamaicensis n. sp., 255, 256, 263, 266,

268, 289, 291, 292, 294, 297, 300,

302. 308

patriciae n. sp. 254, 255, 263, 266, 267,

270, 274, 276, 287, 289, 290, 291,

292, 294, 297, 300, 302, 308

pisonae, 257, 268, 277, 308

politus, 263, 266, 267, 269, 271, 272,

278, 289, 290, 291, 292, 294, 297,

300. 302, 306

pusillus, 263, 266, 268, 277, 284, 292,

300, 302, 307

pygmaeus, 255, 263, 266, 269, 275, 276,

304, 307

roseae n. sp., 255, 263, 265, 266, 269,

279, 289, 290, 291, 292, 294, 297,

300. 303, 306

septentrionalis, 255, 278

Aneurus simplex, 255, 263, 266, 268, 278, 279,

289, 290, 291, 292, 294, 297, 300, 303, 305

slateri, n. sp., 255, 263, 266, 268, 285, 289,

290, 291, 292, 295, 297, 300, 303, 306

tenuis, 263, 266, 270, 272, 273, 289, 290,

291. 292. 295. 297. 300. 303. 308

usingeri, n. sp., 255, 263, 266, 269, 284, 285,

286, 289, 290, 291, 292, 295, 297, 300,

303. 308

vauriei, 256, 263, 266, 268, 273, 274, 275,

289, 290, 291, 292, 295, 298, 301, 303,

308

veracruzensis, n. sp., 255, 263, 265, 266, 269,

275, 285, 286, 289, 290, 291, 292, 295,

298, 307

wygodzinskyi, n. sp., 255, 263, 265, 266, 270,

271, 272, 289, 290, 291, 292, 295, 298,

301.303.308

angiosperm, 106

Anisogamus modes tus, 42

annato, 102

annectens, Lutra, 110, 112

Annelids, 14, 81

ant-eaters, 107

Anthomyidae, 83

Anthon, H. 197, 198,209

Aotus lemurinus, 1 1 1

trivirgatus, 1 1 1

Aphodiinae, 232

aposeridis, Phytomyza, 327 , 334

Aposeris foetida, 334

glauca, 333

apple tree, 282

aquilegiae, Phytomyza, 332

Aradidae, 255—308

Aradus sanquinosus, 281

Archaeopsyllini, 164

archhieracii, Phytomyza, 337 , 338

Archhieracium sp., 337

arcticum, Simulium, 219 — 224, 228, 229,

321-325

Arctopora pulchella 312

Arctopsychidae, 3 1 2

argenteus group, Limnephilus,, 41

argent eus, Limnephilus, 41

arizonensis, n. sp., Aneurus, 255, 263, 266, 269,

275, 279, 280, 289, 290, 291, 292, 293, 295,

298,301,306

armadillos, 107

VI

armatum, Inophleum, 1 04

armiger, Cephalodesmius, 245

Arnason, A.P. 220, 228, 321, 324

Arnason, A.P. (See Fredeen, F.J.H.), 221,

228, 325

Arnason, A.P. (See Rempel, J.G.), 221, 222,

229

Artemisia, 339

Artibeus, cinereus, 112

jamaicensis, 1 1 0

lit ur at us, 110

arvensis, Sonchus, 330, 336

arvensis, uliginosus, Sonchus, 336, 337

aspen, 282

asper, Sonchus, 330, 334, 336

Astereae, 339

asteris, Chromatomyia, 335

atribarba, Crepis, 334

Asynarchus fusorius, 42

lapponicus, 25, 42, 48, 50, 51, 64, 57

modestus, 42

rhanidophorus, 42

Ateles fuscipes, 112, 158, 166

atricornis, Phytomyza, 339

atropurpureus tomentosus, Senecio, 334

Atta mexicana, 238

Attagenini, 309

Attagenus bicolor, 309, 310

audouini, Kenodactylus, 84, 86

Aulacopris maximus, 247

aurantiacum, Hieracium, 331

aureurn, Simulium, 228

aureus, Thomasomys, 110, 117, 121, 166

auripendulus, Eumops, 112

australis, Heteromys, 112, 166

A ustrosimulium ( A ustro simulium)

tillyardianum, 209

autumnalis, Leontodon, 330

autumnalis n. sp., Rhyacophila, 25, 30* 34,

50, 51, 58, 60, 66

Avicennia marina, 104

avocado, 104

Axelrod, D.I. (See Raven, P.H.), 106, 107,

181

azarae, Didelphis, 111, 117, 121

azteca, Ammophila, 315, 316, 318, 320

Babock, E.B., 335, 338

Babers, F.H. and J.J. Pratt, Jr., 185, 186

Baccharis sp., 105

bacteria, 241

bairdii, Tapirus, 112

Baker, R.H., 108, 178

balachowskyi, Eurysternus, 249, 250

Baiba, M.H. (see Fredeen, F.J.H.), 325

balbisiana, Weinmannia, 105

Balduf, W.V., 34, 53

Baldwin, S.J., 309

Ball, G.E., 84, 87

Ball, G.E., and J. Negre, 86, 87

balsa, 102, 104

Banks, N., 34, 42, 44, 46,53

Baranov, N., 197, 198, 209

barbara, Eira, 112, 142, 166

barbarae, Thalassotrechus, 75, 83, 84, 85, 86,

87,88,89,90

Barber, H.G.,271,287

barberi, Aneurus, 263, 268, 286, 289, 290, 291,

292, 298, 301, 308

Barreropsylla, 176

Barrow, J.H. (see Herman, C.M.), 227 , 229

Basak, S.(see Sengupta, R.), 183

Bassaricyon gabbi medius, 112

bats (see Chiroptera)

Beal, R.S., Jr., 309

bears, 107

Beckman, L., 184, 186

beebei, Polygenis roberti, 11.3, 116, 137, 140,

141, 154

beebi, Rhopalopsyllus, 140

Seek, K.J. and D.L. Bramao, 98, 178

beech tree, 282

beetles (see Coleoptera)

beetles, dung, 252, 253

Befaria sp., 105

Beiger, M., 331 , 334, 338

Bembecinus godmani, 315

neglect us, 318

Bembicini, 318

Bendell, J.F., 227, 228

Bennett, G.F., 226, 228, 227

Bennett, G.F. (see Fallis, A.M.) 226, 227, 228

Bennett, G.F. (see Greiner, E.C.), 226, 229

Bennett, G.F. (see Laird, M.), 227, 229

Berck, B. (see Fredeen, F.J.H.)., 325

Bergroth, E., 256, 270, 271, 275, 278, 281, 287

Betten, C., 34, 42, 46, 53

biclavata, Twinnia, 204

bicolor, A ttagenus, 309, 310

Vll

bicolor, Malagoniella, 247

biennis, Crepis, 331

bifila, Rhyacophila, 33

big belly tree, 104

bilobatum, Uroderma b., 112

birch 282

bispinus, Canthon, 247

Bixa orellana, 1 02

blackberries, 105

black rubber, 102

Blatchley, W.S., 271, 272, 275, 278, 281,

287

Blattella germanica, 77

Blepharoceridae, 209

Bocconia frutescens, 104

bogotensis, Sigmodon hispidus, 110, 112

bohlsi, Polygenis bohlsi, 116, 132, 142, 166,

172, 174

bohlsi, Pulex, 1 32

Bombycillidae, 226

Bombyx mori, 13, 15

bonariensis, Eumops, 1 1 0

Bonasa umbellus, 228

bonasae, Leucocytozoon, 228

borealis, n. sp., Aneurus, 225, 263, 266,

270, 282, 289, 290, 291, 292, 293, 295,

298, 301, 305

Bornemissza, G.F., 242, 251, 252

Borrero, J.I., 113, 178

boucardi, Copris, 243

bourgaei, Cicerbita, 331

brachinus, Polygenis, 138

brachiopods, 97

Braconidae, 339

Brady pus griseus, 1 1 2

brasiliensis, Eplesicus, 1 1 1

brasiliensis, Sylvilagus, 110, 111, 112

brasiliensis, Tadarida, 1 1 0

b/eviceps, Ammophila, 319

brevistyla, Rhizophora, 104

Briegal, H., 183, 186

Britton, W.E., 270, 278, 28 1 , 287

Brown, A.W.A. (see Arnason, A.P.), 220

228, 324

Brown, A.W.A. (see West, A.S.), 225, 229

bryozoans, 97

Brunellia sp., 105

brunneus, Zygodontomys brevicauda,

110

Buhr, H., 330, 331,332, 338

Burgl, H., 96, 97, 178

Burgl, H. (see Jacobs, C.H.), 97, 98, 180

buttonwood forest, 104

C. minus cuius, 40

Cabrera, A., 113, 178

Cabrera, A., and J. Yepes, 108, 113

cabuyas, 102

cacicus, Rhopalopsyllus saevus, 113, 115, 158,

162, 166, 172, 174

cactus, 275

Caenolestes, \16

fuliginosis, 124

obscurus, 113, 121, 124, 127, 131, 132, 166

Caenolestidae, 166

caespitosum, Hieracium, 331

Cafius 88

calabash, 102

Calamagrotis, 105

Calandra oryzae, 1 6

Calathus ruficollis, 86

Calhoun, E.H., 183, 186

Cali Virus Laboratory, 93, 178

calmnosus, Oryzomys, 111, 117, 121. 132,

137, 138, 140, 141, 142, 158, 164

caliginosus, Oryzomys (Melanomys), 140, 166

Caluromys derbianus derbianus, 108, 112

camels, 107

Cameron, A.E., 220, 221, 228

canadensis, Lactuca, 337

candezei, Megathoposoma, 247, 253

canaster, Galictis vittata, 1 1 2

cancrivora, Didelphis, 1 40

cancrivorus, Procyon, 1 12

candace, Nasua nasua, 1 1 0

canescens, Lantana, 102

caniceps, Diplomys, 1 1 2

Canidae, 107, 166

Canis familiaris, 142, 166

Canthidium, 241

Canthon bispinus, 247

cyanellus, 241, 243, 248, 252

cyanellus cyanellus, 247, 248

edentulus, 247

muticus, 247

virens, 247

Canthonina, 233, 235, 237, 242, 243, 245,

247

capers, 105

Vlll

capillaris, Crepis, 331

Caprifoliaceae, 339

capucinus, Cebus, llz

Carabidae, 75, 83, 84, 86, 87, 88

Carabidae, Neotropical-Indices, 479-493

caracasana, Euphorbia, 102

caracasana, Wigandia, 104

Carausius (=Dixippus) morosus, 6, 10, 12,

13, 15, 16

carbonero, 105

Carludovia palmata, 104

Carayon, J., 315, 318

carnivores, 107, 164, 166, 175

carolinus, Dichotomius, 238, 240, 242

Carollia castanea, 1 1 2

subrufa, 1 12

cascarillo, 105

cashew, 104

Casida, J.E., 185, 186

Cassia sp., 105

Cassiope mertensiana, 335

Castella elastica, 102, 104

Cathartidae, 226

cattle, 221, 222, 223, 224,238, 231, 242

catus, Felis, 164, 166, 175

caucae, Mormosa improvida, 110

caucensis, Polygenis n. sp., 91, 116, 137,

142, 144, 145, 166, 172, 174

caucenis, Scuirus, 1 1 1

Cavanillesia platani folia, 104

Cebidae, 166

Cebus capucinus, 112

Cecropia spp., 104

cedar, 279

Cedrela spp., 102

cedrillo, 105

Ceiba pentandra, 102

cepa, Allium, 105

Cephalocereus colombianus, 102

Cephalodesmius, 235, 243, 245, 247, 252

armiger, 245

laticollis, 245

quadridens, 245

cephalopods, 97

Ceratophyllidae, 131, 173, 180, 181

Ceratophyllinae, 131

Ceratophylloidea, 1 1 7

Ceratophyllus equatoris, 1 3 1

peripinnatus, 131

Cerdocyon thous, 113, 164, 166

Cespedesia macrophylla, 1 04

Cetraria sp. 104

Chabaud, A.G. 93, 178

Ceratopogonidae, 226

cervids, 241

cervipedis, Onchocerca, 226

Chalcidoidea, 333

Champion, G.C., 270, 271, 272, 275, 278,

285,288

championi, Aneurus, 263, 266, 268, 278, 289,

290, 291, 292, 293, 296, 298, 301, 307

Chance, M.M., 196, 197, 198, 201, 204, 205,

206, 209

cheesmanae, Simulium, 195, 196, 207

chickens, 222, 227

chayote, 104

cheopis, Pulex, 1 58

cheopis, Xenopsylla, 93, 1 14, 158, 164, 166,

167, 172, 174, 175

cherry tree, 105

cinctus, Oniticellus, 237, 251, 252,

cinerascens, Liponeuria, 209

cinqulatus, Dysdercus, 1 3

Chilco, 105

chiloensis, Myotis, 1 1 1

Chilomys, 1 1 7

instans, 124

Chilo stigma alas cense, 46

alascensis, 46

Chimaeropsyllidae, 180

Chiroderma villosum, 1 12

Chironectes minimus, 110, 111, 112, 142, 166

Chironomidae, 322, 323

Chiroptera, 107, 108, 115, 164, 165, 175, 182

chocoensus, Dasyprocta, 1 12

cholinesterases, 75, 183—189

Choloepus hoffmani, 112

Chapman, F.M., 112, 178

Chromatomyia, 327 , 328, 324-337,

Chromatomyia asteris, 335

kluanensis, 335

ixeridopsis n. sp., 327, 335, 336, 344, 345

lactuca, 327, 336, 342, 344, 345

senecionella, 327 , 334

syngenesiae, 327 , 328, 334, 335, 337

chryantha, Tabebuia, 104

Chylomys instans, 1 1 1

Chrysomelidae, 12

IX

Cicerbita ( =Mulgedium), 331, 338

Cicerbita alpina, 330, 331

bourgaei, 331

prenanthoides, 331

Cichorieae, 327, 333, 334, 337, 341

Cichorium intybus, 337

Cicindelinae, 88

cinereiv enter, Thomasomys, 111, 113, 117,

121, 124, 127, 131, 132, 164, 166

cinereoargenteus, Urocyon, 110, 166

cinereus, Artibeus, 1 12

Citharexylum sp., 102

Clarke, C.H.D., 227, 228

Clay, T., 93, 178

Clayton, J.S. (see Mitchell, J.), 321, 325

Cleoptsylla, 173, 175, 176

monticola, 114, 124, 125, 166

Clifford, E.A. (see Smart, J.), 196, 198, 210

Clifford, H.F., 42, 53

Cnephia dacotensis, 200

Coelopidae, 83

Coleoptera, 3,5,6, 11, 12, 13, 14, 15, 16,

75, 78, 81, 83, 87, 88, 231, 232, 253, 309

Collembola, 77, 81, 86

Colombians, Metachiurus nudicaudatus,

111, 176

Colombians, Proechimys, 1 1 2

Colombians, Scolopsyllus, 115, 119, 141,

159, 160, 166, 172, 174, 181

coloradensis, Rhyacophila, 33

coloradensis group, Rhyacophila, 33

columbiana n. sp., Phytomyza, 327, 332,

333, 343, 345

comis, Tetrapsyllus, 91, 1 15, 132, 136,

166, 172, 174

communis, Lapsana (=Lampsana), 330, 331

Compositae, 327, 332, 334, 339, 341

compositus, Onthophagus, 352

concolor, Felis, 1 1 3

concolor, Oryzomys, 112

Condylarthra, 106, 107

Conepatus semistriatus, 1 10

conges tus var. palustris, Senecio, 334

conistylum, Prosimulium, 207, 209

Conley, D.L. (see Jacobs, C.H.), 97, 178

Cono carpus, 104

Coombs, R.F. (see Greiner, E.C.), 226, 229

Coprina, 235,243,245,247

Coprini, 233, 234, 238, 242, 243, 245, 247

Copris, 245

boucardi, 243

Coptopsyllidae, 180

copha, Adoratopsylla intermedia, 116, 117, 121,

166

copha, Stenopsylla intermedia, 1 1 7

Cora pavonia, 105

coralito, 104

Corbett, J.R., 183, 186

cordoncillo, 104,

Coronapsylla, 175

Corvidae, 226

Costa Lima, A. da, 94, 178

Coulson, J.R. (see Stone, A.), 220, 229

Courtney, K.D. (see Ellman, G.L.), 184, 186

Couvert, L., 207, 209

Craig, D.A., 195, 196, 197, 198, 205, 206, 207,

209

Craneopsylla, \16

Craneopsyllinae, 124, 175, 176

Craneopsyllini, 124

Crepidinae, 327 , 330, 336, 337

Crepis atribarba, 334

biennis, 331

capillaris, 331

(Ixeridopsis) elegans, 335, 345

gracilis, 334

jacquini, 331

nana, 335

paludosa, 331

rubra, 331

runcinata, 334

sibirica, 331

tectorum, 334, 337

crepitans, Hura, 102

Crescentia cujete, 102

Cricetidae, 166, 175

Cricetinae, 107

crinoids, 97

Crosby, T.K., 198,209

Crosskey, R.W., 198, 201, 206, 209

Croton sp., 102, 104, 105

crustaceans, 83

cryptoctenes, R. lugubris, 158

Cryptotis medallinius, 1 1 1

thomasi, 1 1 1

Ctenidiosomus, 173, 175

croxtoni, Simulium, 226

croze tensis, Croze tia, 204, 209

X

Crozetia crozetensis, 204, 209

Ctenocephalides, 173

felis, 114, 118, 164, 166, 168, 172, 175

Ctenidiosomus perplexus, 124

rex , 115, 121, 172, 174

traubi, 91, 94, 115, 121, 122, 124, 166,

172, 174

Ctenomys, 176

Ctenophtalminae, 117, 180

Cuculidae, 226

cuipo, 104

cujete, Crescentia, 102

Culicidae, 75, 198, 226

curculionidae, 1 1

Curtis, J. 225,256,288

Curtis, L.C., 220, 221, 228

cyanellus, Canthon, 241, 243, 248, 252

cyanellus cyanellus, Canthon, 247 , 248

Cyclopes didactylus, 1 1 2

Cyclorrhapha, 79

dacotensis, Cnephia, 200

daphnis, Phanaeus, 233, 242, 244

Darlington, P.J., Jr., 84, 85, 87

Dasyprocta candelensis, 1 1 1

chocoensis, 1 1 2

fuliginosa, 1 1 1

punctata, 110, 112, 142, 166

Dasyproctidae, 166

Dasypsyllus, 173

Dasypsyllus gallinulae peripinnatuSj

91, 116, 131, 134, 166, 172, 174, 175

Dasypus novemcinctus, 110, 158

novemcinctus fenestratus, 1 5 8

Davies, D.M., 225, 228,

Davies, D.M., (see Wood, D.M.), 206, 210

Davies, L., 195, 197, 204, 209, 224, 228

deborahae, n. sp., Aneurus, 225, 263, 266,

269, 274, 275, 289, 290, 291, 292, 298,

301,306

decemlineata, Leptinotarsa, 12, 16

decorum, Simulium, 224

deer, 107, 222

DeFoliart, G.R. (see Anderson, J.R.), 226

228

defoliarti, Simulium, 221

delpontei n. sp Polygenis, 91, 116, 138

146, 147, 148, 166, 172, 174

Dendragapus obscurus fuliginosus, 228

Denning, D.G., 27, 30, 33, 37, 39, 40, 46,

53, 54

derbianus, Caluromys derbianus, 108, 112

Dermaptera, 77, 79, 81 j

Dermestidae, 309

Desmanthus virgatus, 102

Desmodontidae, 166

Desmodus rotundus, 112, 164, 166 j

Dewhurst, S.A., 185, 186

Diaemus youngi, 112 j

dichopticus, sp., near; Gymnopais, 204

Dichotomiina, 235, 238, 241, 243

Dichotomius carolinus, 238, 240, 242

tortulosus, 235

Dicosmoecinae, 34

Dicosmoecus, 34, 47

gilvipes, 25, 34, 47, 50, 51, 61, 66 j

jucundus, 35

didactylus, Cyclopes, 1 1 2

Didelphidae, 166

Didelphis azarae, 111, 117, 121, 166

cancrivora, 140

marsupialis, 117, 121, 124, 137, 141, 142, 158,

158, 164, 166, 176

m. marsupialis, 110, 112

diomedes, Sphinctosylla, 91, 94, 1 14, 124, 166,

174

Diplura, 8 1

Diptera (flies), 11,75,77,78,79,83, 182, 195,

209, 210, 228, 229, 325, 327, 328, 339, 340,

341

discolor, Phyllostomus, 1 1 2

dissimilis, Aneurus, 225, 262, 263, 265, 266, 267, j

267, 270, 276, 289, 290, 291, 292, 293, 296,

299, 301, 307

distincta, Sternopsylla, 175

Dobzhansky, T., 86, 87

Dodds, G.S., 34, 54

Dolophilodes aequalis, 69

nora, 69, 71

novusamericanus, 69

Dolophilodes (D.) pallidipes , 69, 70

donaldi, n.sp Rhyacophila, 25, 29, 34, 50, 51,

58, 59, 66

Donvar-Zapolski, D.P., 338

Doratopsyllinae, 180

dorsalis, Vampyrops, 1 1 2

Douglas, J,, 256, 288

Drosophila melanogaster, 1 5

XI

dry as, Marmosa, 113, 124

ducks (disease transmission to), 226, 227,

228, 229

dulce, Pithcellobium, 102

Dumbleton, L.J., 195, 197, 198, 204, 209

Dunn, L.H.,94, 178

dunni, Polygenis, 113, 116, 139, 141, 149,

172, 174

dunni, Rhopalopsyllus, 139

Dyplomys (=Diplomys) caniceps, 1 1 2

Dysdercus cingulatus, 13

ebria, Rhyacophila, 29

echidnophagoides, Juxtapulex, 158

Echimyidae, 140, 166

Echinoprocta rufescens, 1 1 1

echioides, Picris, 330, 334

edentates, 91

edentulus, Canthon, 247

Edmonds, G.F., 238, 252

edule, Sechium, 104

Edwards, F.W., 195, 197, 209

Eidt, R.C., 100, 178

Eira barbara, 1 12, 142, 166

elastica, Castella, 102, 104

elder, 282

elegans, Crepis (Ixeridopsis), 335, 345

elk, 226

Ellman, G.L., 184, 186

encenillo, 105

Encephalitis, Eastern Equine, 226, 228

encimo, 105

engelmanni, Picea, 279

Ephemeroptera, 77, 79, 322, 323

equatoris, Ceratophyllus, 131

equatoris, Pleochaetis equatoris, 117, 131,

172, 174

Equisetum, 39, 105

erigerophila, Phytomyza, 337

esmaralarum, Nectonys alfari, 111, 112

Escoto, J.A.V., 86, 88

espadero, 105

Espeletia, 105

Espinal, T., L.S., 94, 100, 102, 178

Essig, E.O.,34, 56, 54

Etnier, D.A., 43, 54

Eumops auripendulus, 1 1 2

Eumops bonariensis, 1 1 0

Euphorbia caracasana, 102

euryadminiculum, Simulium, 226

Eurysternini, 233, 237, 249

Eurysternus, 241, 249, 251, 252

balachowskyi, 249, 250

magnus, 249

mexicanus, 249, 251

Eusimulium, 227

Evans, H.E., 315, 316, 317, 318

Evans, W.G., 75, 83, 84, 88

Ewing, H.E.,93, 178

excelsum, Anacardium, 104

falcata, Kohlsia, 173

Fallen, C.F., 329, 339

Fallis, A.M., 226, 227, 228

fallisensis, Ornithofilaria, 226, 228

familiaris, Canis , 142, 166

farnesiana, Vachelia, 102

fasciatus, Oncopeltus, 6, 12, 13, 14, 15, 16

Featherston, F.M. (see Ellman, G.L.), 184,

186

Felidae 166

Felis catus, 1 64, 1 66

concolor, 1 1 3

pardalis, 1 1 2

tigrina pardionoides, 110, 112

yagouaroundi, 110, 112

felis, Ctenocephalides, 114, 118, 164, 166, 168,

172, 175

felis, Pulex, 1 64

femoralis, Limnephilus, 41

fenestratus, Dasypus novemcinctus, 158

ferns, 104

Fest, C., 183, 186

Festuca, 105

Fewkes, J.W., 309

Ficus (spp.), 102, 104

Fischer, F.C.J., 27, 30, 33, 34, 37, 39, 42, 43,

44, 46, 54

fish, 321, 322, 325

fiskei, Aneurus, 263, 266, 267, 269, 280, 281, 282

282, 289, 290, 291, 292, 293, 296, 299,

301,306

Fittkau, H.S., 107, 179

flavus, Pot os, 1 12

fleas, 91-94, 96-98, 102, 106-108, 110-117,

121, 122, 124, 127, 130, 131, 132, 134,

137 142, 149, 154, 158, 161-166, 168,

170-182

flea, bird, 165, 175

fleas, helmet

Xll

Flint, O.S., Jr., 34, 54

foetida, Aposeris, 334

Foote, R.H. (see Stone, A.), 220, 229

forests, tropical, evergreen, deciduous,

mangrove, 98

Fortner, G., 196, 201, 206, 209

Fourcraea sp. 104

Fox, I. (see Tamsitt, J.R.), 94, 182

fox tail, 1 04

Fragaria sp., 105

frailijones, 105

frenata, Mustela, 110, 112, 166

Fredeen, F.J.H., 204, 209, 221, 222, 224,

228, 229, 321, 322, 323,

Fredeen, F.J.H. (see Arnason, A.P.), 220,

228, 324

Frey, R., 331, 339

Freyvogel, T.A., 184, 186

Freyvogel, T.A. (see Briegel, H.), 183, 186

Freziera sericea, 1 05

Frick, K.E., 334, 336, 337, 339

frigidus, Petasites, 334

froeschneri, Aneurus, 263, 265, 266, 269,

280, 289, 290, 291, 292, 293, 296, 299,

301, 307

Frost, S., (see Kershaw, W.E.), 325

Frost, S.W., 336, 337, 339

frutescens, Bocconia, 104

Fugatera pterota, 102

fuliginosis, Caenolestes, 124

Fuller, H.S.,94, 179

fulvescens, Sylvilagus, 110, 111

fulvum, Prosimulium, 224

fungi, 241,255,257,259

Furcraea sp., 102

fusca, Phryganea, 42

fuscata, Marmosa, 1 24

fuscatus, Lasiurus ega, 1 1 0

fuscatus, Thomasomys, 110, 117, 121, 124

132, 140, 166

fuscipes, Ateles, 112, 158, 166

fusorius, Anabolia (Asynarchus), 42

fusorius, Asynarchus, 42

fusorius, Stenophylax, 42

fuscum, Prosimulium, 224

gabbi, Bassaricyon medius, 1 12

Galictis allamandi, 1 42

vittata canaster, 112

gallicus, Aneurus, 228

Gardner, J.C.M., 25 1 , 253

Garth, J.S., 85, 88

gasipals, Guilielma, 104

Gast Galvis, A., 94, 179

geese, 227, 228, 229

Geotrupinae, 232, 233

germanica, Blattella, 77

Ghosh, J.J. (see Sengupta, R.), 183, 187

gigantea, Trichantera, 104

Gillaspy, J.E. (see Evans, H.E.), 315, 316, 318

gilvipes, Dicosmoecus, 25, 34, 47, 50, 51, 61,

66

gilvipes, Stenophylax, 34

Glaphyrocanthon subhyalinus, 243

glauca, Aposeris, 333

glauca, Picea, 279

Gleicheniaceas, 104

Glossosomatidae, 52, 69

Glossophaga soricina, 110, 112, 164, 166

Glydenstolpe, N., 113, 179

Glynne-Williams, J. & J. Hobart., 84, 88

godmani, Bembecinus, 3 1 5

gossypiifolia, Jatropha, 1 02

goudotti, Odocoileus, 113

gracile, Hieracium, 333, 335

gracilis, Crepis, 334

Graham, S.A., 309

grandiflora, Lapsana, 331

grasses, 105

gregaria, Schistocerca, 13, 15

Greiner, E.C., 226, 229

Grenier, P., 197, 199, 109

Griffiths, G.C.D., 327, 328, 331, 332, 334,

335, 336, 339

griscescens, Philander, 110, 111, 112

griseipennis, Stenopsyche, 6, 11, 12, 13, 15

griseum, Simulium, 224

griseus, Brady pus, 1 1 2

Groschke, F., 334, 339

grouse, blue, 227, 228

grouse, ruffed, 227, 228

guamo, 104

guanta, Agouti paca, 110

guarumo, 102, 104, 105

guayacan, 104

Guilielma gasipals, 1 04

Guppy, R., (see Schmid, F.), 34, 37, 39, 56

Gust ania superb a, 104

guyannensis, Proechimys, 1 1 2

7\

xiii

gymnurus, Hoplomys , 112, 140, 142, 166

Gymnopais sp. near dichopticus, 204

Gymnopleurus, 247

Gyorkos, H. (see Wood, D.M.), 206, 210

Haemoproteus, 227

Haffer, J., 108, 111, 179

Hagen, H.A., 34, 42, 54

haitiensis, Aneurus, 255, 256, 263, 266,

268, 273, 274, 289, 290, 291, 292, 293

296, 299, 302, 308

Halesus alascensis, 46

Halffter, G., 232, 233, 240, 244, 245, 246,

247, 253

Halffter, G., (see Edmonds, W.D.), 238, 252

Halffter, G., (see Halffter, V.), 238, 253

Halffter, V., 238, 244, 253

Hamelia patens, 104

Hammen, T.V.D., 98, 108, 179

Hanson, R.P., (see Anderson, J.R.), 226,

228

Hanssen, H. (see Mendez, E.), 94, 181

Harlan, T.P., (see Nichols, R.F.), 309, 310

Harper, P.P., (see Roy, D.), 38, 43, 44, 55

Hartig, R., 331, 339

has tat us, Phyllostomus, 1 1 1

Hathaway, C.R., (see Costa Lima, A. da),

94, 178

Hearle, E., 224, 229

Heideman, O, 278, 281, 288

Heliconia, 104

Heliotropium, sp. Ill

helleri, Vampyrops, 112

hematozoa, 219, 220, 226, 227, 229

Heming, B.S., 81

Hemiptera, 11, 16, 81, 225, 284, 287, 288

Hendel, F., 329, 321, 332, 337, 339

Hennig, W., 79, 81

herbivore, 241

Herman, C.M., 227, 229

Hering, E.M., 329, 330, 331, 337, 340

Hering, E.M. (see Groschke, F.), 334, 339

Hering, M., 329, 330 331, 337, 339, 340

Hershkovitz, P.,97, 106, 107, 108, 111,

113, 173, 179

Heteromyidae, 166

Heteromys australis, 112, 166

Heteroptera, 6, 12, 13 15, 16, 287, 288

hieracina, Phytomyza, 329, 330

hieracioides, Picris, 330

Hieracium, 327 , 330, 335, 337, 338

albiflorum, 330

amplexicaule, 331

aurantiacum, 331

caespitosum, 331

gracile, 333, 335

japonicum, 334

lachenalii, 331 , 337

laevigatum, 331

laevigatum var. tridentatum, 331

murorum, 331

prenanthoides, 331

pulmonarioides, 331

sabaudum, 330, 331

schistosiphon, 33 1

silvaticum, 331

thapsoides, 331

transylvanicum, 331

triste, 333, 345

umbellatum, 331

villosum, 331

vulgatum, 330, 331

Hieracium sp., 331

hirtipes, Prosimulium, 224, 225

Hisaw, F.L. (see Dodds, G.S.), 34, 54

hispaniolensis n. sp Aneurus, 255, 256, 263,

265, 266, 268, 273, 274, 289, 290, 291,

292, 299, 302, 308

hispidus, Leontodon, 330

hispidius, Sigmodon, 1 1 2

Hodge, F.W., 309

hoffmani, Choloepus, 112

hogplum, 102

Holdrige, L.R., 102, 179

Holland, G.P., 165, 179

Hominidae, 166

Homoptera, 78

Homo sapiens, 164, 165, 166, 175, 220, 223,

224, 225,226, 227,242,247

Hooper, E.T., 179

Hopewell, W.W. (see Arnason, A.P.), 220, 228

324

Hopkins, G.H.E., 93, 165, 175, 179, 180

hopkinsi n. sp Polygenis, 91, 115, 116, 137,

150, 151, 166, 172, 174

Hoplomys gymnurus, 112, 140, 142, 166

Horn, G.H., 84, 88

horse, 107,221,222,226

horse tail (see Equisetum)

XIV

hrdyi, Aneurus, 257 , 263, 267 , 288

Hulten, E., 328, 340

Hunter, R.L. (see Freyvogel, T.A.), 184,

186

Hum crepitans, 1 02

Hussey, R.F., 272, 288

hyalinata, Rhyacophila, 33

hydrobat es, Ichthysomys, 1 1 1

Hydrochaeris hydrochaeris, 1 1 2

isthmius, 112

hydrochaeris, Hydrochaeris, 112

Hydrophilidae, 83, 88

Hydropsy chidae, 312

Hylepsyche plectrus, 44

hylophilus, Thomasomys, 124

Hymenoptera, 77, 315, 318

Hystrichopsyllidae, 117, 173, 180

Ichthyomys hydrobates, 1 1 1

nicefori, 1 1 1

Imania thomasi, 25, 35, 47, 50, 58, 62, 66

inambari, Akodon pulcherrimus, 141

incisus group, Limnephilus, 39

inconstans, Aneurus, 259, 260, 263, 266,

269, 271, 272, 280, 281, 282, 283, 289,

290, 291, 292, 293, 296, 299, 301, 305

indigo, 104

Indigobera sp. 104

inflatum, Prosimulium, 210

Inga spp. 1 04

Inophleum armatum, 104

Insecta, 14, 15, 16, 80

inseparata, Phytomyza, 329, 330, 332

instans, Chylomys, 111, 124

insularis, Limnephilus, 25, 39, 48, 50, 62,

67

intermedia, Adoratopsylla (T.), 166, 172,

174, 176

intybus, Cichorium, 337

invaria group, Rhyacophila, 33

iraca 104

irritans, Pulex, 93, 114, 164, 166, 169,

172, 174, 175

Isa, J.M. (see Savage, A.), 227, 229

Ischnopsyllidae, 127, 180

Ischnopsyllinae, 127

Isoderminae, 265

Isopoda, 86

isthmica, Marmosa, 112

isthmius, Hydrochaeris, 1 12

isthmius, Microsciurus falviventer, 1 12

Ixeridopsis, 321, 335, 336

ixeridopsis, n. sp., Chromatomyia, 327 , 335,

336. 344. 345

Ixeris alpicola, 336

Jacobs, C.H.,97, 98, 180

jacquini, Crepis, 331

jamaicensis, Artibeus, 1 10

janus, Limnephilus, 40, 63

japonica, Phytomyza, 334

japonicum, Hieracium, 334

Jatropha gossypiifolia, 102

Jeannel, R., 84, 88

johannseni, Simulium, 226

Johnson, F.M. (see Beckman, L.), 184, 186

Johnson, P.T., 94, 113, 140, 165, 175, 180

Johns, P.M., 84, 86, 88

Johnson, R.D. (see Kershaw, W.E.), 325

jucundus, Dicosmoecus, 35

Judulien, F., 247, 253

juliflora, Prosopis, 102

Juxtapulex echidnophagoides, 158

kangaroo, 241

Kaplan, W.D. (see Dewhurst, S.A.), 185, 186

Kapoor, I.P., 323, 325

kappleri, Peropteryx kappleri, 1 1 0

Karl, O., 331, 340

Keast, A., 106, 180

kelp, 83

Kenodactylus audouini, 84, 86

keratus, Lenar chus (Prolenarchus), 43

keratus, Limnephilus, 43

Kershaw, W.E., 323, 325

Kjellgren, B.L, (see Betten, C.), 34, 42, 46, 63

klagesi, Polygenis, 116, 140, 152, 166, 172, 174

klagesi, Pulex, 1 40

kluanensis, Chromatomyia, 335

Knight, K.L., 197, 198,209

Kohlsia 181

Kohlsia falcata 173

tiptoni, 173

kok-sghyz, Taraxacum, 334

Koble, H.J.,42, 54

Kormilev, N., 257, 270, 272, 273, 274, 275,

276, 279, 280, 281, 284, 285, 286, 288

Kristensen, N.P., 81

Kubska, J., 331, 340

Kurtak, D.C., 201, 209,

Kurten, B., 107, 180

labialis, Noctilio, 110, 164, 166

! lackberry, 276

|| lachenalii, Hieracium, 331, 337

lactuca, Chromatomyia, 327 , 336, 342, 344,

345

lactuca, Phytomyza, 336

Lactuca canadensis, 337

scarida var. integrata, 337

serriola, 331, 336, 337, 345

spicata, 331

tatarica, 331

virosa, 331

Lactuca sp., 334

Ladenbergia, 105

laevigatum, Hieracium, 331

laevigatum, var. tridentatum, Hieracium, 331

lagopus, Ochroma, 102, 104

Laguncularia racemosa, 104

Laird, M., 227, 229

lampsanae, Phytomyza, 329, 330, 331

laniger, Thomasomys, 121

Laniidae, 226

Lantana canescens, 102

lapponicus, Asynarchus, 15, 42, 48, 50, 51,

64, 67

lapponicus, Limnephilus, 42

Lapsana (=Lampsana) communis, 330, 33 1

Lapsana grandiflora, 33 1

Lasiurus sp., 113

Lasiurus ega fuscatus, 1 1 0

Lateritics, Reddish-brown , 1 00

Latersol 100

laticollis, Cephalodesmicus, 245

latimanus, Rhipidomys, 110, 117, 131, 140,

141, 166

latipes, Simulium, 226, 228

Lee, V.H. (see Anderson, J.R.), 226, 228

Leech, H.B., 83, 88

Leg worms, 226

lemurinus, Aotus, 1 1 1

Lenarchus (Prolenarchus) keratus, 15, 43,

48, 50, 51, 64, 67

Leonard, F.A. (see Leonard, J.W.), 44, 54,

55

Leonard, J.W., 44, 54 55

Leontodon autumnalis, 330

hispidus, 330

Leontodontinae, 330

Lepidoptera, 11, 13, 88

Leptinotarsa decemlineata, 12, 16

leptocerus, Aneurus, 263, 265, 266, 270, 272,

289, 290, 291, 292, 294, 296, 299, 302, 307

Leptopsylla 173

segnis, 115, 131, 133, 166, 172, 174, 175

Leptopsyllidae, 180

Leptopsyllinae, 131

Lethierry, L., 270, 271, 275, 278, 281, 288

leucocytozoan infection, 219, 226, 227, 229

Leucocytozoon anatip, 226, 229

bonasae, 228

simondi, 227 , 228, 229

Lewis, R.E., 165, 180

Liatongus monstrosus, 238, 239, 241, 252, 253

lichens, 105

lilium, Sturnira, 1 12

Limnephilidae, 25, 26, 34, 43, 47, 50, 51, 312,

Limnephilinae, 38

Limnephilus alvatus, 25, 27, 39, 40, 48, 50, 51

63, 67

argent eus, 41

femoralis, 41

insularis, 25, 39, 48, 50, 62, 67

janus, 40, 63

keratus, 43

lapponicus, 42

modestus, 42

nimmoi, 25, 27, 38, 48, 50, 52, 62, 66

rhanidophorus, 42

secludens, 39, 40

vernalis, 25, 41 , 48, 50, 5 1 , 63, 67

Lindner, E. (see Rubtzov, I.A.), 209

Lindroth, C.H., 84, 88

Ling, S.W., 37, 55

Linsley, E.G., 318

Linsley, E.G. (see Evans, H.E.), 315, 316, 317,

318

Liponeura cinerascens, 209

Listropsyllinae, 180

Litopterna, 106

lituratus, Artibeus, 1 10

lizards, 1 1 1

Locusta migratoria, 10, 12, 13, 14, 16

Loomis, F.G., 106, 180

Lopez, G. (see Halffter, G.), 232, 233, 243, 244,

253

luggeri, Simulium, 219, 223, 321, 323

lugubris, Rhopalopsyllus, 1 13, 115, 158, 163,

166, 172, 174, 176

XVI

lulo, 105

Lundquist, A., 331, 340

Lutra annectens, 110, 112

Lygaeidae, 16

Lyneborg, L. (see Anthon, H.), 197, 209

Lytta viridana, 5, 6, 7, 9, 10, 11, 12, 13,

14, 15, 16

Macchiavello, A., 93, 94, 180, 181

macedero, 104

Machado-Allison, C.E. (see Tipton, V.J.),

113, 124, 139, 158, 182

Macropsyllidae, 180

macrophylla, Cespedesia, 104

Macropocopris, 241

magnus, Eurysternus, 249

Mahowald, A.P., 81

maize, 105

major, Molossus molossus, 110, 112, 164,

165, 166

Malachiidae 88

Malacopsyllidae 180

Malagoniella bicolor, 247

puncticollis tubericeps, 247

mammagua, 104

Mammalia, 75, 91, 92, 93, 97, 98, 102,

106, 107, 108, 110-117, 121, 124, 127,

131, 132,137-142, 158, 164, 165,

172-182, 220

man (see Homo sapiens )

mangrove, red 104

mangrove, black 1 04

mangrove, white 104

Manning, S.A., 331, 340

Mansingh, A. (see Smallman, B.N.), 183, 187

maple tree, 282

marginella, Phytomyza, 327 , 329, 332, 333,

337, 338, 342

marina, Avicennia, 104

marinus, Aepus, 84

Marmosa, 117

dry as, 113, 124

fuscata, 124

isthmica, 112

robinsoni, 112

improvida caucae, 1 10

Marsupiaha, 91, 106, 107, 127, 141, 175,

176, 241, 242, 247

marsupialis, Didelphis, 110, 112, 117, 121,

124, 137, 141, 142, 158, 164, 166, 176

mastodonts, 107

maryae, n. sp., Aneurus, 255, 263, 266, 270,

283, 289, 290, 291, 292, 294, 296, 299, 302,

307

Matchett, R.E. (see Kershaw, W.E.), 325

Matsuda, R. 75

Matsuda, R., (see Usinger, R.L.), 256, 265, 270,

271, 272, 275, 278, 281, 285, 288

Matthews, E.G., 233, 241, 243, 245, 247, 253

Matthews, E.G. (see Halffter, G.), 232, 233,

240, 243,245,246,247,253

Mayr, E., 86, 88

mays, Zea, 105

Mazama americana, 112, 113

americana zetti, 1 1 1

Mazur, J. 331, 340

maximus, Aulacopris, 247

Mecoptera, 77

medallinius, Cryptotis, 1 1 1

megalotus, Pot os flavus, 1 10

megastigmata, Rhynchopsyllus, 91, 94, 165

Megathoposoma candezei, 247, 253

Meijere, J.C.H. de, 329, 330, 331, 340

melanoceras, Acacia, 104

melanogaster, Drosophila, 15

melanurus, Philander, 1 12

Melinis minulti flora, 104

Melilotus, 315

Meloidae, 5, 14, 15, 16

Membrillo, 104

Mendez, E., 75,94, 181

Mendez, E (see Tipton, J.), 113, 121, 122, 130,

134, 139, 142, 149, 154, 158, 161, 162, 163,

165, 168, 170, 171, 173, 182

Menke, A.S., 318

mephistophiles, Pudu, 1 13

meridionale, Simulium, 226

Merkley, D.R., (see Ross, H.H.), 39, 42, 43, 55

mertensiana, Cassiope, 335

mesquite, 102, 276

Metachirus nudicaudates, 1 66

nudicaudates Colombians, 111, 176

Metcalf, R.L. (see Kapoor, I.P.), 325

mexicana, Atta, 238

mexicanus, Eurysternus, 249, 251

mexicanus, Reithrodontomys, 110, 138

micantha, Trema, 104

Michalska, Z., 331, 340

Michna, J., 331, 340

XVII

Miconia albicans, 1 05

micranthum, Ocimum, 102

microfilaria, 226, 227

Microsciurus flaviventer isthmius, 1 1 2

migratoria, Locusta, 10, 12, 14, 16

Millar, J.L., 222, 229

milleri, Reithrodontomys mexicanus, 1 1 1

Milne, L.J., 27, 33, 34, 37, 40, 42, 46, 55

milnei, Rhyacophila, 25, 31, 34, 50, 60, 66

Mills, M.L. (see Kershaw, W.E.), 325

Mimosa, 104

minimus, Chironectes, 110, 111, 112, 142

minusculus, C., 40

minuscula, Phytomyza, 332

minutiflora, Melinis, 104

minutus, Aneurus, 255,263, 266, 269, 274,

275, 276, 277, 278, 280, 289, 290, 291,

292, 294, 296, 299, 302, 304, 307

minutus, Oryzomys, 124, 127, 132

mirae, Tylomys, 1 1 2

Miridae, 288

Mitchell, J., 321, 325

mixtum, Prosimulium, 224

modes ta, Anabolia, 42

modesta, Stenophylax, 42

modestus, Anisogamus, 42

modestus, Asynarchus, 42

modestus, Limnephilus, 42

Mokry, J.E., 207, 209

molitor, Tenebrio, 6,1, 11, 16

Moll, A. A., 94, 164, 181

Molossidae, 164, 166

Molossus molossus, 1 64

Molossus molossus major, 164, 165, 166

Mollusca, 97, 98

mombin, Spondias, 102

monkeys, 107, 243

montanus, Aneurus, 263, 266, 270, 285,

289, 290, 291, 292, 294, 296, 299, 302,

308

Montenegro, E. (see Espinal, T., L.S.), 102,

178

monteno, 105

monticola, Cleopsylla, 114, 124, 125, 166

monstrosus, Liatongus, 238, 239, 241, 252,

253

Moore, I., 85, 88

Moores, E. (see Valentine, J.W.), 106, 182

Moose, 226

mori, Bombyx, 13, 15

mosquito, 209, 210, 225

morosus, Carausius (=Dixippus), 6, 10, 12, 13,

15, 16

mortino, 105

mosquero, 104, 105

Moss, H.C. (see Mitchell, J.), 321, 325

mosses, 105

mouse, house, 131, 175,345

Muller, P., 107, 108, 112, 181

mulgedii, Phytomyza, 329, 330, 331

munchiquensis, Oryzomys, 110

muralis, Mycelis, 330, 331

Muridae, 158, 1 66, 173

murids, 158, 175

murorum, Hieracium, 331

Mus mus cuius, 110, 112, 131, 175,

Mustela frenata, 110, 112, 166

Mustelidae, 166

muticus, Canthon, 247

Mycelis muralis, 330, 331

My otis sp., 113

Myotis chiloensis, 1 1 1

myxomatosis, 226

Maclnnes, C.D., (see Bennett, G.F.), 227, 228

McCaman, R.E. (see Dewhurst, S.A.), 185, 186

186

McLachlan, R., 34, 42, 55

nana, Crepis, 335

narvae, Rhyacophila, 33, 34

Nasua nasua, 1 1 2

nasua candace, 1 1 0

olivacea, 1 1 1

socialis, 140

nasua, Nasua, 1 1 2

nasutus, Aneurus, 263, 268, 286, 289, 290, 291,

292, 294, 296, 308

Navas, L., 33, 55

Neacomys tenuipes tenuipes, 112

Neave, F., 34, 35, 55

Nectomys alfari esmeraldarum, 111,. 112

neglectus, Bembecinus, 3 1 8

Negre, J., 86, 87

Nematocera, 79, 195, 209

Neocanthidium, 241, 252

neojamaicensis, n. sp., Aneurus, 255, 256, 263,

266, 268, 277, 278, 289, 290, 291, 292, 294,

297, 300, 302, 308

Neophylacinae, 36, 47

XV111

Neophylax, 36, 47

pulchellus, 37

rickeri, 25, 37, 47, 50, 51, 58, 61, 66

Neotropical Carabidae, Indices, 479—493

Neotyphloceras, 173

rosenbergi, 116, 117, 120, 166, 172, 174,

176

Neotyphloceratini, 117

Neuroptera, 77

Newell, R.L., 28, 30, 33, 55

nice fori, Ichthyomys, 1 1 1

Nichols, R.F., 309, 310

nigricans, Achat ocarpus, 102

nigrifrons, Oxymycteris p., 141

nigripennis, Thalassotrechus, 84, 85

nigripennis, Thalassotrechus barbarae,

83, 86

Nimmo, A.P., 26, 27, 28, 30, 31, 32, 33,

34, 38, 40, 41, 42, 45, 46, 47, 48, 49,

50, 51, 52, 55, 57

nimmoi, Limnephilus, 25, 27, 38, 48, 50,

52, 62, 66

Noctilionidae, 166

Noctilio labialis, 110, 164, 166

Noel-Buxton, M.B., 323, 325

Nonapsylla, \16

nora, Dolophilodes, 69, 71

norvegicus, Rattus, 3, 11 0, 111, 1 12, 158,

166

Notungulata, 106

novemcinctus, Dasypus, 110, 158

novusamericans, Dolophilodes, 69

Nowak, Z., (see Michalska, Z.), 331, 340

Nowakowski, J.T., 330, 331, 337, 340

Nowicki, J., 331, 340

nudicaudates, Metachirus, 166

Nunberg, M., 331, 340

Nygren, W.E., 97, 111, 181

Nystrom, R.F. (see Kapoor, I.P.),'325

oak, 275

obliqua, Steniolia, 315

O’Brien, R.D., 183, 187

obscurus, Caenolestes, 113, 121, 124, 127,

131, 132, 166

obscurus fuliginosus, Dendragapus, 228

O chroma lagopus, 102, 104

Ocimum micranthum, 102

Odocoileus goudotti, 1 13

tropicalis, 112

Odocoileus virginianus, 110, 112, 113

Odonata 323 I

officinale, Taraxacum, 331, 333, 337

oleraceus, Sonchus, 330, 334, 336

Oligophlebodes, 36

Oligoryzomys, 182

olivacea, Nasua, 1 1 1

O’Leary, S.B. (see Moll, A. A.), 94, 164, 181

Olsson, A.S., 97, 181

onca, Felis, 1 12

Onchocerca (=Wehrdikmansia) cervipedis, 226

Oncopeltus fasciatus, 6, 12, 13, 14, 15, 16

onion, 105

Oniticellini, 233, 235, 237, 238, 251

Oniticellus cinctus, 237 , 251, 252

Onitini, 233, 235, 238

Onitis, 243, 245

Onthophagini, 233, 235, 238, 253

Onthophagus, 238, 243

Onthophagus compositus, 252

parvus, 241

ophyrus, Rhyacophila, 27

opossum, Philander, 110, 111, 112, 117,

121, 166

Orcutt, A.W. (see Betten, C.), 34, 42, 46, 53

orellana, Bixa, 102

ornatus, Tremarctos, 113

Ornithofilaria fallisensis, 226, 228

O’Roke, E.C., 226, 229

Orthoptera, 11, 12, 13, 15, 16,77 79

Orthopteromorpha, 79

oryzae, Sitophilus (=Calendra), 11, 13, 17

Oryzomys albigularis, 91, 110, 117, 124, 127,

131, 129, 140, 141, 166

alfaroi, 117, 132, 166

alfaroi palmirae, 1 10

caliginosus, 91, 111, 112, 117, 121, 132,

137, 138, 140, 141, 142, 158, 164

capito, 1 1 2

concolor, 1 1 2

(Melanomys) caliginosus, 140, 166

minutus, 124, 127, 132

munchiquensis, 110

(Oligoryzomys), 117, 138, 166

physodes, 140

stolzmanni, 141

Osborne, D., 309

Osgood, W.H., 107, 113, 181

oviceps, Simulium, 194, 196, 197, 199, 200, 201,

203, 204, 205, 206, 207, 217, 218

XIX

ovipennis, Trechus, 84

Oxymycteris p. nigrifrons, 141

paca, Agouti, 104, 112, 142, 158, 166

pallidipes, Dolophilodes (D.), 69, 70

palmata, Carludovica, 104

palmirae, Oryzomys alfaroi, 110

paludosa, Crepis, 331

paluster, Sonchus, 330

panamensis, Procyon, 112

paniculatum, Talinum, 102

Panicum barbinode, 96

maximum, 96

paramos, 96, 127

Paraphytomyza, 339

Paridae, 226

Parmenter, L., 331, 340

Parapsylline, 176

Parapsyllini, 132

pardalis, Felis, 1 1 2

pardionoides, Felis tigrina, 1 10

parnassum, Simulium, 224, 226

parvus, Onthophagus, 241

Pascual, R., (see Patterson, B.), 98, 106

107, 108, 176, 181

patens, Hamelia, 104

Patino-Camargo, L., 94, 181

patriciae, n. sp., Aneurus, 254, 255, 263,

266, 267, 270, 274, 276, 287, 289, 290,

291, 292, 294, 297, 300, 302, 308

Patrobini, 87

Patterson, B., 93, 98, 181

Patterson, B. & R. Pascual, 98, 106, 107,

108, 176

pavonia, Cora, 105,

pecari, Tayassu, 112, 142

peccaries, 107

pejibaye, 104

penetrans, Tunga, 113, 165, 166, 171, 172,

174, 175

pentandra, Ceiba, 102

pentaphylla, Tabebuia, 102

peripinnatus, Ceratophyllus, 131

peripinnatus, Dasypsyllus gallinulae, 116,

131, 134, 166, 172, 174, 175

Periplaneta americana, 1 5

Peromyscini 107

Peromyscus, 173

Peropteryx kappleri kappleri, 1 1 0

perplexus, Ctenidiosomus, 124

Persea americana, 1 04

Petasites, 327 , 339

frigidus, 334

Petersen, B., 86, 88

Peterson, B.V., (see Davies, D.M.), 225, 228

Peterson, B.V., (see Wood, D.M.), 206, 210

Peterson, D.G., 225, 229

Peterson, D.G. (see West, A.S.), 225, 229

Peterson, D.G. (see Wolfe, L.S.), 225, 229

Phanaeina, 235, 242

Phanaeus, 243, 253

daphnis, 233, 242, 244

Phasmatodea, 6, 16, 79

Philander griscescens, 110

opossum, 110, 111, 112, 117, 121, 166

melanurus, 112

Philarctus quaeris, 313

Philocasca alba, 25, 45, 49, 50, 65, 67

thor, 45, 49

Phryganea fusca, 42

Phthiraptera 78

phyllisae, Plocopsylla, 91, 1 14, 127, 128, 166,

172, 174

Phyllostomus discolor, 112

hastatus, 1 1 2

Phyllotis, 176

Phytagromyza, 339

Phytomyza albiceps, 327 , 328, 329, 332, 334,

337, 338

aposeridis, 327

aquilegiae, 332

archhieracii, 337 , 338

atricornis, 339

columbiana n. sp., 327 , 332, 333, 343, 345,

erigerophila, 337

hieracina, 329, 330

inseperata, 329, 330, 332

japonica, 334

lactuca, 336

lampsanae, 329, 330, 331

marginella, 327 , 329, 332, 333, 337, 338,

342

minuscula, 332

mulgedii, 329, 330, 331

prenanthidis, 329, 330, 331

robustella, 327

senecionella, 334

sonchi, 329,330,331,332,338

sonchi cicerbitae, 329

XX

Phytomyza sonchi hieracina, 329, 338

sonchi lampsanae, 329

sonchi mulgedii, 329, 338

sonchi prenanthidis, 329

sonchina, 329, 330, 332

syngenesiae, 339

taraxaci, 331, 337, 338

Phytomyza sp., 329, 330, 331

Picea engelmanni, 279

glauca, 279

Picris, 330, 338

echioides, 330, 334

hieracioides, 330

pictipes, Simulium, 206

picturatus group, Limnephilus, 39

pigs fern, 104

Pilosa, Portulaca, 1 02

pinchaque, Tapirus, 1 1 3

Pinus ponder osa, 320

Pinus, sp., 320

phaeus, Phyllotis, 141

Phyllostomatidae, 166

Phyllotis. phaeus, 141

physodes, Oryzomys, 140

Piper aduncum, 104

pisonae, Aneurus, 257 , 268, 277 , 308

Pithcellobium dulce, 102

plantanifolia, Cavanillesia, 104

platanillos, 104

Platycentropus plectrus, 25, 44, 49, 50,

51, 65

Platyphylax alascensis, 46

Plecoptera, 322, 323

plectrus, Hylepsyche, 44

plectrus, Platycentropus, 25, 44, 49, 50, 51,

67

Pleochaetis, 94, 173, 180

equatoris equatoris, 117, 131, 172, 174

dolens quit anus, 131

smiti, 117, 131, 132, 134, 166, 172, 174

Pleridium aquilinum, 104

Philopotamidae, 52, 69

Plocopsylla, 173, 175

phyllisae, 91, 114, 127, 128, 166, 172,

174

scotinomi, 175

thor, 114, 127, 128, 166, 172, 174

Podalonia valida, 3 1 8

Pogonini, 84

politus, Aneurus, 263, 266, 267 , 269, 271, 272,

278, 289, 290, 291, 292, 294, 297, 300, 302,

306

Pollitzer, R., 94, 181

Polytricum sp., 105

Polygenis bohlsi bohlsi, 116, 132, 142, 166, 172

174

brachinus, 138

caucensis n. sp., 91 , 116, 137, 142, 144, 145,

166, 172, 174

delponti, 91, 116, 138, 146, 147, 148, 166,

172, 174

dunni, 113, 116, 139, 141, 149, 172, 174

hopkinsi n. sp., 91, 1 15, 1 16, 139, 150, 166,

172, 174

klagesi, 116, 140, 152, 166, 172, 174

klagesi samuelis, 91, 140

pradoi, 91, 116, 140, 153, 166, 172, 174

roberti, 172, 174

roberti beebei, 113, 116, 137, 140, 141, 154

thurmani, 91, 115, 141, 155, 166, 172, 174

trapidoi n. sp.,91, 115, 116, 141, 156, 166,

172, 174

popayanus, Thomasomys, 1 1 1

Portulaca pilosa, 1 02

potatoes, 105

Potos flavus, 1 12

flavus megalotus, 1 10

Potter, D.S. (see Newell, R.L.), 28, 30, 33, 55

potteri, Rhyacophila, 30

poultry, 226, 227

pradoi, Polygenis, 91, 116, 140, 153, 166, 172,

174

pradoi, Rhopalopsyllus, 140

Pratt, J.J., Jr. (see Babers, F.H.), 185, 186

Prenanthes, 331

purpurea, 330

prenanthidis, Phytomyza, 329, 330, 331

prenanthoides, Cicerbita, 331

primates, 107, 166

prinoides, Quercus, 272

prita, Psychoglypha, 15, 46, 49, 65

Procyon cancrivorus panamensis, 1 1 2

Proechimys, 140

colombianus, 112

guyannensis, 1 1 2

semispinosus, 112, 121, 140, 142, 166

Prolenarchus, 43, 48

prolixus, Rhodnius, 12

XXI

Prosimulium, 228

conistylum, 207 , 209

fulvum, 224

fuscum, 224

hirtipes, 224, 225

inflation, 210

mixtum, 224

Prosopis juliflora, 1 02

Prosympiestinae, 265

Provancher, L., 281, 288

Prunus, 105

Psychodidae, 198, 209

Psychoglypha alascensis, 25, 46, 47, 49, 65

alaskensis, 46, 49

prita, 25 ? 49, 65

schmidi, 49

subborealis, 46, 49

ulla, 46, 47, 49

Psocoptera, 77

pterota, Fugatera, 102

Ptychopteridae, 198, 209

pubescens, Ammophila, 318

pucherani, Sciurus, 1 1 1

Pudu mephistophiles, 113

pulchella, Arctopora, 312

pulchellus, Neophylax, 37

Pulex, 1 73, 1 75

bohlsi, 132

cheopis, 158

felis, 164

klagesi, 140

irritans, 93, 164, 166, 169, 172, 174,

175

segnis, 131

simulans, 91, 114, 164, 166, 172, 174,

175

pulex, Rhynchopsyllus, 91, 94, 1 13, 164,

165, 170, 172, 174, 175

Pulicidae, 158

Pulicinae, 158

Pulicini, 164

Pulicoidea, 158

pulmonarioides, Hieracium, 331

punctata, Dasyprocta, 110, 112, 142, 166

puncticollis tubericeps, Malagoniella, 247

Puri, I.M., 198, 209

purpurea, Prenanthes, 330, 331

pusillus, Aneurus, 263, 266, 268, 277 , 284,

292, 300, 303, 307

Putnam, J.D., 34, 55

Pygiopsyllinae, 121, 180

Pygiopsyllidae, 121, 175

pygmaeus, Aneurus, 255, 263, 266, 269, 275,

276, 304, 307

quaeris, Philarctus, 3 1 3

quadridens, Cephalodesmius, 245

Quercus prinoides, 272

virginiana, 215, 276

Quassia sp., 105

quitanus, Ploechaetis dolens, 1 3 1

quitoense, Solanum, 105

rabbits, 107, 226, 247

racemosa, Langulallaria, 104

Rageau, J. (see Grenier, P.), 197, 199, 209

rainbow trout, 322

Rapnea sp. 105

rat, spiny, 140

Rattus, 3, 158, 175, 176

novegicus, 3, 110, 111, 112, 158, 166

rattus, Rattus, 3, 110, 111, 112, 131, 158, 166

Raven, P.H., 106, 107, 181

Ray, C., 86, 88

redbud tree, 176

Reed, E.B., 322, 323, 325

Reichardia, 330

Reithrodontomys mexicanus, 110, 138

mexicanus milleri, 1 1 1

Rempel, J.G., 221, 222, 229

Rempel, J.G., (see Fredeen, F.J.H.), 221, 228, 325

Rempel, J.G., (see Millar, J.L.), 222, 229

Rempel, J.G., (see Arnason, A.P.), 324

Renjifo-Salcido, S., 165, 181

re x, Ctenidiosomus, 115, 121, 172, 174

rhanidophorus, Asynarchus, 42

rhanidophorus, Limnophilus, 42

Rheomys, 1 17

Rhipidomys latimanus, 110, 117, 131, 140,

141, 166

similis, 117, 124, 166

Rhizophora brevistyla, 104

Rhodnius prolixus, 12

Rhopalopsyllidae, 132, 176, 180

Rhopsalopsyllinae, 132

Rhopalopsyllini, 132

Rhopalopsylloidea, 132

Rhopalopsyllus, 115, 164, 173, 176

australis tupinus, 115, 142, 161, 166, 172,

174, 176

XXII

Rhopalopsyllus beebei, 140

cacicus saevus, 1 13, 115, 158, 162, 166,

172. 174. 176

dunni, 139

lugubris, 1 13, 1 15, 158, 162, 166, 172,

174. 176

lugubris cry ptocteres, 158

pradoi, 140

Rhyacophila acropedes, 28

autumnalis, n. sp., 25, 30, 34, 50, 51,

58, 60, 66

bifila, 33

coloradensis, 33

donaldi, n. sp., 25, 29, 34, 50, 51, 58,

59, 66

ebria, 29

hyalinata, 33

milnei, 25, 31, 34, 50, 60, 66

narvae, 33, 34

ophrys, 27

potteri, 30

rickeri, 30

vepulsa, 33

simplex, 25, 27, 34, 50, 58, 59, 66

unimaculata, 25, 30, 34, 50, 51, 60, 66

vobara, 33

vocala, 33

vao, 25, 27, 28, 34, 50, 51, 59, 66

Rhyacophilidae, 25, 26, 27, 50, 51, 31 1

Rhynchopsyllus megastigmata, 91, 94, 165

pulex, 91, 94, 113, 164, 165, 170, 172

174, 175

Rhypidae, 198, 209

rickeri group, Neophylax, 37

rickeri, Neophylax, 25, 37, 47, 50, 51,58,

61 , 66

rickeri, Rhyacophila, 30

robinii, Aepopsis, 84

roberti, Polygenis, 172, 174

Robineau-Desvoidy, J.-B., 329, 340

robinsoni, Marmosa, 1 1 2

roble, 102

robustella, Phytomyza, 327

Rodentia, 166

rodents, 92, 93, 102, 108, 124, 127, 131,

132, 141, 173, 175, 176, 241

rodents, caviomorph, 107, 175, 176

rodents, cricetine, 92, 106, 124, 127, 132,

141, 175, 181

rodents, myomorph, 107, 176

rodents, oryzomine, 132

rodents, sciuromorph, 107

roseae, n. sp., Aneurus, 255, 263, 265, 266, 269,

279, 289, 290, 291, 292, 294, 297, 300, 302,

306

rosenbergi, Neotyphloceras, 116, 117, 120, 172,

174, 176

rosenbergi, Typhlocerus, 117

Ross, H.H., 26, 27, 30, 33, 34, 37, 39, 40, 42,

43,44, 55

Rothmaler, W., 328, 241

Rothschild, M. (see Hopkins, G.H.E.), 165, 175,

180

rotunda group, Rhyacophila, 28

rotundus, Desmodus, 112, 164, 166

Roy, D. 38,43,44,55

Royer, L.M. (see Fredeen, F.J.H.), 325

rubber, 104

rubra, Crepis, 331

rubrum, Acer, 282

Rubtzov, I.A., 197, 198, 209

Rubus sp., 105

rufescens, Echinoprocta, 1 1 1

ruficollis, Calathus, 86

rugglesi, Simulium, 226

runcinata, Crepis, 334

Ryden, N.S., 329, 330, 331, 332, 341

sabaudum, Hieracium, 330, 331

Sabrosky, C.W. (see Stone, A.), 220, 229

saevus, Rhopalopsyllus cacicus, 158, 162, 166,

172, 174

Saha, J.G., (see Fredeen, F.J.H.), 325

samuelis, Polygenis klagesi, 140

Sanchez, E.H., 94, 181

Sangha, G.K. (see Kapoor, I.P.), 325

sanguinosus, Aradus, 281

sapiens, Homo, 164, 165, 166, 175

Sarkar, D. (see Sengupta, R.), 183, 187

Sasakawa, M., 334, 341

Sauer, C.O., 94, 181

Savage, A., 227, 229

Savage, J.M., 106, 181

Saxifragaceae, 339

Say, T., 271, 278, 288

Scarabaeidae, 231, 252, 253

Scarabaeinae, 231, 232, 233, 241, 242, 245,

249,251,253

Scarabaeini, 233, 235, 237, 243, 245, 247, 248,

249, 253

XX111

scariola var. integrata, Lactuca, 233

Schistocerca gregaria, 13, 15

schisto siphon, Hieracium, 33 1

Schmid, F., 26, 27, 28, 30, 33, 34, 36, 37

39,42,43,44, 55, 56

Schmid, F., (see Denning, D.G.), 30, 54

schmidi, Psychoglypha, 49

Schmidt, K.J., (see Fest, C.), 183, 186

Sciuridae, 166, 175

Sciurus granatensis valdiviae, 108, 166

pucherani caucenis, 1 1 1

Scolopsyllus, 173, 176

colombianus, 115, 119, 141, 159, 160,

166, 172, 174, 181

scotinomi, Plocopsylla, 175

Scott, J. (see Douglas, J.), 256, 288

Scott, W.B., 106, 107, 181

scrofa, Sus, 165, 175

Scudder, G.G.E., 77, 78

Sechium edule, 104

secludens, Limnephilus, 39, 40

segnis, Leptopsyllus, 91, 115, 131, 133,

166, 172, 174, 175

segnis, Pulex, 1 3 1

Sehgal, V.K., 334, 336, 337, 341

Seidel, J., 331, 341

semispinosus, Proechimys, 121, 140, 142,

166

semistriatus, Conepatus, 110

Senecio, 105, 327, 334, 339

atropurpureus tomentosus, 334

congestus var. palustris, 334

Senecioneae, 334, 339

senecionella, Chromatomyia, 327 , 334

senecionella, Phytomyza, 334

Sengupta, R., 75, 183, 187

sensitive-plant, 104

septentrionalis, Aneurus, 255, 278

sericea, Freziera, 105

Serra-Tosio, B., 207, 210

serriola, Lactuca, 331, 336, 337, 345

Severin, G. (see Lethierry, L.), 270, 271,

275,278,281,288

shale, 97

sheep, 222, 241

Sherman, F., 281, 288

Shewell, G.E., 226, 229

sibirica, Crepis, 331

sibirica group, Rhyacophila, 30

Sigmodon sp., 117, 132, 176

hispidus, 1 1 2

hispidus bogotensis, 110, 112

Sigmodontini, 107

silkworms, 16

silvaticum, Hieracium, 331

similis, Rhipidomys, 117, 124, 166

Simon, Jean-Pierre, 183, 187

simile, Simulium, 228

simondi, Leucocytozoon, 227 , 228, 229

simplex, Aneurus, 255, 263, 266, 268, 279,

289, 290, 291, 292, 294, 297, 300, 303, 305

simplex, Rhyacophila, 23, 27, 34, 50, 58, 59, 66

Simpson, G.G., 106, 107, 166, 181

simulans, Pulex, 91, 1 14, 164, 166, 172, 174, 175

Simuliidae, 195, 197, 198, 204, 206, 207, 209,

210, 219, 226, 228, 229, 322, 335

Simulium, 325

arcticum , 221, 222, 223, 224, 228, 229, 321,

322,323,324,325

aureum, 228

cheesmanae, 195, 196, 207

croxtoni, 226

decorum, 224

defoliarti, 221

euryadminiculum, 226

griseum, 224

johannseni, 22 6

latipes, 226, 228

luggeri, 219, 223, 321, 323,

meridionale, 226

oviceps, 195, 196, 197, 199, 200, 201, 203,

204, 205, 206, 207, 217, 218,

parnassum, 224, 226

pictipes, 206

rugglesi, 226

simile, 228

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

203, 204, 205, 206, 207, 21 1, 212, 214,

215,216,217

tuberosum, 224

venustum, 207, 219, 224, 225, 226

vittatum, 196, 200, 202, 206, 207, 224, 225,

Siphonaptera, 77, 77, 81, 91, 94, 122, 130, 134

139, 140, 149, 154, 161, 162, 163, 165, 168,

170, 171, 173, 177, 180, 181

Sitophilus (=Calendra) oryzae, 11,13

Sittidae, 226

Skala, H., 331,341

XXIV

slateri n. sp., Aneurus, 255, 263, 266, 268,

285, 289, 290, 291, 292, 295, 297, 300,

303, 306

Smallman, B.N., 183, 187

Smart, J., 196, 198, 210

Smith, E.L., 78

Smith, E.M. (see Freyvogel, T.A.), 184, 186

Smith, J., 184, 187

Smith, S.D., 27, 33, 56

Smithies, O., 184, 187

smiti, Pleochaetis, 117, 131, 132, 134, 166,

172, 174

socialis, Nasua, 140

Solanum quitoense, 105

tuberosum, 105

Solidago, 330, 332

sonchi, Phytomyza, 329, 330, 331, 332, 338

sonchi cicerbitae, Phytomyza, 329

sonchi hieracina, Phytomyza, 329, 338

sonchi lampsanae, Phytomyza, 329

sonchi mulgedii, Phytomyza, 329, 338

sonchi prenanthidis, Phytomyza, 329

sonchina, Phytomyza, 329, 330, 332

Sonchus arvensis, 330, 336

arvensis uliginosus, 336, 337

asper, 330, 334, 336

oleraceus, 330, 334, 336

paluster, 330

Sdnderup, H.P.S., 331, 341

Sortosa, 69, 70

soricina, Glossophaga, 110, 112, 164, 166

speciosa, Sternopsylla distincta, 115, 127,

130, 166, 172, 174

Spencer, G.J., (see Ross, H.H.), 27, 30, 33,

34, 37, 39, 55

Spencer, K.A., 329, 330, 331, 332, 336,

337, 341

Spencer, K.A. (see Hering, E.M.), 331, 339

Sphecidae

Sphinctopsylla, 17 3, 175, 176

diomedes, 91,94, 114, 124, 126, 166,

174

tolmera, 114, 127, 166, 172, 174

spicata, Lactuca, 331

Spondias mombin, 102

sponges, 97

spurge, 102

squirrels, 107, 173

squirrel, red, 108

Stainer, J., 56

Stal, C., 271,278, 281,288

stalzmanni, Oryzomys, 141

Staphylinidae, 83, 88

Starke, H., 331,341

Stary, B., 331, 341

Stebbins, G.L., 330, 337, 341

Steiner, A.L., 315,318

Steniolia, 316, 318

obliqua, 315

Stenophylax fusorius, 42

gilvipes, 34

modesta, 42

Stenopsyche griseipennis, 6, 11, 12, 13, 15

Stenopsychidae, 15

Stephanocircidae, 124, 175, 180

Stephano circus, 175

Sternopsylla, 173

Sternopsylla distincta, 175

intermedia copha, 117

speciosa, 1 15, 127, 130, 166, 172, 174

Stirton, R.A., 98, 106, 182

Stone, A., 198,210, 220,229

strawberries, 105

Strepsiptera, 77

Sturnia lilium, 1 1 2

Stys, P., 257, 259, 262, 265, 271, 288

subborealis, Psychoglypha, 46, 49

subcentralis group, Limnephilus, 38

subhyalinus, Glaphyrocanthon, 243

subrufa, Carollia, 112

superba, Gustavia, 104

surrumbo, 104

Sus scro fa, 165, 175, 222

Sylvilagus andinus, 112

brasiliensis, 110, 111, 112

fulvescens, 110, 111

syngenesiae, Chromatomyia, 327, 328, 334,

335, 337

syngenesiae, Phytomyza, 339

Tabebuia chrysantha, 104

pentaphylla, 102

Tadarida sp., 127, 166

brasiliensis, 110, 127

tahitiense, Simulium, 195, 196, 197, 199, 200,

201,202, 203, 204, 205, 206, 207, 211, 212,

213,214,215,216,217

tajacu, Tayassu, 112, 142

Talinum paniculatum, 102

XXV

Tamandua tetradactyla, 1 12

Tamsitt, J.R., 94, 182

tapirs, 107

Tapirus bairdii, 1 1 2

pinchaque, 1 1 3

taraxaci, Phytomyza, 331, 327, 338

Taraxacum, 331, 333, 337, 338

kok-sghyz, 334

officinale, 331, 333, 337

Tarshis, I.B. (see Herman, C.M.), 227, 229

tatarica, Lactuca, 331

Tate, G.H.H., 113, 182

Tayassu pecari, 112, 142

tajacu, 112, 142

tectorum, Crepis, 334, 337

Tenebrio molitor, 6, 7, 11, 16

Tenebrionidae, 6

tenuicornis, Aneurus, 272

tenuipes, Neacomys tenuipes, 1 1 2

tenuis, Aneurus, 263, 266, 270, 272, 273

289, 290, 291, 292, 295, 297, 300, 308

testaceus, Thalassobius, 84

Tetraonidae, 226

Tetrapsyllus, 173, 176

com/s, 91, 115, 132, 136, 166, 172, 174

tetradactyla, Tamandua, 1 1 2

Tettigoniidae, 77

Thalassotrechus barbarae, 75, 83, 84, 85,

86, 87,88,89,90

barbarae barbarae, 83, 86

barbarae nigripennis, 84, 85, 89

testaceus, 84

Thallomyscus, 182

thapsoides, Hieracium, 331

Thierry, A. (see Freyvogel, T.A.), 184, 186

thomasi, Crypto tis, 1 1 1

thomasi, Imania, 25, 35, 47, 50, 58, 62, 66

Thomasomys sp., 121, 127, 166, 173, 176

aureus, 110, 111, 117, 121, 166

cinereiv enter, 111, 113, 117, 121, 124,

127, 131, 132, 164, 166

fuscatus, 111, 117, 121, 124, 132, 140,

166

hylophilus, 124

laniger, 121, 124

po pay anus, 1 1 1

vestibus, 124

thor, Philocasca, 45, 49

thor, Plocopsylla, 114, 127, 128, 166, 172,

174

Thorpe, W.H.,317,318

thous, Cerdocyon, 112, 164, 166

Thraupidae, 226

thurmani, Polygenis, 91, 115, 141, 155, 166,

172. 174

Thysanoptera, 8 1

Thysanura, 77, 78

Tiamastus, \16

Tiarapsylla, 176

tillyardianum, Austro simulium ( Austro simulium),

209

Tipton, V.J. (see Wenzel, R.L.), 107, 165, 175,

176, 182

Tipton, V.J. and G.E. Machado-Allison, 1 13,

132, 139, 158, 182

Tipton, J&E. Mendez, 113, 121, 122, 130, 134,

139, 142, 149, 154, 158, 161, 162, 163, 165,

168, 170, 171, 173, 182

tiptoni, Kohlsia, 173

tolmera, Sphinctopsylla, 114, 127, 166, 172, 174

Torre-Bueno, J.R. de la, 281, 288

tortulosus, Dichotomius, 235

transylvanicum, Hieracium, 331

trapidoi, n. sp Polygenis, 91, 115, 116, 141, 156,

157, 166, 172, 174

traubi, Ctenidiosomus, 91, 94, 115, 121, 122,

124. 174

Trechini, 84

Trechus ovipennis, 84

Trema micantha, 104

Tremarctos ornatus, 113

Trichant era gigantea, 1 04

Trichoceridae, 198,209

Trichoptera, 6, 15, 25, 26, 27, 30, 60, 311, 312,

313,322,323,

trilobites, 97

tripunctata group, Imania, 35

triste, Hieracium, 333, 345

Triticum, 105

trivirgatus, Aotus, 1 1 1

trompet, 104

tropicalis, Odocoileus, 1 1 2

Trypanosoma, 226, 227, 228

Tschirnhaus, M. von, 328, 341

tuberosum, Simulium, 224

tuberosum, Solanum, 105

Tunga, 173, 175

Tunga penetrans, 114, 165, 166, 171, 172, 174,

175

XXVI

Tunginae, 164

tupinus, Rhopalopsyllus australis, 115, 142,

161, 166, 172, 174, 176

Turdidae, 226

turkey, 227, 229

Turnera almifolia, 102

turtles, 181

Tussilago, 339

Twinnia biclavata, 204

Tylomys mirae, 112

typhus, murine, 93

Typhloceras rosenbergi, 1 1 7

Tyrannidae, 226

Udvardy, M.D.F., 86, 88

Uhler, P.R., 225, 271, 278, 281, 288

ulla, Psychoglypha, 46, 47, 49

Ulmer, G., 34, 42, 46, 56

umbellatum, Hieracium, 331

Umbelliferae, 332

umbellus, Bonasa, 228

ungulates, 91, 106, 164

unimaculata, Rhyacophila, 25, 30, 34, 50,

51, 60, 66

Urocyon cinereoargenteus, 110, 166

Uroderma b. bilobatum, 1 12

Usinger, R.L., 256, 265, 270, 271, 272,

275,278,281,285,288

usingeri n. sp., Aneurus, 255, 263, 266, 269,

284, 285, 286, 289, 290, 291, 292, 295,

297, 300, 303, 308

Vaccinium sp., 105

Vachelia farnesiana, 102

Vadlamudi, S. (see Anderson, J.R.), 226,

228

vagrita group, Rhyacophila, 31

valdiviae, Sciurus granatensis, 108, 166

Valentine, J.W. and E. Moores, 106

Valeriana, 339

valida, Podalonia, 318

Vampyrops dorsalis, 112

helleri, 1 1 2

Van Duzee, E.P., 272, 275, 278, 281, .288

Van Dyke, E.C., 83, 84, 88

Vanzolini, P.E., 108, 182

vao, Rhyacophila, 25, 27, 28, 34, 50, 51,

59, 60

vaueri, Aneurus, 256, 263, 266, 268, 273,

274, 275, 289, 290, 291, 292, 295, 298,

301,303,308

venestus, Rhipidomys, 124

venestum, Simulium, 207, 219, 224, 225, 226

vepulsa, Rhyacophila, 33

veracruzensis n. sp., Aneurus, 225, 263, 265,

266, 269, 275, 285, 286, 289, 290, 292, 295,

298, 307

Vermipsyllidae, 180

vernalis, Limnephilus, 25, 41, 48, 50, 51, 63,

67

verrula group, Rhyacophila, 30

vestibus, Thomasomys, 124

villosum, Chiroderma, 112

villosum, Hieracium, 331

Vieronidae, 226

virens, Canthon, 247

virgatus, Desmanthus, 102

viridana, Lytta, 5, 6, 7, 9, 10, 11, 12, 13, 14, 15

16

virginiana, Quercus, 275, 276

virginianus, Odocoileus, 110, 112

virosa, Lactuca, 331

vittatum, Simulium, 196, 200, 202, 206, 207,

224, 225

vobara, Rhyacophila, 33

vocala, Rhyacophila, 33

vofixa group, Rhyacophila, 27, 33

Voigt, G., 331, 341

Vuilleumier, B.S., 98, 182

vulgatum, Hieracium, 330, 331

wasps, 315, 316, 317

Waite, E.R.,247,253

Walker, F., 255, 278, 288

Wallengren, H.D.J., 42, 56

Waters, T.F., 323, 325

Weeks, L.G., 98, 182

Wenzel, R.L., 107, 165, 175, 176, 182

West, A.S.,-225, 229

Weinmannia sp., 105

Weinmannia balbisiana, 105

West, R.C., 182

wheat, 105

White, E.M. (see Greiner, E.C.), 226, 229

Wickware, A.B., 226, 229

Wigania caracasana, 1 04

Wiggins, G.B.,311

Wille, A., 247, 253

Williams, T.R., (see Kershaw, W.E.), 325

Wirth, W.W. (see Stone, A.), 220, 229

Wobeser, G., 226, 229

XXV11

Wold, J.L. (see Anderson, N.H.), 28, 37, 53

Wolfe, L.S. (see Peterson, B.V.), 225, 229

Wood, D.M., 206, 210

Wood, D.M. (see Davies, D.M.), 225, 228

worms, leg, 226

wrack, 83

wygodzinskyin. sp., Aneurus, 255, 263, 265

266, 270, 271, 272, 289, 290, 291, 292,

295, 298, 301, 303, 308

Xenopsylla, 173

cheopis, 93, 114, 158, 164, 166, 167,

172, 174, 175

Xenopsyllini, 158

yagouaroundi, Felis, 1 1 0

yaragua, 104

Yepes, J. (see Cabrera, A.), 108, 113

Yin, R-S, L., 197, 198, 210

youngi, Diaemus, 1 1 2

Zavrel, H., 331, 341

Zavrel, H. (see Skala, H.), 331, 341

Zea mays, 105

Zetterstedt, J.W., 42, 56

zetti, Mazama americana, 1 10

Xiphiopsyllidae, 180

Zoraptera, 79

Zygodontomys brevicauda brunneus, 110, 112

ffl-A

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Quaestiones

Entomologicae

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

VOLUME 13

NUMBER 1

JANUARY 1977

QUAESTIONES ENTOMOLOGICAE

ISSN 0033-5037

A periodical record of entomological investigation published at the Department of

Entomology, University of Alberta, Edmonton, Alberta.

Volume 13 Number 1 January 1977

CONTENTS

Editorial — Comments About a Few Words in the Biological Sciences 1

Rempel, Heming and Church — The Embryology of Lytta Viridana LeConte

(Coleoptera: Meloidae). IX. The Central Nervous System, Stomatogastric

Nervous System, and Endocrine System 5

Nimmo — The Adult Trichoptera (Insecta) of Alberta and Eastern British Columbia,

And Their Post-Glacial Origins. I. The Families Rhyacophilidae and

Limnephilidae. Supplement 1 25

Nimmo — The Adult Trichoptera (Insecta) of Alberta and Eastern British Columbia,

And Their Post-Glacial Origins. II. The Families Glossosomatidae and

Philopotamidae. Supplement 1 69

Belicek — Corrigenda on Coccinellidae of Western Canada and Alaska with Analyses

of the Transmontane Zoogeographic Relationships Between the Fauna of

British Columbia and Alberta (Insecta: Coleoptera: Coccinellidae) 73

Editorial — Comments About a Few Words in the Biological Sciences

Editors, perhaps to a greater extent than most individuals who are interested in literature,

find word usage a subject for serious consideration, from which they derive both pleasure and

pain. An editor may develop concepts of the meanings of certain Words, or about how certain

words should be used, with his notions differing from those that seem generally accepted. These

concepts are expressed in modifications to those manuscripts that an editor is invited to consi-

der for publication in his journal.

Of course, each editor is convinced that his concepts are correct and at least worthy of at-

tention, if not emulation, by others. This is the thought that has impelled me to list in alpha-

betical order and discuss some of the words that seem to me to be commonly mis-used in bio-

logical literature generally, and especially in the literature of systematic biology. Because I am

a systematist, I have also listed some commonly used words that I think are inappropriate in

writing about systematics.

ALWAYS (antonym - never). — I think it’s best to avoid this word and its antonym in wri-

ting about fields that rely mainly on observation for information about nature, and hence are

mainly inductive. Further, does the statement “prolegs are always absent” mean anything dif-

ferent from “prolegs are absent”? Unqualified, “absent” means just that. I suppose “always”

gives emphasis to the adjective, but this seems unnecessary.

ANATOMY. — This word refers, literally, to dissection. It is not a synonym of morpholo-

gy (study of structure and form) except by virtue of careless usage. Neither should it be used

as botanists and some entomologists do: that is, to refer to internal structures and organs. I

suppose this unfortunate usage developed because one must dissect (anatomize) to examine

internal organs. The noted arachnologist, T.H. Savory wrote (Browsing Among Words of Sci-

ence, 195 1, page 67): “we read of . . . external anatomy, which if it means anything can only

mean shaving; and of comparative anatomy, which ought to be a contrast between the use of

scissors, scalpels and saws . . . .” Why not use the simple terms “external structure” and “in-

ternal structure” when referring to these aspects of morphology? See also MORPEIOLOGY,

below.

2

BIOLOGY. — This is a term that refers to a field of human endeavor. It means “study of

life”. It seems to me utterly incorrect to use “biology” for the non-morphological, non-class-

ificatory portions of the general field. By implication then, morphology and systematics are

excluded from biology, and in turn, morphologists and systematists are not biologists!

An organism or group of organisms cannot have a biology. An organism has a life, and study j

of its life (in the broadest sense) is the domain of biology. j

Rather than entitling that section of a systematic treatment “Notes on Biology” that deals j

with some special aspects of the living members of a particular taxon, it’s best to be more spe- !

cific, and more accurate. If the notes deal with host associations, then use these words for the

title; if they deal with habitat and life history, then use these words for the title. j

CASE (synonym - situation), as used in the expression “in this case”. — I doubt that this

phrase need be used outside the literature of the legal profession or the brewing or cartage in-

dustries. More specific expressions are available for biological literature.

CLASSIFICATION. - See TAXONOMY.

DIVERGENCE. — This term should be used to express relative amount of difference among

taxa. One group, whose members are removed by appreciable structural differences from mem-

bers of other taxa, is said to exhibit marked divergence, or to be highly divergent. A higher

taxon whose member taxa exhibit considerable difference from one another is said to be high-

ly divergent in comparison with another taxon whose member taxa show slight differences.

See also DIVERSITY.

DIVERSITY. — This term is best restricted to reference to numbers of members of groups

(either individuals or taxa, or both). A higher-ranking taxon including many lower-ranking taxa

is highly diverse, but some exhibit slight divergence. On the other hand, some highly divergent

taxa exhibit low diversity. In any event, it’s best to refer to inclusiveness of a taxon in terms

of amount of diversity or words implying this (that is, “this genus includes few (or many)

species”), rather than as “large” or “small”. These latter terms deal with size, too, but the con-

text is different. Also, as these words are used, a “large genus” can be either one including many

species, or one that includes species whose members are of large size.

DIVISION. — This is not an appropriate word to describe the actions of systematists with

reference to taxa. Activities include grouping, re-grouping, and arranging. Re-grouping of spe-

cies that results in more genera than were previously recognized seems to be described appro-

priately by the word division, but it is not the same thing as, say, cutting a pie, which has a

high degree of physical continuity, or cutting a piece of string. Of course, dissection is a form 1

of division, and when a biologist makes a dissection he is indeed dividing.

LARGE (as applied to taxa). - See DIVERSITY.

MAY. — As used with reference to occurrence or non-occurrence of characters among mem-

bers of a taxon, this word is inappropriate. In fact, in each instance, a given character state is

or is not exhibited. So, rather than writing “structure X may be present”, it’s better to write

“structure X is present in a few individuals; it is absent from most”.

MORPHOLOGY. — Means study of form and structure. An individual (not a taxon) has a

particular structure, or a particular combination of structures. It does not have a morphology.

As with the term “anatomy”, what can “external morphology” possibly mean: “the external

study of form and structure”? When referring to parts of a particular organism, the word struc-

ture is adequate, and does not need the embellishment of the word morphology.

NEVER. - See ALWAYS.

OCCASIONALLY (related words - rarely, sometimes, usually). - This word is not appropr-

iate in description of distribution of static characters states among taxa. What is implied by

most authors is that a particular character is represented by more than one state, and expression

of the less frequent one is described as occasional. It is best to indicate the matter more precisely.

3

without reference to time, as follows: “members of most species with red antennae, those of

a few species with black antennae”; in preference to “antennae usually red, occasionally (or

sometimes, or rarely) black”.

On the other hand, if antennal color of an individual changes periodically and is usually red,

then it would be occasionally (or sometimes, or rarely) black.

RARELY. - See OCCASIONALLY.

SMALL (with reference to taxa). — See DIVERSITY.

SOMETIMES. - See OCCASIONALLY.

SPECIES (and other taxa). — These have members, which are represented by eggs, immatur-

res, or adult males and females. These members have characters — structural, behavioural eco-

logical, genetical, physiological, and so on. Species and other taxa do not have characters. Thus

it is incorrect to write that “A few species deposit their eggs in the nests of other bees”, or

that “the aedeagus of this species has a knob on the distal end”. Rather, one should write that

“Lemales of a few species deposit their eggs . . . ” et cetera', or that “aedeagi of males of this

species” . . . et cetera.

SYSTEMATICS. - See TAXONOMY.

TAXONOMY. — This is the general term for study of the principles of classification, appli-

cation of the principles to formation and ranking of formal groups (taxa), and naming the taxa.

Systematics is the field of organic diversity, and taxonomy is a portion of this field.

Classification is used in two senses: 1. for the process of establishing taxa; 2. for the resul-

ting formal system of arrangement. Thus, one classifies as one constructs a system. The sys-

tem is a classification.

I doubt that it is correct to write, as the title of a paper: “The Taxonomy of the Genus

X-us ”. Genus X-us does not have a taxonomy. Rather, its species are arranged, or classified in

a particular way, according to taxonomic principles.

The word taxonomy also seems to me to be mis-used with reference to study of an existing

classification for the purpose of learning to identify specimens. This type of activity is not tax-

onomy.

TECHNOLOGY. — This word must mean “study of techniques”. It is used by many writers

and speakers in place of technique, so that even the most mundane and inconsequential of pro-

cedures is given some aura when it is referred to as a technology!

THE. — Categorical expressions (as implied by use of “the”) are to be avoided with referen-

ce to selected individuals. For instance, vertebrate physiologists commonly use the expression

“the rat” in published work, when what is meant is groups of rats of the species Rattus norve-

gicus on which some experiments have been conducted. “Rats”, if a more modest expression

than the categorical “the rat”, is also a more accurate expression.

UNIQUE. — Because members of each taxon have some unique features, it is inappropriate

to state that a given taxon is unique without specifying in what way this is so.

USUALLY. - See OCCASIONALLY.

VARIABLE. — This means that a feature of an individual has the capacity to exhibit varia-

tion. However, in descriptive writing, what is meant by most authors is that a particular char-

acter is expressed in one or more states. Each state is either expressed or not expressed in a

given individual, and is non-varying therein. Hence, it seems preferable to use “varied”, or “var-

ious”. For example, I prefer: “The general form of beetle larvae is widely varied” rather than

“ . . . form of beetle larvae is variable”. To me, the second expression implies that a given indi-

vidual at different times might be compodeiform, scarabaeiform, and eruciform. Of course,

when one writes about larvae of meloid beetles (and larvae of some other groups, too), “var-

iable” is indeed the word of choice, because form of an individual larva changes markedly dur-

ing development.

4

Examples of uses of many of these words that are counter to my present views are in my

earlier publications. Hopefully, these past actions will not be held against me, or used to ne-

gate my arguments! I invite the readers of Quaestiones Entomologicae to consider not only

my views on the use of these words, but also the more general question of economy and ac-

curacy of written expression of observations and ideas. I’ll be pleased to receive comments

about these matters.

George E. Ball

THE EMBRYOLOGY OF L YTTA VIRIDANA LE CONTE

(COLEOPTERA: MELOIDAE). IX. THE CENTRAL

NERVOUS SYSTEM, STOMATOGASTRIC NERVOUS

SYSTEM, AND ENDOCRINE SYSTEM 1 2 3

o

J.G. Rempel

Department of Biology

University of Saskatchewan

Saskatoon, Saskatchewan

Canada

B.S. Heming ^

Department of Entomology

University of Alberta

Edmonton, Alberta

Canada T6G 2E3

N.S. Churchy-

Research Station

Agriculture Canada

Saskatoon, Saskatchewan Quaestiones Entomologicae

Canada 13: 5-23 1977

The central nervous system of Lytta viridana arises in the usual way from ventral, longitu-

dinal files of neuroblasts in the head, thorax, and first 10 abdominal segments of the embryo.

Median nerve strand cells have two fates: clumped, inter segmental cells shift cephalad and

differentiate into ganglion cells ; intrasegmental cells probably develop into glial elements of

the lateral nerve cords. The perilemma appears to originate from modified outer ganglion

cells.

The stomatogastric nervous system develops from three evaginations in the roof of the

stomodaeum: the frontal ganglion from the first, the hypo cerebral ganglion and corpora car-

diaca from the second, and the ventricular ganglion from the third. It is suggested that these

three ganglia are not serially homologous with the intersegmental clumps of the post-oral me-

dian nerve strand.

A corpus allatum invaginates inward from the anterior face of each maxillary base and

eventually fuses with a corpus cardiacum below the brain. Probable prothoracic glands pro-

liferate inward between the inter ganglionic connectives from the ventral, labial-pro thoracic

intersegmental ectoderm. Their cells eventually spread over the surface of a transverse trach-

eal commissure emanating from branch 2 of the mesothoracic spiracular tracheal system.

Observations are discussed in relation to findings on the origin and function of compara-

ble structures in other insects.

1. Previous papers in this series were published in the Canadian Journal of Zoology.

Dr. Rempel stipulated that he wished all subsequent ones to appear in Quaestiones

Entomologicae.

2. Deceased.

3. Requests for reprints should be sent to this address.

6

Rempel, Heming & Church

Le systdme nerveu central de la Lytta viridana se forme d’une facon normale a partir de files ventrales et longitudinales

de neuroblastes dans la tdte, le thorax, et les dix premiers segments abdominaux de I’embryon. Les cellules nerveuses et

dtroites apparaissent en deux groups: les cellules intersegmentaires groupdes s’assemblent antdrieurement et se diffdrencient

en cellules ganglionaires et les cellules intrasegmentaires probablement se develop pent en dldments gliaux de la chaine ner-

veuse la t dr ale. Le perilemme semblent dtre formd de cellules ganglionaires externes et modifides.

Le systdme nerveux stomatogastrique se ddveloppe a partir de trois evaginations dans la partie supdrieure du stomodeum:

du premier vient le ganglion frontal, du second le ganglion hypocdrdbral et les corpora cardiaca, et du troisieme le ganglion

ventriculaire. Nous suggdrons que ces trois ganglions ne sont pas des homologues en sdrie avec les groupes de cellules inter-

segmentaires du ruban nerveux median et post-oral.

Un corpus allatum pdndtre a Vinterieur a partir de la face antdrieure de la base de chaque maxillaire et dventuellement se

fusionne avec un corpus cardiacum sous le cerveau. Les glandes prothoraciques probables proliferent vers Vintdrieur entre

les connectifs rdunissant les ganglions subesophageaux et prothoraciques Vectoderme intersegmentaire labial-prothoracique

du cdtd ventral. Leurs cellules dventuellement se repandent au-dessus de la surface d’une commissure trachdaire transverse

dmanant de la branche 2 du systdme trachdaire et spiraculaire de mdsothorax.

Les observations sont discutdes en relation aux ddcouvertes sur Vorigine et la fonction de structures comparables chez

d’autres insectes.

INTRODUCTION

Early stages in the embryonic development of blister beetles of the species Lytta viridana

LeConte, have been described in previous papers in this series (Rempel and Church 1 965,

1969 a, b, 1971; Church and Rempel 1971). Organogenesis and differentiation of individual

organ systems are being treated in separate papers and the first of these, on the respiratory

system, has already appeared (Rempel and Church 1972). This paper is devoted to the ner-

vous and endocrine systems, to which brief reference has already been made (Rempel and

Church 1969b; Church and Rempel 1971). Since Ullmann (1967) described development

of the nervous system in Tenebrio molitor L. (Coleoptera, Tenebrionidae) in considerable

detail, and because events in L. viridana are very similar, we here concentrate on aspects

that supplement her publication.

Embry ogenesis of the brain and lateral nerve cords of insects is well known (Edwards 1 969;

Anderson 1972 a, b, 1973) and quantitative analysis of the events involved has begun (Bate

1976; Kankel and Hall 1976). However, there is still controversy regarding the development

and ultimate fate of the median cord or nerve strand. Although authors agree that the median

cord arises from a continuous strip of median ectoderm between the lateral cords, few studies

describe its development throughout embryogenesis.

Ontogeny of the stomatogastric nervous system has been thoroughly described recently

for T. molitor by Ullmann (1967), for Carausius (= Dixippus) morosus Br. (Phasmatodea)

by Scholl (1969), for Oncopeltus fasciatus Dallas (Heteroptera) by Dorn (1972, 1975 a, b)

and for Stenopsyche griseipennis MacLachlan (Trichoptera) by Miyakawa (1974). Like ear-

lier workers, these authors believed the system to develop from three evaginations in the

roof of the stomodaeum. According to Ullmann ( 1967), the most anterior evagination gives

rise to the frontal ganglion, the second to the hypocerebral ganglion, and the third to the

ventricular ganglion. Scholl (1969) also followed development of the frontal ganglion back

to the first evagination, but he considered the hypocerebral and ventricular ganglia to come

from the third evagination and the corpora cardiaca from the second. Miyakawa ( 1974) clai-

med that the recurrent nerve arose from the second evagination. All three interpretations

differ slightly from the traditional one of Roonwal (1937) where evagination 2 gives rise to

the hypocerebral (occipital) ganglion and corpora cardiaca (pharyngeal ganglia).

Although many reviews are available about the structure and function of insect endocrine

organs (e.g. Cazal 1 948; Pflugfelder 1958; Herman 1967; Dorn 1972; Gilbert and King 1973;

Slama, et al. 1974; Novak 1975), their embryonic development has received little attention.

In this paper, we describe the embryogenesis of the central nervous system, stomatogastric

Embryology of Lytta viridana IX

7

nervous system and endocrine system of L. viridana. Limited comment is made concerning

the neuropile and neurosecretory cells since stains appropriate for these structures were not

used. We have described all our methods in previous papers (Rempel and Church 1965, 1969b).

OBSERVATIONS

Central Nervous System

In Lytta viridana , segmentation precedes neurulation (Rempel and Church 1969b). The

former process is initiated at approximately 36 h, with coelom formation being well advan-

ced by 50 h. At this time, the ectoderm (ect), as seen in parasagittal section (Fig. 1), consists

of a single layer of columnar cells with large nuclei and conspicuous vacuoles at their inner

ends. In median sagittal section (Fig. 2), the cells appear to form an irregular double layer.

Neurulation begins at 56 h. The sequence of the two processes involved is similar to events

in T. molitor embryos (Ullmann 1964, 1967) and is probably general in insect development.

Many ectodermal cells (Fig. 3. nbl) enlarge in both their nuclei and cytoplasm and begin to

stain more strongly than neighbouring cells. Gradually, they withdraw from the surface and

move inward. As this process continues, the germ band soon separates into an outer dermato-

gene layer (dg. 1) and an inner neurogene layer (ng: 1). The latter consists of neuroblasts (nbl)

which, by successive but unequal, vertical, teloblastic division, give rise to small pre-ganglion

and ganglion cells (Fig. 4, ggl.c; see also Fig. 3 and 16 in Church and Rempel 1971). Pre-gang-

lion cells divide at least once, equally, and often at right angles to the neuroblasts, and gen-

erate ganglion cells. This process occurs in continuous, longitudinal files of cells extending on

either side of the midline from the region of the stomodaeum to the proctodaeum.

Simultaneously, along the midline, cells in intrasegmental regions continue, by equal divi-

sion, to form an apparent multi-layered strand of cells (m.n.c), while those in intersegmental

regions develop into clumps of large, neuroblast-like, darkly-staining cells (Fig. 5, m.c.c; see

also Fig. 17 and 18 in Church and Rempel 1971). At first, the dermatogene layer does not

cover these clumps ventrally. These two classes of cells comprise the median nerve strand.

This strand begins anteriorly in the intercalary segment (intc. seg) and extends posteriorly

to a region behind the tenth lateral cord ganglia (Fig. 7, m.n.c). The first clump of cells is

located in the intersegmental region between the mandibular and maxillary segments (Fig.

7, 11, 47), not, as was incorrectly stated earlier (Church and Rempel 1971, but see their

Fig. 2), between the intercalary and mandibular segments.

Cephalic ganglia develop in the same way as do the lateral nerve cords. Large neuroblasts

separate from the dermatogene layer, and, by repeated, unequal, teloblastic division, give rise

to pre-ganglion and ganglion cells. The first pair of ganglia arise pre-orally and ultimately

form the protocerebrum (Fig. 8, protc; as in T. molitor (Ullmann 1967) these ganglia are

tri-lobed, the optic ganglia (op. 1) arising as separate, ectodermal invaginations — (Fig. 42));

the second pair form paraorally and develop into the deutocerebrum (deutc) (Fig. 8); and

the third pair arise postorally and later move into a pre-oral position to form the tritocere-

brum (Fig. 9, 18, trite). Simultaneously, the stomatogastric nervous system (stmg. n.s) arises

from three evaginations in the roof of the stomodaeum (Fig. 10; in 1969, Rempel and Church,

incorrectly referred to these as invaginations, as did Ullmann (1967) . Although they are

mvaginations of the body wall, they are evaginations of the stomodaeum). The cells surround-

ing the evaginations have large, light-staining nuclei and resemble neuroblasts of the central

nervous system (Fig. 46). However, as Ullmann ( 1967) pointed out, they divide equally not

teloblastically.

By 64 h, the lateral nerve cord ganglia have enlarged and have moved mesad. Meanwhile,

the clumps of median nerve strand cells (m.c.c) have shifted cephalad from a strictly inter-

Quaest. Ent., 1977 13 (1)

8

Rempel, Heming & Church

segmental position into an intrasegmental one in the preceding segment (Fig. 6, 39). Here,

they later contribute to the posterior gangliomere of each ganglion.

By 88 h (see Fig. 1 in Rempel and Church 1971), neuroblasts are prominent in the proto-

cerebrum which, by active division, produce columns of pre-ganglion and ganglion cells. The

innermost of these, beginning at 72 h, have begun to grow out as axons (Fig. 12). Thus, by

this time, a neuropile (npl; often called neuropil) is evident, both here and in all central nerve

cord ganglia and connectives (Fig. 40). In each antenna, an ectodermal invagination has arisen

which will eventually differentiate into an antennal sense organ (Fig. 12, ant. s.o). Its ontogeny

will be the subject of a future contribution.

At 88 h, the tritocerebrum (tr. com) is still post oral and is still attached to the intercalary

ectoderm (Fig. 11, 13, intc. seg). The mandibular (md. ggl), maxillary (mx. ggl) and labial

ganglia (lb. ggl) have moved closer to each other, foreshadowing formation of the subesoph-

ageal ganglion (Fig. 1 1) but the 10 abdominal ganglia are still separate at this time. Cells of

the median nerve strand clump (m.c.c) are still larger than their neighbours and their cyto-

plasm still stains more darkly (Fig. 1 1). They have become oval and have developed clearly

discernible axons, which join in a bundle and extend forward and upward, reaching the dor-

sal region of the ganglion midway between the anterior and posterior cross commissures (Fig.

1 1, 47, com). These cells maintain this appearance until the end of embryogenesis, although

not as obviously.

In cross sections through the tritocerebrum (Fig. 13), cells of the median strand differ

little from those of ordinary, body wall ectoderm. In the mandibular region (Fig. 14), the

strand appears as a cluster of cells having faintly stained nuclei. Sections through the poster-

ior cross commissure (com), of the maxillary segment (Fig. 1 5) show numerous axonal ex-

tensions into the neuropile. Here, the median strand cells are small, rectangular and more

lightly staining than neighbouring ganglion cells. Sections through the median nerve strand

clump (m.c.c) of the labial ganglion (Fig. 16, 41) show its cells to be distinctly separate from

those of the lateral cord ganglia. Finally, in the prothoracic-mesothoracic interganglionic re-

gion (Fig. 17), median strand cells are indistinguishable from those of ordinary, body wall

ectoderm, suggesting that, at this time, the median strand has become discontinuous in inter-

segmental regions.

By 120 h(see Fig. 18 and 19 in Rempel and Church 1971), cephalization of the embryo

has become more pronounced, and the head has become clearly set apart from the rest of the

body. The gnathal ganglia have fused to form the subesophageal ganglion (Fig. 19, sb. ggl) and

abdominal ganglia 9 and 10 have amalgamated (see Fig. 6 in Rempel and Church 1972). The

lateral nerve cord ganglia have moved to the midline and have fused to form a single ganglion-

ic mass in each segment. Ganglion cells have encroached ventrally upon the median strand,

restricting it to the dorsal region of each ganglion (Fig. 20, 43, m.n.c). The strand seemingly

has disappeared from intersegmental regions (Fig. 21) of the ventral nerve cord but it and its

clumps (m.c.c) are retained in intrasegmental regions. For example, three clumps are clearly

visible in the subesophageal ganglion. (Fig. 19).

A characteristic feature of this stage in development, is the presence of long, cytoplasmic

strands (cyt. std) in the intersegmental regions, extending from the body wall (bd. w) to the

interganglionic connectives (int. cn) and from there to the developing midgut (Fig. 21, mdgt).

We do not know what significance they have and in embryos older than 132 h, they are no

longer present.

By 132 h, a perilemma has appeared. It is best developed around the interganglionic con-

nectives (Fig. 22) and consists of a layer of cells, the perineurium (prn), and its secretion pro-

duct, the neural lamella (nr. 1ml; often termed neurilemma or neural lemma). Here and there,

the interface between ganglion cells and neuropile (npl) is occupied by cells (gll. c) that have

Embryology of Lytta viridana IX

9

small, light-colored nuclei (Fig. 20, 43). These cells first appear at about 104 h, and apparently

originate from the median strand. We believe them to be glial. Springer (1967) and Springer

and Rutschky (1969) referred to them as the inner sheath.

By 180 h, the entire central nervous system is enclosed by a well-developed perilemma. It

is especially prominent about the interganglionic connectives (Fig. 25, 28, 56) and dorsally

in each ganglion (Fig. 44). By this time, most neuroblasts have disappeared, although a few

dividing in the brain remain active until hatching. The sheath of glial cells (gll.c) separating

neuropile from ganglion cells within each ganglion is much more pronounced (Fig. 44) and a

few small, dark-staining glial cells are scattered among ganglion cells throughout the central

nervous system. Although inadequate staining makes their details impossible to sort out, fibre

tracts and glomeruli are now clearly visible within the brain and ventral nerve cord ganglia.

These first begin to take shape at 120 h (Fig. 43) and become steadily more complex until

hatching at 250-264 h. (Fig. 44, 48).

Between 1 80 and 200 h, most changes affect the distribution of glial elements enclosing

the neuropile (Fig. 23). The cells of the median strand clump (m.c.c) are still recognizable

but can be confused easily with other ganglion cells. In parasagittal sections of the nerve cord

(Fig. 24), the glial cells (gll. c) appear to be stacked up vertically at each end of each intergang-

lionic connective, but transverse sections through these regions (Fig. 45) show they are not.

In cross sections (Fig. 25-28) made at the points indicated by lines in Fig. 23 and 24, differ-

ences in distribution of glial cells can also be seen. In interganglionic connectives, the perineu-

rium (prn) is thick dorsally and very thin ventrally, whereas the opposite is true of the neural

lamella (nr. 1ml) (Fig. 24, 25, 28, 56). Except for continued differentiation of neuropile cen-

tres, no other significant changes were observed in nervous system development up to the time

of hatching (250 to 264 h).

Stomatogastric Nervous System (stmg. n.s.)

In Lytta viridana embryos, when the stomodaeum begins to invaginate at 56 h, the three

evaginations in its roof — the anlagen of the stomatogastric nervous system (stmg. n.s) - are

already evident (see Fig. 50-53 in Rempel and Church 1969b). At 64 h (Fig. 10), nuclei of

cells surrounding the evaginations become enlarged and, from 72 to 80 h (Fig. 1 1, 46), strands

of cells arise behind each evagination and begin to stream forward over the roof of the stomo-

daeum. This movement is accompanied by considerable mitotic activity. By 96 h, the frontal

ganglion (frt. ggl) has begun to enlarge and neuropile to differentiate within it (Fig. 47) (the

latter actually commences at about 88 h). Simultaneously, fibres of the recurrent nerve (ret.

nv) become apparent just behind the frontal ganglion. At 104 h (Fig. 29), forward streaming

of nerve cells from the three stomodaeal evaginations is still evident but, by 1 20 h (Fig. 19),

evaginations 1 and 2 have begun to disappear, and the three cell streams are no longer separ-

able.

By 120 h, the frontal ganglion (frt. ggl) is well-developed (Fig. 19) and by 132 h, the hypo-

cerebral ganglion (hyp. ggl) has appeared as a slight swelling in the recurrent nerve (ret. nv)

below the pars intercerebralis (prs. intr) of the brain (Fig. 33). Evaginations 1 and 2 have dis-

appeared and all cell proliferation now seems to arise from evagination 3 — an indication that

this is a source of most cells comprising the recurrent nerve (ret. nv). The tritocerebral (= fron-

tal) connectives of the frontal ganglion form at 1 20 h, (see Fig. 31, C, D in Rempel and Church

1971). By 156 h, the system is essentially complete, and from here on most changes involve

ganglionic enlargement and an increase in length of the recurrent nerve to match continued

growth of the stomodaeum. Fig. 38 shows the system as it appears at 216 h and Fig. 48 at

264 h.

Quaest. Ent., 1977 13 (1)

10

Rempel, Heming & Church

Endocrine System

The endocrine system of L. viridana consists of neurosecretory cells, which we will not

consider, and paired corpora cardiaca, corpora allata, and prothoracic glands.

The corpora cardiaca (crp. crd) first become recognizable at 96 h. They originate from cells

emanating from evagination 2 in the roof of the stomodaeum, also the source of the hypocere- 1

bral ganglion (Fig. 29, 47). These cells move laterally around the stomodaeum (Fig. 31 c, 49)

and, by 1 1 2 h, have been carried forward by the cell streams to the region of the pars inter-

cerebralis (prs. intr). As this movement occurs, the cells comprising the cardiaca, become more

and more loosely arranged (Fig. 50). Later, each gland rudiment shifts laterally and attaches

to the posterior surface of the brain. By 132 h, the glands have established contact posterior-

ly with the corpora allata (crp. all) (Fig. 37). Differentiation of the glands’ cells first becomes

evident at about 1 20 h, both secretory(s) and glial elements being present (Fig. 50). The former j

resemble ganglion cells in their dark-staining, homogeneous, cytoplasm; the latter form a loose, j

parenchymatous network. By 250 h (Fig. 37, 51, 54), this network has contracted, so that

the fully-formed glands are not much larger than the allata.

The corpora allata (crp. all) first become apparent at 56 h, as small, ectodermal invagina-

tions of the anterior base of each maxilla (Fig. 34, 52). Each invagination grows inward and

dorsad until, at 96 h (Fig. 53), it reaches a position below the caudal extension of the anter-

ior tentorial arm (a. tent). It now shifts slightly laterally and then dorsally over the tentorial

arm, maintaining, throughout this movement its connection with the body wall (Fig. 35, 36).

At 1 1 2 h, this connection breaks.

In the meantime, the ventral tracheal trunk of each side (2 in Fig. 6 in Rempel and Church

1972) has extended forward from the mesothoracic spiracle over the posterior tentorial arm

(p. tent). At 120 h, this trunk sends one branch (2b) to the gnathal appendages and another

(2a) to the brain. The second branch becomes closely applied to the corpus cardiacum-corpus

allatum complex (Fig. 13 and 17 in Rempel and Church 1972). At 120 h, each corpus allatum

reaches its final position and, at 132 h, re-establishes its connection with the body wall via a

clear, tendon-like strand (Fig. 37, 54). The connective tissue sheath of the corpus allatum,

described by Weismann ( 1 926) in C. morosus and by Roonwal (1937) in L. migratoria , was

not evident in our preparations, even under oil immersion (Fig. 54).

At 96 h, two, small protuberances proliferate inward from the ventral, labial-prothoracic

intersegmental ectoderm (Fig. 30, 55, pthr. gl). They originate close together and grow dor-

sad between the developing interganglionic connectives, (int. cn). At the same time, tracheae

2 (Fig. 6, 9 and 17 in Rempel and Church 1972) each give off a medially-directed branch

(tracheae 9) to form a commissure above the ventral nerve cord between the subesophageal

and prothoracic ganglia (Fig. 55, 56, trch). The proliferated cells become associated with the

tips of these branches as they advance, but maintain their connections with the body wall

until 156 h (Fig. 32, 33). Gradually, the cells from the proliferations spread over the surface

of the tracheal commissure (Fig. 25, 38, 56). At later stages, they are very difficult to see be-

cause they so closely resemble cells of the tracheal epithelium (Fig. 25, 56).

We believe these cellular proliferations to constitute the prothoracic glands. We have no

idea how far they spread over tracheae 1, 2 and 9 (Rempel and Church 1972, Fig. 6, 9, 17),

since, even in mature larvae of other beetles, such glands are difficult to trace (Svrivastava

1959). Additionally, the area between the subesophageal-prothoracic interganglionic connec-

tives is eventually occupied by a complex array of tracheal branches (Fig. 56). This also causes

difficulties in distinguishing between glandular and tracheal tissues.

Embryology of Lytta viridana IX

11

DISCUSSION

Central and Stomatogastric Nervous Systems

Embryogenesis of the brain, lateral nerve cords and stomatogastric nervous system in

Lytta viridana has been described very briefly because the sequence of events is so similar

to that occurring in Tenebrio molitor L. (Ullmann, 1967) and Sitophilus [= Calendra) oryzae

(L.) (Coleoptera, Curculionidae) (Tiegs and Murray 1938).

The median strand - Authors agree generally about the origin of this system, and the des-

cription we have presented outlines the usual pattern. The median strand arises from ectoder-

mal cells along the mid-ventral line of an embryo in a region extending from behind the in-

tercalary segment to the end of the tenth abdominal segment. Early in its development, two

types of cells are recognized. In intrasegmental regions, cells of the median strand resemble

those of the body wall ectoderm; in intersegmental regions, they assume the appearance of

neuroblasts. These latter cells divide teloblastically, like typical neuroblasts, and in fact, seem

to be neuroblasts. Nonetheless, we prefer to use the word “clump” for groups of these cells,

as did Springer (1967) and Springer and Rutschky ( 1 969). The forward shift of each clump

of the median strand from an intersegmental to an intrasegmental position is in agreement

with descriptions of this process for other insects (Springer 1 967; Springer and Rutschky

1969).

General agreement about development of the median strand gives way to disagreement

and controversy about fate and role of the components of this system (Edwards 1969; An-

derson 1972a and b and 1973). Details are provided below.

Fate and role of the clumps of the median strand — Ullmann ( 1 967) and Springer and

Rutschky ( 1 969) summarized information presented by earlier workers about this topic.

It was claimed by Ullmann and others that clumps in embryos of S. oryzae and T. molitor

contribute to development of the definitive ganglia, whereas in the insects studied by Sprin-

ger and Rutschky (various hemipterans, orthopterans, beetles, moths and dipterans), the

clumps disappear after katatrepsis. In embryos of L. viridana , the clumps are present from

their first appearance (56 h) until after hatching (±250 h). We agree with Ullmann (1967)

that these cells probably act as ganglion cells because they develop axons (Fig. 11, 19, 47).

Although, for most insects studied, the clump cells seem to be involved with development

of the nervous system, Miyakawa ( 1 974) reports that in embryos of S. griseipennist the inter-

ganglionic portions of the median cord of the thoracic segments develop into furcae, and thus

do not have a nervous function.

Clumps of the median strand and segmentation of the insect head.— Because the clumps

are intersegmental in all post-oral segments, we assume that in ancestral hexapods they were

present also in those segments that arise post-orally but which, in more highly evolved stocks,

become pre-oral. If one could recognize the clumps in heads of extant insects, one would have

additional evidence for deducing the number of segments involved therein. (See Malzacher

1968; Scholl 1969; Rempel and Church 1971 ; and Rempel 1975 for comprehensive discus-

sions of head segmentation). Although it is tempting to suggest that the stomatogastric ner-

vous system with its three ganglia (frontal, hypocerebral, and ventricular) is the forward con-

tinuation of the median strand, and that the ganglia represent respectively the clumps of the

preantennal (labral), antennal, and tritocerebral segments, this is probably not so, for the fol-

lowing reasons: As cephalization occurred, the floors of these segments supposedly contribu-

ted to the floor of the stomodaeum. However, each post-oral clump is situated behind the pos-

terior cross commissure of its ganglion. How then could the clump of, for instance, the tri-

tocerebral segment, get onto the roof of the stomodaeum to form the ventricular ganglion

while its commissure remained post-oral? It seems developmental^ impossible. Thus, it also

Quaest. Ent., 1977 13 (1)

12

Rempel, Heming & Church

seems impossible that these ganglia are the homologues of clumps of the median strand. Pro-

bably in extant insects, the segments in question are without a median strand and without

clumps, and this is probably the result of atrophy occurring in the extinct ancestral stock.

Role of the intraganglionic portions of the median strand. - Springer (1967) and Springer

and Rutschky (1969) showed that cells in this region do not become functional ganglion cells,

and this seems to be generally accepted. However, authors disagree about what these cells do.

Ullmann (1967) claimed that those near the periphery contribute to formation of the perilem-

ma in dorsal portions of the ganglia. On the other hand, Miyakawa ( 1 974) reported that, in

S. griseipennis embryos, the perilemma over most of each ganglion seemed to arise from mod-

ified outer ganglion cells. Our observations for L. viridana embryos support the conclusion of

Miyakawa.

Cells of the intraganglionic portions of the median strand are involved in production of

glial cells. We believe that glial elements associated with axons of the interganglionic connec-

tives also originate from the median strand. We conclude, therefore, that the principal role

of the median cord is to form glial cells associated with nerve fibres of the lateral nerve cords.

More specifically, in embryos of L. viridana at 104 h, some median strand cells move lat-

erally between neuropile and innermost ganglion cells to form an inner sheath of glial cells

(Fig. 20, 43, gll. c), a process very similar to that occurring in embryos of S. griseipennis (Miy-

akawa 1974). We disagree with Springer and Rutschky (1969) who claimed that the inner

sheath develops from the innermost ganglion cells.

Glial cells. - Three of the four types of glial cells described by Wigglesworth (1972) from

specimens of Rhodnius prolixus Stal (Heteroptera) are evident in larvae of L. viridana that

are ready to hatch (prolarvae). These cells are illustrated in Fig. 44: type i (perineurium); type

ii (cells scattered among ganglion cells); and type iv (neuropile sheath). Type iii cells having

giant nuclei are not present, although some type iv cells have quite large nuclei. All three types

of glial cells are also evident in ganglia of the stomatogastric nervous system.

Endocrine System

This includes the corpora cardiaca, corpora allata, and prothoracic glands. Each of these

paired glands is discussed below.

Corpora cardiaca. — Embryogenesis of these glands has been studied by a number of work-

ers, notably Weismann ( 1 926, in C. morosus), Roonwal (1937, in Locusta migratoria (L.)

(Orthoptera)), Pflugfelder ( 1 937, in C. morosus ), and Dorn (1972, and 1975a, in O. fasciatus).

All agree that ontogenetically, these glands originate with the hypocerebral ganglion, by cell

migration from the roof of the stomodaeum.

Dorn (1975a) followed embryogenesis of the corpora cardiaca in eggs of O. fasciatus, using

the transmission electron microscope. In these insects, the glands appear at 56 h, and begin

to differentiate into glandular and glial components at 96 h. The glandular cells assume a

spherical distribution about a lumen (his Fig. 4 and 8) into which grow cell projections, pro-

bably axons of the nervi corporis cardiaci. This occurs when the glands attach to the aorta.

Evidence of protein synthesis appears at 96 h, and by 1 1 1 h, neurosecretory granules are

evident. During hatching (at 124 h) the cells of the glands appear to be secretory.

Secretory cells in the cardiaca of embryos of L. viridana are probably homologous with

the “instrinsic secretory cells” described by Schoonveld (1970) in the cardiaca of adult speci-

mens of Leptinotarsa decemlineata Say (Coleoptera, Chrysomelidae).

Phylogenetically, Hanstrom ( 1 942) assumed the corpora cardiaca to have evolved from a

stomodaeal ganglion. Because these glands occur in all apterygote insects which have been

examined for them, and because the corpora allata do not, Novak (1975) assumed that the

former glands are evolutionarily older than the latter glands.

Embryology of Lytta viridana IX

13

Corpora allata. — In embryos of different taxa of insects, these glands seem to originate

from different germ layers, and in different positions. For example, in embryos of L. virid-

ana, O. fasciatus (Dorn 1972), and S. griseipennis (Miyakawa 1974), they arise as ectodermal

invaginations at the anterior base of each maxilla. On the other hand, in L. migratoria (Roon-

wal 1937) and C. morosus embryos (Pflugfelder 1937), the glands arise as paired, ectodermal

invaginations between the mandibular and maxillary segments. And, in embryos of S. oryzae,

they originate from mesoderm of the antennal coelomic sacs (Tiegs and Murray 1938). Accor-

ding to Pflugfelder (1937), some authors have even reported corpora allata as being of endo-

dermal origin. Probably some of these observations are incorrect. Certainly the question of

ontogenetic origin of these glands should be investigated using a wide taxonomic spectrum

of pterygote insects and a diversity of approaches.

The ultrastructure and function of the developing corpora allata of O. fasciatus embryos

have been studied by Dorn (1975 b) who showed that high titers of juvenile hormone present

just before hatching are probably the result of activity of these glands.

Pro thoracic glands. - For Coleoptera, Srivastava (1959) described the prothoracic glands

of larvae of 1 5 species (none were meloids). These glands occur in the head, cervical region

and prothorax as thin cords or sheets of cells closely associated with one or both of two large

tracheae extending from the prothoracic spiracles into the head. Embryogenesis of the pro-

thoracic glands was previously unknown for Coleoptera, and has been little studied in any

insect species.

Differences of opinion among authors suggest i) that the prothoracic glands of different

insects are not homologous, or ii) that some accounts of their origin are in error, or iii) that

some tissues referred to as prothoracic glands are some other structure. We cannot resolve the

problem, but we review the different viewpoints, below.

According to Gilbert and King ( 1 973), Toyama ( 1 902) identified the prothoracic glands

in embryos of Bombyx mori F. (Fepidoptera) as epithelial invaginations of the labial segment

of the head. A similar origin was postulated for the glands of embryos of Dysdercus cingula-

tus (Fab.) (Heteroptera) by Wells ( 1 954), and for those of O. fasciatus embryos by Dorn ( 1 972).

In embryos of Schistocerca gregaria (Forskal) (Orthoptera), the prothoracic glands invaginate

before katatrepsis from lateral ectodermal regions between the maxillary and labial segments

(Micciarelli and Sbrenna 1972). Novdk(1975) suggested that the prothoracic glands of most

pterygote insects originate from the ventral margin of the prothoracic segment, basing his con-

clusion on their innervation from the prothorax, and on their supposed homology with the

cephalic nephridia of apterygote insects. The proposed labial-prothoracic origin for these glands

in L. viridana embryos, if proved, would support Novak’s conclusion.

We at first thought that the labial diverticula (Fig. 19, lb. div), not the intersegmental pro-

liferations suggested here, might be the progenitors of the prothoracic glands, because these

diverticula originate from the labial segment, as do the glands of some of the insects named

above. The diverticula grow caudad under the suboesophageal ganglion where they bend ab-

ruptly upward to end in the labial-prothoracic region. Ullmann (1967) detailed evidence to

suggest that these diverticula develop into “maxillary glands”. We have followed developmen-

tally the “glands” of L. viridana embryos to similar but very indistinctly developed structures

in prolarvae of this species.

Publications also present conflicting evidence concerning the function of embryonic pro-

thoracic glands (see discussions and refs, in Dorn 1972; and Micciarelli and Sbrenna (1972)).

Dorn (1 972) noted that the glands of O. fasciatus showed three cycles of activity (as did the

neurosecretory cells and corpora cardiaca) that correlated well with secretion of three pre-

hatch cuticles. A similar correlation was noted by Micciarelli and Sbrenna (1972) in the two

moults of S. gregaria embryos. However, these authors showed that isolated embryonic abdo-

mens, lacking these glands, were also capable of secreting two cuticles and concluded that

Quaest. Ent., 1977 13 (1)

14

Rempel, Heming & Church

!

embryonic apolyses are not under control of the prothoracic glands. Since much recent evi-

dence suggests that ecdysone can be synthesized in organs other than the prothoracic glands

(Nakanishi, et al. 1 972; Romer, et al. 1 974; Hsiao, et al. 1 975), action of this hormone in em-

bryonic moults has still not been ruled out.

Embryos of L. viridana produce a single, delicate embryonic cuticle between 1 20 and 132

h, shortly after katatrepsis, about the time that secretory cells become evident in the corpora

cardiaca. However, there is no obvious change at this time in cells of the corpora allata or pro- '

thoracic glands. Neither is any change noted in these cells at 168 to 1 80 h, when deposition

of larvae cuticle begins. Thus, we have no positive evidence of endocrine function in embryos

of L. viridana.

Writing the above discussion on the origin and function of embryonic insect endocrines was

frustrating because of the conflicting results presented in the literature. We do not believe that

structures as fundamental to insect development as the corpora allata and prothoracic glands

can have so many different ontogenetic origins. Diversity in site of origin implies multiple in-

dependent origin during insect evolution. This conflicts with the proven similarity in biochem-

ical function of these glands in representative insects of most orders (Gilbert and King 1 973;

Slama, et al. 1 974; Novak 1975). What is required to resolve the conflict are detailed compara-

tive embryological studies of the most critical kind using all methods available. Dorn’s (1972,

1975 a, b) studies in which conventional, ultrastructural and experimental methods are com-

bined, provide a start in the right direction. It would also help if embryological studies the

quality of Dorn’s were carried out on individuals of species for which experimental proof of

gland function exists.

ACKNOWLEDGEMENTS

An early draft of this manuscript was read and constructively criticized by G.E. Ball, D.A.

Craig and B.K. Mitchell. Dr. Craig also offered suggestions to B.S. Heming on photomicrogra-

phy. J. Scott prepared the photographic plates, Mrs. Patricia Cookson typed the manuscript

and H. Goulet translated the abstract into French. We thank all these individuals for their

generous assistance. Publication costs were met by grants from the Strickland Memorial Trust

Fund and the National Research Council of Canada (A5756 to B.S. Heming).

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Waddington (eds). Developmental Systems: Insects. Vol. I. Academic Press, London and N.Y.

pp. 95-163.

Anderson, D.T. 1972b. The development of holometabolous insects. Ibid. pp. 165-242.

Anderson, D.T. 1973. Embryology and Phylogeny in Annelids and Arthropods. Pergamon

Press, N.Y. and Oxford 495 p.

Bate, C.M. 1976. Embryogenesis of an insect nervous system. I. A map of the thoracic and

abdominal neuroblasts in Locusta migratoria. Journal of Embryology and Experimental

Morphology 35; 107-123.

Cazal, P. 1 948. Les glandes endocrines retrocerebrales des insectes. Etude morphologique.

Bulletin biologique de la France et de la Belgique. Supplement 32: 1-228.

Church, N.S. & J.G. Rempel 1971. The embryology of Lytta viridana LeConte (Coleoptera:

Meloidae). VI. The appendiculate 72-h embryo. Canadian Journal of Zoology 49: 1 563-1570.

Dorn, A. 1972. Die endokrinen Drusen im Embryo von Oncopeltus fasciatus Dallas (Insecta,

Embryology of Lytta viridana IX

15

Heteroptera). Morphogenese, Funktionsaufnahme, Beeinflussung des Gewebewachstums

und Beziehungen zu den embryonalen H&utungen. Zeitschrift fur Morphologie und 6kologie

der Tiere 71 : 52-104.

Dorn, A. 1 975a. Electronenmikroskopische Studien iaber Differenzierung und Funktionsauf-

nahme der Corpora cardiaca im Embryo von Oncopeltus fasciatus Dallas (Insecta, Heteroptera).

Cytobiologie 10: 235-248.

Dorn, A. 1975b. Struktur und Funktion des embryonalen Corpus allatum von Oncopeltus fas-

ciatus Dallas (Insecta, Heteroptera). Verhandlungen der Deutschen zoologischen Gesellsch-

aft 67: 85-89.

Edwards, J.S. 1969. Postembryonic development and regeneration of the insect nervous system.

Advances in Insect Physiology 6: 97-137.

Gabe, M. 1966. Neurosecretion. Pergamon Press, N.Y. 872 p.

Gilbert, L.I. & D.S. King 1973. Physiology of growth and development: endocrine aspects.

In Rockstein, M. (ed.) The Physiology of Insecta. 2nd Ed. Vol. 2. Academic Press. N.Y.

pp. 249-370.

Hanstrom, B. 1942. Die Corpora cardiaca und Corpora allata der Insekten. Biologia generalis

15: 485-531.

Herman, W.S. 1967. The ecdysial glands of arthropods. International Review of Cytology 22:

269-347.

Hsiao, T.H., C. Hsiao & J. de Wilde 1 975. Moulting hormone production in the isolated abdo-

men of the Colorado beetle. Nature 255: 727-728.

Kankel, D.R. & J.C. Hall 1 976. Fate mapping of nervous system and other internal tissues in

genetic mosaics of Drosophila melanogaster. Developmental Biology 48: 1-24.

Malzacher, P. 1968. Die Embryogenese des Gehirns paurometaboler Insekten. Untersuchungen

an Carausius morosus und Periplaneta americana. Zeitschrift fhr Morphologie und Okologie

der Tiere 62: 103-161.

Micciarelli, A.B. & G. Sbrenna 1972. The embryonic apolyses of Schistocerca gregaria (Orth-

optera). Journal of Insect Physiology 18: 1027-1037.

Miyakawa, K. 1 974. The embryology of the caddisfly Stenopsyche griseipennis MacLachlan

(Trichoptera, Stenopsychidae). III. Organogenesis: Ectodermal derivatives. Kontyu 42:

305-324.

Nakanishi, K., H. Moriyama, T. Okauchi, S. Fujioka & M. Koreeda 1972. Biosynthesis of

a and (5 ecdysones from cholesterol outside the prothoracic gland in Bombyx mori. Science

176: 51-52.

Novak, V.J.A. 1975. Insect Hormones. 2nd English Ed. Chapman & Hall, London. 600 p.

Pflugfelder, O. 1937. Bau, Entwicklung, und Funktion der Corpora allata und cardiaca von

Dixippus morosus Br. Zeitschrift fur wissenschaftliche Zoologie 149: 477-512.

Pflugfelder, O. 1958. Entwicklungsphysiologie der Insekten. 2 Auflage. Akad. Verlag. Geest

& Portig. K - G., Leipzig. 490 p.

Rempel, J.G. 1975. The evolution of the insect head: the endless dispute. Quaestiones Ento-

mologicae 11: 7-25.

Rempel, J.G. & N.S. Church 1965. The embryology of Lytta viridana Le Conte (Coleoptera:

Meloidae). I. Maturation, fertilization and cleavage. Canadian Journal of Zoology 43: 915-

925.

Rempel, J.G. & N.S. Church 1969a. The embryology of Lytta viridana LeConte (Coleoptera:

Meloidae). IV. Chromatin elimination. Ibid. 47: 351-353.

Rempel, J.G. & N.S. Church 1969b. The embryology of Lytta viridana LeConte (Coleoptera:

Meloidae). V. The blastoderm, germ layers, and body segments. Ibid. 47: 1 1 57-1 1 71 .

Rempel, J.G. & N.S. Church 1971. The embryology of Lytta viridana LeConte (Coleoptera:

Quaest. Ent., 1977 13 (1)

16

Rempel, Iieming & Church

Meloidae). VII. Eighty-eight to 132 h: the appendages, the cephalic apodemes, and head

segmentation. Ibid. 49: 1571-1581.

Rempel, J.G. & N.S. Church 1972. The embryology of Lytta viridana LeConte (Coleoptera:

Meloidae). VIII. The respiratory system. Ibid. 50: 1547-1554.

Romer, F., H. Emmerich & J. Nowock. 1974. Biosynthesis of ecdysones in isolated prothora-

cic glands and oenocytes of Tenebrio molitor in vitro. Journal of Insect Physiology 20:

1975-1987. I

Roonwal, M. L. 1 937. Studies on the embryology of the African migratory locust Locusta

migratoria migratorioides, R & F. II. Organogeny. Philosophical Transaction of the Royal

Society. Series B 227: 175-244. j

Scholl, G. 1 969. Die Embryonalentwicklung des Kopfes und Prothorax von Carausius morosus \

Br. (Insecta, Phasmida). Zeitschrift fur Morphologie und Okologie der Tiere 65: 1-142.

Schoonveld, H. 1970. Structural aspects of neurosecretory and corpus allatum activity in the

adult Colorado beetle, Leptinotarsa decemlineata Say, as a function of day length. Nether-

lands Journal of Zoology 20: 1 51-237.

Slama, K., M. Romanuk & F. Sorm. 1974. Insect Hormones and Bioanalogues. Springer Verlag

N.Y. and Wien. 477 p. j

Springer, C.A. 1 967. Embryology of the thoracic and abdominal ganglia of the large milkweed

bug, Oncopeltus fasciatus (Dallas) (Hemiptera, Lygaeidae). Journal of Morphology 122: 1-1 8. j

Springer, C.A. & C.W. Rutschky, III. 1969. A comparative study of the embryological develop-

ment of the median cord in Hemiptera. Ibid. 1 29: 375-400.

Svrivastava, U.S. 1959. The prothoracic glands of some coleopteran larvae. Quarterly Journal

of Microscopical Science 100: 51-64.

Tiegs, O.W., & F.V. Murray 1938. The embryonic development of Calandra oryzae. Ibid. 80: j

159-271.

Toyama, K. 1902. Contributions to the study of silkworms. I. On the embryology of the silk-

worm. Bulletin of the College of Agriculture. Tokyo Imperial University 5: 73-1 18.

Ullmann, S.L. 1964. The origin and structure of the mesoderm and the formation of the

coelomic sacs in Tenebrio molitor L. (Insecta, Coleoptera). Philosophical Transactions of

the Royal Society. Series B 248: 245-277.

Ullmann, S.L. 1967. The development of the nervous system and other ectodermal derivatives j

in Tenebrio molitor L. (Insecta Coleoptera). Ibid. 252: 1-25.

Wells, M. J. 1 954. The thoracic glands of Hemiptera-Heteroptera. Quarterly Journal of Micro-

scopical Science 95: 231-244.

Weismann, R. 1926. Entwicklung und Organogenese der Colomblasen. pp. In Leuzinger, H.,

R. Weismann und F.E. Lehmann (eds.). Zur Kenntnis der Anatomie und Entwicklungeschich-

te der Stabheuschrecke Carausius morosus Br. G. Fisher Verlag, Jena.

Wigglesworth, V.B. 1972. The Principles of Insect Physiology. 7th Ed. Chapman and Hall.

London. 827 p.

Embryology of Lytta viridana IX

17

ABBREVIATIONS

Quaest. Ent., 1977 13 (1)

18

Rempel, Heming & Church

Fig. 1. Body wall and coelomic sacs at 50 h, parasagittal section. Fig. 2. Body wall at 50 h, median sagittal section. Fig. 3.

Same, at 56 h, parasagittal section, showing dermatogene (dg. 1) and neurogene (ng. 1) layers, the latter with neuroblasts (nbl).

Fig. 4. Same, at 60 h, showing formation of pre ganglion and ganglion cells (ggl. c). Fig. 5. Same, median sagittal section, show-

ing “clumps” of median strand cells (m.c.c). Fig. 6. Same, at 64 h, showing forward shift of median strand clump. Fig. 7. Diagram

of central nervous system, showing cephalic ganglia, lateral nerve cords (l.n.c) and median nerve strand (m.n.c). Fig. 8. Parasagit-

tal section through protocephalic lobe and antenna (ant) at 56 h, showing formation of protocerebral (protc) and deutocerebral

(deutc) neuroblasts. Fig. 9. Same, through stomodaeum, showing formation of tritocerebral (trite) neuroblasts from intercalary

ectoderm (intc. seg). Fig. 10. Median sagittal section through stomodaeum at 64 h, showing formation of stomatogastric nervous

system (stmg. n.s). Fig. 11. Same, at 88 h, showing median strand clumps (m.c.c). Fig. 12-17. Transverse sections through dev-

eloping nerve cord at 88 h, taken at points indicated by lines in Fig. 11: 12, Through antenna and stomodaeum, showing devel-

oping ganglion cells, stomatogastric nervous system, (stmg. n.s) and antennal sense organ (ant. s.o); 13, Through tritocerebral

ganglion; 14, Through mandibular ganglion; 15, Through posterior commissure of maxillary ganglion; 16, Through median strand

“clump” (m.c.c) of labial ganglion; 17, Through prothoracic-mesothoracic intersegmental region and inter ganglionic connectives

(int. cn). Fig. 18. Parasagittal section through tritocerebral ganglion (trite) and anterior tentorial arm (a. tent).

Quaest. Ent., 1977 13 (1)

20

Rempel, Heming & Church

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Embryology of Lytta viridana IX

21

Fig. 39-56. Photomicrographs, scale 10 /Ltm except where indicated. Fig. 39. Median sagittal section of 72 h embryo, showing

forward shift of maxillary median strand clump (m. c. c). Fig. 40. Transverse section through maxillary median strand clump

(m. c. c) and interganglionic connectives (int. cn) of 72 h embryo, showing neuroblast (nbl) and developing neuropile (npl).

Fig. 41. Same, of 96 h embryo through median strand clump (m. c. c) of labial ganglion. Fig. 42. Frontal section through head

of 96 h embryo, showing three lobes of protocerebrum (protc), lobe 1 (op. 1) developing as an invagination. Note optic cup

(op. c). Fig. 43. Transverse section through labial commissure (com) of subesophageal ganglion of 120 h embryo. Note dev-

eloping glial cells (gll. c) and perineurium (prn). Fig. 44. Same, through subesophageal ganglion of 264 h embryo, showing

glial cells (gll. c), perineurium (prn) and neural lamella (nr. 1ml).

Quaest. Ent., 1977 13 (1)

22

Rempel, Heming & Church

Fig. 45. Transverse section through anterior ends of subesophageal-prothoracic interganglionic connectives of 264 h embryo,

showing glial cells in neuropile. Fig. 46. Median sagittal section through stomodaeum (stom) of 72 h embryo, showing evag-

inations of stomatogastric nervous system (stmg. n. s). Fig. 47. Same, of 96 h embryo, showing tritocerebral commissure (tr.

com) and those of subesophageal ganglion (com). Note also the mandibular median strand clump (m. c. c). Fig. 48. Same, of

264 h embryo, showing frontal (frt. ggl), hypocerebral (hyp. ggl), and ventricular ganglia (vnt. ggl). Fig. 49. Transverse section

through 96 h embryo, showing corpora cardiaca (crp. crd) and hypocerebral ganglion (hyp. ggl). Fig. 50. Same, through 120

h embryo. Note secretory cells (s) in corpus cardiacum (crp. crd).

Embryology of Lytta viridana IX

23

Fig. 51. Transverse section through corpora cardiaca (crp. crd) of 264 h embryo. Fig. 52. Parasagittal section through base of

maxilla (mx) of 88 h embryo, showing invaginating corpus allatum (crp. all). Fig. 53. Same, 96 h. Fig. 54. Same, through corpus

cardiacum - corpus allatum complex of 252 h embryo. Note delicate strand extending from apex of corpus allatum (crp. all).

Fig. 55. Transverse section through labial-prothoracic intersegmental region at 96 h, showing origin of prothoracic glands (pthr.

gl). Fig. 56. Same, 264 h.

Quaest. Ent., 1977 13 (1)

THE ADULT TRICHOPTERA (INSECTA) OF ALBERTA

AND EASTERN BRITISH COLUMBIA, AND THEIR POST-GLACIAL

ORIGINS. I. THE FAMILIES RHYACOPHILIDAE AND LIMNEPHILIDAE.

SUPPLEMENT 1.

ANDREW P. NIMMO

Department of Entomology

The University of Alberta Quaes tiones Entomologicae

Edmonton, Alberta T6G 2E3 13: 25-67 1977

Six species are described as new: Rhyacophila simplex, R. donaldi, and R. autumnaliso/

the Rhyacophilidae; Imania thomasi, Limnephilus vernalis, and Philocasca alba of the Limne-

philidae. Previously known species here recognised as members of the study area fauna are:

Rhyacophila vao Milne, R. unimaculata Denning of the Rhyacophilidae; and Dicosmoecus

gilvipes (Hagen), Neophylax rickeri Milne, Limnephilus nimmoi Roy and Harper, L. insularis

Schmid, L. alvatus Denning, Lenarchus keratusRoss, and Pla ty centropus plectrus Ross of the

Limnephilidae. Asynarchus lapponicus (Zetterstedt) ( Limnephilidae), is more fully described

and illustrated. The female of Rhyacophila milnei Ross is described and illustrated for the first

time; and Rhyacophila sp.3, and sp.4 and Limnephilus sp.2 are described and illustrated from

single females only. Females o/Psychoglypha prita (Milne) and P. alascensis (Banks) are figur-

ed and described. Rhyacophilidae total 29 species, and Limnephilidae 102 species, in the study

area. The range type of each species is determined, and the probable post-glacial source for

each is considered. The proportions of the fauna of these two families from each source area

are now ( combining data from this paper, and that to which it is supplementary): from Cordil-

lera south of the ice, 58. 7%; from Alaska, 4.5%; from central plains, 6.8%o; from eastern North

America, 8.4%; from North America as a whole, south of the ice, 19.8%; and indeterminate,

1.5%.

Dans ce travail nous ddcrivons six nouvelles especes: Rhyacophila simplex, R. donaldi, et R. autumnalis des Rhyacophil-

idae; Imania thomasi, Limnephilus vernalis, et Philocasca alba des Limnephilidae. Nous y ajoutons des especes dd ja ddcrites

mais nouvelles pour notre faune: Rhyacophila vao Milne, R. unimaculata Denning des Rhyacophilidae; Dicosmoecus gilvipes

(Hagen), Neophylax rickeri Milne, Limnephilus nimmoi Roy et Harper, L. insularis Schmid, L. alvatus Denning, Lenarchus

keratus Ross, et Platycentropus plectrus Ross des Limnephilidae. Nous illustrons et ddcrivons plus completement Asynarchus

lapponicus (Zetterstedt) (Limnephilidae), les femelles de Rhyacophila milnei, Ross, Psychoglypha prita (Milne), et P. alascen-

sis (Banks), et le seules femelles de Rhyacophila sp.3 et sp.4 et Limnephilus sp.2. Nous reconnaissons prdsentement pour not-

re territoire d’dtude 29 Rhyacophilidae et 102 Limnephilidae. Le type de distribution est determine et nous discutons V ori-

gin e probable depuis les temps glaciaires de chaque espe'ce. Nous considdrons que I’origine de la faune des deux families dans

notre region se portage comme suit (nous combinons les r 4 suit at s de cette publication et de celles qui lui sont supplementai-

res): de la Cordilliere au sud des glaciers, 58. 7%; de V Alaska, 4.5%; des plaines centrales, 6.8%; de Vest de VAmdrique du nord,

8.4%; de VAmdrique du nord au sud des glaciers, 19.8%; et demeurant indetermind, 1.5%.

CONTENTS

Introduction 26

The Family Rhyacophilidae 27

Notes on Rhyacophila spp previously recorded 33

Placement of species in keys 34

The Family Limnephilidae 34

Placement of species in keys 47

Post-glacial origins, and affinities 49

Conclusions 52

26

Nimmo

Acknowledgements 52

References 53

Illustrations 57

Corrigenda to Nimmo, 1971a.

These corrigenda are additional to those listed in Nimmo, 1971b, and 1974.

p. 1 8 Couplet 2, choice ‘a’ - Fig. 65 should read Fig. 64.

p. 85 Couplet 26, choice ‘a’ - ‘Cercus Triangular . . .’ should read ‘Clasper triangular . . .’.

p. 1 1 1 Tine 10. \ . . segment X distinguished from segment X . . . ’ should read \ . . segment

IX distinguished from segment X . . .’.

p. 129 Key to Asynarchus females, choice Ta’, \ . . enlarge . . should read \ . . enlarged . . .

p. 162 Fig. 1 59. Apparently I made this drawing from a detached aedeagus. The illustration

given is presented upside-down and backwards. For proper appearance it should be

viewed with the page inverted, with the distal end of aedeagus pointed to the obser-

ver’s left.

INTRODUCTION

Although the thought was unstated in my 1971(a) paper, it is implicit in a faunal survey

that more remains to be discovered. This paper is proof of that. But, apart from finding males

for unassociated females, and females for males for which they are yet unknown, neither can

this supplement be considered the final word. While I hope to keep supplements to a minimum

there will, nevertheless, be further instalments.

The title of this supplement differs from that of my 1971(a) paper, but is equivalent to it,

and conforms to that of my series on the Trichoptera fauna, of the study area.

Figure numbering in papers of the main series will be consecutive throughout, except that

there will be, if required, a ‘Fig. 1 ’ in each paper, followed by a lower case letter. This figure

will present collection localities in the study area which are new to the series as a whole. The

use of a lower case letter after the T ’ follows on in sequence from the suite of locality maps

presented in 1971(a). Figure numbering in the supplements to each main paper, on the other

hand, will be independent of the main series, but will be consecutive within the supplement

series for each main paper. Remarks above on ‘Fig. 1’ apply equally to the supplements. Fig. Id

serves this paper.

Full collection data are available on file, either from myself, or from the Department of

Entomology, University of Alberta, Edmonton. I am adding to these records through contin-

ued collecting.

For those species already recorded from the study area, but for which I here record the fe-

male, the only literature reference in this paper is to my original record (1971a), giving page

and figure numbers.

For species described elsewhere, and recorded here from the study area, I give literature to

date, so far as it is known to me.

The genera and species are set out here in the same order as in my 1 971(a) paper. Those taxa

recorded as new to the area are inserted in the sequence, relative to other taxa, used by Ross

( 1956) for Rhyacophilidae, and by Schmid (1955) for Limnephilidae.

At the ends of the sections on Rhyacophilidae and Limnephilidae respectively are guides

that locate the species, or newly associated sexes in my original (1971a) keys.

In the legend to each plate illustrating genitalia, abbreviations of certain commonly used

Rhyacophilidae and Limnephilidae

27

words are as follows: ‘lateral’ to ‘lat. ‘posterior’ to ‘post.’; ‘ventral’ to ‘ventr.’; ‘dorsal’ to

‘dors.’; ‘partial’ to ‘part.’; and ‘segment’ to ‘Seg. ’.

Previousdntroductory material is in Nimmo (1971a) and Nimmo ( 1 974). Information found

therein applies to this paper also.

One problem in the taxonomy of Trichoptera is that, on occasion, females which cannot be

associated with known species are encountered. Most authors ignore these, and do not illustra-

te them. Also, females may be associated with known species, and mentioned in the literature,

but not illustrated. Neither practice is sound. An example of the usefulness of illustrating un-

associable females is the case of Limnephilus nimmoi Roy and Harper, presented in this paper.

An example of how failure to illustrate, somewhere, the known female of a known species can

lead to confusion is presented by L. alvatus Denning in this paper. I illustrate all unassociable

females known to me from the study area in the hope that somewhere, someone will recogni-

se the species to which it belongs, and may either publish on the association, or inform me

of it.

THE FAMILY RHYACOPHILIDAE STEPHENS

The Genus Rhyacophila Pictet

The vofixa group

Rhyacophila simplex Nimmo new species

(Fig. 2a-b, 10-13, 111)

This species is very close to R. ophrys Ross. Males may be distinguished from those of ophrys

by blunter, more angular distal clasper segment (Fig. 10); by simpler segment X with sloped anal

sclerite; by apparently non-erectile lateral arms of aedeagus (Fig. 13); and by blunt, recurved

horn on postero-dorsal edge of aedeagal sheath.

Description. — Antennae deep red-brown; annular sutures paler. Vertex of head almost black, with lateral sutures

and postero-lateral warts paler. Thorax dark brown dorsally, paler to almost white laterally. Spurs brown; formula 3,4,4.

Fore-wing length of male 7.2 mm; uniform red-brown except hyaline areas as in Fig. 2a; dorsal surface with patterned dis-

tribution of golden hairs, especially on posterior areas. Hind-wing grey-brown, with hyaline areas. Venation as in Fig. 2 a-b.

Male genitalia. (Specimen from Ruby Ck, nr Ruby Lk, at 6,750( Waterton National Park, Alberta). Segment IX basically

rectangular in lateral aspect, with postero-dorsal edge expanded posterad (Fig. 10); anterior rim partly black-edged. Clasper

with parallel sides except antero-ventral portion angled sharply anterad; mesal face with slight meso-dorsal ridge, not quite

extended to base; distal segment four-sided, angles rounded, mesal face setose. Segment X ventrad of postero-dorsal edge of

segment IX (Fig. 10), with ventral extensions laterally, each flanged posterad by weakly coloured screen; in dorsal aspect

segment deeply cleft mesally, bilobed (Fig. 12). Tergal strap inverted-v-shaped anterad of segment X, lightly sclerotised. Anal

sclerite tear-shaped in lateral aspect (Fig. 10), partly enclosed by ventral extension of segment X; bifid, flared laterad distal-

ly (Fig. 11). Aedeagus short, stout, main body as lightly sclerotised tube (Fig. 13); postero-dorsal extremity projected blunt-

ly dorsad; main shaft very small, with expanded, button-like tip; lateral arms enclosed by main body of aedeagus, roughly

triangular in lateral aspect, with thick fringe of setae along distal edge.

Female. Unknown, or unassociated.

Note on habitat. - While I am unfamiliar with the exact location, the single known locality

for this species is probably a small, turbulent, stream with rocky bottom.

Holotype. — Male. Ruby Creek, near exit from Ruby Lake, at 6;750', Waterton National

Park, Alberta (Fig. Ill); June 29, 1 975; D.B. Donald.

The holotype is in the Canadian National Collection, Ottawa, with type number 1 5,160.

This species is named with reference to the simplicity of the genital capsule.

The aero pedes group

Rhyacophila vao Milne, 1936

(Fig. 23-25, 110)

Rhyacophila vao Milne, 1936:93, 102, 111, Fig. (no number). (Type locality: Cultus Lake, British Columbia). Ross, 1944:

291. Denning, 1948b: 105, plate 2, Fig. 9 - 9b. Ross and Spencer, 1952: 45. Ross, 1956: 117, Fig. 184A-B. Denning,

1956b:74. Fischer, 1960: 150. Smith, 1965: 243. Smith 1968: 658, 670, 673, Table 1. Schmid, 1970: 87, 132. Fischer,

Quaest. Ent., 1977 13 (1)

28

Nimmo

1971: 132. Anderson and Wold, 1972: Table 1. Newell and Potter, 1973: 14. Schmid, 1974: 933, Fig. 1.

This species is very similar to R. acropedes Banks (see Nimmo, 1971a: 23, Fig. 39-41).

Males of R. vao are distinguished by lack of acuminate process on postero-dorsal edge of seg-

ment IX; by presence of lip along disto-ventral area of mesal face of clasper distal segment; and

by abrupt constriction of mid-point of basal segment of clasper (Fig. 23). Females are distin-

guished by carinate postero-ventral edge of segment VIII curved postero-dorsad (Fig. 25).

Description. — Antennae deep yellow-brown, with paler annular sutures. Vertex of head dark brown anteriorly, paler

to yellow-brown posteriorly; more uniformly dark brown in female. Thorax brown dorsally, pale straw to almost white lat-

erally. Spurs dark brown. Male fore-wing length 10.08 mm; tinted brown, with faint irrorations; with hyaline areas. Hind-wing

hyaline basally, faintly tinted grey-brown distally. F emale fore-wing warm reddish-brown, with faint distal irrorations, plus

hyaline areas; hind-wing as in male. Venation indistinguishable from that of R. acropedes Banks (see Nimmo, 1971a: Fig. Sa-

bi.

Male genitalia. (Specimen from Bauerman Bk, nr Twin Lk, 6,400', Waterton National Park, Alberta). Segment IX fairly

uniformly long in lateral aspect (Fig. 23), with anterior black border broadly indented ventrally; posterior border irregular,

with pronounced dorsal process with minute postero-dorsal projection immediately dorsad of segment X base. Clasper with

deep base, narrowly constricted posterad then broadly widened distad; distal segment with deep base, abruptly narrowed to

roundly tapered disto-ventral portion with heavily setose mesal face; setose portion set off by rounded ventral ridge. Segment

X with antero-dorsal bump on each lateral half; each half tapered postero-dorsad; with slight concavity on antero-ventral angle.

Aedeagus small, with paired, large, fleshy, ventral lateral arms with lightly sclerotised, semi-circular tips heavily setose (Fig. 24);

median shaft sclerotised from base, with even basal 0.66 followed by tapered distal portion curved dorso-posterad.

Female genitalia. (Specimen from Cameron Ck, 5,480', Waterton National Park, Alberta). Sclerotised portion of segment

VIII evenly tapered posterad, with slight taper at anterior edge (Fig. 25); retractor rod firmly attached at mid-point of lat-

eral edge, with dark, linear, tapered extension ventrad within body of segment wall; postero-ventral surface pinched in mesally,

carinate, the keel curved postero-dorsad.

Notes on habitat and activity period. — This species appears to favour the smaller hill streams

with gravel or small boulder bottoms, and with medium, at times turbulent, flow. The Alberta

flight season extends from about mid-July to late September.

Geographical distribution. — The known range extends from Alaska to Montana and Oregon

(Fig. 1 10), largely confined in Alberta to foothills of the Rocky Mountains, with one record

from the south-west flank of the Swan Hills. The known altitudinal range is from 2,500' to

6,400'.

I examined 203 specimens ( 133 males, and 73 females) from the study area.

Note. — When, in my 1 976 summer collecting, many specimens of this species were taken,

I checked back to the material recorded in my previous paper (Nimmo, 1 971a), for R. acrope-

des Banks. Little material of R. vao was found there. That material has been recorded here, and

corrections made to the detailed record of acropedes. This elision of material from the acrope-

des record does not require changes to my original distribution map (Nimmo, 1971a: Fig. 108).

The rotunda group

This group is here recorded for the first time from the study area. The following character-

isation is abridged in translation from Schmid ( 1 970:43).

Dorsal processes of segment IX and anal appendages fused throughout their length, projec-

ted posterad almost as far as claspers. Segment X a long, curved band; vertical; usually within

segment IX. Anal sclerites large, fused on basal half, distinct from apical band. Apical band

very large, strongly sclerotised, articulated basally with segment X; free, bifid distally, appar-

ently almost an independent appendage. Aedeagus very large, heavy, deeply concave dorsally;

distally bifid with reduced dorsal process. Lateral arms membranous, basally erectile. Claspers

short, with distinct relief on mesal face.

Of the six species known from this group, one is here recorded from the study area, and

described as new.

Rhyacophilidae and Limnephilidae

29

Rhyacophila donaldi Nimmo new species

(Fig. 3 a-b, 4 a-b, 14-22, 114)

This species is very similar to R. ebria Denning. Males of R. donaldi are distinguished from

males of R. ebria by a variety of minor details. However, the priminent distinguishing feature

is the high angular dorsal edge of distal segment of clasper in lateral aspect (Fig. 14). In R. eb-

ria this edge is concave ventrad. Females of R. donaldi are characterised by dorsal surface of

sclerotised portion of segment VIII sharply concave (Fig. 1 9), and by hyaline ‘lobe’ immediat-

ely posterad of attachment point of retractor rod. Spermathecal sclerite (Fig. 21, 22) membran-

ous dorsally, with complexly folded ventral plate with pair of ventro-lateral apertures.

Description. — Antennae very dark brown, except inter-annular sutures paler. Vertex of head very dark brown, al-

most black, with warts very slightly paler. Thorax very dark brown, with pleura slightly paler. Spurs brown; formula 3,4,4.

Fore-wing length of male 7.84 mm; uniform red-brown, veins darker, with hyaline areas as in Fig. 3a. Venation of male as

in Fig. 3 a-b; of female as in Fig. 4 a-b.

Male genitalia. (Specimen from Rowe Bk, 6,550', Waterton National Park, Alberta). Segment IX with long dorsal area;

narrowed rather abruptly ventrad to form short ventral area (Fig. 14); dorsal area, in dorsal aspect, with deep cleft in poster-

ior edge to receive dorsal process of segment IX (Fig. 15); this process rounded, curved slightly postero-ventrad, situated im-

mediately dorsad of, and apparently fused to dorsal lobe of segment X. Clasper directed postero-dorsad, with ridged and fol-

ded mesal face (Fig. 14), with tuft of long, black setae on disto-dorsal angle; distal segment angular except rounded postero-

ventrad directed extremity with mesal face setose. Segment X, other than dorsal process, of two long, narrow straps depen-

dent from antero-ventral corners of process to engage antero-lateral edges of distal portion of tergal strap (Fig. 16). Distal

portion of tergal strap deeply cleft, each half with edges folded dorsad to enclose ventro-lateral lobes of anal sclerite (Fig.

17) . Aedeagus with lateral arms dependent from baso-ventral portion; basal two-thirds of arms membranous, erectile (Fig.

18) ; distal third weakly coloured, sclerotised, curved dorsad; spinate along edges and mesal face; median shaft scoop-like,

with upturned lateral edges; with heavily sclerotised distal portion curved, tooth-like; ejaculatory duct terminated at tip of

recurved, tapered, dorsally directed process.

Female genitalia. (Specimen from Bertha Bk, 6,000', Waterton National Park). Segment VIII with sclerotised portion tap-

ered, mid-portion of dorsal surface sharply concave (Fig. 19); postero-dorsal edge indistinctly merged with membranous area;

retractor rod fused to anterior edge, adjacent to short, rounded hyaline area (Fig. 19). Spermathecal sclerite complex, with

membranous dorsal area, very distinct, folded ventral sclerite with lateral apertures at mid-point of length (Fig. 21, 22); scler-

ite deeply cleft along ventral mid-line from anterior edge; small angular sclerite located antero-dorsad of ventral sclerite, one

on each side.

Notes on habitat and activity period. — I am not directly familiar with the localities recor-

ded for this species, but they appear to be in the sub-alpine area of Waterton National Park,

and are very likely small, shallow streams flowing over gravel, sand, or small rock bottoms,

with occasional rapids or small falls. Flight season is known to extend at least over the period

August 10 to September 8.

Holotype. — Male. Rowe Bk, 6,550', Waterton National Park, Alberta (Fig. 1 14); August

10, 1975; D.B. Donald.

Allotype. - Female. Bertha Bk, 6,000', Waterton National Park, Alberta; August 30, 1975;

D.B. Donald.

Paratypes. - Bertha Bk, 6,000', Waterton National Park, Alberta; August 30, 1 975; D.B.

Donald; six males, two females. Lineham Bk, 7,050', Waterton National Park, Alberta; Sep-

tember 7, 1 975; D.B. Donald; two males, one female. Bertha Bk, 6,000', Waterton National

Park, Alberta; August 25, 1976; D.B. Donald; nine males, three females.

The holotype, allotype, and Bertha Bk (30/8/75) paratypes are in the Canadian National

Collection, Ottawa, with type series number 15,161. The Bertha Bk (25/8/76) paratypes are

in collections of the following institutions: four males, one female to Department of Entomo-

logy, University of Alberta; five males, two females to Royal Ontario Museum, Toronto. The

Lineham Bk paratypes are in United States National Museum, Washington D.C., U.S.A.

This species is named for David B. Donald, Canadian Wildlife Service, University of Calgary,

collector of the type series.

Quaest. Ent., 1977 13 (1)

30

Nimmo

The sibirica group

Rhyacophila unimaculata Denning, 1941

(Fig. 30-31, 111)

Rhyacophila unimaculata Denning, 1941: 198 - 199, Fig. 7. (Type locality: Robson, British Columbia). Ross, 1944: 291.

Ross and Spencer, 1952: 45. Ross, 1956: 120, Fig. 234A, G. Ross, 1965: 591. Schmid, 1970: 65, 126, Plate 15, Fig. 4 -

8. Fischer, 1971: 130. Newell and Potter, 1973: 13.

In the study area this species is quite distinctive. The most prominent features are: long, dis-

tally flared dorsal lobe of segment X; and long, horizontal, stout ventral lobe, in lateral aspect

(Fig. 30).

Description. — (Taken from pinned specimen). Antennae deep red-brown. Vertex of head dark chocolate-brown,

almost black, warts slightly paler. Thorax overall virtually black. Legs red-brown to straw-colored, somewhat banded mid-

dle legs; spurs formula 3,4,4. Fore-wing length of male 10.24 mm; translucent deep grey-brown, faintly irrorate, with larger

hyaline areas at distal ends of marginal cells ft, f3, f4, f5, and between M3+4 and wing edge basad of termination of A. Hind-

wings translucent grey-brown disto-anterad, remainder hyaline. Stigma evident, not prominent, on both wings. Venation

little different from that of R. rickeri Ross (Nimmo, 1971a: Fig. 12 a - b) except fore-wing cross-vein rl - r2 present; no

drawing is available as only holotype examined, which is pinned, with wings attached and rumpled somewhat.

Male genitalia. (Specimen from Robson, British Columbia. Holotype). Segment IX massive, longer dorsally, constricted

toward ventral area (Fig. 30), antero-ventral edge paler. Clasper with irregular, slightly tapered proximal segment with con-

cave mesal face; distal segment tapered distad, curled disto-mesad, with concave disto-mesal face; disto-mesal surface clothed

in Fine, very short pilosity. Segment X large, in lateral aspect like V on side; dorsal lobe projected postero-dorsad, with com-

plexly folded anterior portion, and distal doubly cleft tip (Fig. 31), with paired, flared lateral flaps setose on mesal face; ven-

tral lobe massive, horizontal, distally slightly cleft mesally, black distally, with dorsal groove. Aedeagus massive, complex

(Fig. 32); dorsal lobe with concave dorsal surface, tip curved dorsad with shallow mesal cleft; ventral lobe originated from

large, bulbous, membranous mass, long, sclerotised, wide with disto-dorsal surface concave, all with dorsal surface with min-

ute, posteriorly directed, triangular dentitions, fewer distally; median shaft sclerotised, with paired lateral flanges with free

distal spines, basal two-thirds arched dorso-laterad then ventrad; ejaculatory duct on long, thin tube, sinuate, directed postero-

dorsad, with slight distal flare. Sclerotised tubular base of aedeagus with dorsal process from antero-dorsal edge, which abuts

on antero-ventral edge of ventral lobe of segment X.

Female. Unknown, or not yet associated.

Notes on habitat. — None of the three recorded localities is directly known to me; I have

not been to western Montana, and the Fernie and Robson, British Columbia localities are

insufficiently precise, being, presumably, the nearest named places. However, all three local-

ities are at relatively low altitudes, in areas along the eastern periphery of the Rocky Mount-

ain Trench, and higher than the floor of the trench except perhaps in western Montana. The

only collecting date known to me, April 16, at Robson, B.C., probably explains why this spe-

cies is so rarely taken, as it is well in advance of the great majority of other Trichop tera spe-

cies in the area, and therefore of the usual field season.

Geographical distribution. — This species is presently known only from localities at low

altitude, along the southern half of the Rocky Mountain Trench. Altitudes known to me are

(approximately) 3,000' and 3,300'. See Fig. 111.

The verrula group

Rhyacophila autumnalis Nimmo new species

(Fig. 5 a-b, 26-29, 112)

This species is similar to R. potteri Denning and Schmid (1971: 1556). Male R. autumnalis

are distinguished by distal clasper segment strongly tapered to fairly fine tip (Fig. 26), basal

segment of fairly uniform height; dorsal lobe of segment X complexly folded in lateral aspect,

smoothly curved postero-dorsal lobe of segment IX with dorsal, ventral sides, and tip concave

and dorsal concavity extended anterad as deep trough to dorsal surface of main body segment.

Description. — Antennae uniform dull-brown. Vertex of head chocolate-brown except warts, lateral sutures, and

postero-mesal area deep yellow-brown. Thorax deep brown to pale dorsally, generally pale yellow-brown laterally. Fore-leg

spurs straw-yellow; remainder dark brown; formula 3,4,4. Fore-wing length of male 9.44 mm; pale grey-brown; heavily

Rhyacophilidae and Limnephilidae

31

irrorate, with solid colouration less than 50% of wing area. Hind-wing hyaline generally, with dusky grey areas at terminations

of veins R2 to Culb. Venation of male as in Fig. 5 a-b.

Male genitalia. (Specimen from Rowe Bk, 6,350', Waterton National Park, Alberta). Segment IX massive (Fig. 26), with

long dorsal strap and ventral surface, longer lateral walls; notable is process of postero-dorsal edge, directed postero-dorsad,

with concave dorsal and ventral surfaces, and tip; black border to anterior edges of segment. Clasper with large, irregular, par-

allel-sided basal segment with proximal and distal areas of mesal face membranous; distal segment triangular, strongly tapered

to tip. Segment X large, projected posterad of segment IX dorsal process; complexly folded dorsal surface, broadly cleft pos-

tero-mesally (Fig. 28); with paired lateral straps dependent from antero-lateral corners. Anal sclerite large, sinuately curved

ventro-anterad then ventrad to lower portion of segment IX and clasper base area (Fig. 26); concave on posterior face to en-

close similarly sinuate, darker, tergal strap (Fig. 26, 27). Aedeagus with membranous base and clear, sclerotised lobe dorsad

of median shaft base (Fig. 29); base of median shaft a membranous lobe with sclerotised dorso-anterad hook; distad of hook

a thin, membranous tube followed by thin, irregular, sclerotised tube; lateral arms with common, long, membranous base or-

iginated ventrad of other structures; separate sclerotised distal arms broadly blade-like, tapered to blunt tip, with dorsal por-

tion of median face heavily setose.

Female. Unknown, or unassociated.

Notes on habitat and activity period. — This species appears to favour small gravel or rocky

creeks close to or in the sub- alpine region. Flight date records extend from August 24 to Sep-

tember 12.

Holotype. — Male. Rowe Bk, 6,350', Waterton National Park, Alberta; September 12, 1975;

D.B. Donald. (Fig. 112).

Paratypes. — Same data as holotype; four males. Ruby Bk, 6,750', Waterton National Park,

Alberta; 24/8/75; D.B. Donald; one male.

The holotype and two of the Rowe Bk paratypes are in the Canadian National Collection,

Ottawa, with type series number 15,162. The Ruby Ck paratype is in the Department of Ent-

omology, University of Alberta, and one each of the remaining Rowe Bk paratypes is in the

Royal Ontario Museum, Toronto, and the United States National Museum, Washington, D.C.,

U.S.A.

This species is named with reference to the autumnal flight period.

The vagrita group

Rhyacophila milnei Ross, 1950

(Fig. 33-34, 112)

Rhyacophila milnei ; Nimmo, 1971a: 36 - 37, Fig. 93 - 95, 119.

Description. — The females are like males, except that the female antennae are very dark brown, almost black. Also,

while the middle and hind-leg spurs are longer than those of the fore-leg, and stout, they are not remarkably long.

Female genitalia. (Specimen from Twin Ck, Marmot Basin, west of Kananaskis River). Sclerotised portion of segment

VIII evenly tapered posterad (Fig. 33), concave dorsally for most of length; retractor rod firmly, broadly attached to anter-

ior edge, with less darkly sclerotised, rounded area extended along segment side to enclose elliptical, hyaline spot. Sperma-

thecal sclerite minute, sinuate in lateral aspect, ends acuminate; in ventral aspect (Fig. 34) sclerite bifid, cleft from posterior

end; lobes arcuate, tapered, acuminate posteriorly; sclerite with small, rounded process anterad.

Notes on habitat and activity period. — I have not visited the collection site, which is a

sub- alpine, or close to sub-alpine area in which T win Ck may be expected to be small, bubbl-

ing, with gravel to small boulder bottom. Combining capture date information with that given

for the male paratype in my 1971(a) paper, the flight period for this species is increased to at

least one month, from September 5 to October 5.

Geographical distribution. — The locality recorded here is about 25 miles southeast of the

type locality (Fig. 1 1 2), at about 6,500'.

I examined three specimens of this species, two males, one female, all from the same local-

ity and date. These were obtained by R. Mutch of the University of Calgary.

Unassociated females

I record here two further species of Rhyacophila from the study area, each represented by

Quaest. Ent., 1977 13 (1)

32

Nimmo

single females. They do not approximate in characters to any species of which I have know-

ledge. These two species are numbered onward consecutively from the two species recorded

in my 1971(a) paper, represented only by females.

Rhyacophila species 3

(Fig. 6a-b, 35-37, 1 13)

This female is distinguished by retractor rod unattached to anterior edge of segment VIII,

which is produced anterad as small stubby process to meet rod (Fig. 35). Anterior border of

segment VIII tapered shortly anterad.

Description. — Antennae chocolate-brown, inter-annular sutures paler. Vertex of head deep chocolate-brown with

lateral sutures almost white; warts slightly paler. Thorax deep chocolate-brown dorsally, with mixed darker and paler areas

laterally. Spurs deep yellowish brown; formula 3,4,4. Fore-wing length of female 14.92 mm; brown, generally irrorate. Ven-

ation of female as in Fig. 6 a-b.

Male. Unknown, or not yet associated.

Female genitalia. (Specimen from Bertha Bk, 6;000', Waterton National Park, Alberta). Sclerotised portion of segment

VIII tapered posterad in lateral aspect (Fig. 35) with ventral surface curved gently postero-dorsad; anterior end tapered slight-

ly anterad, with sharp ridge demarcating anterior and posterior tapers. Retractor rod ‘socketed’ into small process of an-

tero-lateral edge, posterad of which is a small oval hyaline area. Spermathecal sclerite complex (Fig. 36, 37); of two main

lateral parts, darker interior sclerite centrally, and smaller posteriorly -hooked anterior sclerite partly internal; main body of

spermathecal sclerite long, tapered, acuminate in lateral aspect, tapered, blunt, rounded in dorsal aspect. Antero-ventral an-

gles of each half of main sclerite produced antero-ventrad as broad flaps which enclose yet another anterior sclerite attached

to membranous anterior tube; in dorsal aspect this sclerite is compositely triangular.

Notes on habitat and activity period. - The single locality for this species is a small stream

with pebble bottom, with scattered larger rocks, overhung by willows. Date of capture was

June 25.

Geographical distribution. — Only one locality is known for this species, in extreme south-

western Alberta, in the mountains (Fig. 1 13). Elevation 6,000'.

Rhyacophila species 4

(Fig. 7 a-b, 38-41, 113)

This female is distinguished by retractor rod firmly attached to anterior edge of segment

VIII (Fig. 38), with three tapered, black, acuminate lines produced at right angles to each

other from point of attachment. Spermathecal sclerite single, wedge-shaped in lateral aspect

(Fig. 41).

Description. — Antennae, head, and thorax black. Spurs dull grey-brown; formula 3,4,4 . Fore-wing length of fe-

male 8.32 mm; dark grey-brown with paler patches in distal cells, costal area, and termination of cubital and anal veins. Ven-

ation of female as in Fig. 7 a-b.

Male. Unknown, or not yet associated.

Female genitalia. (Specimen from small stream on north side Scalp Ck road, north boundary of Ya Ha Tinda Ranch,

Alberta). Sclerotised portion of segment VIII tapered postero-dorsad (Fig. 38); with dorso-lateral lobe at posterior edge;

postero-ventral edge acuminate in lateral aspect, shallowly cleft mesally in ventral aspect (Fig. 39); retractor rod firmly at-

tached to lateral edge, with three black, acuminate ‘arms’ radiated at right angles from point of attachment; median ‘arm’

terminated adjacent to elliptical hyaline spot. Spermathecal sclerite in lateral aspect wedge-shaped (Fig. 41), very dark about

edges, with acuminate posterior end; merged anterad to rectangular anterior portion from ventral surface of which depends

a lightly sclerotised, irregular tube; in ventral aspect (Fig. 40) sclerite with curved lateral edges, tapered to rounded poster-

ior end; anterior end semi-circular.

Notes on habitat and activity period. — This specimen was taken adjacent to a very small

(maximum width about 12”), cold stream flowing in pools and riffles, often under moss floor

of an aspen forest. Date of capture was July 16.

Geographical distribution. — The Ya Ha Tinda Ranch is located in hill country on north

side of the Red Deer River, between first and second ranges of the eastern Rocky Mountains

(Fig. 1 13). Elevation of collection point about 5,800'.

Rhyacophilidae and Limnephilidae

33

Notes on species of Rhyacophila

previously recorded from Study Area (Nimmo, 1971a)

These notes are derived from Schmid’s ( 1970) revision of the genus Rhyacophila.

Rhyacophila narvae Navas, 1926

Rhyacophila vepulsa Milne. Nimmo, 1971a: 28-29.

Rhyacophila narvae Nav^s, 1926: 57, PI. 1 Fig. 7. (Type locality: Vladivostok). Schmid, 1970: 125; ( vepulsa as junior syn-

onym of narvae). (See Fischer, 1960: 107, and 1971: 103 for palaearctic literature).

The range of this species includes eastern Siberia and western Canada.

Rhyacophila vobara Milne, 1936

Rhyacophila vobara Milne. Nimmo, 1971a: 32-33.

Schmid (1970) places this species in the vofixa group, removing it from its own nominal

group.

Rhyacophila bifila Banks, 1914

Rhyacophila bifila Banks. Nimmo, 1971a: 25.

Schmid ( 1970) removes this species from the invaria group and places it in the coloradensis

group.

Rhyacophila coloradensis Banks, 1904

Rhyacophila coloradensis Banks. Nimmo, 1971a: 26.

Schmid (1970) removes this species from the invaria group and places it in the coloradensis

group.

Rhyacophila vocala Milne, 1936

Rhyacophila hyalinata Banks: Nimmo, 1971a: 17, 18, 27, 203, 205, 214, Fig. 11 a-b, 51 -53, 111.

D.G. Denning (pers. comm.) pointed out that my identification of this species is incorrect.

As a result, the bibliography presented under R. hyalinata Banks, 1905, on p. 27 (Nimmo,

1971a) is to be deleted. Following is the correct bibliography for R. vocala Milne, 1936:

Rhyacophila vocala Milne, 1936: 100, 102, Fig. (unnumbered). (Type locality: Cultus Lake, British Columbia). Ross, 1944:

291. Denning, 1948b: 106, PI. 4 Fig. 16 - 16B. Ross and Spencer, 1952: 45. Ross, 1956: 118. Denning, 1956a: 235, Fig.

10, lie. Fischer, 1960: 152. Smith, 1968: 658, 663, 672, 673, Tab. 1. Schmid, 1970: 59, 124, PI. 15 Fig. 1 - 3, PI. 50

Fig. 4- 5. Fischer, 1971: 135. Newell and Potter, 1973: 14.

Deleting the distribution for hyalinata from Fig. 1 1 1 (Nimmo, 1971a), the distribution of

vocala extends from Alberta and southern British Columbia to California.

Quaest. Ent., 1977 13 (1)

34

Nimmo

Placement of Species of Rhyacophila

new to Study Area, in Keys previously constructed

Males. Refer to key on p. 16 of Nimmo, 1971a.

R. simplex Nimmo n. sp. keys to 1 8a, wherein it may be distinguished by simple

lateral lobe of segment X, horizontal in lateral aspect (Fig. 10).

R. vao Milne keys to 14b wherein it may be distinguished by anal sclerite absent

(Fig. 23).

R. donaldi Nimmo n. sp. keys to 7a wherein it may be distinguished by postero-dor-

sal lobe of segment IX fused dorsally to segment X(Fig. 14).

R. unimaculata Denning keys to 16a wherein it may be distinguished by segment

X V-shaped in lateral aspect (Fig. 39), with dorsal lobe narrow, flared dis tally.

R. autumnalis Nimmo n. sp. keys to 10a, wherein it may be distinguished by tergal

strap not dish-like; segment X of one, horizontal, convolute sclerite (Fig. 26).

Females. Refer to key on p. 18 of Nimmo, 1971a.

R. vao Milne keys to 1 la wherein it may be distinguished by keel curved, postero-

dorsad(Fig. 25).

R. donaldi Nimmo n. sp. keys to 8, wherein it may be distinguished by short sperm-

athecal sclerite complexly folded within itself, with paired ventro-lateral apertures

(Fig. 21).

R. milnei Ross keys to 14 wherein it may be distinguished by spermathecal sclerite

deeply bifid posteriorly, in ventral aspect (Fig. 34).

R. species 3 keys to 14 wherein it may be distinguished by spermathecal sclerite

composed of complex of four sclerites intricately folded within each other (Fig. 36).

R. species 4 keys to 9a wherein it may be distinguished by postero-lateral edge of

segment VIII with posterior edge sinuate in lateral aspect (Fig. 38).

Finally, couplet 10 should be reworded as follows:

10a. (9a) Anterior border of sclerotised segment VIII without hyaline spot laterally

(Fig. 100 (Nimmo, 1971a)) R. species 1

1 0b. Anterior border of sclerotised segment VIII with hyaline spot laterally

(omitted in Fig. 61 of Nimmo, 1971a). . . R. narvae Navas ((= vepulsa Milne))

THE FAMILY LIMNEPHILIDAE KOLENATI

The Subfamily Dicosmoecinae Schmid

The Genus Dicosmoecus McLachlan

Dicosmoecus gilvipes (Hagen), 1875

(Fig. 42-46, 114)

St enophy lax gilvipes Hagen, 1875: 601-602, 605. (Type locality: Colorado). Putnam, 1876: 205.

Dicosmoecus gilvipes', McLachlan, 1875: 113. Banks, 1892: 364. Ulmer, 1905: 20. Ulmer, 1907: 60. Banks, 1907: 38.

Dodds and Hisaw, 1925a: 125, Fig. 6. Dodds and Hisaw, 1925b: 386. Essig, 1926: 176. Neave, 1929: 189. Betten et al,

1934: 53, 318, Textfig. 271, m, q. Milne, 1935: 36, 50. Ross, 1938b: 30. Balduf, 1939: 122, Fig. 62b. Banks, 1943:

360, PI. 4 Fig. 89, PI. 5 Fig. 99, 107, 110, 111. Ross, 1944: 297. Schmid and Guppy, 1952: 42. Ross and Spencer,

1952: 47. Schmid, 1955: 36, Fig. 3e, 9-10. Flint, 1960: 23-24, Fig. 4. Flint, 1966: 367, Fig. 2 i, h. Anderson, 1967:

Table II. Fischer, 1967: 65-66. Fischer, 1973: 12-13.

This is the third species of Dicosmoecus recorded from the study area. Males are distingui-

tinguished by presence of small, slightly clavate lobe originated just ventrad of clasper base

(Fig. 42); by notch in dorsal edge of distal clasper segment, at base; and by several other

minor characters. However, the best character is the form of the mesal ridge of the basal

clasper segment viewed from posterior (Fig. 43); with straight, rounded ventral ridge at 45°

Rhyacophilidae and Limnephilidae

35

to vertical, dorso-laterad of which is a distinct, thin- walled, black tooth directed postero-lat-

erad.

Females are distinguished by apparent absence of supragenital plate (Fig. 46); by massive

vulval scale fused dorso-mesally; by inverted- triangular median lobe of vulval scale; by short,

triangular segment X(Fig. 45); and by sharply triangular ventro-lateral lobes of segment IX.

Description. — Antennae very dark grey, annuli anterior faces pale; scape antero-mesal face cream-colored, glabrous.

Vertex of head black, faded posterad to red-brown. Frons clothed in golden hair; warts of vertex with black or white hairs;

hairs of antennal scape golden. Thorax very dark brown, faded to red-brown and yellow, dorsally; laterally straw-colored to

red-brown, with some very dark brown areas. Thoracic hairs very fine, white except at fore-wing base heavier (setose), black.

Legs yellow except hind-legs darker straw-colored. Spurs red-brown; formula 1,3,4. Fore-wing length of male 24.8 mm; deep

grey-brown, semi-translucent, with veins deep brown, almost black; hyaline areas basad of divergence of Ml+2 and M3, at

termination of Cu2 and anal veins, and adjacent to termination of C. Hind-wing uniform pale grey. Venation as for Dicosmoe-

cus jucundus Banks (Nimmo, 1971a: Fig. 122 a-b).

Male genitalia. (Specimen from Oldman R., Hwy 922, nr Maycroft, Alberta). Segment VIII tergum with paler posterior

border. Segment IX high, irregularly slender (Fig. 42); with projection posterad of poster o-ventral edge in lateral aspect; with

basal portion of antero-lateral edge black, the black line curved postero-dorsad parallel to clasper base. Clasper with massive,

slightly tapered proximal segment in lateral aspect; distal segment with broad base narrower than distal edge of proximal seg-

ment, distinct notch at baso-dorsal angle, distal process long, tapered, distally acuminate, curved postero-mesad. Mesal face

of proximal segment of clasper concave, with baso-mesal edge with oblique ridge slanted latero-ventrad (Fig. 43); ridge with

dorso-lateral, black, sharp-edged tooth ventro-mesad of which is a long, rounded ridge. Dorsal strap of segment IX high, nar*-

row, indistinguishable from segment X. Cerci long, parallel-sided in lateral aspect, concave on mesal face, with fringe of setae

distally. Median lobes of segment X dorsad of anus bifid, short, rounded, closely appressed; ventro-lateral lobes long, slender,

slightly clavate distally, each with short, thumb like subsidiary lobe slightly dorso-mesad of base; ventro-lateral lobe pair con-

nected with median bridge arched dorsad just below anus; posterior edge of bridge projected postero-dorsad, parallel-sided,

rounded distally, dark. Postero-ventral angle of segment X projected postero-ventrad as large ledge which bears above assem-

blage of lobes. Aedeagus long, slender, parallel-sided, with rounded distal head slightly delimited; lateral arms originated basad

of head, on ventro-lateral face; not quite extended to tip of median shaft, tapered, acuminate, with fringe of short, heavy,

black spines on dorsal edge, a few such spines scattered on ventral edge (Fig. 44).

Female genitalia. (Specimen from Clatsop County, Oregon). Posterior edge sternum VII widely bordered with fine hairs.

Vulval scale massive, with lateral lobes fused dorso-mesally (Fig. 46), with meso-ventral faces deeply recessed; median lobe

with narrow base, triangularly expanded distally, disto-lateral angles housed in lateral lobe recesses. Segment IX in lateral

aspect with irregular dorsal portion ventrally concave; dependent from antero-lateral angles a triangular sclerotised lateral

process with narrow bridge to main segment body (Fig. 45), laterad of vulval scale, embedded in membrane. Segment X small,

triangular, demarcated from IX by curved, abrupt declivity; mesally cleft in ventral aspect (Fig. 46); with mesally directed,

narrow, rounded lobes from antero-lateral angles. No evident supragenital plate.

Notes on habitat and activity period. — The single specimen taken by myself was under a

bridge high above the Oldman River at a point where it emerges from the mountains, through

Foothills region. There, the river is roughly 1 50' wide, forming shallow rapids over large rocks

fragmented from cliffs on either side. Neave (1929) records the species from two lakes in Jas-

per National Park, but I suspect these were adults that flew in from surrounding streams which,

in that area, are the smaller, mountain type. Known dates of capture are July 24, August 27,

and September 5.

Geographical distribution. — This species is known from British Columbia south to Califor-

nia and Colorado (Fig. 1 14), with three records from Alberta. Known altitudes are 4,000'.

4,900', and 5,490'.

I have examined two specimens, one male, one female.

The Genus Imania Martynov

The tripunctata group

Imania thomasi Nimmo new species

(Fig. 8 a-b, 56-58, 1 15)

Males of this species, compared to others of tripunctata group, are distinguished by clasper

distal segment trilobed, dorso-lateral lobe deeply cleft longitudinally as dorsal, ventral lobes

(Fig. 56, 57); segment X distal edge triangularly cleft (Fig. 57); aedeagus basal portion with

Quaest. Ent., 1977 13 (1)

36

Nimmo

four, paired, lobes (Fig. 58), aedeagus distal portion elongated, tip concave dorsally, edges

flared dorso-laterad.

Description. — Antennae brownish black. Vertex of head black. Thorax black dorsally, with slightly paler areas lat-

erad. Spurs brown; formula 1,3,3. Fore-wing length of male 8 mm; translucent purplish brown; with hyaline areas as in Fig.

8a. Venation of males as in Fig. 8 a-b.

Male genitalia. (Specimen from Rowe Bk, 6,350', Waterton National Park, Alberta). Segment IX of uniform width, except

ventral surface expanded posterad; curved gently dorso-posterad; with black bordered anterior edge (Fig. 56). Clasper massive,

with large, roughly elliptical in lateral aspect, basal segment; antero-mesal angle developed meso-posterad as long, sinuate,

slender, acuminate process (Fig. 57). Distal clasper segment trifid; in lateral aspect lateral face expanded broadly distad;

deeply, broadly cleft to dorsal and ventral lobes; mesal lobe developed from meso-basal area of segment, thin-walled dorso-

ventrally (Fig. 57), finger-like in lateral aspect (Fig. 56); all three lobes toothed as in Fig. 57. Segment X uncertain in nature;

possibly ventral portion fused to long, slender, down-curved process developed from postero-dorsal edge of segment IX (Fig.

56); ventral portion v-cleft distally (Fig. 57), with bridge between lateral halves, and aperture anterad of bridge. Aedeagus

(Fig. 58) very long, with sclerotised, tubular base; middle portion with two pairs of dorsal arms; proximal pair relatively

straight, black, originated dorsally; distal pair lightly sclerotised, distally curved postero-ventrad to acuminate tip, thin blades.

Distal portion of aedeagus dependent postero-ventrad below base of distal arms, long, bulbed two-thirds distance from tip;

bulb with aperture of ejaculatory duct; distad of bulb a constriction, flared to dorsally concave, thin-walled tip.

Female. Unknown, or not yet associated.

Notes on habitat. — The two localities for this species are both at, or above, the sub-alpine

region. The streams adjacent to which specimens were taken are both small, relatively shallow,

with gravelled or small boulder bottoms, with occasional rapids or small falls.

Geographical distribution. — The localities are in mountainous country, close to the alpine

region, and about 200 miles apart, north to south (Fig. 1 1 5), in Alberta only. Altitudinal range

is from 6,350' to 6,500'.

I examined three males of this species.

Holotype. — Male. RoweBk, 6,350', Waterton National Park, Alberta; 12/6/75; D.B. Donald.

Paratypes. — Twin Ck, Marmot Basin, west of Kananaskis R., Alberta; 12/5/75; R. Mutch;

one male. Twin Ck (as above); 26/6/76; R. Mutch; one male.

The second paratype has wings fragmented, from time of collection.

The holotype and Twin Ck (26/6/76) paratype are in the Canadian National Collection,

Ottawa, with type series number 1 5,163. The Twin Ck (1 2/5/75) paratype is in the Depart-

ment of Entomology, University of Alberta.

This species is named for my late father-in-law, Thomas Finlay Hird, M.D.

The Subfamily Neophylacinae Schmid

The Genus Neophylax McLachlan

This genus is recorded for the first time from the study area, where it is represented by one

species. The following characterisation is abridged in translation from Schmid (1955: 93-96).

Synopsis of characters. — Medium to small, slender, with wings strongly irrorate, hairy.

Male maxillary palps very large, first article short, second at least as long as scape. Meso-apical

spur of male hind-leg modified. Specimens of most species with process directed posterad

from sternum VI and/or VII. Posterior borders of abdominal segments of both sexes often

with fine granular pattern. Wing from various species with smaller members with anterior wings

distally angular; in species with larger members, anterior wings elongated. Venation much as

in Fig. 9 a-b (this paper), with variations between species, and minor sexual dimorphism. Male

genitalia with segment IX massive, with postero-ventral process; dorsal process often very long,

tapered, fused at least basally. Median lobes of segment X primitively small, weakly concave;

large, curved in advanced species; never fused to segment IX. Aedeagus small, located dorsally,

between dorsal processes of segment IX; lateral arms absent. Claspers not projected; basal ar-

ticle much reduced, composed of irregular field ventrad of aedeagus; apical article often bifid,

complex, with points or dentitions. Female genitalia scarcely distinguished from Oligophlebo-

des except on basis of vulval scale which is primitively very small, lightly sclerotised, flanked

Rhyacophilidae and Limnephilidae

37

laterad by ventral lobes of segment IX fused to it. In more advanced species vulval scale more

strongly sclerotised, generally bifid; segment IX ventral lobes almost completely fused later-

ally.

The rickeri group

Neophylax rickeri Milne, 1 935

(Fig. 9a-b, 47-55, 1 15)

Neophylax rickeri Milne, 1935: 22, 52. (Type locality: Cultus Lk, British Columbia). Ross, 1944: 300. Denning, 1948a: 122,

PL 7 Fig. 7-7 A. Schmid and Guppy, 1952: 42. Ross and Spencer, 1952: 50. Schmid, 1955: 97, Fig. 58-59, 61c, 62c.

Denning, 1956a: 262, Fig. 10.29i. Fischer, 1967: 141-142. Anderson and Wold, 1972: 197, 198, 199, 200. Fischer, 1973:

37.

Neophylax pulchellus Ling, 1938: 68-69. (Type locality: Oakland, California). Ross, 1944: 300. Fischer, 1967: 142.

Males of this species may be distinguished by postero-dorsal lobes of segment IX extended

well posterad of remainder of genitalia (Fig. 47); and by form of postero-median plaque (Fig.

50) in posterior aspect.

Description. — Antennae straw-coloured; scape, pedicel, dark brown; up to ninth annulus with dark brown band

around posterior half of distal edge, near inter-annular suture. Vertex of head yellow-brown, with areas bounded by lateral

ocellus, frontal wart, and median wart grey-brown; warts grey-brown. Thorax with ground of straw colour, shaded to grey-

brown in parts. Spurs red-brown; formula 1,3,3; disto-mesal spur of male hind leg modified as in Fig. 55; distal edge of spur

sheath apparently fibrous. Male fore-wing 13.92 mm; dark grey, with scattered irrorations, especially in costal area; pattern

augmented by areas of silver and gold hairs; postero-distal edge markedly sinuate, with unusually distinct angle on posterior

edge of termination of Cu2 and anal veins. Hind-wing translucent grey. Venation of male as in Fig. 9 a-b.

Male genitalia. (Specimen from Galwey Bk, Highway 6, just north of Waterton National Park boundary, Alberta). Sternum

VII with posterior projection, broad, rounded process (Fig. 52); posterior area of sternum with minute ovoid markings an-

terad of which is a narrower band of linear markings. Posterior area of sternum VIII with similar linear markings. Segment

IX with area of linear markings (not illustrated). Segment IX large, bulky, with anterior edge folded somewhat upon itself

(Fig. 47); with postero-ventral area angled dorsad at about 45°, biconcave, each concavity with tuft of long setae directed

postero-dorsad (Fig. 47, 49); separated by median keel; dorsad of concavities a broadly based, tapered, black process with

irregular, transverse ridges with edges directed distad. Dorsal strap of segment IX laterally narrow; dorsally expanded postero-

ventrad in median line as bifid, tapered process, more widely separated proximally than distally (Fig. 47, 48, 50). Claspers

short in lateral aspect; exposed part inverted triangular; irregularly rounded in posterior aspect, with latero-dorsal ridge,

median black-bordered declivity within which disto-lateral flanges of segment X are housed. Segment X massive, of two parts,

one either side of segment IX dorsal strap processes; with very large, conical dorsal part hollow anteriorly, tapered to dis-

tinct, black tip with flanges developed latero-anterad; lobes of segment directed postero-ventrad. Aedeagus minute, housed

between lobes of segment IX dorsal strap; membranous, with ejaculatory duct sclerotised within; duct tubular, straight, or-

iginated from expanded, indistinct bulb (Fig. 51).

Female genitalia. (Specimen from Galwey Bk, Highway 6, just north of Waterton National Park boundary, Alberta).

Tergum VII posterior border with ovoid, closely spaced markings (Fig. 54); sternum VII with similar, linear, markings on

posterior border. Segment VII with massive, rectangular in lateral aspect, tergum, and small, triangular, indistinct sternum;

sternum, in ventral aspect, broadly dished (Fig. 53). Ventral portion of segment IX of two triangular lobes, dark, with ven-

tro-lateral folds; dorsal portion short, sclerotised, free of ventral lobes, attached to postero-dorsal edge of tergum VIII; with

segment X imperceptably attached distally. Segment X membranous, ventrally concave, with disto-ventral hook; triangular

in ventral aspect with thin lateral walls. Vulval scale set dorso-mesad of ventro-lateral lobes of segment IX; black, with lat-

ero-anterior pockets directed antero-ventrad within segment VIII; distally bifid, with lateral processes hooked mesad in ven-

tral aspect; mesad of lobes a circular cleft; ventrally concave.

Notes on habitat and activity period. — All specimens available to me were taken from three

localities on two consecutive days. The localities were: a quiet, small, gravel-bottomed creek

draining large sedge bogs; a larger stream (about 30' wide) with gravel and larger rock bottom;

and a full-sized river (about 1 50') running through a gorge, over boulder bottom. This species

appears to have a wide latitude in its choice of streams. Dates of capture were September 4 and 5.

When collecting specimens of this species, I encountered a habit previously unknown to me

in Trichoptera. The adults rested on the substrate (concrete culverts) with all feet on the sur-

face. However, the body, and consequently the wings, were raised from the surface at an angle

of about 20° or more, with head down. This simplified picking them up by hand, as they also

had the unusual habit, in culverts at least, of resting low down, close to running water.

Quaest. Ent., 1977 13 (1)

38

Nimmo

Geographical distribution. — This species is reported from British Columbia and Alberta

south to California in the west, and to Idaho and Montana in the east (Fig. 115). In Alberta

it is known only from the south-west corner of the Province, from the Oldman River south

to Waterton National Park. It appears to be a species of the eastern fringes of the Foothills.

Known altitude range is from 4,100' to 5,000'.

I have examined 1 1 specimens of this species from the study area: four males; seven females.

The Subfamily Limnephilinae Ulmer

The Tribe Limnephilini Schmid

The Genus Limnephilus Leach

The sub centralis group

Limnephilus nimmoi Roy and Harper, 1975

(Fig. 64-70, 116)

Limnephilus species 1; Nimmo, 1971a: 94-95, 203, 208, Fig. 296-297, 612.

Limnephilus nimmoi Roy and Harper, 1975: 1083, 1085-1088, Fig. 1-4, 7-8. (Type locality: Lac Monroe, Mt. Tremblant

Park, Quebec).

Males of this species are distinguished by segment X lobes long, projected posterad of cerci,

straight, tapered distally (Fig. 64); by cercus with distinct ventral edge beyond body of segment

IX; by substantial (in lateral aspect) dorsal strap; and by distinct black tooth on dorso-lateral

face of clasper tip. Females are distinguished by large dorsal body of segment IX; by long, dis-

tally up-turned segment X; and by long, slender, tapered cerci not appressed dorsally to Seg-

ment X(Fig. 69).

Description. — Antennae yellow-brown. Vertex of head, thorax, legs yellow-brown; warts paler, to white. Fore-legs

of male with brush a single row of minute dark spines along basal portion of postero-mesal edge of femur. Middle, hind-legs

with strong, dark setae on tibiae. Fore-wing length of male 11.52 mm; yellow-brown overall, with hyaline areas in pattern

typical of the sublunatus group. Stigma virtually indistinguishable. Venation typical as for Limnephilus generally.

Male genitalia. (Specimen from Lac Monroe, Mt. Tremblant Provincial Park, Quebec. Paratype). Tergum VIII with slight

postero-dorsal spinate process (Fig. 64). Main body of segment IX with acuminate antero-ventral and posterior edges; dorsal

edge produced postero-dorsad, with postero-ventral fold just dorsad of clasper; dorsal strap stout, angled posterad. Clasper

delineated from segment IX, with short postero-dorsal projection tapered distad, with dorso-mesally directed black tooth

(Fig. 64, 65). Cercus large, broad, curved posterad, with dorsal and ventral edges parallel in lateral aspect (Fig. 64); with

straight distal edge; with various stout, strongly sclerotised teeth on mesal face (Fig. 65). Segment X of two black, stout

(in lateral aspect) blades slightly longer than cerci; each blade with complexly folded ventro-lateral projections directed ven-

trad of cereal bases (Fig. 64, 65); with minute tooth just laterad of tip (Fig. 65, 66). Aedeagus very stout, short (Fig. 67);

with median shaft slightly shorter than lateral arms, strongly tapered, with distinct head; lateral arms massive, with fleshy,

erectile distal one- third fringed dorsally by heavy, dark spines; with massive dorsally directed, triangular blade at dorso-distal

angle of sclerotised portion of arm (Fig. 68).

Female genitalia. See Nimmo, 1971a: 95 for full description; and Fig. 69, 70 of this paper.

Notes on habitat and activity period. — I have taken a second female of this species in

Alberta. The first record was from a small muskeg lake in the Athabasca R. valley south of

Jasper. The second is from a similar lake, but well out in the lower foothills to the east, about

35 miles north of Nordegg, on the Forestry Trunk Road. The collection dates for Alberta are

now July 1 8, and September 10. Roy and Harper (1975) give Quebec collection dates rang-

ing from July 19 to August 28.

Geographical distribution. — To date this species is only known from Canada (Fig. 116),

with records only from Alberta and Quebec. The Alberta distribution suggests that the species

is confined to lower altitudes, whether the mountain ranges, or in the Foothills.

Rhyacophilidae and Limnephilidae

39

The picturatus group

Limnephilus insularis Schmid, 1950

(Fig. 59-63, 1 17)

Limnephilus insularis Schmid, 1950: 47-48, Fig. 1-4. (Type locality: Wellington, Vancouver Island). Schmid and Guppy,

1952: 42. Ross and Spencer, 1952: 48. Ross and Merkley, 1952: 449. Schmid, 1955: 136. Fischer, 1968: 198. Fischer,

1973: 85.

This species is distinguished by lack of spinate postero-dorsal edge of tergum VIII (Fig. 59);

by long, triangular cerci, in lateral aspect, at angle of 45° ; by lobes of segment X almost as

long as cerci, acute triangular, parallel to but ventrad of cerci in lateral aspect; and by clasper

with minute dorsal process.

Description. — Antennae reddish brown, scape and pedicel markedly darker. Vertex of head deep chocolate-brown;

frons slightly paler, with setal bases pale reddish cream; remainder of head pale yellowish brown, especially pair of warts

along postero-dorsal edge. Thorax generally deep chocolate-brown, especially dorsally; paler laterally; warts and setal bases

deep reddish straw coloured. Spurs yellow. Brush of male foreleg fairly discrete row of spines of mixed larger and smaller

sizes, along only basal two-thirds of femur. Fore-wing length of male 9.92 mm; virtually hyaline, with faint brownish tinge

faintly patterned to irrorate at distal quarter of wing; costal area quite clear; stigma a reddish brown thickening. Hind-wings

about hyaline. Venation of male typical of Limnephilus species.

Male genitalia. (Specimen from bog at highest crossing of James R.by Forestry Trunk Road, Alberta). Tergum VIII un-

modified. Segment IX main body in lateral aspect with slightly curved anterior edge (Fig. 59); widest dorsally, tapered broad-

ly basad; dorsal strap substantial (Fig. 59, 61), with considerable dorso-ventral depth. Clasper delineated from segment IX,

along full height of posterior edge, with slight, rounded, postero-dorsal process (Fig. 59,60). Cercus, in lateral aspect, acute-

triangular, directed postero-dorsad at 45°; with small, conical tooth just basad of tip, on mesal face; with postero-ventral an-

gle folded mesad (Fig. 60). Segment X of two sclerites, each with basal plate directed laterad, slightly convoluted; postero-

dorsal process long, narrow, tapered distad, almost acuminate, with minute dentitions along distal 0.25 of dorsal edge; with

slight lateral hook to tip (Fig. 61); generally darkly sclerotised, black on distal quarter. Small fleshy lobe meso-ventrad of

segment X (Fig. 60, 61). Aedeagus with almost rectilinear median shaft (Fig. 62), with distinct distal head; lateral arms

slightly shorter, broad in lateral aspect, with less coloured, expanded, leaf-like tip fringed with fine setae; with long, heavy

setae directed meso-dorsad from area two-thirds distance from arm base (Fig. 62, 63), in part originated from small lobe

projected mesad of distal expansion (Fig. 63).

Female. Unknown, or not yet associated.

Notes on life history and activity period. — The single specimen was taken by sweeping a

bog of sedge in which there are areas of standing water of various depths. The bottom is gen-

erally firm, not quaking. Through the bog extends the upper reach of James R, which is deep,

smooth-flowing, quite fast, with silt or clay bottom. Equisetum is scattered commonly through

the sedge. Date of capture was September 22.

Geographical distribution. — This species is known from Vancouver Island to central British

Columbia to the Alberta Foothills, south to Idaho (Fig. 1 1 7). Altitude of the Alberta locality

is 4,900'.

I have one male of this species.

The incisus group

Limnephilus alvatus Denning, 1968

(Fig. 71-77, 118)

Limnephilus alvatus Denning, 1968: 21-22, Fig. 7-7B. (Type locality: ‘near Lethbridge’, Alberta).

Males of this species are distinguished primarily by long, rounded process of postero-mesal

angle of cercus (Fig. 71); also by long, very slender aedeagal lateral arms originated dorso-lat-

erad on aedeagal base (Fig. 74). Females are very similar to those of L. secludens Banks, but

may be distinguished by cereal lobes projected dorsad of segment X in lateral aspect (Fig. 76);

and by ventral portion of segment IX distinct from dorsal portion.

Description. — Antennae reddish brown, as is remainder of body except vertex of head, legs paler. Spurs red-brown

except fore-leg spur black. Brush of male fore-leg largely of fine, black spines intermingled with finer, shorter spines at edges;

extended along distal three-quarters of femur, perpendicular to surface; tibia with corresponding brush of shorter, stouter

Quaes t. Ent., 1977 13 (1)

40

Nimmo

black spines directed distad. Male fore-wing length 9.76 mm; golden brown, extensively irrorate on distal, posterior portions;

clear along costal area.

Male genitalia. (Specimen from ‘near Lethbridge’, Alberta. Holotype). Tergum VIII unmodified. Segment IX high, narrow

in lateral aspect (Fig. 71), widest laterally, with dorsal strap heavy, angled posterad, black in continuation of black-bordered

anterior edge of segment. Clasper inverted acute-triangular in lateral aspect; with wide base, rounded in posterior aspect (Fig.

73); claspers almost contiguous ventrally, converged at acute angle. Cercus prominent, horizontal, with postero-dorsal angle

produced as rounded, tapered, finger-like lobe (Fig. 71, 72, 73); mesal face concave. Segment X bipartite, each part with bas-

al plate from which originates a thin-bladed projection directed postero-laterad (Fig. 73), with small hook directed anterad

in lateral aspect (Fig. 71). Aedeagus with sclerotised tubular base with strap originated from postero-dorsal edge directed pos-

tero-anterad towards segment X (Fig. 74); median shaft long, slender, bowed ventrad, with distinct, large head (Fig. 74); lat-

eral arms originated dorso-laterad of base of median shaft, almost as long as median shaft; thin, pale rods with small tuft of

setae at tip (Fig. 75).

Female genitalia. (Specimen from about 10 miles southeast of Fairview, Alberta). Vulval scale lobes of equal length; me-

dian lobe fairly narrow, with rounded tip; lateral lobes rectangular, directed postero-laterad, away from median. Supragenital

plate small, lightly sclerotised, with slight median extension posterad (Fig. 77). Segment IX with dorsal and ventral portions

separate (Fig. 76); ventral portion triangular in lateral aspect; both sides broadly joined ventrally (Fig. 77); dorsal portion in-

distinguishably fused with segment X. Segment X tubular, with disto-lateral edges bowed anterad. Cerci represented by short,

broad-based, triangular lobes fused to dorso-lateral sides of segment X.

Notes on life history and activity period. — As I have not yet taken specimens of this species

myself, I can say little. However, the known range adheres closely to the distribution of prairie

in Alberta, wherein there are prairie outliers in the northwest and north centre of the Province.

With the exception of the holotype from Lethbridge, however, all other material was obtained

from light traps set out by Alberta Department of Agriculture to survey pest species of insects.

These traps were located in agricultural areas, which occur predominantly in former prairie

areas so that this apparent prairie distribution may be an artifact.

Geographical distribution. — To date known only from Alberta, from Lethbridge in the

south, to Fort Vermilion in the far north (Fig. 118), and confined strictly to the Great Inter-

ior Plain. Altitude ranges from 900' to 3,000'.

Apart from the male holotype, I have examined a total of 40 specimens from the study

area: 31 males; nine females.

Note on taxonomy. — When Denning described this species ( 1 968) he included a female

as allotype, taken at the same time and place. This female belongs to L. janus Ross. I examin-

ed the allotype of L. alvatus and illustrate it here (Fig. 78, 79). Comparison with my illustra-

tion of L. janus (Nimmo, 1971a: Fig. 383-384) will confirm the identity of the two. Until

my 1971(a) paper, the female of L. janus had not been illustrated, though it was known — Mil-

ne ( 1 935: 42) keyed what is clearly the female of the species known as L. janus (=C. minu-

sculus ), and Ross ( 1 938b: 37) mentions the female of the species now known as janus. Finally,

in my own collecting, I have independently associated the two sexes of L. janus. While L. alva-

tus and L. janus are placed in the same species group, so also is L. secludens Banks. The males

of L. alvatus and L. secludens are very much closer to each other in appearance than either is

to that of L. janus , though superficially these three species are very similar in general habitus.

As indicated above, I found the correct female for L. alvatus by examination of light trap

collections in which both sexes were found from the same collecting event, the female of L.

alvatus very closely resembling that of L. secludens, as could be expected.

Limnephilus janus Ross, 1 938

(Fig. 78-79)

Limnephilus alvatus', Denning, 1968: 21-22, Fig. 8-8c. (Female of L. janus Ross as allotype of L. alvatus Denning).

Rhyacophilidae and Limnephilidae

41

The argent eus group

Limnephilus vernalis Nimmo, new species

(Fig. 80-84, 118)

Males of this species may be distinguished from those of L. argenteus Banks, the other

species in the group, by clasper trapezoidal in lateral aspect (Fig. 80); by median lobes of seg-

ment X projected prominently posterad of cercus; by elliptical main body of segment IX in

lateral aspect.

Description. — Antennae deep, warm, reddish brown; scapes darker, antero-mesal face pale, almost glabrous. Vertex

of head varied from deep red-brown, to paler, with warts, setal bases very pale. Thorax generally red-brown, with darker ar-

eas. Brush of male fore-leg absent. Spurs deep straw-coloured; formula unknown, hind femora missing. Male fore-wing length

14.92 mm; warm red-brown, with scattered irrorations, larger hyaline areas especially at bases of cells f2, f3, and cell between,

and around vein Ml+2, and on either side of M at mid-point of wing length. Irrorations of costal area fused to produce lar-

ger hyaline areas. Hind wing hyaline except distal cells from costa to f3 very pale brown. Venation typical as for Limnephilus

species.

Male genitalia. (Specimen from Grande Prairie, Alberta). Tergum VIII with slight projection of postero-dorsal edge as

spinate lobe (Fig. 80). Segment IX roughly elliptical in lateral aspect, with main body merged dorsad with sharp-edged dorsal

strap, tapered gently ventrad. Clasper attached only at base of process, at point half height of posterior edge of segment IX;

short, trapezoidal in lateral aspect; with toothed edges to concave mesal face (Fig. 81). Cercus rectangular in lateral aspect,

with curved distal edge; bulky in posterior aspect (Fig. 81); with dorso-ventral ridge on concave mesal face. Segment X bip-

artite; each sclerite with long, tapered, up-turned tip on median lobe, with tip slightly bulbous, curved dorso-laterad (Fig. 80,

81, 82); brown; lateral plaque with convolute lateral edge (Fig. 80), and ventrally directed acuminate process (Fig. 81). Ae-

deagus originated from tubular sclerotised base; median shaft curved gently dorsad, with distinct head; lateral arms originated

from dorso-lateral portion of membranous basal portion, with narrow proximal portion, expanded distad of mid-point as fla-

red, thin, mesally concave blade (Fig. 83, 84); blade with distal acuminate point setose on dorsal edge; with other, dorsal,

angle small, bulbous, setose; mesal face of blade with four stout, black setae.

Female. Unknown, or not yet associated.

Geographical distribution. - This species is known only from a single locality in Alberta

(Fig. 118) at about 2, 150'. Additional data are not available.

Holotype. — Male. Grande Prairie, Alberta; 25/6/75; Alberta Department of Agriculture

light trap.

The holotype is in the Canadian National Collection, Ottawa, with type number 1 5,164.

Unassociated females

Limnephilus species 2

(Fig. 91-92, 119)

The single female is similar to females of L. femoralis (Kirby), but may be distinguished

by cerci and distal projections of segment X tube in lateral aspect (Fig. 91) well separated

vertically; and by segment IX in lateral aspect parrallelogram-like. Illustrations of L. femoral-

is for comparison are to be found in Nimmo, 1971a: Fig. 335-336.

Description. — Antennae warm red-brown; scapes, pedicels darker. Vertex of head very dark brown, almost black.

Thorax very dark brown dorsally, with dark and paler brown areas laterally. Spurs dark straw-colour; formula 1,3,4. Fore-

wing length of female 13.12 mm; warm red-brown, with hyaline areas of various sizes from simple irrorations to larger, hya-

line areas in bases of cells f2, f3, the cell between these, on either side of vein Ml+2, and on either side of M about half way

along wing length; costal area clear, with pale red-brown cast. Hind wing clear, but with very pale red-brown cast. Venation

typical as for Limnephilus species.

Male genitalia. Unknown, or not yet associated.

Female genitalia. (Specimen from Three Hills, Alberta). Vulval scale (Fig. 92) with three lobes of equal length; lateral

lobes curved, from common base with median lobe to oppose median lobe tip; median lobe truncated acute-triangular. Seg-

ment IX in lateral aspect (Fig. 91) parallelogram-like, dorsad of large membranous lobe; ventral portion wide (Fig. 92), or-

ientated vertically under segment X (Fig. 91). Supragenital plate short (Fig. 91), narrow, weakly sclerotised (Fig. 92). Cercus

acute-triangular in lateral aspect. Segment X with deeply separated acuminate dorsal lobes; with scoop-like single ventral lobe.

Notes on habitat and activity period. — The single specimen is another product of the Al-

berta Department of Agriculture light trap, and I can, therefore say little about the probable

Quaest. Ent., 1977 13 (1)

42

Nimmo

habitat. The Three Hills area is in the open plains of Alberta, with scattered bodies of water

(small lakes and sloughs, and occasional small streams). Date of capture was 23/6/76.

The locality of collection of the single female is shown in Fig. 1 1 9, at about 2,800'.

The Genus Asynarchus McLachlan

Asynarchus lapponicus (Zetterstedt), 1 840

(Fig. 85-90, 119)

Phryganea fusca var. b lapponica Zetterstedt, 1840:10. (Type locality: Lapland).

Asynarchus lapponicus', Schmid, 1954: 78-81, Fig. 15-17. Schmid, 1955: 154. Clifford, 1969: 582. Fischer, 1969: 47-50.

Nimmo, 1971a: 132. Fischer, 1973: 111. (See Fischer, 1969: 47-48, and 1973: 111 for palaearctic literature).

Limnephilus lapponicus ; Ross and Merkley, 1952: 443.

Anabolia modesta Hagen, 1861: 265. (Type locality: Labrador). Hagen, 1864: 804. McLachlan, 1876: 9. Banks, 1892: 363.

Banks, 1897: 28. Ulmer, 1905: 20. Ulmer, 1907: 48. Kolbe, 1912: 41. Betten etal, 1934: 354-355. Milne, 1935: 43,49.

Stenophylax modesta ; Banks, 1907: 39.

Anisogamus modestus-, Banks, 1930: 128

Limnephilus modestus', Ross, 1938b: 37, PI. 7 Fig. 65-65A. Ross, 1944: 298.

Asynarchus modestus ; Fischer, 1969: 49. (See Fischer, 1969: 49 for palaearctic literature).

Stenophylax fusorius McLachlan, 1875: 114, 116-117, PI. 12 Fig. 1-3. (Type locality: Lapland). (See Fischer, 1969: 48 for

palaearctic literature).

Anabolia (Asynarchus) fusorius', (See Fischer, 1969: 48 for palaearctic literature).

Anabolia fusorius-, Banks, 1916: 122.

Asynarchus fusorius-, Fischer, 1969: 48-49. (See Fischer, 1969: 48-49 for palaearctic literature).

Limnophilus rhanidophorus Wallengren, 1879: 274-275. (Type locality: Scandinavia).

Asynarchus rhanidophorus; Fischer, 1969: 50.

I reported the presence of this species in the study area in paper I of this series (Nimmo,

1971a: 132), as a brief note. Specimens are still not available from the study area, but I ob-

tained specimens from Baffin Island from which to make the illustrations.

Males are distinguished by segment X without projected median lobes (Fig. 89); and by ven-

tral process of cercus curved smoothly into poster-ventral edge of clasper in lateral aspect. Fe-

males are distinguished by segment X dorsal lobes as vertical, thin plates projected little pos-

terad of segment IX (Fig. 89,90); and by anal tube not cleft in horizontal plane (Fig. 89).

Description. — Antennae red-brown, paler inter-annular sutures; scape, pedicel darker. Vertex of head very dark

red-brown, warts slightly paler. Thorax very dark red-brown overall. Brush of male fore-leg much reduced, of scattered, fine

setae on basal quarter of femur. Spurs yellow. Fore-wing length of male 12.46 mm; deep yellow-brown; costal area much

paler than remainder; very faint irrorations throughout remainder of wing. Hind-wing hyaline with faint reddish tinge. Vena-

tion typical as for Limnephilus spp.

Male genitalia. (Specimen from Lake Harbour, Baffin Island, N.W.T.). Tergum VIII unmodified; posterior border paler

than remainder (Fig. 85). Segment IX large; anterior edge black-bordered; segment longest laterally, smoothly constricted

dorsad to vertical, stout dorsal strap. Clasper not clearly distinguished from ventro-lateral portion of posterior edge of seg-

ment IX; dorsal process in lateral aspect distally cleft to two black, acuminate processes; ventral process directed dorso-mesad

(Fig. 86). Cercus massive, tapered dorso-posterad, with ventral pad-like lobe or tooth (Fig. 85, 86); with disto-mesal tooth

(Fig. 87); entire posterior edge black; dorsal edges of claspers parallel, with opposed straight portions mid-way from base

(Fig. 87). Segment X small, bipartite (Fig. 86), with thin, tenuous mesal bridge; lateral sclerites orientated in line from dorso-

lateral to ventro-mesal. Aedeagus with median shaft curved ventrad then dorsad (Fig. 88), with distinct head mounted below

centre-line of curve; lateral arms curved as median shaft, distally bifid, with dorso-anteriorly directed dorsal lobe; with round-

ed distal lobe projected posterad, with fringe of long setae; dorsal lobe acuminate, with two stout distal spines.

Female genitalia. (Specimen from Lake Harbour, Baffin Island, N.W.T.). Sternum VIII with ventrolateral process concave

(Fig. 89). Vulval scale triangular in ventral aspect (Fig. 90); median lobe slightly longer than lateral ldbes, rectangular; lateral

lobes triangular. Supragenital plate minute, broadly bell-shaped in outline. Segment IX,in lateral aspect,of triangular dorsal

part, massive, irregular ventral part, with indistinct anterior edges; ventral parts fused narrowly ventro-mesally (Fig. 90). Seg-

ment X fused solidly to IX, vertical (Fig. 89), slightly grooved down posterior face (Fig. 90); with anal tube clearly segrega-

ted ventrally, simple.

Notes on habitat. — Clifford (1969) describes the single known Alberta locality as a brown-

water stream of low velocity, with silt, mud, or grit bottom. To judge from the known distri-

bution, this is a species of the northern forest and tundra, probably living in the slower streams,

and muskeg ponds and lakes. Altitude of the single locality is about 2,800'.

Rhyacophilidae and Lirnnephilidae

43

Geographical distribution. — I have three records for this holarctic species in Canada: Nova

Scotia; Alberta; and Baffin Island (Fig. 1 1 9).

I examined only a male and a female of this species, from Baffin Island.

The Genus Lenarchus Martynov

The Subgenus Prolenarchus Schmid

Three subgenera of Lenarchus are now known from the study area with the discovery of

L. (P.) keratus Ross. The following characterisation of the subgenus Pro lenarchus is abridged

in translation from Schmid, 1955: 162.

Fore- wings large, rounded apically; uniformly finely speckled grey; hind- wings little larger

in area. Male segment IX dorsally convex, short, not overhanging appendages; ventrad of dor-

so-posterior border two small, strongly sclerotised, spur-like hooks mesad of cerci, fused to

each other basally. Cerci characteristic, concave on mesal face, uniformly sclerotised. Male

segment X large, projected posterad, directed postero-dorsad; lateral flanges massive, solidly

joined to median processes. Claspers fused to segment IX, not projected, appearing as part of

segment postero-ventral edge. Aedeagus slender, sclerotised; lateral arms slender, distally ex-

panded as concave, setose blade, with small disto-ventral process. Female segment IX short,

of one piece; appendages long, slender, cylindrical. Segment X well developed, with two small

processes dorso-laterad, similar to those of segment IX, but more slender. Supragenital plate

large, horizontal. Vulval scale large, with oblique lateral lobes.

Lenarchus keratus (Ross), 1938

(Fig. 93-97, 117)

Limnephilus keratus Ross, 1938a: 165-166, Fig. 104. (Type locality: Thunder Bay, Ontario). Ross, 1944: 298.

Lenarchus keratus', Ross and Merkley, 1952: 437. Schmid, 1952: 172-173, Fig. 9-10. Schmid, 1955: 162. Etnier, 1968:191.

Fischer, 1969: 68. Fischer, 1973: 115. Roy and Harper, 1975: 1083.

There is only one other species recognised in this subgenus, which is found only in northern

Europe. L. keratus is distinguished immediately from other Lirnnephilidae in the study area

by paired, black horns projected posterad from ventrad of Segment IX dorsal surface (Fig. 93).

Description. — Antennae dark red-brown with basal portion of annuli paler. Vertex of head dark red-brown, warts

paler. Thorax dark red-brown dorsally, paler laterally. Brush of male fore-leg a single line of fine, even hairs except widened

basad. Spurs dark red-brown; formula 1,3,4. Male fore-wing length 15.36 mm; dark chocolate-brown, costal area clear ex-

cept faint patches of colour; remainder finely irrorate. Venation typical as for Limnephilus species.

Male genitalia. (Specimen from about 10 miles south-east of Fairview, Alberta). Segment IX massive, with very long ter-

gal area in lateral aspect (Fig. 93), with narrow sternal area; anterior border lined with black. Clasper triangular in lateral

aspect, dorsal, distal angles rounded; base solidly fused to segment IX; each clasper connected ventrally by small humped

bridge, dorsally by faint sclerotised strap (Fig. 95). Cerci ventrad of postero-dorsal projection of segment IX; short, irregular,

rounded in lateral aspect; in posterior aspect (Fig. 95) concave, with black, well defined ‘wall’ around lateral, dorsal edges;

ventral edge straight, thin-walled. Segment X basal sclerites ventrad of cerci; large, rounded, dorsally concave; median lobes

large, directed dorsad, tip hooked anterad, black; membranous anal lobe meso-ventrad of segment X. A pair of black, slight-

ly divergent, tapered, somewhat blunt horns located under projection of segment IX, dorsad of dorso-mesal angles of cerci

(Fig. 93, 94, 95); curved postero-ventrad. Aedeagus fairly massive, with thick median shaft with distal head partly recessed

into tip (Fig. 97); lateral arms heavy, originated dorso-laterad of aedeagal base, expanded distally to concave, thin-walled

tip; with dorsal edge of expansion with fringe of heavy, black setae (Fig. 96, 97); with stout, long, dentate process origina-

ted from mesal surface.

Female. Unknown, or not yet associated.

Note on habitat. — I am unfamiliar with the single Alberta locality except that it is in the

Peace River outlier of the Prairie, which is heavily agricultural and flat except for the incised

river valleys. The type of water body it inhabits cannot be determined at present. Altitude of

collection point is about 2,000'.

Geographical distribution. — For many years after Ross described it, this species was known

Quaest. Ent., 1977 13 (1)

44

Nimmo

only from the area around Lake Superior. Roy and Harper (1975) recently recorded a male

from southern Quebec. The Alberta record represents a westward extension of the range of

about 1,500 miles (Fig. 117).

I examined a single male of this species.

The Genus Platy centrop us Ulmer

This is the second limnephilid genus recorded in this paper as new to the study area. The

following characterisation is abridged in translation from Schmid, 1955: 164-165.

Head short, eyes large, prominent; cephalic warts very small. Maxillary palpi long, massive;

in male first article half length of second, which is as long as scape. Spur formula 1,3,3; meso-

apical spurs of hind leg modified. Wings large; hind- wings little smaller than fore. Fore- wing

with large brown areas. Venation of fore- wings with large discoidal cell clearly longer than pet-

iole; chord disrupted, concave towards abdomen, roughly parallel to it; cell f5 often pointed;

vein A 2 with basal half absent. Male segment VIII unmodified. Genital capsule generally not

prominent, a rigid, massive assemblage due to conspicuous development of segment IX; dor-

sally segment IX with single process, or two strongly sclerotised points dorsad of remainder

of genitalia; laterally segment IX strongly constricted, considerably enlarged latero-ventrad,

such that distal aperture and appendages directed postero-dorsad. Cerci in most males as mas-

sive, horizontal spurs; thick, heavily sclerotised. Lateral extensions of segment X large, hori-

zontal spurs; thick, heavily sclerotised. Lateral extensions of segment X large, horizontal, of-

ten concave ventrad. Claspers small, not projected, with no free part. Aedeagus not large, me-

dian shaft slender, often wrinkled basally; lateral arms varied, in some males shortly spiniform,

in others long, setiform. Female tergum IX long, stout, continuous with segment X which

has free appendages. Ventral lobes of segment IX small, slender, as short, transverse plates.

Vulval scale with long, thin, median lobe; with narrow, sub qua dr angular lateral lobes. Supra-

genital plate small, short.

Platy centropus plectrus Ross, 1 938

(Fig. 98-101, 120)

Platycentropus plectrus Ross, 1938a: 169-170, Fig. 111. (Type locality: Honor, Michigan). Ross, 1944: 297. Leonard and

Leonard, 1949a: 4, Pl. 3 Fig. 3. Leonard and Leonard, 1949b: 16, Fig. 1. Schmid, 1952: Fig. 3. Schmid, 1955: 166.

Fischer, 1969: 71. Fischer, 1973: 116.

Hylepsyche plectrus-, Banks, 1943: 350. Fischer, 1969: 71.

Males of this species are distinguished by black horns at lateral angles of postero-dorsal edge

of segment IX (Fig. 98, 99, 100); by short, triangular lateral arms fused immovably to base of

aedeagus (Fig. 101).

Description. — Antennae dull yellow-brown; scapes, pedicels paler. Vertex of head yellow-brown, mottled with

darker brown patches. Thorax dark red-brown dorsally, yellow-brown laterally; legs yellow-brown. Brush of male fore-leg

of straight, thin, black spines along basal half of femur mesal face, thinned out distdd. Spurs dark yellow-brown. Hind-leg

meso-apical spur flattened slightly, with fibrous flanges along opposite edges, not quite extended to spur tip. Male fore-wing

length 12.55 mm; pale straw-coloured; with grey-brown areas posterad of M and Ml+2, and R5, except area around basal

half of M2 straw-coloured, and area between Cu2 and A straw-coloured. Hind-wing hyaline, with faint red-brown tinge to

veins. Venation typical as for Limnephilus species, except male fore-wing with A2 very short, extended only to crossvein

al+a2.

Male genitalia. (Specimen from Hartley Ck, east side Athabasca R., opposite Ft. MacKay, Alberta). Tergum VIII with

only sparse patch of long setae parallel to posterior border, set anterad of border (Fig. 98). Segment IX wide ventro-laterally,

narrowly constricted dorso-laterally, markedly widened dorsad (Fig. 98); dorsal strap postero-dorsal edge with black horn at

lateral angles; horns curved slightly mesad, strongly sclerotised (Fig. 98, 99, 100). Segment IX in posterior aspect inverted-

triangular, with dorsal area bulged dorsad; posterior edge between horns sharply declivous, high; anterior edges in lateral as-

pect (Fig. 98) folded meso-posterad for most of height. Clasper short, rectangular in lateral aspect, ventro-laterad on poster-

ior edge of segment IX; angled latero-ventrad in posterior aspect, both connected meso-ventrally by narrow strap (Fig. 99).

Rhyacophilidae and Limnephilidae

45

Cerci small, roughly rectangular in lateral aspect, directed posterad; oriented vertically in posterior aspect (Fig. 99); slightly

concave on mesal face; with membranous base. Median lobes of segment X large, directed postero-laterad, tapered distad,

with tips curved latero-dorsad; each with complexly folded basal sclerite, postero-ventral portion of which is deeply concave.

Aedeagus small, short, stout, with large head telescoped into median shaft; lightly sclerotised; lateral arms reduced to small,

triangular, immobile stubs set dorso-laterally on aedeagalbase (Fig. 101).

Female genitalia. This sex is known for this species, but I was unable to obtain a specimen for illustration and description

in time for this paper. See Leonard and Leonard (1949a: PI. Ill Fig. 3) for an illustration.

Note on habitat. — I am not familiar with the single known study area locality. The single

male from Alberta was taken in an emergence trap set in Hartley Ck, a locality well within the

northern boreal forest. Date of capture was July 16.

Geographical distribution. — Recognition of this species represents a range extension of some-

thing over 1,000 miles to the west. The species was formerly known only from just south of

the Great Lakes (Fig. 1 20). I also indicate the presence of the species in Saskatchewan. This

is based on a male which I identified for D.H. Smith, University of Saskatchewan, in 1 975,

and which he took at some point north of Saskatoon, again in the boreal forest region. Alberta

altitude 950'.

I examined two males from the study area.

The Tribe Stenophylacinae Schmid

The Genus Philocasca Ross

Philo casca alba Nimmo new species

(Fig. 102-105, 121)

This is the second species of Philocasca recorded from the study area.

This species is similar to P. thor Nimmo (1971a: 1 47), but males are distinguished by cerci

angular, not projected beyond base of segment X median lobes, in lateral aspect (Fig. 102);

by segment median lobes sinuate, not projected posterad in smooth curve; and by aedeagal

lateral arms located along dorso-lateral surfaces of median shaft, massive, blade-like (Fig. 105).

Description. — Antennae pale yellow-brown; antero-mesal face of scape glabrous, paler; postero-mesal face white.

Vertex of head pale red-brown, warts cream-coloured. All setae hyaline except black setae in row parallel to poster o-lateral

margin of compound eye. Thorax reddish brown to straw-coloured in parts dorsally, with cream-coloured warts; pale straw-

colour laterally. Spurs red-brown; formula 1,2,4. Male fore-wing length 17.76 mm; milky opaque grey, with faint irrorations

except costal area. Hind-wing uniformly opaque white. Venation little different than for P. thor Nimmo (1971a: Fig. 143 a-b),

except discoidal cell closed distally on hind-wing by cross-vein r2-r3.

Male genitalia. (Specimen from Rowe Bk, 6,350', Waterton National Park, Alberta). Segment IX with broad, angular

basal half tapered irregularly dorsad to substantial dorsal strap (Fig. 102); with black-bordered anterior and postero-dorsal

edges; dorsal portion of segment interior sclerotised as paired, deep, anteriorly concave cups (Fig. 104) separated by rounded

median ridge of medium height. Clasper short, button-like, fused ventrally with opposite clasper (Fig. 103), with dark band

round basal area. Cercus short, angular, concave mesally (Fig. 102, 104); with postero-ventral edge free of segment IX, dor-

sad of membranous area (Fig. 102). Segment X paired sclerites apparently confluent laterally with cerci (Fig. 104); dorso-

median process sinuate, tapered to thin, rounded tip projected well clear of remainder of segment; mesal faces of processes

parallel. Aedeagus stout, with antero-lateral straps attached to sclerotised base (Fig. 105); median shaft short, stout, mostly

membranous with distinct sclerotised head concave dorsally at which point ejaculatory duct opens; lateral arms very dark,

curved dorso-posterad to lie along dorso-lateral surfaces of median shaft; tapered. Antero-dorsad of lateral arms a high, hood-

like expansion of base, with posterior face concave, black.

Female. Unknown, or not yet associated.

Notes on habitat and activity period. — This is a species of higher altitudes, from the sub-

alpine, and probably alpine, regions of the study area. I do not yet know if it frequents stand-

ing or running water areas as larvae; the possibility exists that, like other species, this is terres-

trial as larvae. Dates of capture are May 1 1, May 27, and June 1 2.

Geographical distribution. — Known only from the southern mountains of Alberta (Fig.

1 21), close to the continental divide. Altitudes are 6,350', and 6,500'.

Holotype. — Male. Rowe Bk, 6,350', Waterton National Park, Alberta; 1 2/6/75; D.B. Donald.

Paratypes. — Twin Ck, Marmot Basin, west of Kananaskis R., Alberta, 1 1/5/76; R. Mutch;

Quaest. Ent., 1977 13 (1)

46

Nimmo

two males. Same as preceding; 27/5/75; R. Mutch; three males.

I examined six males of this species. Wings, legs and antennae of the Twin Ck paratypes are

damaged from the time of collection.

The holotype, and Twin Ck(l 1/5/76) paratypes are in the Canadian National Collection,

Ottawa, type series number 1 5, 165. The remaining three paratypes (Twin Ck, 27/5/76) are in

the Department of Entomology, University of Alberta.

This species is named with reference to the overall greyish white colour, especially of the

wings.

The genus Psycho glyp ha Ross

Two changes of name are noted with reference to my 1 971(a) paper, necessitated by Denning

(1970), as follows:

P. alaskensis (Banks); Nimmo, 1971a: 153-154, 212, Fig. 572-577, 659, Tab. 2, 3.

is changed to

P. subborealis (Banks), 1924. The references in Nimmo (1971a: 153) to Platyphylax alascensis

Banks should be deleted as they refer to quite another species and genus. On p. 1 54, the ref-

erences to Psychoglypha alascensis and Psychoglypha alaskensis should be placed after those

to subborealis , as alaskensis and alascensis are now the synonyms.

P ulla (Milne); Nimmo, 1971a: 154, 212, Fig. 578-581, 660, Tab. 2, 3.

is changed to

P. alascensis (Banks), 1900. References under P. ulla (Nimmo, 1971a: 155) are retained on

basis of synonomy, but it is necessary to insert, prior to these, the following earlier refer-

ences under the present name:

Halesus? alascensis Banks, 1900: 471, PI. 28 Fig. 19, 20.

Halesus alascensis', Ulmer, 1905: 21. 1907: 70.

Chilostigma alascensis; Banks, 1907: 40. Banks, 1924: 441. Essig, 1926: 176.

Chilostigma alascense; Ulmer, 1932: 215, 217. Betten et al, 1934: 368. Milne, 1935: 35, 50. Fischer, 1969: 319.

Psychoglypha alascensis; Denning, 1970: 16, 17, 20, 22, Fig. 13, 13A, 14, 14A.

Psychoglypha prita (Milne), 1935

(Fig. 106-107)

Psychoglypha prita; Nimmo, 1971a: 152-153, Fig. 562-565.

Denning (1970) described and illustrated a female of this species. I provided description

and illustrations for the male in my 1971(a) paper, and here provide description and illustra-

tions of the female derived from Denning’s specimen.

Females distinguished by postero-ventral lobe of segment IX projected posterad, ventrad

of segment X (Fig. 1 06); and by lobe ventrad of segment IX indistinguishable from sternum

VIII.

Description. — Habitus of female not significantly different from that of male, except body colouration pale yellow-

ish brown, with no darkening except marginally on thorax.

Female genitalia. (Specimen from Rice Ck two miles above Salmon R., Idaho County, Idaho). Sternum VIII posterior edge

produced posterad as large, rounded lobe in lateral aspect (Fig. 106). Vulval scale in ventral aspect (Fig. 107) massive, lateral

lobes rectangular, with much smaller, acute-triangular median lobe only half length of laterals. Supragenital plate long in

lateral aspect (Fig. 106), with lateral angles curved dorsad, with relatively straight posterior edge (Fig. 107). Segment IX main

body rectangular in lateral aspect with antero-dorsally directed invagination from lower half of antero-lateral edge; lower half

of postero-lateral edge produced as relatively acute-triangular lobes, finger-like in ventral aspect, ventro-laterad of segment

X. Segment X small, tapered, with abrupt taper disto-ventrad; arched dorsad, with wide, trapezoidal cleft in distal edge, in

ventral aspect; with prominent (Fig. 106), rounded lobe ventrad, between posterior lobes of segment IX.

Rhyacophilidae and Limnephilidae

47

Psychoglypha alascensis (Banks), 1 900

(Fig. 108-109)

Psychoglypha ulla ; Nimmo, 1971a: 155, Fig. 578-581.

The female was illustrated and described previously. Here I provide description and illus-

trations of a female from the study area taken in 1976. The male was described in my 1971(a)

paper.

Females distinguished by massive postero-lateral lobes of segment IX projected posterad

of remainder of genitalia; and by segment X almost totally enclosed by them in lateral aspect

(Fig. 108).

Description. — Habitus not significantly different from that of male.

Female genitalia. (Specimen from Coppermine Ck, Red Rock Canyon Rd, Waterton National Park, Alberta). Sternum VIII

projected posterad of tergum, ventrad of segment IX (Fig. 108). Vulval scale lateral lobes roughly trapezoidal, large, poster-

ior edge angled postero-laterad, with minute, triangular median lobe (Fig. 109). Segment IX large, of one peice as roof-like

structure over remainder of genitalia, with postero-lateral angles produced posterad to flank segment X; these lobes slightly

demarcated from body of segment IX by shallow lateral declivity, abruptly narrowed dorsad to beak-like, tapered termina-

tion in lateral aspect (Fig. 108). Segment X ventrad of IX, trapezoidal in lateral aspect, concave ventrally, with large, semi-

circular cleft in ventral aspect. Supragenital plate lunate in ventral aspect, prominent in lateral aspect, dominated by large

fleshy lobe dorsad.

Placement of Genera and Species

of Limnephilidae new to Study Area, in Keys

previously constructed

The Genus Dicosmoecus McLachlan

Male. Refer to key on p. 51 of Nimmo, 1971a. Add

lc. Mesal ridge of clasper base in posterior aspect not vertical, with clearly

visible ventral ridge and dorso-lateral ridge irregular, thin-walled, high,

black (Fig. 43) D. gilvipes (Hagen).

Female. Refer to key on p. 52 of Nimmo, 1 971 a. Add

1 c. Median lobe of vulval scale less than half length of laterals (Fig. 46),

expanded disto-laterad, the projections housed in large concavities of

lateral lobe mesal face; lateral lobes massive, fused dorso-mesally.

Segment X short, triangular in lateral aspect (Fig. 45), shallowly cleft

in ventral aspect (Fig. 46), with ventro-mesal lobes near junction with

segment IX, directed mesad D. gilvipes (Hagen).

The Genus Imania Martynov

Male. Refer to key on p. 56 of Nimmo, 1971a.

I. thomasi Nimmo, n. sp. keys to 2a wherein it may be distinguished by aedeagal

base with second pair of processes basad of lateral arms, set dorsad, long, tapered,

black (Fig. 58).

The Genus Neophylax McLachlan

Refer to key to genera of Neophylacinae in study area on p. 71 of Nimmo, 1971a.

This key encompases both sexes. Add

lc. Segment X of male of two or three pairs of elongated appendages (Fig. 47).

Claspers projected, not telescoped into IX. Female segment IX lobes elon-

gate, fused to lateral borders of vulval scale (Fig. 53)

Neophylax McLachlan (N. rickeri Milne).

Quaest. Ent., 1977 13 (1)

48

Nimmo

Males.

Females.

Male.

Female,

Males.

Male.

The Genus Limnephilus Leach

Refer to key on p. 83 of Nimmo, 1971a.

L. nimmoi Roy and Harper keys to 22a wherein it may be distinguished by mesal

faces of cerci heavily toothed (Fig. 65, 66).

L. insularis Schmid keys to 25b wherein it may be distinguished by cercus long, tri-

angular in lateral aspect (Fig. 59), with segment X median lobes long, tapered, acum-

inate, directed postero-dorsad.

L. alvatus Denning keys to 26 wherein it may be distinguished by aedeagal lateral

arms as long as median shaft, sclerotised, with minute tuft of setae at tip; clasper

triangular, vertically high (Fig. 71, 74).

L. vernalis Nimmo n. sp. keys to 1 5b wherein it may be distinguished by clasper

trapezoidal in lateral aspect (Fig. 80). Also, segment X median lobes projected pos-

terad of clasper extremity; segment IX spindle-shaped in lateral outline.

Refer to key on p. 86 of Nimmo, 1971a.

L. nimmoi Roy and Harper keys to 27a, where it is designated L. species 1 (Fig. 69,

70, this paper).

L. alvatus Denning keys to 30a, wherein it may be distinguished by lateral lobes of

segment X projected dorsad of segment in lateral aspect (Fig. 77).

L. species 2 keys to 9b wherein it may be distinguished by segment X dorsal lobes

visually discrete from dorso-lateral lobes, acuminate (Fig. 91). Segment IX ventral

junction vertical in lateral aspect (Fig. 91).

The Genus Asynarchus McLachlan

Refer to key on p. 1 29 of Nimmo, 1971a.

A. lapponicus (Zetterstedt) keys to lb wherein it may be distinguished by segment

X median lobes not evident in lateral aspect (Fig. 85), completely enclosed in seg-

ment IX

Refer to key on p. 1 29 of Nimmo, 1971a.

A. lapponicus (Zetterstedt) keys to lb wherein it may be distinguished by segment

X vertical in lateral aspect (Fig. 89), of two parallel ridges (Fig. 90).

The Genus Lenarchus Martynov

The Subgenus Prolenarchus Schmid

Refer to key to genera and subgenera of Limnephilini on p. 83 of Nimmo, 1971a.

This key refers primarily to males. This subgenus keys to 7 wherein it may be dis-

tinguished by segment IX dorsal strap produced only slightly posterad; with paired,

black, curved hooks ventrad of edge (Fig. 93).

Refer to key on p. 135 of Nimmo, 1971a.

L. keratus Ross keys to 1 wherein it may be distinguished by no dorsal plate present

— postero-dorsal edge of segment IX produced only slightly posterad (Fig. 93). Ven-

trad of segment IX postero-dorsal edge a pair of curved spines or hooks, black, stout,

slightly joined basad, projected posterad (Fig. 93, 94).

The Genus Platycentropus Ulmer

Males.

Refer to key to genera of Limnephilini in study area on p. 83 of Nimmo, 1971a.

Rhyacophilidae and Limnephilidae

49

This key refers primarily to males.

This genus keys to 2 wherein it may be distinguished by meso-apical spur on hind-

leg modified; flattened, with thin-walled flanges along edges, not quite to tip of

spur.

P. plectrus Ross is the only species known from the study area.

The Genus Philocasca Ross

Males. la. Cerci almost as long as segment X median lobes, smoothly rounded

distally (Fig. 545 (Nimmo, 1971a)); lateral arms of aedeagus narrow,

laterad of median shaft (Fig. 547 (Nimmo, 1971a))

P. thor Nimmo.

lb. Cerci short, angular; segment X median lobes sinuate, projected well

posterad of cerci (Fig. 102); lateral arms of aedeagus stout, blade-like,

curved dorso-posterad along dorso-lateral faces of median shaft

(Fig. 105, this paper) P. alba Nimmo n. sp.

The Genus Psy choglypha Ross

Females. My original key (Nimmo, 1971a: 1 52) was to females of only two species. It is

inadequate to provide separation for the four now known. Therefore, I here pre-

sent a new key.

la. Vulval scale lateral lobes posterior edges in line with each other

(Fig. 571 (Nimmo, 1971a)) (Fig. 107, this paper) 2

1 b. Vulval scale lateral lobes posterior edges irregular, not in line

(Fig. 577 (Nimmo, 1971a)) (Fig. 109, this paper) 3

2a. (la) Segment IX postero-lateral edges with small, rounded process; seg-

ment X mostly recessed into IX (Fig. 570 (Nimmo, 1971a))

P. schmidi Nimmo.

2b. Segment IX postero-lateral edges with long, triangular process;

segment X mostly projected clear of IX (Fig. 106, this paper)

P. prita (Milne).

3a. (lb) Segment X dorsad of segment IX postero-lateral lobes, which

are short, triangular, rounded (Fig. 576 (Nimmo, 1971a)) ....

P. subborealis ((=alaskensis)) (Banks).

3b. Segment X ventrad of segment IX postero-lateral lobes, almost

completely enclosed by them (Fig. 108, this paper); Segment IX

lobes massive, with beak-like dorsal extension

P. alascensis (Banks) (( =ulla (Milne))).

POST-GLACIAL ORIGINS, AND AFFINITIES OF THE FAUNA

As this paper presents oddments additional to the fauna recorded previously, I refrain from

examining pre-Wisconsin affinities of the groups represented here.

Table 1 presents visually the known study area altitudinal distribution of the species recor-

ded herein. This may be compared with the similar table for these two families in my previous

paper (Nimmo, 1971a: Tab. 3). I refrain from drawing conclusions at this time as the mater-

ial on which these distributions are based is, at best, scanty. Also, I hope, in the near future,

to carry out a more detailed study along these lines.

Quaest. Ent., 1977 13 (1)

50

Nimmo

Table 1. Altitudinal distribution of the species recorded in this paper, in the study area,

based on adult records.

Species

Limnephilus alvatus

Platycentropus plectrus

Lenar chus keratus

Limnephilus vernalis

Rhyacophila vao

Limnephilus sp. 2

Asynarchus lapponicus

Rhyacophila unimaculata

Neophylax rickeri

Rhyacophila milnei

Limnephilus insularis

Dicosmoecus gilvipes

Rhyacophila sp. 3

Rhyacophila sp. 4

Rhyacophila donaldi

Imania thomasi

Philo casca alba

Rhyacophila autumnalis

Rhyacophila simplex

Altitude

Range

Pattern

Post-Glacial origins of the Species recorded, relative to Study Area

Nimmo, (1971a: 201-204) presented a list of 12 range patterns exhibited by study area

species, and a list of postulated post-glacial dispersal routes (p. 218). Relating the species

dealt with here, to these lists, produces the following table:

Table 2. Species of Rhyacophilidae and Limnephilidae recorded in this paper from the

study area, listed under range pattern to which each is assigned, and giving postu-

lated dispersal routes.

Rhyacophilidae and Limnephilidae

51

Most of the probable dispersal routes assigned in table 2 are tentative, and the best that can

be assigned in the circumstances. Apart from species here described as new, most of the re-

mainder are too poorly known.

Dicosmoecus gilvipes, Neophylax rickeri, Limnephilus alvatus, and Platy centro pus plectrus

are the most certain, and require no further comment.

Rhyacophila vao is uncertain as either route ‘IT or V could be involved, although I favour

the route (‘a’) assigned.

Asynarchus lapponicus presents difficulties as, in North America it appears to belong to

range pattern 1 2, but the species as a whole is holarctic. It may be that the species was split

during the glaciation, with one part south of the ice in North America. It appears to be, from

the isolated North American records, a species of the boreal forest and tundra.

Limnephilus nimmoi is assigned to range 1 1 primarily for convenience at this time. Probab-

ly it exists, unrecognised, in collections under different names as it is easily confused with

others of its species group. It may be northern boreal as suggested by Roy and Harper (1975),

and belong to range pattern 8, or even 7.

Limnephilus vernalis cannot be placed as yet.

Range pattern 6 is primarily a catch-all category for those species known only from the

study area, and immediately adjacent areas to the south and west, but which are poorly known.

Many included species are isolated, high altitude forms and, thus, poorly collected. Rhyacoph-

ila unimaculata is peculiar in appearing to follow the Rocky Mountain Trench. Ross’ (1965)

conjecture that this species is the product of a glacial refugium in the Mt. Robson area now

appears to be confounded — not only on the basis of its presently known range, but also due

to the low altitudes which it appears to favour.

However, accepting table 2 at face value, and recognising the small number of species in-

volved, the following percentages of species from each source area prevail:

These figures are close to those determined in my 1 971(a) paper, which provides the follow-

ing figures for the known fauna of Rhyacophilidae and Limnephilidae, of the study area. The

Quaest. Ent., 1977 13 (1)

52

Nimmo

1971 figures are given first, followed by the present total (for space reasons figures are pre-

sented thus — species numbers: %):

* (1) refers to Limnephilus sp. 1 (Nimmo, 1971a), now recognised as L. nimmoi Roy and

Harper in this paper. The former total for 1971(a) was 66. The percentage value is left as ori-

ginally determined.

Finally, taking the figures from my 1 974 paper, on the Glossosomatidae and Philpotamidae

of the study area, we obtain the following tabulation, which includes the percentage values

calculated for the four families so far worked up for the study area; values from the 1 974 pap-

er are given first, followed by cumulated values for the four families:

From Cordillera south/west of study area —

From Alaska —

From central plains

From eastern North America

From North America as a whole, S. of ice —

Indeterminate

While most values are seen to oscillate about those determined in 1 971(a), the notable

changes with additional information are the decrease in proportion of the fauna attributable

to the Cordillera south of the ice, and the increase in proportion attributable to transcontin-

ental species. This trend may be expected to continue with the addition of information from

families remaining to be dealt with, as they are, by and large, concentrated east of the Cordil-

lera.

CONCLUSIONS

1. Compared with data in Nimmo (1971a), proportions of species, recorded in this paper,

derived from various source areas post-glacially do not differ drastically, except that there

are proportionately fewer derived from eastern North America.

2. Combining species numbers derived from these source areas, from the earlier paper, and

this, results in little change in relative proportions except that relatively fewer are derived

from the Cordillera south of the ice, and more are derived from eastern North America.

3. Addition of species of Glossosomatidae and Philopotamidae, considered in another pap-

er produces much the same results, with accentuation of decrease in Cordilleran species, and

increase of eastern North American species.

4. It is expected that this trend will continue with further work on other, as yet untreated,

families in the study area, which are predominently eastern (east of the Cordillera).

ACKNOWLEDGEMENTS

I am inestimably grateful to G.E. Ball, Chairman, and to the Department of Entomology,

University of Alberta generally, for allowing me space and facilities wherewith to continue

this work. Publication costs were met by funds from NRC grant A- 13 99, held by G.E. Ball,

Rhyacophilidae and Limnephilidae

53

and from the Strickland Fund of this Department. G.E. Ball read and commented upon the

manuscript.

Little of the material recorded here was collected by myself and I am happy to acknowled-

ge my debt to the following persons who contributed specimens and records as a result of

their own work: D.B. Donald, Canadian Wildlife Service, Biology Department, University of

Calgary; R. Mutch and R. Crowther, graduate students in the same department; and H.G. Philip,

Alberta Agriculture, Edmonton.

No small measure of praise and gratitude is due to my wife, Susan, for her patience with

my constant absences on collecting trips, or in the office on evenings and weekends.

Many thanks are due to D.G. Denning, California, F. Schmid, Ottawa, and O.S. Flint, Jr.,

Washington, D.C., for making available specimens required for this study.

I wish to thank, also, the following institutions for loan of specimens: Dept, des Sciences

biologiques, Universite de Montreal; Entomology Department, U.C.L.A., Davis, California;

Department of Entomology, University of Minnesota, St. Paul.

G.C.D. Griffiths of this department assisted with latinisation of new species names.

J.E.H. Martin, Canadian National Collection, Ottawa, provided labels for type material,

and the C.N.C. type series numbers recorded herein.

Western Region, Parks Canada, kindly continues to issue permits to collect in the western

National Parks.

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new species. Entomological News 79: 188-192.

Fischer, F.C.J. 1960. Trichopterorum Catalogus: 1, Necrotauliidae, Prosepididontidae, Rhy-

acophilidae. iv + 168 pp. Nederlandsche Entomologische Vereeniging, Amsterdam.

Fischer, F.C.J. 1967. Trichopterorum Catalogus: 8, Goeridae, Limnephilidae pars 1. iv + 263

pp. Nederlandsche Entomologische Vereeniging, Amsterdam.

Fischer, F.C.J. 1968. Trichopterorum Catalogus: 9, Limnephilidae pars 2. iii + 363 pp. Neder-

landsche Entomologische Vereeniging, Amsterdam.

Fischer, F.C.J. 1 969. Trichopterorum Catalogus: 10, Limnephilidae pars 3. iii + 332 pp. Ned-

erlandsche Entomologische Vereeniging, Amsterdam.

Fischer, F.C.J. 1971. Trichopterorum Catalogus: 12, Supplement — Micropterygidae to Sten-

opsychidae. vii + 3 1 1 pp. Nederlandsche Entomologische Vereeniging, Amsterdam.

Fischer, F.C.J. 1973. Trichopterorum Catalogus: 1 5, Supplement — Goeridae to Helicopsychi-

dae. vii + 1 66 pp. Nederlandsche Entomologische Vereeniging, Amsterdam.

Flint, O.S., Jr. 1960. Taxonomy and biology of nearctic limnephilid larvae (Trichoptera),

with special reference to species in eastern United States. Entomologica Americana 40: 1-117.

Flint, O.S., Jr. 1966. Notes on certain nearctic Trichoptera in the Museum of Comparative

Zoology. Proceedings United States National Museum 1 18: 373-390.

Hagen, HA. 1861. Synopsis of the Neuroptera of North America, with a list of the South Am-

erican species. Smithsonian Miscellaneous Collections, Washington, D.C. 347 pp.

Hagen, H.A. 1 864. Phryganidorum synopsis synonymica. Verhandlungen Zoologisch-botanis-

chen Gesellschaft, Vienna 14: 794-890.

Hagen, H.A. 1875. Report on the Pseudo-neuroptera collected by Lieut. W.L. Carpenter in

1 873 in Colorado. United States Geological Survey Territorial Report for 1 873, pp. 571-606.

Kolbe, H.J. 191 2. Glazialzeitliche Reliktenfauna im Hohen Norden. Deutsche Entomologische

Zeitschrift 7: 33-63.

Leonard, J.W. & F.A. Leonard. 1949a. Noteworthy records of Caddis Flies from Michigan,

with descriptions of new species. Occasional Papers Museum of Zoology, University of

Michigan 520: 1-8, 5 plates.

Rhyacophilidae and Limnephilidae

55

Leonard, J.W. & F.A. Leonard. 1949b. An annotated list of Michigan Trichoptera. Occasional

Papers Museum of Zoology, University of Michigan 522: 1-35.

Ling, S-W. 1938. A few new Caddis Flies in the collections of the California Academy of Sci-

ences. Pan-Pacific Entomologist 14: 59-69.

McLachlan, R. 1875. Monographic revision and synopsis of the Trichoptera of the European

fauna. 3: 109-144, 4 plates. London.

McLachlan, R. 1 876. Monographic revision and synopsis of the Trichoptera of the European

fauna. 5 (& Supplement, Part 1) 221-280, i- xiii, 8 plates.

Milne, L.J. 1935. Studies in North American Trichoptera. 2: 20-55. Cambridge, Mass.

Milne, L.J. 1936. Studies in North American Trichoptera. 3: 56-128, 2 plates. Cambridge, Mass.

Navas, L. 1926. Trichoptera, Megaloptera und Neuroptera aus dem Deutschen Entomologischen

Institut (Berlin-Dahlem). II. Series Entomologischen Mitteilungen 1 5: 57-63, 1 plate.

Neave, F. 1 929. Report on the Jasper Park lakes investigations 1 925-1926, IV. Aquatic

insects. Contributions to Canadian Biology and Fisheries 4: 1 87-195.

Newell, R. L. & D.S. Potter. 1973. Distribution of some Montana Caddisflies (Trichoptera).

Proceedings Montana Academy of Sciences 33: 1 2-21.

Nimmo, A.P. 1971a. The adult Rhyacophilidae and Limnephilidae (Trichoptera) of Alberta

and eastern British Columbia, and their post-glacial origin. Quaestiones Entomologicae 7:

3-234.

Nimmo, A.P. 1971b. The adult Rhyacophilidae and Limnephilidae (Trichoptera) of Alberta

and eastern British Columbia, and their post-glacial origin: Corrigenda. A. Nimmo 1971.

Quaestiones Entomologicae 7: 406.

Nimmo, A.P. 1974. The adult Trichoptera (Insecta) of Alberta and eastern British Columbia,

and their post-glacial origins. II. The families Glossosomatidae and Philopotamidae. Quaest-

iones Entomologicae 10: 315-349.

Putnam, J.D. 1 876. Report on the insects collected in the vicinity of Spring Lake Villa, Utah

Co., Utah, during the summer of 1 875. Davenport Academy of Sciences 1 : 1 93-205.

Ross, H.H. 1938a. Descriptions of nearctic Caddis Flies. Bulletin Illinois Natural History Sur-

vey, Urbana 21: 101-183.

Ross, H.H. 1938b. Lectotypes of North American Caddis Flies in the Museum of Comparative

Zoology. Psyche 45: 1-61.

Ross, H.H. 1944. The Caddis Flies, or Trichoptera, of Illinois. Bulletin Illinois Natural History

Survey, Urbana 23: 1-326.

Ross, H.H. 1956. Evolution and classification of the mountain Caddisflies. vii + 213 pp. Un-

iversity of Illinois Press, Urbana.

Ross, H.H. 1965. Pleistocene events and insects, pp. 583-595. In: The Quaternary of the Uni-

ted States, H.E. Wright Jr. & D.G. Frey, eds.

Ross, H.H. & D.R. Merkley. 1 952. An annotated key to the nearctic males of Limnephilus

(Trichoptera, Limnephilidae). American Midland Naturalist 47: 435-455.

Ross, H.H. & G.J. Spencer. 1952. A preliminary list of the Trichoptera of British Columbia.

Proceedings Entomological Society of British Columbia (1951) 48: 43-51.

Roy, D. & P.P. Harper. 1975. Nouvelles mentions de Trichopteres du Quebec et description

de Limnephilus nimmoi sp. nov. (Limnephilidae). Canadian Journal of Zoology 53: 1080-

1088.

Schmid, F. 1950. Trois nouveaux Limnophilides (Trichopt.). Mitteilungen Schweizerischen

Entomologischen Gesellschaft 22: 47-54.

Schmid, F. 1952. Le groupe de Lenarchus Mart. (Trichopt. Limnophil.). Mitteilungen Schweiz-

erischen Entomologischen Gesellschaft 25: 1 57-210.

Schmid, F. 1954. Le genre Asynarchus McL. (Trichopt. Limnoph.). Mitteilungen Schweizer-

ischen Entomologischen Gesellschaft 27: 57-96.

Quaest. Ent., 1977 13 (1)

56

Nimmo

Schmid, F. 1955. Contribution a l’etude les Limnophilidae (Trichoptera). Mitteilungen Sch-

weizerischen Entomologischen Gesellschaft 28: 1-245.

Schmid, F. 1970. Le genre Rhyacophila et la famille des Rhyacophilidae (Trichoptera). Mem-

oirs Entomological Society of Canada 66: 3-230, 52 plates.

Schmid, F. 1974. Un Rhyacophila nearctique meconnu (Trichoptera, Rhyacophilidae). Le

Naturaliste Canadienne 101: 933-934.

Schmid, F. & R. Guppy. 1952. An annotated list of Trichoptera collected on southern Van-

couver Island. Proceedings Entomological Society of British Columbia (1951) 48: 41-42.

Smith, S.D. 1965. Distributional and biological records of Idaho Caddisflies (Trichoptera).

Entomological News 76: 242-245.

Smith, S.D. 1968. The Rhyacophila of the Salmon River drainage of Idaho with special ref-

erence to larvae. Annals Entomological Society of America 61 : 655-674.

Stainer, J. 1937. Some observations on a collection of Trichoptera from Lapland, with a note

on the synonymy of Asynarchus modestus Hagen. Entomologist 70: 139-141.

Ulmer, G. 1905. Uber die geographische Verbreitung der Trichoptera. Zeitschrift Wissenscha-

ften Insekten-Biologie 1: 16-32.

Ulmer, G. 1907. Trichoptera. Genera Insectorum 60: 259 pp, 41 plates.

Ulmer, G. 1932. Die Trichopteren, Ephemeropteren und Plecopteren der arktischen Gebietes.

Fauna Arctica 6: 207-226.

Wallengren, H.D.J. 1 879. Description of a new species of Trichoptera from Skandinavia. Ento-

mologist’s Monthly Magazine 15: 274-275.

Zetterstedt, J.W. 1840. Insecta Lapponica, Neuroptera. 5: 1025-1074. Leipzig.

Rhyacophilidae and Limnephilidae 57

1971), or Fig. lc (Nimmo, 1974).

Quaest. Ent., 1977 13 (1)

58

Nimmo

5MM

5MM

A 4

A 5

Fig. 2-9. Fore (a) and hind (b) wings of: 2. Rhyacophila simplex Nimmo n. sp. (male). 3. R. donaldi Nimmo n. sp. (male).

4. R. donaldi Nimmo n. sp. (female). 5. R. autummlis Nimmo n. sp. (male). 6. R. species 3 (female). 7. R. species 4 (female).

8. Imania thomasi Nimmo n. sp. (male). 9. Neophylax rickeri Milne (Male).

Rhyacophilidae and Limnephilidae

59

Fig. 10-25. Genitalia of: Rhyacophila simplex Nimmo n. sp. 10. Male, lat. aspect. 11. Seg. X, post, aspect. 12. Seg. X, dors,

aspect. 13. Aedeagus, lat. aspect. R. donaldi Nimmo n. sp. 14. Male, lat. aspect. 15. Dors, aspect (Seg. IX, X). 16. Seg. X,

anal sclerite, lat. aspect. 17. Anal sclerite, post, aspect. 18. Aedeagus, lat. aspect. 19. Female Seg. VIII, lat. aspect. 20. Seg.

VIII, dors, aspect. 21. Spermathecal sclerite, lat. aspect. 22. Ventr. aspect. R. vao Milne 23. Male, lat. aspect. 24. Aedeagus,

lat. aspect. 25. Female Seg. VIII, lat. aspect.

Quaest. Ent., 1977 13 (1)

60

Nimmo

pect. 28. Seg. IX, X, dors, aspect. 29. Aedeagus, lat. aspect. R. unimaculata Denning. 30. Male, lat. aspect. 31. Seg. X dors,

lobe, dors, aspect. 32. Aedeagus, lat. aspect. R. milnei Ross 33. Female Seg. VIII, lat. aspect. 34. Spermathecal sclerite, ventr.

aspect. R. species 3. 35. Female Seg. VIII, lat. aspect. 36. Spermathecal sclerite, lat. aspect. 37. Dors, aspect. R. species 4.

38. Female Seg. VIII, lat. aspect. 39. Seg. VIII post, edge, ventr. aspect. 40. Spermathecal sclerite, dors, aspect. 41. Lat.

aspect.

Rhyacophilidae and Limnephilidae

61

Fig. 42-55. Genitalia of: Dicosmoecus gilvipes (Hagen). 42. Male, lat. aspect. 43. Clasper basal seg. mesal face, post, aspect.

44. Aedeagus, lat. aspect. 45. Female, lat. aspect. 46. Ventr. aspect. Neophylax rickeri Milne. 47. Male, lat. aspect. 48. Dors,

aspect. 49. Ventr. aspect. 50. Post, aspect. 51. Aedeagus, lat. aspect. 52. Sternum VII post, border, ventr. aspect. 53. Female,

ventr. aspect. 54. Lat. aspect. 55. Male hind-tibia, distal spurs.

Quaest. Ent., 1977 13 (1)

62

Nimmo

Fig. 56-70. Genitalia of: Imania thomasi Nimmo n. sp. 56, Male, lat. aspect. 57. Ventr. aspect. 58. Aedeagus, lat. aspect.

Limnephilus insularis Schmid. 59. Male, lat. aspect. 60. Post, aspect. 61. Dors, aspect. 62. Aedeagus, lat. aspect. 63. Dors,

aspect (part.). L. nimmoi Roy and Harper. 64. Male, lat. aspect. 65. Post, aspect (part.). 66. Dors, aspect, (part.). 67. Aedea-

gus, lat. aspect. 68. Dors, aspect (part.). 69. Female, lat. aspect. 70. Ventr. aspect.

Rhyacophilidae and Limnephilidae

63

Fig. 71-84. Genitalia of: Limnephilus alvatus Denning. 71. Male, lat. aspect. 72. Dors, aspect, (part.). 73. Post, aspect. 74.

Aedeagus, lat. aspect. 75. Dors, aspect. 76. Female, lat. aspect. 77. Ventr. aspect. L. janus Ross. 78. Female, lat. aspect. 79.

Ventr. aspect. L. vernalis Nimmo n. sp. 80. Male, lat. aspect. 81. Post, aspect. 82. Dors, aspect, (part.). 83. Aedeagus, lat.

aspect. 84. Dors, aspect.

Quaest. Ent., 1977 13 (1)

64

Nimmo

Fig. 85-97. Genitalia of: Asynarchus lapponicus (Zett.). 85. Male, lat. aspect. 86. Post, aspect. 87. Dors, aspect. 88. Aedeagus,

lat. aspect. 89. Female, lat. aspect. 90. Ventr. aspect. L. species 2. 91. Female, lat. aspect. 92. Ventr. aspect. Lenarchus ker-

atus Ross. 93. Male, lat. aspect. 94. Dors, aspect. 95. Post, aspect (part.). 96. Aedeagus, dors, aspect. 97. Lat. aspect.

Rhyacophilidae and Limnephilidae

65

Fig. 98-109. Genitalia of: Platycentropus plectrus Ross. 98. Male, lat. aspect. 99. Post, aspect. 100. Dors, aspect. 101. Aedea-

gus, lat. aspect. Philocasca alba Nimmo n. sp. 102. Male, lat. aspect. 103. Ventr. aspect. 104. Dors, aspect. 105. Aedeagus,

lat. aspect. Psychoglypha prita (Milne). 106. Female, lat. aspect. 107. Ventr. aspect. P. alascensis (Banks). 108. Female, lat.

aspect. 109. Ventr. aspect.

Quaest. Ent., 1977 13 (1)

66

Nimmo

Fig. 110-115. Maps of distribution in Alberta & North America of: 110. Rhyacophila vao Milne. 111. R. simplex Nimmo

n. sp., R. unimaculata Denning. 112. R. milnei Ross, R. autumnalis Nimmo n. sp. 113. R. species 3, R. species 4. 114. R.

donaldi Nimmo n. sp., Dicosmoecus gilvipes (Hagen). 115. Imania thomasi Nimmo n. sp., Neophylax rickeri Milne.

Rhyacophilidae and Limnephilidae

67

Fig. 116-121. Maps of distribution in Alberta & North America of: 116. Limnephilus nimmoi Roy and Harper. 117. L.

insularis Schmid, Lenarchus keratus Ross. 118. Limnephilus vernalis Nimmo n. sp., L. alvatus Denning. 1 19. L. species 2,

Asynarchus lapponicus (Zett.). 120. Platycentropus plectrus Ross. 121. Philocasca alba Nimmo n. sp.

Quaest. Ent., 1977 13 (1)

-

THE ADULT TRICHOPTERA (INSECTA) OF ALBERTA

AND EASTERN BRITISH COLUMBIA, AND THEIR POST-GLACIAL

ORIGINS. II. THE FAMILIES GLOSSOSOMATIDAE AND PHILOPOTAMIDAE.

SUPPLEMENT 1.

ANDREW P. NIMMO

Department of Entomology

University of Alberta Quaestiones Entomologicae

Edmonton, Alberta T6G 2E3 13: 69-71 1977

Dolophilodes nora Nimmo (Philopotamidae) is described as new, is recorded from Bertha

Brook, Waterton National Park, Alberta 6,000' , and post-glacial origin of the Alberta popula-

tion is briefly considered. This species is assumed to have entered the study area from imme-

diately south of the cordilleran ice sheet.

Nous ddcrivons une nouvelle Dolophilodes nora Nimmo (Philopotamidae) , localite-type: Bertha Brook, 6,000', Parc

National de Waterton, Alberta). Nous supposons que cette espece dans Cette region soit venue du sud de la masse glaciaire

des Rocheuses.

Corrigenda to Nimmo, 1974

The species Dolophilodes aequalis (Banks) and D. novusamericanus (Ling) are members of

subgenus Dolophilodes, not of subgenus Sortosa Ulmer. The name Dolophilodes should be

substituted for Sortosa as follows: p. 316, Table 1 ; p. 330, lines 3 and 12 from bottom; p. 335,

Table 2; and p. 336, line 8 from top.

INTRODUCTION

Previous publications in this series — Nimmo, 1971, 1974, 1977 — should be consulted.

Normally averse to publication of very short papers, I publish this for three reasons: as

part of an already established series; two additional years of collecting have failed to disclose

more than this one species as new to the study area fauna; and the species described here is

new.

The Family Philopotamidae Stephens

The Genus Dolophilodes Navas

The Subgenus Dolophilodes Ulmer

Dolophilodes nora Nimmo, new species

(Fig. 1-4)

Males of this species most resemble those of D. (D.) pallidipes (Banks), but may be distin-

guished by clasper basal segment base parallel, close to segment IX, ventro-lateral edge (Fig.

1); by segment IX, ventral edge in lateral aspect only slightly distinguishable from almost con-

tinuous postero-ventral curve of lateral walls; and by clasper, distal segment with slight down-

ward curve distally, of uniform width throughout, except rounded distally.

Description. — Antennae pale yellow-brown. Vertex of head yellow-brown, warts paler. Thorax yellow-brown dor-

sally, warts paler; pale yellow-brown laterally. Spurs dark brown; formula 2,4,4. Fore-wing length of male 7.84 mm; opaque

grey-brown. Hind-wing opaque white with tinge of red-brown distally. Male fore-wing lacking bifurcation of R2+3.

Male genitalia. (Specimen from Bertha Brook, 6,000', Waterton National Park, Alberta). Segment IX large, bowed an-

terad in lateral aspect (Fig. 1); anterior edges bordered by thin black line; sternal area long; tergal area non-existant (Fig. 2).

70

Nimmo

Clasper massive, very long, high, directed postero-dorsad; both segments of almost equal height at junction, distal segment

tapered only distally to broadly rounded tip; distal segment slightly concave on mesal face. Cerci projected beyond postero-

dorsal angle of segment IX, rounded distally, with broad base extended to anterior edge of segment IX; shielded laterally by

dorso-lateral wall of segment IX. Segment X small, deeply, finely cleft mesally (Fig. 2); each lobe broad, blunt; in lateral as-

pect tapered distad, slightly bent ventrad. Aedeagus short, totally membranous, with long, bilobed sclerite on left side (rela-

tive to anterior end of insect) (Fig. 3); dorsal lobe short, rounded; posterior lobe long, slender, acuminate.

Female. Unknown.

Note on habitat and flight season. — The single known locality is close to the sub-alpine

zone. A small creek draining Bertha Lake, flowing over small pebble and boulder bottom; ov-

erhung by willow. The specimen was taken on August 30.

Holotype. — Male. Bertha Brook, 6,000', Waterton National Park, Alberta; 30/8/75; D.B.

Donald. Locality shown on Fig. 4.

The holotype is in the Canadian National Collection, Ottawa, with type number 15,166.

This species is named for my elder daughter, Alice Nora.

Males of this species key to couplet 2 (Nimmo, 1974: 330), wherein they are distinguished

by clasper with distal segment curved slightly postero-ventrad distally (Fig. 1); by distal seg-

ment with mesal face membranous basally; and by base of segment X not visible anterad of

tergal area of segment IX in lateral aspect.

POST-GLACIAL ORIGIN

According to my scheme of range patterns exhibited by the study area fauna, and postula-

ted dispersal routes into the study area post-glacially (Nimmo, 1971 : 201, 218), this species

belongs to range pattern 6, and dispersal route d was used. Such conclusions, based on mini-

mal data, are tentative. Further data may exist in existing collections for the reason that D.

nora males are very similar to those of D. pallidipes (Banks), and could easily be confused

with males of D. pallidipes. Discovery of this species does not alter significantly my previous

conclusions (Nimmo, 1974: 338).

ACKNOWLEDGEMENTS

I am most grateful to G.E. Ball, Chairman, and to the Department of Entomology, Univer-

sity of Alberta, generally, for space and facilities, whereby I am enabled to continue my study

of the Alberta fauna. Publication costs were met by National Research Council Grant A-1399

held by G.E. Ball, and by the Strickland Memorial Fund, Department of Entomology, Univer-

sity of Alberta. G.E. Ball also read, commented most usefully upon the manuscript.

0. 5. Flint, Jr., United States National Museum, pointed out the confusion of subgeneric

names Sortosa and Dolophilodes.

Once again, I express my appreciation to D.B. Donald, Canadian Wildlife Service, Univer-

sity of Calgary, for passing on this material for my examination.

J.E.H. Martin, Canadian National Collection, Ottawa, provided the type number, and suit-

able labels.

REFERENCES

Nimmo, A.P. 1971. The adult Rhyacophilidae and Limnephilidae (Trichoptera) of Alberta

and eastern British Columbia and their post-glacial origin. Quaestiones Entomologicae 7:

3-234.

Nimmo, A.P. 1974. The adult Trichoptera (Insecta) of Alberta and eastern British Columbia,

and their post-glacial origins. II. The families Glossosomatidae and Philopotamidae. Quaest-

iones Entomologicae 10: 315-349.

Nimmo, A.P. 1977. The adult Trichoptera (Insecta) of Alberta and eastern British Columbia,

and their post-glacial origins. I. The families Rhyacophilidae and Limnephilidae. Supplement

1. Quaestiones Entomologicae 13: 25-67.

Glossosomatidae and Philopotamidae

71

Fig. 1-4. Genitalia of Dolophilodes nora Nimmo new species. 1. Male, lat. aspect. 2. Seg. IX & X, dors, aspect. 3. Aedeagus,

lat. aspect. Fig. 4. Distribution map of D. nora Nimmo new species.

Quaest. Ent., 1977 13 (1)

COCCINELLIDAE OF WESTERN CANADA AND ALASKA WITH ANALYSES OF

THE TRANSMONTANE ZOOGEOGRAPHIC RELATIONSHIPS BETWEEN THE

FAUNA OF BRITISH COLUMBIA AND ALBERTA

(INSECTA: COLEOPTERA: COCCINELLIDAE)

JOSEPH BE LICE K

NORCO R Engineering and

Research Limited

Yellowknife, Northwest Territories Quaestiones Entomologicae

X0E1H0 1 3: 73 1 9 77

Corrigenda. J. Belicek 1976, Quaest. Ent. 12(4): 283-409.

change “ quattuordecimguttata” to quatuordecimguttata

change “ quattuordecimguttata” to quatuordecimguttata

change “ quattuordecimguttata ” to quatuordecimguttata

change “ quattuordecimguttata ” to quatuordecimguttata

change “Bilberg” to Billberg

change “numer” to number

change “ quattuordecimguttata ” to quatuordecimguttata

change “ quattuordecimguttata ” to quatuordecimguttata

change “ quattuordecimguttata ” to quatuordecimguttata

change “ novendecimpunctata ” to novemdecimpunctata

change “ novendecimpunctata ” to novemdecimpunctata

change “ quattuorodecimguttata ” to quatuordecimguttata

change “ quattuorodecimguttata ” to quatuordecimguttata

change “ quattuorodecimguttata ” to quatuordecimguttata

change “ Oxynchus” to Oxynychus

Additional References

Crotch, G.R. 1871. A list of Coccineilidae. Cambridge. 8 p.

d’Orbigny, C.D. 1841-1849. Dictionnaire universel d’Histoire naturelle. Paris. 13 volumes of

text, 3 volumes of plates.

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 comprehensive

accounts of entomological research of greater than average length. Page

charges are normally levied, the rate determined by printer’s charges. For

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Copy for all types of papers should conform to the Style Manual for

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I

13

rft.

Quaestiones

Entomologicae

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

VOLUME 13

NUMBER 2

APRIL 1977

QUAESTIONES ENTOMOLOGICAE

ISSN 0033-5037

A periodical record of entomological investigation published at the Department of

Entomology, University of Alberta, Edmonton, Alberta.

Volume 13 Number 2 April 1977

CONTENTS

Book Review — Matsuda, R. 1976. Morphology and Evolution of the Insect

Abdomen 75

Evans — Geographic Variation, Distribution and Taxonomic Status of the Intertidal

Insect Thalassotrechus barbarae (Horn) (Coleoptera: Carabidae) 83

Mendez — Mammalian-Siphonapteran Associations, The Environment, and

Biogeography of Mammals of Southwestern Colombia 91

Sengupta - Changes in Acetycholinesterases and Cholinesterases During Development

of Aedes aegypti (L.) (Diptera, Culicidae) 183

BOOK REVIEW

MATSUDA, R. 1976. Morphology and Evolution of the Insect Abdomen - with special refer-

ence to developmental patterns and their bearings upon systematics. International Series in

Pure and Applied Biology. Volume 56. Pergamon Press, Oxford and New York, viii + 532 pp.,

155 text- figures, 1 table, author, subject and taxonomic indices. Cloth $34.00 (U.S.).

In 1965, Matsuda published “Morphology and Evolution of the Insect Head” (Memoirs of

the American Entomological Institute, Volume 4), the first of a series of works in which he

proposed to review more recent studies on insect morphology. His second volume “Morpholo-

gy and Evolution of the Insect Thorax” (Memoirs of the Entomological Society of Canada,

Volume 76) appeared in 1970 (see review in Quaestiones Entomologicae 7(2): 284-286, 1971)

while the third (and last?), the subject of this review, came out in 1976.

This book is larger (532 pp) than the first two (respectively 334 and 43 1 pp) and its cover-

age of the literature apparently more extensive (2173 vs. 586 and 744 refs.), although these

differences could reflect the relative research activity directed towards each tagma of the in-

sect body. As before, Matsuda’s goal is to “determine the homologies of structures in the light

of recently accumulated facts (p. 1, 1965) . . . using as many kinds of evidence as possible.”

(p. 2, 1965). The evidence he marshalls in this volume has its origin in descriptive and experi-

mental embryology and postembryology, developmental genetics, comparative morphology,

palaentology, and, to a lesser extent than in his previous works, phylogeny. He omits discus-

sion of musculature (an important component of his previous volumes) because (p. vii) “homo-

logies of the kinds of structures treated in this work can safely be established without referen-

ce to the associated musculature.” This may be true but it results in this work being of less

value than previous ones to biologists interested in how structures work. As before, his conclu-

sions are based on his reading, interpretation and digestion of published work - not on the re-

sults of his own investigations.

The book is in three parts: the principles of structural evolution (45 pp), a general discus-

sion of the abdomen (56 pp) and comprehensive treatments of the abdomen and its appenda-

ges in representatives of each order (3 1 8 pp).

In part I, Matsuda provides the theoretical and philosophical framework for his analyses of

homology of structure which follow. Homology is the inheritence, through natural selection,

of structures of descendants from those of a common ancestor. The development of these is

often heterochronic in different insects, ie. the timing of morphogenesis of homologous struc-

tures has often become retarded or accelerated in different evolutionary lines. Metamorphosis,

76

neoteny, caenogenesis, hypermorphosis, the “biogenetic law”, and “law of deviation” are all

manifestations of or generalities on heterochronic development. With substitution , one devel-

opmental process is replaced by another in producing a homologous structure in different ani-

mals.

Matsuda emphasizes that the concept of homology applies primarily to the end product of

morphogenesis, ie. to the functioning, differentiated structure be it embryonic, larval, or ima- \

ginal, because it is on the finished structure that selection works. He also provides a list of cri-

teria that one can use to establish homology between any two structures (eg. position of the

structure relative to another, special features of the structure (often the only criterion used by

palaentologists), developmental sequence of complex structures, development of muscles and i

exoskeleton and innervation) and discusses the various kinds of homology presently recogni-

zed.

Homology of a complex organ must be determined from study both at the level of the

whole and at the level of its components. The penis and ovipositor, for example, have been

largely maintained as homologous organs throughout insect evolution (complete homology)

but their components, in different evolutionary lines, have undergone frequent modification

due to reduction, loss, interiorization, fusion of preexisting components and addition of new j

components (incomplete homology). Serial Homology , the structural correspondence among

repetitive or serial structures within a single individual, is a concept that features prominently

in controversies concerning the sternal or appendicular origin of insect genitalia and larval ab-

dominal legs; these are discussed by Matsuda in part II.

When considering homology, one must be aware of the havoc that convergent evolution can

wreak on one’s conclusions. Convergence is the development of similar structures separately

in two or more lineages without a common ancestor but involving adaptation to similar ecolo-

gical status. The analogous structures which result thus have functional similarity not related

to common ancestry.

Part I is an original and valuable contribution because it expands upon numerous topics ig-

nored by most recent students of insect structure. Unfortunately, it is sorely marred by foggy

exposition. Matsuda’s hope (expressed in his acknowledgements) that “I have a right amount

of “the” in the manuscript.” is - alas - not realized. Rather, he usually has “the’s” where they

shouldn’t be and leaves them out where they should be. Here is Matsuda’s definition of the

biogenetic law: “The essence of the biogenetic law or Haeckel’s law, is that ontogeny consists

of the stage of development in which the adult structures of the ancestor are recapitulated by

the function of heredity (palingenesis) and the stage in which the reconstruction of the adult

structures of the ancestor does not take place (caenogenesis); the latter falsifies the aspect of

palingenesis.” This statement has a higher “fog index” than “ontogeny recapitulates phylogeny.’

There are numerous additional examples - all pointing to a lack of editorial attention to the

manuscript by those having English as their native tongue. This is not a criticism of Matsuda

but of the editorial board of Pergamon Press.

In part II, Matsuda summarizes the conclusions of his detailed, order-by-order-analysis of

the insect abdomen (part III). He has chapters on segmentation, abdominal appendages, male

and female external genitalia, efferent ducts and abdominal ganglia. He believes the ancestral

number of segments in the insect abdomen to be 12 (including the telson) but shows how this

number is reduced in members of most taxa through loss or fusion of segments during subse-

quent development. This reduction makes recognition of segment number in larval or adult

abdomens difficult because of the possibility of originally non-adjacent segments becoming

juxtaposed.

The criteria by which he defines a segment are the presence of paired 1) segmental ganglia,

2) coelomic sacs and 3) appendages in the embryo, of 4) intersegmental sutures, and of

77

a segmental pattern of 5) chaetotaxy, 6) innervation, 7) musculature, and 8) spiracles. Based

on presence of these, the telson is not a segment in spite of Matsuda’s statement to the contrary

(p. 52) because only 4 and 5, in a few insects hold true for it.

According to Matsuda, paired segmental appendages develop on abdominal segments 1-11 in

embryos of most species in most, less-derived, evolutionary lines. In numerous groups one or

more pairs of these develop directly into larval prolegs and/or into other larval appendages. Mat-

suda uses this developmental continuity to oppose Hinton’s (1955. Transactions of the Royal

Entomological Society of London 106: 455-545) widely-accepted theory of multiple, indepen-

dent, secondary origin for these structures. However, Matsuda does not do this as convincingly

as he could. If one plots the presence of embryonic appendages and larval prolegs on a dendo-

gram showing the presently-accepted cladistic relationships of insect orders, one finds prolegs

to be absent from members of many orders (eg. Orthoptera) whose embryos have appendages

and present in members of other orders (eg. many Diptera) whose embryos do not. Even with-

in lower monophyletic taxa, abdominal appendages are present in larvae of some groups, and

are absent from larvae of other groups.

Because of this variation, prolegs can be considered homologous throughout the Insecta

only if they can be traced directly to embryonic abdominal cells in appendage - less embryos

having a latent potential to develop embryonic appendages in Scudder’s sense (1964. Canadian

Entomologist 96: 405-417). Matsuda implicitly accepts this idea to interpret data on prolegs.

However, he does not accept Scudder’s reasoning when considering the presence or absence of

direct developmental continuity between embryonic abdominal appendages and imaginal male

and female genitalia (pp. 90-91 - see below).

Experimental evidence is available to support the existence of latent homology. The “not-

ched sternite” mutant of Blattella germanica which has many of the abdominal characteristics

typical of Thysanura (pregenital styli, division of abdominal sterna into median and lateral ster-

nites and paired abdominal membranous outgrowths comparable to the eversible sacs) reveals,

in this species, the presence of relict genetic mechanisms which may have influenced or contro-

lled development of abdominal segments in some Thysanura - like ancestral form (Ross, 1966.

Annals of the Entomological Society of America 59: 473-484; 1 160-1 162). I see no reason why

similar genetic mechanisms could not exist in most insects.

Matsuda also uses this evidence to support Berlese’s theory in which larvae are claimed to

be free-living embryos that hatch at earlier (protopod) or later (polypod, oligopod) stages of

development.

The appendages of abdominal segment 1 develop into the purely embryonic pleuropodia

in investigated representatives of orders other than Collembola, Ephemeroptera, Dermaptera,

Psocoptera, Neuroptera, Hymenoptera, Mecoptera, Siphonaptera, Diptera and Strepsiptera.

Their role in embryogenesis is mostly unknown but appears to vary greatly in different insects

(their contributions to embryonic nutrition, hatching and electrolyte transport have been demon-

strated experimentally in embryos of different insects).

In females of Acrididae and Tettigoniidae (Orthoptera) the ovipositor valves of segments 8

and 9 develop directly from the embryonic appendages of these segments and in males the ex-

ternal genitalia from those of segment 10. These structures have a proven appendicular

origin only in members of these families. In most investigated representatives of other orders,

the external genitalia of both sexes appear to arise independent of the embryonic appendages

from sternal epidermis of the genital segments. This occurs either after the embryonic appen-

dages have withdrawn into the abdominal epidermis (in members of those taxa having such ap-

pendages) or de novo (in members of those taxa lacking them). Matsuda thus refuses to accept

a leg origin for insect genitalia and criticizes workers such as G.G.E. Scudder and E.L. Smith

for using leg terms for genitalic structures.

78

As mentioned above, Matsuda uses the evidence of direct developmental continuity be-

tween embryonic abdominal appendages and larval prolegs to support homologizing the pro-

legs of all immature insects having them. If he accepted Scudder’s (1964) thesis of latent ho-

mology between embryonic appendages or equivalent cells and insect genitalia, he could do

the same for insect genitalia. He does, in fact, homologize the valvular ovipositor in female in-

sects and the penis in males throughout the Insecta - but not on this basis. Instead, he consi-

ders the apparent appendicular origin of genitalia in acridids and tettigoniids to have arisen

secondarily through acceleration and substitution in the cells determined to form these struc-

tures. Additional evidence of a sternal origin for genitalia exists in Thysanura where the ovi-

positor of females arises between the coxopodites of the eighth and ninth segments, only the

latter developing from embryonic abdominal appendages of these segments.

Until further evidence is accumulated, I am inclined to agree with Matsuda on both counts

even though the evidence for his concept of “homology through substitution” is not strong.

In most insects, Matsuda considers the penis (phallus of Snodgrass) and its accessory struc-

tures to arise postembryonically from a pair of primary phallic lobes in the sternal epidermis

at the posterior margin of segment 9. In those insects (some species of Phthiraptera, Homop-

tera, Coleoptera and Diptera) in which they appear to arise on other segments (7, 8, or 10),

one or more adjacent segments have probably been lost through segmental reduction or fus-

ion. The later differentiation of these lobes into the diversity of structures so loved by insect

systematists provides good examples of Matsuda’s law of deviation (the developmental process

whereby similar rudiments in different animals become increasingly dissimilar in later devel-

opment) and of complete (the primary phallic lobes themselves) and incomplete (the diverse

external genitalia they become) homology. Because deviation occurs, E.L. Smith’s (1969.

Annals of the Entomological Society of America 62: 1051-1079) piece-by-piece homologi-

zing of each component of male and female external genitalia is unacceptable as is Snodgrass’

(1957. Smithsonian Miscellaneous Collections 136(6): 1-60) of parts of the male genitalia.

The ovipositors of most females are considered by Matsuda to be gonapophyseal in origin

(paired apophyses of the 8th and 9th abdominal sterna) but are formed secondarily by modi-

fication of posterior abdominal segments after the gonapophyseal ovipositor has been lost.

Matsuda hence concludes that the valvular ovipositor is archaic and arose early in insect evo-

lution. It has since become modified or lost independently in members of each evolutionary

line. Therefore, although the ovipositor is completely homologous in most insects, its parts

are incompletely homologous.

In chapter 1 1 , Matsuda critically analyzes the 1 1 theories that have been developed to ex-

plain the evolutionary origin of insect external genitalia. These are grouped into three categor-

ies: 1) those based only on developmental information, 2) those based on comparison of adult

structures, and 3) those based on consideration of both kinds of information. Insect genital

appendages have not only been homologized with abdominal legs but with diplopod gonopods,

crustacean biramous limbs and with the eversible sacs of thysanuran pregenital segments. Mat-

suda believes both male and female genitalia to be sternal.

The efferent ducts and associated structures of the reproductive systems of insects of both

sexes in most species consist of primary and secondary parts. The primary exit system is pair-

ed and usually originates from posterior extensions of genital ridges of the mesodermal coelo-

mic sacs during embryogenesis. The secondary exit system is not paired and usually takes form

after hatching by invagination of sternal ectoderm of the genital segments (7, 8 and 9).

The primary exit system of males begins as a pair of rudimentary vasa deferentia each ter-

minating anteriorly in a testicular rudiment and posteriorly, in segment 9, as a swollen genital

ampulla. With subsequent differentiation, these primordia give rise to the vasa deferentia, sem-

inal vesicles, and accessory glands (mesadenes) and to a greater or lesser part of the ejaculatory

79

bulb. The primary exit system of females is similar but each primordial lateral oviduct termin-

ates in segment 7 and, at most, differentiates into the imaginal lateral oviduct. Female embryos

of some species of Phasmida and Orthoptera and male embryos and larvae of Diptera-Nemato-

cera have rudiments of the primary exit system of both sexes, one or the other degenerating

with subsequent development (a phenomena similar to the sexually indifferent period exper-

ienced by vertebrate embryos).

The secondary exit system of most male insects ultimately comprises the ejaculatory duct,

ejaculatory bulb and additional accessory glands (ectadenes). These usually first appear as a

single, forward invagination of ectoderm from between the bases of the primary phallic lobes

shortly before or after hatching. In most females, the secondary exit system consists of a com-

mon oviduct, spermatheca, vagina and accessory glands. Most begin to develop post embryon-

ically, the common oviduct as an invagination behind the 7th sternum (often continued to the

back of the 8th as a longitudinal groove whose lips subsequently fuse), the spermatheca from

sternum 8 and the accessory glands from sternum 9. The openings of the spermathecal duct

and accessory glands are then carried in by invagination of what remains of the 8th and 9th

sterna to form the vagina.

The above developmental sequence characterizes most insects (in females of investigated

species of Ephemeroptera only the primary system develops to any extent, each lateral ovi-

duct having its own opening at the back of the seventh sternum. In males of most investigated

species of Ephemeroptera and Dermaptera, two ejaculatory ducts arise, each invaginating for-

ward from the apex of a primary phallic lobe). In endopterygote lines, a greater or lesser num-

ber of steps have been omitted due to heterochrony and substitution. There has been a gener-

al tendency for the secondary efferent system to substitute for the primary one and for the

imaginal discs of adjacent segments in the female abdomen to amalgamate. This trend culmin-

ates in members of the Diptera - Cyclorrhapha in which the genital disc(s) in a larva differen-

tiate not only into the complete male or female reproductive system and external genitalia

(except gonads) but also into part of the hindgut and into the terminal abdominal segments of

adults.

Part III is an almost overwhelming detailed analysis of abdominal structure and development

in representatives of 30 orders. The chapter on Diptera is the longest (28 pp) because of the

vast amount known about abdomens of members of this order. In it are treated 1) embryonic

and 2) larval abdominal segmentation; 3) female and 4) male imaginal segmentation; 5) theor-

ies of abdominal segmentation; 6) abdominal appendages; 7) postembryogenesis of male ex-

ternal genitalia; 8) torsion of the post-abdomen; 9) the male and 10) female terminalia; 1 1)

germ cells; 12) postembryogenesis of the male efferent system; 13) male internal reproductive

system; 14) postembryogenesis of the female efferent system, and 15) female internal repro-

ductive system. One or more of these discussions is omitted from the treatments of other or-

ders because of lack of information. For example, the chapter on Zoraptera is only 1 1/2 pages

long and considers only 3, 4, 9, 10 and 13 above. Matsuda’s book is thus not only an impor-

tant source of information on abdominal characteristics for members of each order, but also

a good place to find research problems.

Development of the ovary and testis and oogenesis and spermatogenesis are not covered be-

cause of the reviews of these subjects already available. Matsuda does, however, provide detail-

ed access to these literature sources.

My principal criticism of this book concerns Matsuda’s decision not to consider musculature

(already alluded to), phylogeny, or Hennig’s (1966. “Phylogenetic Systematics”) principles of

cladistic analysis. I am convinced that the best way to relate everything in a complicated work

of this kind is through the use of phylogenetic diagrams and through knowledge and use of

sister group relationships between higher taxa. Except for included taxa of the Orthopteromorpha

Entognothd Orthopteromorpha Acercaria Panorpoidea

80

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81

(knowledge of whose relationships is still inadequate) accepted sister group relations of the

insect orders are becoming stabilized (see Hennig, W. 1969. “Die Stammesgeschichte der In-

sekten” and Kristensen, N.P. 1975. Zeitschrift fur zoologische Systematik und Evolutions-

forschung 13: 1-44). If details of particular abdominal structures or developmental sequences

are plotted on a phylogenetic dendrogram of the insect orders, one sees quickly whether they

arose independently several times or only once in the common ancestor of monophyletic ass-

emblages of orders. For example, if one does this for ovariole type (Fig. 1) one sees at a glance

that the panoistic type is plesiomorphic but has been lost and secondarily regained either once

(Siphonaptera) or twice (Thysanoptera and Siphonaptera), that the polytrophic type has in-

dependently arisen four times (in Collembola, Diplura, Dermaptera, and in the common an-

cestor of all remaining orders), and that the acrotrophic type has arisen independently twice

(Hemiptera and Coleoptera) from ancestors having the polytrophic type. If such a plot is then

superimposed on an identical diagram in which species diversity per order is indicated, one sees

that the evolution of trophocytes (nurse cells) has probably contributed directly to the evolu-

tionary success of those orders whose members’ ovarioles are characterized by their presence.

The selective advantage of meroistic (nurse cell-containing) ovarioles is that their oocytes grow

faster and are produced in greater numbers than those of panoistic ovarioles because of the

large amount of template DNA (up to 1024C) available for oogenesis in the polyploid nuclei

of their nurse cells (only 4C amounts of DNA are available for RNA synthesis in the germinal

vesicles of panoistic oocytes - see Mahowald, A.P. 1972 in Vol. Developmental Systems: In-

sects).

Use of Hennig’s methods would add rigor to Matsuda’s analysis and would make more ob-

vious structural and developmental dines. This would result in Matsuda’s book being of use

to a larger number of workers than it is now. It would also enable readers to make conpari-

sons more easily and to use the book in making predictions about structure of the abdominal

appendages in unstudied taxa.

The volume is well-produced but there are many typographical errors and some of the print

of my copy ended up on my fingers and on the pages of the first draft of this review. Some fig-

ures, though fully-labeled, are difficult to interpret because they were sloppily done or repro-

duced with too much reduction. They compare unfavourably with those of Anderson’s recent

treatise (1973. “Embryology and Phylogeny in Annelids and Arthropods”) in the same series.

In spite of these criticisms, I consider Matsuda’s book to be a major contribution because

of its comprehensive bibliography and because of the vast amount of information it contains.

ACKNOWLEDGEMENT

I thank G.E. Ball for constructive criticism and discussion during the writing of this review.

B.S. Heming

Department of Entomology

University of Alberta

Edmonton, Alberta T6G 2E3

GEOGRAPHIC VARIATION, DISTRIBUTION AND TAXONOMIC STATUS

OF THE INTERTIDAL INSECT THALASSOTRECHUS BARBARAE

(HORN) (COLEOPTERA: CARABIDAE)

WILLIAM G. EVANS

Department of Entomology

University of Alberta

Edmonton, Alberta T6G 2E3

Quaestiones Entomologicae

13: 83-90 1977

Populations of Thalassotrechus barbarae (Horn), a nocturnal, flightless, intertidal carabid

beetle species, are distributed linearly from Point St. George ( Crescent City), California to Bah-

ia Magdalena, Baja California, Mexico. Mean elytral lengths vary clinally from 2.68 mm in the

extreme southern part of the range to 3.54 mm in the northern part. Elytral color also varies

clinally with northern populations being darker. These data are not consistent with recogni-

tion of subspecies. Consequently, T. b. barbarae (Horn, 1892; type locality - Santa Barbara,

California), and T. b. nigripennis (Van Dyke, 1918; type locality - Moss Beach, San Mateo

County, California) are consubspecific, and their names are synonyms. North-South temper-

ature and rainfall gradients may be implicated in selection of local variants along the linear

dine of elytral length and color.

Les populations de Thalassotrechus barbarae (Horn), une especes de carabique nocturne, et apte're se trouvent dans la zone

des maries et se rencontrent de la pointe St. George (Crescent City), Californie jusqu’a Bahia Magdalena, Basse Calif ornie, Mex-

ique. La longeur moyenne des ilytres varie graduellement a partir de 2.68 mm dans V extreme sud de leur distribution jusqu’a

3.54 mm dans le nord. La couleur des ilytres varie de la mime facon avec les populations les plus foncies dans le nord. Ces

donnies ne sont pas compatibles avec notre concept de la sous-espece. Done T.b. barbarae (Horn, 1892; localite type - Santa

Barbara, Californie) et T. b. nigripennis (Van Dyke, 1918; localiti type - Moss Beach, San Mateo County, Californie) ne repri-

sentent qu’une seule sous-espece, et leurs noms deviennent synonymes. Le changement graduel de la tempirature et de la pri-

cipitation entre le nord et le sud pourrait expliquer la silection de variations locales en fuonction de changement liniaire de

la couleur et de la longeur des elytres.

On most coastlines of the world certain insects are conspicuous on rocky shores and sandy

beaches, particularly in those areas characterized by extensive beds of offshore kelp and by

diverse intertidal algal communities. During daytime low tide in such places, flies are very evi-

dent among cast-up algal masses (wrack) and occasionally swarms of beetles (staphylinids and

hydrophilids) are encountered (Leech, 1971); during periods of low tide at night predatory

and scavenging beetles are active on sand or rock surfaces (Evans, 1977). However, the major-

ity of intertidal insects, particularly their larval stages, are not easily observed since they inha-

bit subsurface or rock crevice habitats or algal vegetation.

In general, similar kinds of intertidal insects occur in similar habitats all over the world. Kelp

flies of the families Anthomyidae and Coelopidae, for instance, are found wherever wrack ac-

cumulates and an extensive beetle fauna is also associated with this food base either as preda-

tors or scavengers. Some beetles eat other insects or small crustaceans stranded by receding

tides, and members of Thalassotrechus barbarae (Horn) are such predators. Adults and larvae

of this species live in cracks in rocks in the intertidal zone; adults are brachypterous, and there-

fore flightless. Larvae are restricted to crack habitats but adults emerge at night and walk over

the rock surfaces; feeding and mating takes place at this time.

INTRODUCTION

84

Evans

Ecological equivalents of this species are undoubtedly found in other parts of the world and

probably include carabid beetles of the tribe Trechini such as Kenodactylus audouini (Guerin) I

occurring in the subantarctic islands of New Zealand (the Antipodes Islands, Auckland Islands, j

Campbell Island, Snares Island, Stewart Island), the Falkland Islands and Patagonia (Darlington, !

1964; Johns, 1974), the European Aepus marinus (Strom) and Aepopsis robinii (Laboulbene)

(Glynne-Williams and Hobart, 1952) and Thalassobius testaceus Sober from Chile (Jeannel,

1926).

S

GEOGRAPHICAL DISTRIBUTION OF THALASSOTRECHUS BARBARAE (HORN)

The geographic range of Thalassotrechus barbarae extends from Point St. George, (near Cre-

scent City) Del Norte County, California to Punta Belcher, Bahia Magdalena, Baja California

(Fig. 1), a straight-line distance of approximately 2400 km that spans 17° 10' of latitude and

12° 03' of longitude. Populations of this species occur on rocky shores or on rocky outcrops

of sandy beaches along this range and conceivably south of Bahia Magdalena as far as Cabo

San Lucas, some 185 km away at the tip of Baja California. A part of this range is in the mar-

ine biogeographic zone (based on sea water surface temperature) called Cold-Temperate which ,

extends from Point Conception, California to Alaska (Abbott, 1966) while the remaining part

coincides with the Warm-Temperate zone between Point Conception and Bahia Magdalena. So j

the latter locality is most likely to be very close to the southern limit of the range but as far as

I know no collections have been made between Bahia Magdalena and the southern tip of the

peninsula. The northern limit of the range is more exactly defined because north of Crescent

City, California, where collecting sites are more accessible than in Mexico, I (and presumably

other collectors) have failed to find any specimens.

TAXONOMIC CONSIDERATIONS AND GEOGRAPHICAL VARIATION

Taxonomic history of Thalassotrechus barbarae

T. barbarae , though trechine in general form and habits, has been placed in the Poginini on

the basis of general structure and particularly on characters of the mouthparts (Van Dyke, 1918) 1

and genitalia (Darlington, 1938). Pogonines are halobiontic (Ball, 1968), that is, they inhabit

alkali soils with high concentrations of sodium chloride, but T. barbarae is the only member of

this predominantly Palaearctic group that is found in the rocky intertidal zone, whereas mem-

bers of the Trechini are commonly found in this habitat. Adults and larvae of Trechus ovipen-

nis Motschulsky, for instance, occur in crevices and under stones of the high intertidal zone on

rocky shores from central California to southern Alaska (Evans, 1977; Lindroth, 1961) and the

trechine species mentioned above have similar habits. Thus, T. barbarae is probably a relict gen-

us derived from a stock that gave rise to the present day halobiontic pogonines (Ball, pers. comm.)

but because it has become adapted to living in the rocky marine littoral it has convergently be-

come similar in color and form to marine trechine beetles (Darlington, 1938).

Horn (1892) described Trechus barbarae from specimens collected at Santa Barbara, Califor-

na, the type locality. 1 Van Dyke (1918) transferred this species to the Pogonini, based on it

the new genus Thalassotrechus, the name indicating resemblance of its members to members

of Trechus. He also described a second species, T. nigripennis, from northern California (type

locality - Moss Beach, San Mateo County; see Fig. 1), adults of which were larger and darker,

and with proportionately broader prothoraces (Van Dyke probably judged this last feature, as

taxonomists generally did at that time, rather than taking measurements). Apparently, however,

1. During 1969, specimens could not be found anywhere near Santa Barbara presumably because of the massive oil spill

that occurred early in the year (Evans, 1970).

Thalassotrechus barbarae (Horn)

85

he later came to regard T. barbarae and T. nigripennis as subspecies, a conclusion amplified

by Darlington (1938) on the basis of comparison of adults from Santa Barbara with adults

from the San Francisco area. The elytra of adults of the southern population examined by

Darlington were rufotestaceous or castaneous in contrast to the black and dull-textured ely-

tra of adults of the northern group. Also, the anterior prothoracic angles of adults of barbar-

ae were less flattened than those of the northern population. Moore (1956) also agreed with

this arrangement even though he found no consistent structural difference between series of

both populations.

Geographic variation of T. barbarae

Since clinal variation is expected to occur in Thalassotrechus over the predominantly NW-

SE trend of the coastline on which it is found I examined adult specimens loaned by the Cali-

fornia Academy of Sciences as well as collected by myself. With the use of an ocular microme-

ter, measurements of elytral length (base to apex along the suture) and maximum prothoracic

width were made of individuals obtained from localities shown in Table 1 . These localities re-

present collecting sites that vary in size from individual rocks (at Bahia Magdalena) to indivi-

dual beaches (Pismo Beach) to long stretches of coastline (Del Norte, Humboldt and Mendoc-

ino Counties).

The data, shown in Fig. 1, clearly indicate that the mean elytral length of Thalassotrechus

populations increases progressively in a north-westerly direction agreeing with Bergman’s Rule

(Mayr, 1942) which states that the smallest sized individuals of a species are found in the sou-

thern part of the range and the largest in the northern part. This clinal progression consists of

consecutive, overlapping ranges of elytral length, and of standard deviations of elytral length

(except for San Diego County and Bahia Magdalena) with the largest step occurring between

Bahia Magdalena, Mexico and San Diego, California, a straight-line distance of approximately

1060 km. Because of this distance I would expect that further sampling between these two

localities would give populations that would reduce this large step to a series of overlapping

smaller ones. A population, then, from any locality would not differ significantly in size from

an adjacent population. Prothoracic width/elytral length ratios do not differ significantly be-

tween adjacent localities or between the most northerly and the most southerly localities (Table

1) indicating that the samples were drawn from a population of individuals of different sizes

but not of different form.

As shown in Fig. 1 , the rate of change of mean elytral length appears to be greater in the

middle of the range than at the ends. Theoretically, this may be expected because there would

be a decreasing tendency for selection of variants that reflect submarginal environmental con-

ditions that presumably exist towards the ends of the range. Alternatively, the greater rate of

change may be due to an equally greater rate of change of some environmental variable such

as sea water temperature. An abrupt change such as this probably occurs in the middle of the

range near Point Conception, California (see Fig. 1) a well-known locus of marine inshore fau-

nal discontinuity (Abbott, 1966; Garth, 1955).

Adult specimens were also examined for variation in elytral color. Five color categories

could be distinguished and individuals were assigned to a category as accurately as possible by

this subjective method. The proportions of the populations from each locality (not identical

to the above localities because all the specimens were not available at the time of this analy-

sis) in each of these color categories are shown in Fig. 2 where the suspected color cline is sat-

isfactorily demonstrated. Elytral color progresses from pale testaceous in the extreme south

to black in the northern part of the range. It is interesting to note, however, that color varies

even at the ends of the range so that the elytra of individuals from Bahia Magdalena are not

Quaest. Ent., 1977 13 (2)

all pale testaceous and individuals from Point St. George do not all have black elytra. Fin-

ally, color variation is apparently greater in the middle part where the elytral color of indi-

viduals may vary from the lightest to the darkest.

Because variation in supposed diagnostic features is either non-existent (ratio to prothora-

cic width/elytral length) or clinal (elytral length and color), I conclude that recognition of

subspecies would be wholly arbitrary, and would serve no useful purpose. Consequently, T. j

b. barbarae and T. b. nigripennis are consubspecific, and their names are synonymous. The j

species is thus monobasic, and a binomial name is sufficient.

Discussion of Pattern of Variation of T. barbarae

Clines result from gene flow between populations and from selection of individuals that

are adjusted to the local environment (Mayr, 1963). Since T. barbarae is distributed along a

northwest-southeast trending coastline, that is, linearly (or unidimensionally, Udvardy, 1969),

environmental factors that show a north-south gradient are more likely to be implicated in the

selection of local varients. One such factor, temperature, is an obvious example, and its role in

influencing the body size or body proportions of poikilotherms has been demonstrated by Ray

(1960). A north-south gradient for the California coastline is well-documented for air temper-

ature (U.S. Department of Commerce, 1968) and sea water temperature (U.S. Department of

Commerce, 1967) both of which must influence metabolism of T. barbarae but other factors

such as salinity, rainfall, humidity, wave action, and availability and kind of food, are proba-

bly also involved. Johns (1974) suggests that body size of marine carabids of the species Keno-

dactylus audouini may be related to the amount of exposure to sea water since the largest spe-

cimens of this species are in supralittoral habitats where sea water is much diluted and where

prey such as Collembola and Isopoda, and larvae and eggs of various arthropods are more avai-

lable. A rainfall gradient exists between the ends of the range of T. barbarae with Point St. Geo-

rge (Crescent City), California having the highest average annual rainfall, about 178 cm (U.S.

Department of Commerce, 1968) and Bahia Magdalena, Mexico receiving less than 25 cm a

year (Escoto, 1964). The increasing dilution of sea water in a northerly direction may very

well be implicated in the dine of body size in T. barbarae but perhaps the rainfall gradient,

expressed as a humidity gradient, may be implicated in the clinal variation in elytral color.

Discussing Gloger’s Rule, Dobzhansky (1937), concludes that pigmentation in insects increa-

ses in humid and cool and decreases in dry and hot climates with humidity being more effect-

ive than temperature. If, indeed, this is correct, the correlation between elytral color and hum- 1

idity.in T. barbarae could be explained but the underlying mechanism for this phenomenon

is unknown. In all likelihood, several factors are involved in geographic variation of T. barbar-

ae making it difficult to determine the actual cause of this variation even though the distribu-

tion of this insect is linear, therefore, seemingly much more simple to analyze than two-or

three-dimensional distributions (Udvardy, 1969). In the latter case, for example, a plot of a

single character gradient such as color on a distribution map results in, at best, a series of non-

parallel, crude isophenes (see Petersen, 1947). Or, when geographic variation in body length

of a nonlinearly distributed insect is plotted on a Hubbs-Hubbs diagram, as Ball and Negre

(1972) did for a xerocolous carabid, Calathus ruficollis Dejean, the resulting pattern can be

very complex indeed. But this pattern merely reflects complex gradients of environmental var-

iables. In order to elucidate relationships between biological and environmental variation pre-

cise measurements of such parameters as temperature and rainfall or humidity are needed in

addition to data on several character states of populations of linearly distributed species. This

study suggests that such an approach is possible.

Thalassotrechus barbarae (Horn)

87

ACKNOWLEDGEMENTS

I am grateful to H.B. Leech and D.H. Kavanaugh for the loan of specimens. Bahia Magdal-

ena specimens were collected by the author during Cruise 1 8 of Stanford Oceanographic Ex-

peditions which was supported by NSF grants GB 6870 and GB 6871 to Stanford University.

My thanks are extended to G.E. Ball for his encouragement and for his critical review of the

manuscript and to J.S. Scott for drafting the figures.

Table 1. Prothoracic width/elytral length ratios of populations of Thalassotrechus barbarae

collected at various localities.

LITERATURE CITED

Abbott, D.P. 1966. Factors influencing the zoogeographic affinities of the Galapagos inshore

marine fauna, pp. 108-122. In Bowman, R.I. (editor), The Galapagos. University of Califor-

nia Press. Berkeley, California xvii + 318 p.

Ball, D.E. 1968. Carabidae, Fascicle 4, pp. 55-174. In Arnett, R.H. The beetles of the United

States (a manual for identification). Catholic University of America Press, Washington, D.C.

xi + 1 1 12 p.

Ball, G.E. and J. Negre. 1972. The taxonomy of the nearctic species of the genus Calathus

Bonelli (Coleoptera: Carabidae: Agonini). Transactions of the American Entomological

Society, 98: 412-533.

Darlington, P.J., Jr. 1938. The American Patrobini (Coleoptera, Carabidae). Entomologica

Americana, 18: 135-187.

Darlington, P.J., Jr. 1964. Insects of Campbell Island. Coleoptera: Carabidae. Pacific Insects

Monograph 7: 335-339.

Dobzhansky, T. 1937. Genetics and the origin of species. Columbia University Press, New York,

xvi + 364 p.

Quaest. Ent., 1977 13 (2)

88

Evans

Escoto, J.A.V. 1964. Weather and climate of Mexico, pp. 187-215. In Wauchope, R. and R.C.

West (editors). Handbook of Middle American Indians, Volume 1. Natural environment and

early cultures, University of Texas Press, Austin, Texas. 570 p.

Evans, W.G. 1970. Thalassotrechus barbarae (Horn) and the Santa Barbara oil spill. The Pan-

Pacific Entomologist, 46: 233-237.

Evans, W.G. 1977. Insects and relatives. In Morris, R.H. and D.P. Abbott (editors). Marine

invertebrates of the California shore. Stanford University Press, Palto Alto, California (In Press).

Garth, J.S. 1955. The case for a warm-temperate marine fauna on the west coast of North

America, pp. 19-27. In Essays in the natural sciences in honor of Captain Allan Hancock. j

On the occasion of his birthday, July 26, 1955. University of Southern California Press,

Los Angeles, California xii + 345 p.

Glynne- Williams, J. and J. Hobart. 1952. Studies on the crevice fauna of a selected shore in

Anglesey. Proceedings of the Zoological Society of London, 122: 797-825.

Horn, G.H. 1892. Random studies in North American Coleoptera. Transactions of the Ameri-

can Entomological Society, 29: 40-48.

Jeannel, R. 1926. Monographic des Trechinae. Morphologie comparee et distribution geogra-

phique d’un groupe de Coleopteres. Part I. L’Abeille, Tome 32: 221-550.

Johns, P.M. 1974. Southern New Zealand, Patagonian and Falkland Island Carabidae. Arth-

ropoda of the Subantarctic Islands of New Zealand (I) Coleoptera: Carabidae. Journal of

the Royal Society of New Zealand, 4: 283-302.

Leech, H.B. 1971. Nearctic records of flights of Cafius and some related beetles at the sea-

shore. (Coleoptera: Staphilinidae and Hydrophilidae). Wasman Journal of Biology, 29: 65-

70.

Lindroth, C.H. 1961. The ground beetles (Carabidae, excl. Cicindelinae) of Canada and Alas-

ka. Part 2. Opuscula Entomologica, Supplement 20: 1-200.

Mayr, E. 1942. Systematics and the origin of species. Columbia University Press, New York,

334 p.

Mayr, E. 1963. Animal species and evolution. The Belknap Press of Harvard University Press,

Boston, Mass, vii + 797 p.

Moore, I. 1956. Notes on some intertidal Coleoptera with descriptions of the early stages

(Carabidae, Staphylinidae, Malachiidae). Transactions of the San Diego Society of Natural

History, 12(11): 207-230.

Petersen, B. 1947. Die geographische variation einiger Fennoskandischer Lepidopteren. Zoo-

logiska bidrag fran Uppsala, 26: 330-531.

Ray, C. 1960. The application of Bergmann’s and Allen’s rules to the poikilotherms. Journal

of Morphology, 106: 85-108.

Udvardy, M.D.F. 1969. Dynamic zoogeography. With special reference to land animals. Van

Nostrand Reinhold Publishing Co. New York, xviii + 445 p.

United States Department of Commerce. Environmental Science Services Administration, Coast

and Geodetic Survey. 1967. Surface water temperature and density. Pacific Coast, North and

South America and Pacific Ocean Islands. Publication 31-3, 2nd edition.

United States Department of Commerce. Environmental Science Services Administration.

Environmental Data Service. 1968. Climatological Data, California, Volume 72, No. 13.

Vay Dyke, E.C. 1918. New inter-tidal rock-dwelling Coleoptera from California. Entomolo-

gical News, 29: 303-308.

Thalassotrechus barbarae (Horn)

89

Fig. 1. Mean (vertical line), standard deviation (black bar) and range (horizontal line) of elytral lengths of populations of

Thalassotrechus barbarae collected from localities given in Table 1. A- Type locality for T. nigripennis. •- Type locality for

T. barbarae.

Quaest. Ent., 1977 13 (2)

90

Evans

68

55

12

®00

0 0 0

0000''

/

125°

/

120°

/

115°

Fig. 2. Proportions of populations of Thalassotrechus barbarae collected from various localities in five elytral color categor-

ies. Black portion of circle represents the proportion of a population in that particular color category. Locations: 10. Men-

docino, Humboldt and Del Norte Counties, Calif.; 11. Marin, Sonoma and Contra Costa Counties, Calif.; 12. Pacific Grove,

Calif.; 13. Pismo Beach, Calif.; 14. Santa Barbara and Ventura Counties, Calif.; 15. Punta Belcher, Bahia Magdalena, Baja

California, Mexico.

MAMMALIAN-SIPHONAPTERAN ASSOCIATIONS, THE ENVIRONMENT,

AND BIOGEOGRAPHY OF MAMMALS OF SOUTHWESTERN COLOMBIA

EUSTORGIO MENDEZ

Gorgas Memorial Laboratory

P.O. Box 2016

Balboa Heights

Panama Canal Zone

Quaestiones Entomologicae

13: 91-182 1977

A synopsis of the fleas and their mammalian hosts in southwestern Colombia, with parti-

cular reference to the Departamento del Valle is presented and information about the Siph-

onaptera of Colombia is reviewed.

Descriptions and illustrations of diagnostic features characterize the following new species

o/Polygenis Jordan: P. caucensis (Type locality : Alto Anchicay a, Depto. del Valle, COLOM-

BIA; type host: Oryzomys caliginosusj, P. delpontei (Type locality: Quebrada Honda near

Pichinde, Municipio de Cali, Depto. del Valle, COLOMBIA; type host: Oryzomys caliginosus/

P. hopkinsi (Type locality: Pena del Cerro, Cerro Munchique, Depto. del Cauca, COLOMBIA;

type host: Oryzomys albigularisA P. trapidoi (Type locality: Valle del Rio Pichinde, Munici-

pio de Cali, Depto. del Valle, COLOMBIA; type host: Oryzomys caliginosusj. In addition, des-

criptions are given of the male of Ctenidiosomus traubi Johnson, and the female o/Sphincto-

sylla diomedes Johnson. The former species was previously known from the female, and the

latter species was originally described from the male. The following taxa are reported for the

Republic of Colombia for the first time: Plocopsylla phyllisae Johnson, Leptopsylla segnis

(Schonherr), Dasypsyllus gallinulae peripinnatus (Baker), Tetrapsyllus comis Jordan, Polygen-

is pradoi (Wagner), P. thurmani Johnson, P. klagesi samuelis (Jordan and Rothschild), and

Pulex simulans ( Baker).

The taxon Rhynchopsyllus megastigmata Traub and Gammons, 1958, is considered to be

conspecific with Rhynchopsyllus pulex Haller, 1895.

Records contained herein bring the total number of known Colombian Siphonaptera to at

least 44 species and subspecies. Of this number, more than two-thirds are reported from the

southwestern part of the country.

A key to the fleas of southwestern Colombia, and illustrations of diagnostic features of

most of the taxa are included.

An account is given of the history and zoogeography of the mammalian fauna of south-

western Colombia, as well as comments on host-parasite relationships. The history of the mam-

malian fauna of South America was probably initiated at the beginning of the Tertiary. This

early fauna included marsupials, certain edentates and primtive ungulates that came from

North America. Subsequent isolation of North America from South America interrupted, but

not completely stopped, an interchange of animals by island-hopping. The faunal interchange

was re-established when the Isthmian land bridge appeared at the end of the Tertiary. It is also

possible that during the Cretaceous and early Tertiary, South America received faunal elements

from Africa and from Australia.

The majority of families and genera of South American mammals originated during periods

of total or incomplete isolation. Presently, in the Pacific Coastal Lowlands of Colombia ubi-

quituous mammals outnumber endemic forms. This fauna has strong affinities with those of

the Amazon Region and the coastal lands of Panama and Ecuador. The remaining taxa of the

southwest portion of Colombia are more concentrated in the Andes and include some forms

that probably originated in the lowlands.

Evolution and radiation of the fleas of this territory probably was correlated to evolution

92

Mendez

of their hosts. This phenomenon is more apparent in fleas parasitizing small mammals, such

as cricetine rodents. The lowland flea fauna of the Pacific sector is particularly poor, both in

numbers of taxa and endemism, while that of the Andean mountains displays more local ele-

ments in addition to being significantly diversified. The Southwestern Colombia flea fauna ex-

hibits little specificity and various species may parasitize the same host. On the other hand,

there are related and unrelated host species that harbor the same flea taxon. It is evident that

the flea fauna of the Colombian Pacific lowlands is related to those of Panama and the Amazon

and Orinoco basins. The strong mammalian relationship existing throughout the Andean Cor-

dillera in Colombia and Ecuador is reflected in the affinity observed in the flea fauna.

Geographic aspects, such as topography, geology, soil, climate and vegetation concerned

with the pertinent biota are discussed. Southwestern Colombia is characterized by a diversity

of ecological situations resulting from a complex topography and the concomitant climatic

regimes. The Pacific Coastal lands, and the Andean highlands, in addition to the Cauca Valley,

represent the major geographic areas of this territory.

Se presenta un sumario de la fauna de sifondpteros y sus mamiferos hospederos en la regidn suroccidental de Colombia,

principalmente en el Departamento del Valle; al mismo tiempo, se revisa la informacidn existente sobre las pulgas de Colom-

bia.

Se incluyen descripciones e ilustraciones de aspectos diagndsticos que caraceterizan las siguientes especies nuevas del gdn-

ero Polygenis Jordan: P. caucensis (Localidad tipo: Alto Anchicaya, Depto. del Valle, COLOMBIA; hospedero tipo: Oryzo-

mys caliginosus^, P. delpontei (Localidad tipo: Quebrada Honda, cerca de Pichindd, Municipio de Cali, Depto. del Valle,

COLOMBIA; hospedero tipo: Oryzomys caliginosus^, P. hopkinsi (Localidad tipo: Pena del Cerro, Cerro Munchique, Depto.

del Cauca, COLOMBIA; hospedero tipo: Oryzomys albigularisy, P. trapidoi (Localidad tipo: Valle del Rio Pichindd, Munici-

pio de Cali, Depto. del Valle, COLOMBIA: hospedero tipo: Oryzomys caliginosus/ Adema's, se describen en este trabajo el

macho de Ctenidiosomus traubi Johnson y la hembra de Sphinctopsylla diomedes Johnson. De la primera especie unicamen-

te se conicia la hembra, mientras que la segunda especie se describid originalmente del macho. Los siguientes taxa son citados

por vez primera para la Repiiblica de Colombia: Plocopsylla phyllisae Johnson, Leptopsylla segnis (Schonherr), Dasypsyllus

gallinulae perpinnatus (Baker), Tetrapsyllus comis Jordan, Polygenis pradoi (Wagner), P. thurmani Johnson, P. klagesi samue-

lis (Jordan and Rothschild), and Pulex simulans (Baker).

El taxon Rhynchopsyllus megastigmata Traub & Gammons, 1958, es considerado sindnimo de Rhynchopsyllus pulex

Haller, 1895.

Los registros contenidos en este trabajo permiten estimar que cerca de cuarenta y cuatro especies y subespecies de pulgas

son conocidas de Colombia y que de este numero mas de dos tercios esta'n citadas para la parte suroccidental del pai's.

Se incluye una clave para separar las pulgas del soroeste de Colombia y se ilustran aspectos diagndsticos de la mayoria de

los taxa.

Se expone un bosquejo de la historia y zoogeografi'a de la fauna de mamiferos de la parte suroccidental de Colombia,

asi' como tambien comentarios sobre las relaciones entre pardsitos y hospederos. La historia de la fauna de mamiferos de

Sur Amdrica probablemente se inicid al comienzo del peri'odo Terciario. Esta fauna primitiva consistid de marsupiales, cier-

tos edentados y ungulados primitivos que vinieron de Norte Amdrica. El posterior aislamiento entre Norte Amdrica y Sur

Amdrica logrd interrumpir pero no detener completamente un intercambio de animates mediante su traslado de isla a isla.

El intercambio fauni'stico fue restablecido cuando aparecid el puente Istmeno al final del Terciario. Tambidn es posible que

durante el peri'odo Cretacico y el comienzo del Terciario, Sur Amdrica recibid elementos fauni'sticos de Africa y de Australia.

La mayoria de las familias y gdneros de mamiferos suramericanos se originaron durante periodos de parcial o total aisla-

miento. En los tiempos actuates, en las tierras bajas de las costas colombianas del Pacifico aquellos mamiferos de amplia dis-

tribucidn sobrepasan las formas enddmicas. Estas faunas tienen una fuerte afinidad con aquellas de la regidn amazdnica y

de las tierras costeras de Panamd y Ecuador. Las restantes formas de la porcidn suroeste de Colombia estdn mds concentradas

en los Andes y contienen algunos elementos que probablemente se originaron en las tierras bajas.

La evolucidn y radiacidn de las pulgas de este territorio probablemente estaban correlacionadas con aquellas de sus hos-

pederos. Este fendmeno parece ser mds aparente en las pulgas que parasitan mamiferos pequenos, tales como los roedores

cricetinos. La fauna de pulgas del sector Pacifico es particularmente pobre, tanto en composicidn como en endemismo; mien-

tras que aquella de las montanas Andinas muestra mds elementos locales ademds de estar apreciablemente diver sificada. La

fauna de pulgas del suroeste de Colombia exhibe poca especificidad y varias especies pueden parasitar el mismo hospedero.

Por otra parte, existen especies de hospederos relacionados y no relacionados que comparten el mismo taxon de pulga. Es

evidente que la fauna de pulgas de las tierras bajas del Pacifico colombiano estdn relacionados con las de Panamd y las cuencas

de los rios Amazona y Orinoco. La fuerte relacidn de los mamiferos exist entes a traves de las cordilleras Andinas en Colombia

y Ecuador, se refleja en la afinidad que se observa en la fauna de pulgas.

Se discuten tambidn en este trabajo aspectos geograficos tales como topografia, geologia, suelo, clima y vegetacion que

conciernen al pertinente biota. El suroeste de Colombia se caracteriza por una diversidad de situaciones ecologicas que

Mammalian-Siphonapteran Associations

93

resultan de una topografih compleja y los regi'menes climaticos concomitantes. Las tierras costeras del Pacifico y las elevadas

tierras Andinas, adema's del Valle del Cauca, representan las principales areas geogrdficas de este territorio.

CONTENTS

Introduction 93

Physical Environment of the Departamento del Valle 94

Vegetation Formations of the Departamento del Valle 102

Historic and Zoogeographic Summary of the Mammals of

Southwestern Colombia 106

Key to the Fleas of Southwestern Colombia 113

Taxonomic Treatment of the Fleas of Southwestern Colombia . 117

Analysis of Host-Parasite Relationships 165

Acknowledgements 177

References 178

INTRODUCTION

The Departamento del Valle, one of several political divisions of the Republic of Colombia,

comprises an area of about 20,430 km2, located in the southwest sector of that country. It

contains contrasting ecological situations, including very wet rainforest of the Pacific lowlands,

western and eastern slopes of the Cordillera Occidental, basin of the Rio Cauca in the rainsha-

dow of Cordillera Occidental at 1000 meters, and the ascending western slope of the high Andean

Cordillera Central. This territory is bordered on the north by the departamentos del Choco and

Risaralda, on the south by the Departamento del Cauca, on the east by the departamentos del

Tolima and Quindio, and on the west by the Pacific Ocean.

The Departamento del Valle contains a great variety of plant and animal life, representing

more than half of the total number of species of mammals known to occur in Colombia. There

is little information about the fauna and the importance that many of these animals have as

positive or negative elements in the health of humans and domesticated animals. This need for

basic research in biology is indicative of the whole of Colombia as well as other areas of South

America.

During the past two decades, the Universidad del Valle, in conjunction with the Rockefeller

Foundation and the Tulane University International Center for Medical Research, have contri-

buted significantly to knowledge of arthropods and mammals associated with zoonoses in Col-

ombia (Cali Virus Faboratory, 1965, 1968, 1970). Through the study of ectoparasites collect-

ed by these research units, an interest developed for investigation of fleas and their mammalian

hosts in the Departamento del Valle and other regions of Colombia.

The importance of fleas in the epidemiology of plague, murine typhus and other diseases

is well known, thus primary consideration of these medically important arthropods is warran-

ted. In some instances, knowledge of the ectoparasitic fauna of an area may also provide un-

derstanding of relationships among animal host groups (Clay, 1951 ; Hopkins, 1942; Patterson,

1957). Although plague, which remains a major concern, has not yet been found in Colombia,

it is present in the adjacent countries of Ecuador, Venezuela, Peru and Brazil. There is, how-

ever, a strong possibility that the disease could become established in Colombia, since ideal

climatic factors occur in many areas of the country. In addition, both wild rodent species and

those commensal with man are present; these have been implicated elsewhere either as actual

or potential reservoirs of plague. Several species of fleas occurring in Colombia, such as Xeno-

psylla cheopis, the classical vector of plague, as well as Pulex irritans and certain species of

Polygenis, have been implicated in transmission of plague in other countries (Chabaud, 1947;

Quaest. Ent., 1977 13 (2)

94

Mendez

Macchiavello, 1948, 1954, 1958; Moll and O’Leary, 1945; Panamerican Health Organization,

1956; Pollitzer, 1954).

Presently, the Siphonaptera fauna of Colombia is poorly known. In a survey prior to be-

ginning this study, I found that Colombian material was not well represented in collections

of several institutions, including the British Museum (Natural History) and the United States

National Museum of Natural History.

The literature contains few reports concerning Siphonaptera from Colombia. Dunn (1929)

and Patiho-Camargo (1940) briefly mention some common species. Fuller (1942) and Macch-

iavello (1948) gave several Colombian records of fleas. Costa Lima and Hathaway (1946) pre-

sent information on several Colombian species. Gast Galvis (1950) in his list of fleas from Col-

ombia considered 19 species and subspecies. Johnson (1954 and 1957) has most significantly

contributed to our knowledge of the fleas of Colombia. In 1954, she described a new species

of Pleochaetis from that country. Her outstanding monograph “Fleas of South America”,

published in 1957, contains several new distributional records of species for Colombia and,

in addition, the description of six new species. Mendez (1968) described a new genus and spe-

cies of Colombian flea, and more recently, Mendez and Hanssen (1975) reported a new Col-

ombian taxon discovered in the Departamento del Meta. Tamsitt and I. Fox (1970) recorded

Rhynchopsyllus pulex Haller.

This work is largely based on information obtained from collections made in the Departa-

mento del Valle and neighboring political divisions of southwestern Colombia, such as Nar-

ino, Cauca, Putumayo and Huila. In view of the ecological affinities of these territories and

the similarity of their mammalian fauna, it is likely that these areas have virtually the same

ectoparasitic fauna.

The present account contains descriptions of four new species of Polygenis Jordan. The fe-

male of Sphinctopsylla diomedes Johnson, known previously from the male, and the male of

Ctenidiosomus traubi Johnson, are also described. A key to species of fleas known or presump-

tively existing in the southwestern portion of Colombia, and new records for the country, are

also presented. Rhynchopsyllus megastigmata Traub and Gammons is considered here to be a

synonym of R. pulex Haller. Conventional figures for the majority of the species concerned

are included.

The nucleus of the material was from collections made by Harold Trapido while engaged

in virus research sponsored by the Rockefeller Foundation. Additional specimens were collec-

ted personally or obtained from the Universidad del Valle and other sources.

PHYSICAL ENVIRONMENT OF THE DEPARTAMENTO DEL VALLE

Topography

Inasmuch as most of the siphonapteran material used in this study is from the Departamen-

to del Valle, the following geographical discussion is limited to this territory. No other poli-

tical division of Colombia displays such a diversity of geographic features and, consequently,

ecological conditions. The topographic information given below is derived mainly from Es-

pinal (1968), Sanchez (1965) and Sauer (1950).

Generally, the physiographic and faunistic elements of this area are characteristic of the

southwestern zone of Colombia, which consists of the Cauca Valley territory, surrounding

mountains, and Pacific coastline within the limits of the Departamentos del Valle, Cauca,

Huila, Narino, and Putumayo.

The landscape is dominated by the Western and Central ranges of the Andean mountain

system. These mountains contain some of the higher peaks of Colombia. The Western Cor-

dillera extends north-northeastward parallel to Central Cordillera and is almost parallel to the

Mammalian-Siphonapteran Associations

95

REPUBLIC OF COLOMBIA

Fig. 1. Map of Colombia.

Quaest. Ent., 1977 13 (2)

96

Mendez

Pacific coast. It is the lowest of the three ranges forming the Andean system. The Central

Cordillera in the eastern sector of El Valle contains high peaks such as the mountains of Huila

and Barragan, which exceed 3000 meters.

Several rivers and numerous streams form a Pacific watershed which originates in the moun-

tains and empties into the Pacific Ocean. The more important rivers are the Anchicaya, Dagua, J

Naya and Cauca. This last river is the largest and courses through the Departamento from south :

to north.

Mangrove swamps flooded by high tides are along the shoreline of the Pacific coasts. Warm

tropical rain forests follow the swamps and occupy extensive inland zones of dense vegetation,

which is impenetrable in many areas. The rich plant life, water, and cover, offer optimal con-

ditions for an abundance of animal forms. The pocket-like arid valley of Dagua, however, with

its grassy and brushy cover, introduces a contrasting zone in this lowland territory.

A subtropical zone of humid temperate climate follows the lowland cloud forests. This tem-

perate zone is characteristic of the middle slopes of the mountains, where vegetation is rich

but not very luxuriant. Many of the faunal elements of these subtropical forests are similar to

those found in lowlands, since many species evolved from ancestors which moved to the upper

zones. Apparently climate and other geographical conditions have been limiting factors in es-

tablishment of some species that have more restricted tolerances.

The Cauca Valley, an agriculture area of the temperate zone, is a narrow isolated region con-

sisting of approximately 400,000 hectares between the Western and Central Cordillera. It is

nearly 160 kilometers long and only 12 kilometers wide, and is drained by the Cauca River.

Although much of its territory is occupied by pasture, mainly Para ( Panicum barbinode ) and

Guinea {P. maximum ) grasses, the Cauca Valley represents the most fertile and productive

agricultural land of Colombia.

Lush montane cloud forests occupy extensive zones of moderate and high elevations (from

below 1000 to about 3000 meters) throughout most of the Central and Western Cordilleras.

The uppermost reaches of the mountains, from over 3000 meters, are largely unforested, con-

sisting primarily of extensive grass plains characteristic of paramos (such as Las Hermosas,

Chinche, Miraflores and Barragan). These areas support more selected types of plants and ani-

mals; indeed the paucity of animal life in the paramos is directly correlated to the poor diver-

sity of cover.

As in other areas of Colombia, the Departamento del Valle is suffering from much defores-

tation, due primarily to agricultural development, hydroelectric projects and raising of cattle.

Geology

By virtue of its rock composition, Colombia comprises two geological regions: the Oriental

plains; and the Andean geosynclinal regions (Biirgl, 1961). Only the latter region is in south-

western Colombia, including the Departamento del Valle. The vast Oriental plains are in east

and southeastern Colombia.

The Andean geological region was apparently submerged during long periods from the be-

ginning of the Cretaceous, and accumulated large deposits of marine, continental and volcan-

ic sediment. The complicated tectonic movements experienced by these lands were responsi-

ble for formation of the present Andean Mountains, which constitute the “backbone” of

Colombia.

The Andean geosynclinal region was consolidated before the Cambrian period, and includes

the following mountain systems: 1. Central Cordillera, 2. Western Cordillera and Coastal Cor-

dillera (Serrania de Baudo), and 3. Eastern Cordillera. The Lower and Middle Magdalena Val-

ley basins separate the Central and Eastern Cordillera. The Lower Valley contains non-marine

Mammalian-Siphonapteran Associations

97

Tertiary rocks, while, the Tertiary reservoirs of the Middle Valley are non-marine. Western and

Central Cordilleras are separated by the valleys of the Upper Cauca and Upper Patia rivers. Other

mountains of Colombia, such as the Santa Marta Mountains, Perija Mountains and the Pacific

Coast Range, have affinities with the Andean system.

The coastal zone of the Andean Region is represented by the Bolivar Geosyncline, a lowland

area of Tertiary Marine formation, west of the Andean mountains and extending from South-

ern Ecuador to the Gulf of Uraba in northern Colombia. This strip of land has been interpre-

ted as a seaway which apparently permitted movement of terrestrial animals during periods

ranging from upper Cretaceous times to Recent (Nygren, 1930; Hershkovitz, 1966). No marine

formations from the Upper Cretaceous have been found in this zone.

The Eastern Cordillera displays abundant deposits of Cretaceous and Tertiary rocks, with

little or no recent volcanic elements. In addition, Jurassic, Triassic and Paleozoic rocks are

found in these mountains. Western and Central Cordilleras, as well as the Pacific Coast Range,

are primarily formed of igneous and metamorphic rocks, having only subordinate sedimentary

beds. Each Cordillera has several high volcanic peaks.

The Precambrian sedimentary history of Colombia is not known (Jacobs, Biirgl and Conley,

1963). According to these authors, during a great part of Cambrian and Ordovician time, the

actual territory of the Eastern and Central Cordilleras, and at least the western aspect of the

Llanos and Putumayo-Caqueta lowlands, were occupied by seas. Later, the area came under

the influence of volcanic and diastrophic activity, which destroyed the Cambrian-Ordovician

marine deposits. Rock shields, characteristic of the Andean system during the Cambrian, ap-

parently were greatly disturbed by erosion and their detritus filled marine and terrestrial de-

pressions of the region. The most important fossils from the lower Paleozoic in Colombia are

brachiopods, trilobites and graptolites. The territory west of Central Cordillera shows no in-

dication of Paleozoic marine sediment.

Some Middle Ordovician fossils have been found in Colombia. However, Upper Ordovician

and Silurian apparently are not represented by fossiliferous layers in this country.

Fossils from the Devonian of Colombia are represented by brachiopods, bryozoans and tri-

lobites, among other invertebrates. Many plant and animal fossils, primarily marine forms, are

known from the Carboniferous.

Evidence obtained from fossils indicates that the lower Carboniferous sea invaded the eas-

tern part of the Andean Region, which was covered by sediment thereafter. During the upper

Carboniferous, the sea gradually retreated and large semi-swampy forests moved to sections

previously occupied by water. Abundant Permian fossils have been found in only a few areas

of Colombia, primarily at Serrania de Perija. They represent sponges, crinoids, brachiopods,

gasteropods, cephalopods, and other marine animals. The eastern portion of the Central Cor-

dillera contains Marine Upper Triassic rock (the Payande Formation), and represents a com-

bination of sandstone, limestone and shale. The Eastern and Central Cordilleras contain low-

er Jurassic sedimentary rocks, principally of continental origin.

Evidently much igneous activity occurred in the Andean Region during the Mesozoic. Also,

it is apparent that the major part of western Colombia was occupied primarily by seas during

most of Late Jurassic and Cretaceous time. Fossils from these periods are scant and consist

mainly of ammonites and other molluscs. The Cretaceous Colombian and Peruvian faunas dis-

played a strong relationship with those of Southern Europe (Olsson, 1956). Considerable tec-

tonic movements that occurred during Late Cretaceous and early Tertiary, produced constant

changes in the landscape of this region and in the formation of marine and non marine depo-

sits. The Tertiary Continental deposits contain an excellent vertebrate fauna, including num-

erous mammals. There is also evidence of considerable faulting and folding during the late

Paleocene.

Quaest. Ent., 1977 13 (2)

98

Mendez

Throughout the Eocene, land forms of this area were somewhat different from the present.

However, the principal elements of the Andes and the Pacific Coast basin originated during this

period, and Weeks (1947) points out that the Bolivar Geosyncline introduced important chan-

ges in Western Colombia and Ecuador. Much tectonic and volcanic activity took place during

the Eocene and Oligocene (Hammen, 1961; Vuilleumier, 1971). Very few fossil vertebrates of

Eocene age are known from Colombia (Stirton, 1953). Layers corresponding to the Eocene and

Oligocene are richer in Foraminifera, molluscs and other marine fossils. The Cauca Valley is

particularly rich in Oligocene marine rocks, mainly consisting of algal, and foraminiferal lime-

stone. This area also has coal-bearing Tertiary jocks, that probably originated during the Mio-

cene or before.

A large number of fossil vertebrates has been discovered in Miocene deposits. The fossil mam-

mal fauna of this period is very interesting and contains some relics of families believed to have

disappeared in earlier times. Numerous findings of mammalian Pleistocene fossils have taken

place in Colombia (Stirton, 1953; Patterson and Pascual, 1968). Many of these fossils represent

species that became extinct at the end of the Pleistocene. Jacobs, Biirgl and Conley (1963) con-

sider that the most recent stage of tectonic movement and volcanic activity was initiated in

the early Miocene and continued into Recent time, particularly affecting the Central Cordil-

lera and the southernmost part of the Western Cordillera. Some of the volcanoes existing to-

day are still active.

Soils

The following considerations of soil distribution in the Departamento del Valle are based

primarily on two major sources: the FAO/UNESCO Soil Map of South America (1961), and

the account by Beek and Bramao (1968).

A great deal of work has been done in South America to determine distribution of major

soils as important patterns in development of agricultural zones and exploitation of minerals

and other natural resources significant to the general economic progress of the continent. How-

ever, despite the knowledge that has been accumulated during many years, the study of struc-

ture, composition, and distribution of South American soils is far from complete.

The nature of the soils of the Departamento del Valle is linked to patterns of ecological

factors that govern the life zones, such as climate, geomorphology, topography, and vegeta-

tion. Departamento del Valle, and other areas of southwestern Colombia, from which mater-

ial used in the present study has been collected, consist of Lowlands and Andes soils, two of

the major structural elements of soil distribution that have been established for South Ameri-

ca. Uplands soils, the other element considered in this segregation, are found in the eastern

part of South America. Criteria assumed for establishment of these general categories are based

on a complex association of factors such as geography, climate, vegetation, physiography, etc.

The Pacific Coastal Lowlands form a high portion of the western side of the Departamento

and are primarily characterized by contiguous areas of tropical evergreen and deciduous for-

ests. The alluvial soils predominating in this territory are stratified with little organic matter

and little profile development. The Pacific littoral is particularly dominated by dense mangro-

ve forests, interspersed with swamp forests. This zone contains Quaternary marine and fluvio-

marine deposits with alluvial plains and terraces, estuarine and delta alluvial deposits, and lo-

cal areas of coastal sand dunes.

The Northern Andes is the other soil region of the Departamento del Valle. The soil com-

position is more diverse and the profile of the area interrupted by western and central ranges

of the Andean mountains. These are separated by the Cauca Valley, a strip of land which also

includes areas of the Departamento de Cauca, Narino, Caldas and Antioquia.

RAIN SHADOW IN TRANSECT OF DEPARTAMENTO DEL VALLE

Mammalian-Siphonapteran Associations

99

Quaest. Ent, 1977 13 (2)

Fig. 2. Rainshadow in transect of Departamento del Valle.

100

Mendez

In the western sector of the Northern Andes region of the Departamento del Valle, volcan-

ic rocks are the most important elements of the soil structure. This condition reflects the vol-

canic origin of these mountains. Most soils in areas of moderate elevation in the northern An-

des are Laterosols, derived from volcanic material. Laterosols are dark colored surface soil with

lighter subsoil, and, in addition to being slightly acid, contain a high amount of organic matter.

Because of their low fertility level, they have limited use for farming. Extensive areas of the

northern Andes are represented by Andosols. Such soils consist primarily of volcanic ash with

dark surfaces, combined with organic matter and minerals such as nitrogen, phosphorus, cal-

cium and potassium. Those that are not too acid are relatively fertile and excellent for grow-

ing different crops. Sectors of the Departamento del Valle, dominated by Andosols, have ex-

cellent pasture lands and areas devoted to agriculture.

Dark paramo soils, which may be derived from heavy clays, perhaps of glacial origin are

found adjacent to Andosols. They consist of some volcanic ash and are characterized by a high

degree of acidity and paucity of nutrients. The paramo lands have high humidity, low temper-

ature, the vegetation is poorly diversified and consists of pastures and forests of secondary

growth.

The northern part of the Departamento del Valle is characterized by Reddish Brown Lat-

eritics. These soils contain dark, reddish brown, granular clay surface soil, with yellowish-brown,

clay subsoil. Aluminum silicate, its principal mineral compound, is mixed with iron and other

ingredients of inorganic and organic origin. They occur below 2000 meters on which are pri-

mary and secondary forests in addition to pastures, coffee plantations and other cultivated

areas.

Climate

The complexity of its geography contributes to the variety of climates existing in the De-

partamento del Valle, since these elements are intimately associated. It is interesting to note

that all four of the climatic regions outlined by Eidt (1969) for South America, are represen-

ted in the Departamento del Valle, namely, tropical rain, temperate, arid and tundra.

Tropical rain climates are confined to the Pacific territory, from sea level to the mountain

bases reaching an elevation of close to 1 ,000 meters. These areas are hot and humid, with tem-

perature ranges from 24°C to 30°C. The annual rainfall is heavy, exceeding 760 cm a year. It

rains almost daily, thus this area does not have a true dry season. Humid warm winds of the

Pacific Ocean and Andean chain contribute to the heavy precipitation; Figure 3, adapted

from Espinal (1968), illustrates the rainshadow influence. According to Espinal (1968), with-

in the Western and Central Cordillera there are two rainy periods during the year: the first

from April to June, and the second from September to November. Between these wet sea-

sons there is notably decreased precipitation.

Temperate climates are characteristic of the subtropical areas extending from 1,000 m to

2,000 m, which have a temperature range of 18°C to 24°C. These areas are the first table lands

above the lowlands, and are influenced by higher masses of the Andes and by action of two

kinds of winds, interpreted as mountain-valley breezes and land-sea breezes. These winds af-

fect climate and precipitation of the Cauca Valley system, and rivers, and other subtropical

areas of this territory.

The eastern part of the Departamento maintains a cooler climate typical of areas above

2,000 meters, (in which various paramos are found). The northern sector of the Central Cor-

dillera contains an arid zone, subjected to strong winds. This zone contrasts with the cloudy

and rainy forests dominating the Andean landscape. Some of the higher peaks of Central Cor-

dillera are cool and damp paramos where the temperature is below 12°C.

Mammalian-Siphonapteran Associations

101

SOIL MAP OF THE DEPARTAMENTO DEL VALLE

Fig. 3. Soil map of the Departamento del Valle.

Quaest. Ent., 1977 13 (2)

102

Mendez

VEGETATIONAL FORMATIONS OF THE DEPARTAMENTO DEL VALLE

Most of the information presented below, including Fig. 4, is based on Espinal (1968), and

Espinal and Montenegro (1963). These studies were made according to the classification esta-

blished by Holdridge (1947), which considers aspects of temperature and humidity in analysis

of life zones.

Ecological parameters of rodents and other mammals, as well as of the ectoparasites affect-

ing them are highly dependent on biotic conditions, soils, climates and other factors. Vegeta-

tional formations are therefore of primary importance in understanding habitat requirements

and distribution patterns of these animals. However, such formations in time and space usu-

ally grade into one another and therefore are not permanent. In addition, many animals and

plants are able to tolerate a moderate range of climatic conditions.

A brief description of all of the vegetational zones outlined for the Departamento del Valle

follows.

Tropical Very Dry Forest

This formation is in two areas. One is Loboguerrero, at the upper part of Rio Dagua in the

center of the Departamento. The other region represents a flat belt of forest platform in the

Central Cordillera, extended from Cali to near San Francisco at the left shore of the Cauca

River, reaching an elevation of 1200 to 1400 m. The prevalent climate of this zone is dry and

the mean temperature is over 24°C. Mean rainfall during the year ranges from 500 to 1000 mm.

Characteristic plants of this zone are figs {Ficus spp.), Vachelia farnesiana, spurge {Euphorbia

caracasana ), Pithcellobium dulce, Desmanthus virgatus, Achatocarpus nigricans, mesquite {Pro-

sopis juliflora ), Jatropha go ssypii folia, Fugatera pterota, Ocimum micranthum, Heliotropium

sp., Lantana canescens, Citharexylum sp., Portulaca pilosa and Talinum paniculatum.

Tropical Dry Forest

It is distributed along the Central Valley crossed by the Cauca River and along a strip of

land surrounding the Tropical Very Dry Forest of Loboguerrero. Mean temperature of the

area is over 24°C and rainfall fluctuates between 1000 and 2000 mm. The characteristic vege-

tation of this zone grows in lands not over 1000 m high and consists of cabuyas {Furcraea sp.),

Croton sp., Turnera almifolia, Cephalocereus colombianus and other plants.

Tropical Moist Forest

This zone occurs in three areas below 1 000 m in the upper half of the Departamento. They

are the canyon of Rio Garrapatas near El Cairo and Versalles, a strip near Rio Dagua and pro-

bably an area in the mid portion of Rio Calima. The mean temperature is higher than 24°C

and the mean rainfall is from 2000 to 4000 mm. Some plants found in this type of forest are

the balsa {Ochroma lagopus ), roble {Tabebuia pentaphylla ), and hogplum {Spondias mombiri).

Tropical Very Wet Forest

This formation extends from the basin of Rio Anchicaya as a belt that runs from south to

north along the Western Cordillera. The area is dominated by a variety of plants such as ceiba

{Ceiba pentandra), balsa {Ochroma lagopus), black rubber {Castilla elastica), guarumos {Cec-

ropia spp.), calabash {Crescentia cujete), cedars {Cedrela spp.), annato {Bixa orellana ), sand-

box {Hura crepitans), and others. These lands are not above 1000 m. The temperature exceeds

24°C and there is a mean rainfall of 4000 to 8000 mm per year.

Mammalian-Siphonapteran Associations

103

VEGETATIONAL FORMATIONS

TROPICAL VERY DRY FOREST

tropical dry forest

III TROPICAL MOIST FOREST

TROPICAL VERY WET FOREST

[ | TROPICAL RAIN FOREST

| SUBTROPICAL DRY FOREST

jT' : F 1 SUBTROPICAL MOIST FOREST

j;- SUBTROPICAL VERY WET FOREST

j ' "-"j SUBTROPICAL RAIN FOREST

LOWER MONTANE DRY FOREST

jjfeSI LOWER MONTANE MOIST FOREST

h I LOWER MONTANE VERY WET FOREST

!££££§ LOWER MONTANE RAIN FOREST

MONTANE VERY WET FOREST

MONTANE RAIN FOREST

PACI F

OCE A

DEPARTAMENTO DEL VALLE

Fig. 4. Vegetational formations of Departamento del Valle.

Quaest. Ent., 1977 13 (2)

104

Mendez

Tropical Rain Forest

This is the largest of the vegetational formations of the Departamento, extending from the

very wet lowland coast line to the margins of the Western Cordillera. Dense mangrove forests

consist primarily of red mangrove ( Rhizophora brevistyla) in association with black mangrove

(. Avicennia marina ), white mangrove ( Laguncularia racemosa), and buttonwood ( Conocarpus ).

Inland forests, beyond the Pacific littoral, contain a diversity of plants such as guarumo ( Cec -

ropia sp.),wild figs (Ficus), mammagua (Inophleum armatum), membrillo ( Gustavia superba),

Acacia melanoceras, sensitive-plant (Mimosa), cashew (Anacardium excelsum), balsa (O chroma

spp.), and many others. The tropical rain forests are below 1000 m, in a climate with mean

temperature above 24°C and mean annual rainfall exceeding 8000 mm.

Subtropical Dry Forest

This includes isolated areas on the eastern half of the Departamento. The major area is

along the oriental baseline of the Western Cordillera from Cali to San Francisco. One patch

is in El Dovio’s ravine and two other patches occur in the Central Cordillera. Typical plants

in this zone are figs (Ficus spp.), fiques (Fourcraea sp.), mosquero (Croton sp.), big belly tree

(Wigandia caracasana), indigo (Indigobera sp.), cuipo (Cavanillesia platanifolia), among others.

It registers a mean temperature of 17°C to 24°C and the mean annual rainfall is between 500

to 1000 mm.

Subtropical Very Humid Forest

This formation encompasses very humid areas of the Western and Central Cordillera having

an elevation between 1 100 to 1900 m. The mean temperature is from 17°C to 24°C and the

annual rainfall fluctuates between 2000 to 4000 m. The vegetation of this zone is represented

by guamo (Inga spp.), guayacan (Tabebuia chrysantha), balsa (Ochroma lagopus), guarumo (Cec-

ropia), yaragua (Melinis minulti flora), pigs fern (Pteridium aquilinum), fox tail (Andropogon

sp.), and others.

Subtropical Moist Forest

It is distributed as an extensive area surrounding the dry Rio Cauca valley and extending

to the edges in the Western and Central cordilleras. Other patches of this zone are located in

areas surrounding El Dovio and in the upper portion of the rivers Dagua, El Carmen and El

Treinta. The zone reaches an altitude between 1 100 to 2000 m. The mean temperature ranges

from 17°C to 24°C, while the annual rainfall is between 1000 to 2000 mm. Typical plants of

this zone are the macedero (Trichantera gigantea), surrumbo (Trema micrantha), cordoncillo

(Piper adunCun), balsa (Ochroma lagopus), iraca (Carludovica palmata), trompet (Bocconia

frutescens), coralito (Hamelia patens), rubber (Ficus sp.), chayote (Sechium edule) and many

others.

Subtropical Rain Forest

It is particularly represented by a strip of watered land of the Western Cordillera with ele-

vations between 900 to 1900 m. The humidity in this vegetation formation is very high and

the temperature reaches from 17°C to 24°C. The annual rainfall exceeds 4000 mm. This zone

combines some virgin forests as well as agriculture lands. Some of its characteristic plants are

platanillos (Heliconia), ferns (Gleicheniaceas), guarumo ( Cecropia ), rubber (Castilla elastica),

paco (Cespedesia macrophylla), pejibaye (Guilielma gasipals), and avocado (Persea americana).

Mammalian-Siphonapteran Associations

105

Lower Montane Dry Forest

This formation is limited to an area of high elevations (from 2000 to 3000 m) of Barragan

in the Central Cordillera. The region possesses a dry climate with a mean temperature of 1 2°C

to 17°C, and an annual rainfall of 500 to 1000 mm. The native vegetation is poor and consists

primarily of capers ( Cassia sp.) and mosqueros ( Croton sp.); cultivated species include maize

( Zea mays), onion {Allium cepa), wheat {Triticum), and potatoes {Solanum tuberosum).

Lower Montane Moist Forest

This region includes two areas in the Central Cordillera, where the altitude is from 1 800 to

3000 m. It is a cold formation, humidity is high and mean temperature varies from 12°C to 17°C.

Annual rainfall is over 4000 mm. Some of the plants found in this territory are the following:

carbonero {Befaria sp.), chilco {Baccharis sp.), capers {Cassia sp.), and mortino {Miconia albi-

cans).

Lower Montane Very Wet Forest

This formation occupies extensive subtropical lands of the Western and Central Cordillera

at elevations between 1800 and 3000 m. The cool climate registers a mean temperature of 12°C

to 17°C. Annual precipitation is calculated to be between 2000 to 4000 mm per year. These

forests contain many different plants such as guarumos {Cecropia spp.), lichens {Cora pavonia,

Cetraria sp.), mosses {Polytricum sp.), horse tail {Equisetum sp.), encimo {Weinmannia balbi-

siana), cascarillo {Ladenbergia), etc.

Lower Montane Rain Forest

This is a wet formation distributed as occasional areas in the Western and Central Cordillera.

The elevation ranges from 1800 to 2900 m and the mean temperature varies from 12°C to

1 7°C. The annual rainfall registers over 4000 mm. Typical plants of this zone are the white

guarumo {Cecropia), capers {Cassia sp.), quassia {Quassia sp.), cherry {Freziera sericea), ced-

rillo {Brunellia sp.), lulo {Solanum quitoense), horse tail {Equisetum bogotense), berries {Ru-

bus sp.), strawberries {Fragaria sp.), and others.

Montane Very Wet Forest

This formation occupies a broad portion of the typical paramo distributed from Santa Lucia

to Barragan in the Central Cordillera. It has an elevation above 3000 m and a mean temperature

from 6°C to 12°C. The annual rainfall fluctuates between 1000 to 2000 mm. The plant-life is

represented by frailejones {Espeletia), capers {Cassia sp.), espadero {Rapanea sp.), charcoal mak-

er {Befaria sp.), morteno {Hesperomeles sp.), grasses such as Calamagrotis and Festuca, etc.

Montane Rain Forest

These areas are very humid, over 3000 m in altitude, localized in the Western and Central

Cordillera. Mean temperature is between 6°C and 12°C and rainfall is over 2000 mm. This type

of vegetation formation contains cultivated territories as well as undisturbed forests. Among

the plants found in these lands are frailejones {Espeletia sp.), blackberries {Rubus sp.), encen-

illo {Weinmannia sp.), Senecio sp., Vaccinium sp., cherry-tree {Prunus sp.) and Bacharis sp.

Quaest. Ent., 1977 13 (2)

106

Mendez

HISTORIC AND ZOOGEOGRAPHIC SUMMARY OF THE MAMMALS OF

SOUTHWESTERN COLOMBIA

The ecological parameters of mammals and their ectoparasites in Colombia and other South

American countries is of interest to investigators concerned with a variety of zoonoses linked

to those animals. However, little has been reported on their biology and distribution. Even sys-

tematic studies of these vertebrates are not complete and names and taxonomic status of a

number of forms are not settled.

To understand the origin and distribution of mammals presently occurring in southwest

Colombia one must read an array of discussions by authorities on South American mammals.

To fit the limited scope contemplated here, I have tried to be selective in interpretation of

possible events that, in my opinion, have a logical foundation. References concerning evolu-

tion and zoogeography of South American mammals are Hershkovitz (1962, 1966, 1969, 1972);

Keast (1968); Loomis (1914); Patterson and Pascual (1963, 1968); Savage (1974); Scott (1937);

Simpson (1940, 1943, 1945, 1950, 1969); Stirton (1950).

The theory of plate tectonics has introduced a new argument for exploring the origin and

distribution of Latin American fauna and flora (Raven and Axelrod, 1975; Valentine and

Moores, 1974). According to this theory, movement of land masses or plates constituting the

earth’s crust produces cataclysms and volcanic action with subsequent complicated modifica-

tions of the elements involved. As applied to the southern continents, and particularly to

South America, some intriguing perspectives occur with regard to the origin and affinities of

the biota. In view of the geological facts, during the Middle Cretaceous South America was

connected with Africa, and via Antarctica, with Madagascar, India and Australia. It was sep-

arated from those eastern lands over an extended period of time. Apparently the connection

with Australia lasted longer. It has been speculated on this basis that an exchange of fauna,

at least in part by island-hopping, took place between Australia and South America until early

Oligocene. On the other hand, a more effective connection between South America and Afri-

ca existed until the end of the Cretaceous.

Probably during late Eocene, interchange of tropical and warm-temperate animals between

South America and Africa was greater than between Africa and North America. Some groups

of animals, such as the river turtles, family Pelomedusidae, which occur in South America,

Madagascar and Africa, also seem to afford a logical basis for this assumption. It is also belie-

ved that routes for the movement of cool-temperate organisms between southern South Am-

erica and Australia existed until nearly 4 million years ago. The argument appears to be sub-

stantiated by certain elements of the flora of angiosperms common in Africa and South Am-

erica.

Events of a different nature, primarily geological and climatic, occurring in the millions of

years since separation of the southern continents during the Cretaceous, have interrupted fau-

nal development and geographical ranges of taxa in South America. This has introduced many

gaps that obscure the entire historical sequence.

Geological changes during the Cenozoic, are very important in attempting to understand

some of the general history of the mammals of the Departamento del Valle, and other parts

of western Colombia.

The mammalian faunal structure of South American began to evolve with primitive forms

recorded from the Paleocene, such as marsupials, members of the orders Condylarthra, Not-

ungulata, Litopterna and other ungulates (Hershkovitz, 1968; Patterson and Pascual, 1968;

Simpson, 1950).

Apparently during the beginning of the Tertiary, the Middle American bridge permitted

mammals to move to and from North and South America. The many fossil discoveries from

Mammalian-Siphonapteran Associations

107

southern South America have revealed that, early in the Paleocene, the most important nuclei

of South American mammals were concentrated in this portion of the continent. The earliest

mammal records from northern South America and Middle America are represented only by

marsupials and condylarths.

During the Eocene and Oligocene epochs, and presumably during most of Cenozoic, North

America was isolated from South America by a sea barrier since the connection represented at

present by the Isthmian link did not exist. Fossils of monkeys and caviomorph rodents have

been found in Oligocene beds of South America. However, the evolution and dispersal of these

mammals prior to this time remain a mystery. Hershkovitz (1968) and Simpson (1943, 1950)

believe that these mammals invaded South America probably as island-hoppers in the early

Oligocene. According to Raven and Axelrod (1975), it is more probable that primates and cav-

iomorph rodents reached South America and Africa during the Oligocene. It is also probable

that the ancestors of those mammals arrived in this Southern continent during the Cretaceous.

However, fossils from this epoch that would support this theory have not been found in South

America.

From the fossil history, it is evident that armadillos, ant-eaters and ground sloths inhabited

South America since the Tertiary and were probably distributed in areas of mild climate, at

least close to our present tropical conditions. It is possible that they originated from a Creta-

ceous stock. According to Simpson and other authors, they gradually moved into South Am-

erica when the Middle American land bridge was re-established in the late Pliocene. There is

no indication that these mammals migrated between South and North America before this

time, with the probable exception of the ancient armadillos. Fossil armadillos have been dis-

covered in early Tertiary beds of North America (Kiirten, 1969). Hershkovitz (1972) discus-

sed the probability that migration of terrestrial animals started during the period of isolation

of the continents. Other authors (Scott, 1937; Simpson, 1940, among them) maintain that

during the period of isolation of the continents, the South American rodent fauna was appar-

ently represented only by caviomorphs. They also believe that the North American fauna then

contained myomorph and sciuromorph rodents, but no caviomorphs. According to Hershkov-

itz (1972) the tribe Sigmodontini, of the murid subfamily Cricetinae, distinguished by the

possession of a complicated glans penis with a three digitated baculum, originated in this con-

tinent from ancestors that rafted there from Africa. From South America they subsequently

spread northward into North America and Eurasia. On the other hand, the Peromyscini, char-

acterized by a simple glans penis, with an unbranched and normally elongated baculum, is

essentially North American and apparently replaced the Sigmodontini, when they probably

disappeared from boreal America during Oligocene. The complex-penis type Cricetinae seem

to have more recently invaded Central and North America. This hypothesis appears to be sup-

ported by some ectoparasites, such as fleas, occurring on these mammals. The flea genus Poly-

genis, which has moved into Middle America with its complex-penis type Cricetinae and Cavi-

omorph hosts from South America, is a good indicator of such an event (Wenzel and Tipton,

1966).

In South America, as well as in other areas of the world, the early history of bats is obscure

because of the paucity of fossils. However, it is estimated that they probably appeared in this

continent after the Paleocene (Patterson and Pascual, 1972). Hershkovitz (1969) believes that

bats were probably well established in north-western South America and the Isthmian region

during most of the Tertiary. Storms may have played an important role in interchange of bats

between the Americas when they were separated.

During Pleistocene times, South America received from North America a diversity of mam-

mals such as horses, mastodonts, tapirs, peccaries, camels, deer, some cricetine rodents, squir-

rels, rabbits, canids, bears and other carnivores. Faunal interchange between North and South

Quaes t. Ent., 1977 13 (2)

108

Mendez

America during the late Tertiary, as well as radiation of many mammal groups has been the

subject of much speculation and even today no causal explanation is generally accepted. How-

ever, it appears that the majority of the families and genera of South American mammals or-

iginated on this continent when it was completely or partially isolated.

At the present time the northern sector of South America contains the major portion of the

mammalian fauna, while the southern sector is exceedingly poor in number of species (Fittkau, j

1969; Osgood, 1942). Information based on Baker (1967), Cabrera and Yepes (1940) and other j

authors, as well as on personal data, reveals that the major portion of southwestern Colombia, j

corresponding to Pacific Coastal Lowlands, maintains a number of ubiquitious mammal spe-

cies in addition to those forms possessing a more restricted distribution. The remaining mam-

malian fauna of the southwest region of Colombia is fundamentally represented by a limited

number of Andean elements.

The recent biogeographical approach for the Neotropical region presented by Muller (1973)

is followed for the brief zoogeographic description of mammals of the Departamento del Valle

and the rest of southwestern Colombia. Muller based his interpretation of zoogeographical pat-

terns on dispersal centers devised from comparative studies of plant and animal distribution.

Colombia and several other territories of tropical America belong to the Brazilian Subregion

of the Neutropical Region (Hershkovitz, 1958). The complexity of ecological situations in

this nation has provided for the existence of a diverse mammalian fauna. Many areas, particu-

larly those humid zones rich in vegetation and not yet disturbed by man, offer optimal con-

dition for some populations of mammals, especially rodents and bats. They possess good re-

productive potential and have been able to occupy a number of ecological niches through the

development of specialized features. Those lands characterized by drier conditions and a low

productivity of plants, particularly of fruit trees, as well as those somewhat disturbed by man,

show, as a rule, less diversity and smaller populations of mammals. In a broad sense, sylvan

species are not too selective in their food and habitat requirements, while pastoral species seem

to display a higher level of adaptation by their food habits and locomotor organs.

Among the Neotropical dispersal centers outlined by Muller, the following directly concern

the Departamento del Valle and other major portions of southwestern Colombia: 1. the Cauca

Center, 2. The Colombian Montane Forest Center, 3. The Colombian Pacific Center, and 4.

The North Andean Center. For this discussion it is convenient to follow these zoogeographic

divisions. Figure 5 has been prepared using as a source the map given by Muller (1973).

The Cauca Center. — As defined by Muller, this zone is represented by the Cauca and Patia

valleys, between the west and central Cordilleras of Northern Colombia, and are separated from

each other by the Popayan plateau at an elevation of 1750 meters. Like many other areas of

Colombia, this territory is being progressively deforested; however, it still harbors many mam-

malian species, some of which are widely distributed in other areas of the Continent. Some of

the faunal elements that previously existed in this territory probably disappeared many years

ago. Faunal indicators of this zone are the wooly opossum, Caluromys derbianus derbianus,

and red squirrel, Sciurus granatensis valdiviae, which are found in the Cauca and Patia valleys.

The ancestors of these mammals evidently came from the north. According to Muller, the

slight differentiation between the populations of these subspecies in the two valleys suggests

that the Popayan plateau is not a strong barrier to species adapted to open habitats. In addi-

tion, the forest biome of the southern part of the Cauca Valley did not prevent a spread of

fauna during post-glacial times.

Available information indicates that South America experienced drastic climatic changes

during the Quaternary (Hammen, 1961 ; Patterson and Pascual, 1963; Vanzolini, 1973). Haffer

(1967) considers that climatic changes during the Pleistocene and post-Pleistocene had more

influence than orogenic events on the fauna west of the Andes. He also states that isolation

Mammalian-Siphonapteran Associations

109

Fig. 5. Dispersal Centers related to the southwest Colombian fauna.

Quaes t. Ent., 1977 13 (2)

110

Mendez

and differentiation of most Pacific species occurred after the early Pleistocene uplift of the

Northern Andes, probably under orographic conditions similar to those existing today.

Ecological relationships of mammal species inhabiting this area are indicated in Table 1.

Table 1. Ecological Relationships of Some Mammals of the Cauca Center

Meadow Zygodontomys brevicauda

brunneus

Sigmodon hispidus bogotensis

Sylvilagus brasiliensis

fulvescens

The Colombian Montane Forest Center. — This dispersal center seems to be confined to the

forest biomes of Colombia (with the exception of the Sierra Nevada de Santa Marta), Ecuador

and Venezuela. It includes two subcenters, namely, the West Andean and the East Andean.

Inasmuch as affinities have been found between the animal populations of the Central Andes

and those of the West Andean Subcenter, the Central Andes is included with the West Andean

Subcenter (Muller, 1973).

The fauna of the Colombian Montane Forest Center may be considered subtropical and dis-

plays similarities with those of the Colombian Pacific Center, which is essentially tropical. This

condition results from most of the species of the Colombian Montane Forest Center having

evolved originally from ancestors that came from the lower lands.

Characteristic mammals of the Colombian Montane Forest Center and their ecological re-

lationships are indicated in Table 2.

Mammalian-Siphonapteran Associations

111

Table 2. Ecological Relationships of Some Mammals of the Colombian Montane Forest Center

Forest: Cryptotis thomasi

open or medallinius

close canopy

Oryzomys caliginosus

monticola

Nectomys alfari

esmeraldarum

Thomasomys aureus

popayanus

Thomasomys cinereiventer

Chylomys instans

Dasyprocta fuliginosa

candelensis

Sylvilagus brasiliensis

fulvescens

Nasua olivacea

Mazama americana

Domestic Rattus rattus

or semi- Rattus norvegicus

domestic Mus musculus

Chironectes minimus Eptesicus brasiliensis Didelphis azarae

andinus

Ichthyomys hydrobates Metachirus nudicaudatus

nicefori Myotis chiloensis colombianus

Philander opossum

griscescens

Sciurus pucherani

caucensis

Reithrodontomys

mexicanus milled

Echinoprocta rufescens

Aotus trivirgatus

lemurinus

The Colombian Pacific Center. - The extensive territory of lowlands in the western part of

Colombia involves the major portion of the center. This territory continues to the north along

the base of the mountains and ends on lands watered by the Magdalena River. To the south,

this center encroaches upon a portion of the northern part of Ecuador. This center includes

two subcenters: the Nechi, which encompasses the area between the Rio Sinu and the lower

reaches of the Rio Cauca; and the Choco, which includes the area west of the Andes.

Possibly during the Tertiary, the gap separating Central and South America divided the ter-

ritory now known as the Colombian Pacific Center. This thesis stems from the existence at

that time, of a seaway south of Panama and the Gulf of Uraba, which connected the Caribbean

Sea and the Pacific Choco basin of Western Colombia (Haffer, 1967). This seaway was closed

during the late Pliocene and seems to correspond to the Bolivar Geosyncline discussed by Ny-

gren (1950) and Hershkovitz (1968).

Glacial and interglacial periods of the Pleistocene evidently influenced the Pacific lowlands

of Colombia. Glaciation of the mountains produced considerable temperature reduction and

high humidity. At this time, sea level lowered about 1 00 m and extensive movements of fauna

occurred between the Amazonian region and the trans-Andean area of western Colombia and

Central America. The interglacial periods were particularly drier in the northern part of Col-

ombia, when the humid forest moved southward due to influence of winds, and sea level rose

about 30 to 50 m. As a result of this condition the Maracaibo basin and other parts of the

Northern Colombian plains were flooded (Haffer, 1967).

Studies of birds, lizards and amphibians presently inhabiting this center, have indicated the

origin and distribution patterns of some of these vertebrates. A strong relationship of the Pac-

ific lowland fauna of Colombia to that of the Amazon region suggests the interchange of ani-

mals during remote times of the Pleistocene. Today, the Pacific mammalian fauna of the

Quaes t. Ent., 1977 13 (2)

112

Mendez

Departamentos del Valle, Choco, Cauca and Narino, inhabiting dense forests of the coastal

lands, are, with some exceptions, similar to those of the Pacific lands of Eastern Panama and

Northern Ecuador.

Some elements of the mammal fauna of the Colombian Pacific Center and their ecological

The North Andean Center. — This area is the Central and Eastern Cordillera in Colombia

and the Andean mountains in Ecuador and Peru. The most characteristic biome of this center

is the paramos with its selected plant and animal life. To this center belong such mammals

as whose names and ecological relationships are indicated in Table 4.

According to Muller, it is apparent that 28 species of the North Andean faunal elements

(more than 75%) belong to families of North and Central American origin.

Chapman (1917), referring to birds from the paramo zone, indicated that the majority were

Mammalian-Siphonapteran Associations

113

derived from the sea level equivalent of this zone in southern South America. It seems logical

to assume that the same circumstance occurred with the mammals also.

Table 4. Ecological Relationships of Some Mammals of the North Andean Center

FLEAS OF SOUTHWESTERN COLOMBIA

In this presentation descriptions are limited to those forms that are either new to science

or have been previously described from one sex. Only citation of the original description for

every taxon is given. For complete synonymy, type data and other distributional records the

reader is referred to Johnson (1957), Tipton and Mendez (1966) and Tipton and Machado-

Allison (1972). The principal sources of information of Colombian mammal hosts have been

Allen (1912, 1913, 1915, 1916), Borrero (1967), Cabrera (1958 and 1961), Cabrera and Yepes

(1940), Hershkovitz (1941, 1947, 1948, 1949, 1960, 1962), Gyldenstolpe (1932), Tate (1932a,

b, 1935), and Osgood (1912).

The following report pertains to taxa from southwestern Colombia available to us for study

and forms that are not represented in our material but have been reported before from that

territory or are likely to occur there. The order followed is that of Johnson (1957).

To conserve space, I have abbreviated locality data for material of previously described

species by omitting collection number, name(s) of collector(s), and designating only month

of collection by Roman numeral. These details are available on request. For new species des-

cribed here, complete data are given for each specimen. The acronym “HTC” preceding a num-

ber in parentheses represents Harold Trapido Collection. Other numbers in parentheses refer

to material in the collection of the Gorgas Memorial Laboratory.

3

KEY TO SPECIES OF FLEAS OF SOUTHWESTERN COLOMBIA

1 Thorax very reduced or compressed; antesensilial bristles absent; female

burrowed into skin 2

1' Thorax not reduced or compressed; antesensilial bristles present; females

not burrowed into skin 3

3. The following papers were helpful in the preparation of this key: Johnson (1954 and 1957), Ewing (1929), and Tipton

and Mdndez (1966). Rhopalopsyllus lugubris, R. cacicus saevus, Polygenis dunni, and P. roberti beebei have not been re-

ported from southwestern Colombia; however, they are likely to occur there and are included in this key.

Quaest. Ent., 1977 13 (2)

Mendez

Head frons angular (Fig. 44); anterior lower margin of hind coxa with

tooth-like projection; host not bat Tunga penetrans (Linnaeus), p.

Head frons rounded; hind coxa without tooth-like projection; host-bat . .

Rhynchopsyllus pulex Haller, p.

Sword-like ridge of mesocoxa absent; mesonotum always lacking pseudo-

setae; metanotum rectangular, not markedly broader dorsally than ven-

trally; not more than one row of bristles on abdominal terga II- VI; meta-

coxa with patch of spinelets on inside

Sword-like ridge of mesocoxa present in most specimens; mesonotum with

or without pseudosetae; metanotum much broader dorsally than ventrally;

metacoxa without patch of spinelets on inside ,

Genal and pronotal comb present (Fig. 41)

Ctenocephalides fells (Bouche), p.

Genal and pronotal comb absent

Mesothoracic pleural rod present; tergum VIII of female complete dor-

sally; bulga of spermatheca (Fig. 40E) not globular, heavily pigmented . .

Xenopsylla cheopis (Rothschild), p.

Mesothoracic pleural rod absent; tergum VIII of female divided dorsally;

bulga of spermatheca globular, lightly pigmented

Dorsal aedeagal sclerite broad throughout; crochets small and elongate,

rodlike; sternum VII of most females with 7-9 bristles

Pulex simulans Baker, p.

Dorsal aedeagal sclerite relatively long and slender; crochets expanded

apically, not rodlike; sternum VII of most females with 4-5 bristles ....

Pulex irritans Linnaeus, p.

Head with true helmet (Fig. 1 1A, 1 19A); ctenidia on helmet, gena and

pronotum

Head without helmet (Fig. 16A); ctenidia absent or present, limited to

gena and/or pronotum

Helmet separated from head dorsally; vertical comb posteromarginal ....

Helmet not separated from rest of head (Fig. 1 1 A); vertical comb mesad .

Cleopsylla monticola Smit , p.

Two large genal bristles anterior to cibarial pump; tergum VIII of male

with apodeme; female spermatheca barrel-shaped, without internal tub-

ercle, and hilla not enlarged basally

One or two large genal bristles posterior to or vertically in line with cib-

arial pump; tergum VIII of male without apodeme; female spermatheca

not barrel-shaped, with internal tubercle or with hilla enlarged basally . .

Genal comb teeth reduced, slightly longer than broad (Fig. 1 19A)

Plocopsylla phyllisae Smit , p.

Genal comb teeth normal, more than twice as long as broad (Fig. 14) ... .

Plocopsylla thor Johnson, p.

Pronotal comb spines about 18, not pointed (Fig. 12A); male with

vertical row of four close-set spiniforms on sternum VII; female with

subcaudal row of heavy spiniform setae on sternum IX (Fig. 12A)

Sphinctopsylla diomedes Johnson, p.

Pronotal comb spines about 30, pointed; male only with normal bristles

on sternum VII; female without subcaudal row of heavy spiniform setae

on sternum IX Sphinctopsylla tolmera (Jordan), p.

165

164

. 4

. 7

164

. 5

158

. 6

164

164

8

12

9

124

10

11

127

127

124

127

Mammalian-Siphonapteran Associations

115

Anterior margin of head conical, with two short spiniform bristles

(Fig. 16A); tibia with dorsal comb of short, stout bristles

Leptopsylla segnis (Schonherr), p. 131

Anterior margin of head not conical, without short spiniform bristles;

tibia without dorsal comb of short, stout bristles 13

Several abdominal terga with well developed combs 14

Abdominal terga without combs 15

Crochets with convex margin dorsad; female sternum VII with caudal

margin truncate Ctenidiosomus rex Johnson, p. 121

Crochets with convex margin ventrad; female sternum VII with caudal

margin not truncate Ctenidiosomus traubi Johnson, p. 121

Anteroventral margin head with two large genal spines (Fig. 15A); on

bats Sternopsylla distincta speciosa Johnson, p. 127

Anteroventral margin of head with more than two or without genal spines;

not on bats 16

With combination of: anterior tentorial arm present, inserted anterior to

eyes; mesopleural rod not dorsally bifurcated; ventral margin of pronotum

not bilobed; tarsal segment V with four pairs of plantar bristles 17

Combination not as above 31

Antennal club symmetrical; without apical spinelets on metanotum ....

Tetrapsyllus comis Jordan, p. 132

Antennal club asymmetrical; apical spinelets on metanotum 18

Frontoclypeal margin of head subconical; pronotal comb present

(Fig. 36A) Scolopsyllus colombianus Mendez, p. 142

Frontoclypeal margin of head rounded; pronotal comb absent 19

Prosternosome projected downward between coxae, mesocoxa rectangular,

margins parallel Rhopalopsyllus 20

Prosternosome not projected downward between coxae; mesocoxa asym-

metrical, obviously broadest basally Polygenis 4 22

Spiracle of metepimere oblong, prolonged dorsally; bulga of spermatheca

globular (Fig. 39D) . . Rhopalopsyllus lugubris Jordan and Rothschild, p. 158

Spiracle of metepimere rounded or ovoid; bulga not globular 21

Labial palpus extended to apex of coxa I or beyond (Fig. 38A); movable

process of clasper longer than distal arm of sternum IX (Fig. 38B); sperm-

atheca somewhat boomerang-shaped (Fig. 38C)

Rhopalopsyllus cacicus saevus Jordan and Rothschild, p. 158

Labial palpus not extended to apex of coxa I (Fig. 37A); movable process

of clasper about as long as distal arm of sternum IX (Fig. 37B). Sperma-

theca strongly S-shaped (Fig. 37C)

Rhopalopsyllus australis tupinus Jordan and Rothschild, p. 142

Distal arm of sternum IX much shorter than proximal arm; heel of sternum

IX strongly angular; inner tube of aedeagus reflexed dorsally, not coiled

apically 23

Distal arm of sternum IX about equal to length of, or longer than, proxi-

mal arm; heel of sternum IX weakly angular; inner tube of aedeagus re-

flexed ventrally, coiled apically 24

Clypeal tubercle above eye level (Fig. 32A); posterior margin of fixed

process convex; spermatheca sinuate, without separation between bulga

and hilla (Fig. 32E) Polygenis thurmani Johnson, p. 141

4. Females of Polygenis are difficult to identify and the characters used in this key pertain almost entirely to males. The

females of P. trapidoi, n. sp. and/*, hopkinsi, n. sp., are unknown.

Quaest. Ent., 1977 13 (2)

116

Mendez

23'

24

24'

25

25'

26

26'

27

27'

28

28'

29

29'

30

30'

31

31'

32

32'

33

Clypeal tubercle at eye level (Fig. 29 A); posterior margin of fixed pro-

cess sinuate; spermatheca humped, with distinct separation between bulga

and hilla (Fig. 29C) Polygenis klagesi (Rothschild), p. 140

(22') Distolateral lobes of aedeagus with angular projection; sternum VIII of

male divided in half ventrally, with short dorsocaudal extension

Polygenis trapidoi, new species, p. 141

Distolateral lobes of aedeagus without angular projection; sternum VIII of

male not divided in half ventrally, without dorsocaudal extension 25

(24') Apex of distal arm of male sternum IX with distinct group of stout

bristles Polygenis bohlsi bohlsi (Wagner), p. 132

Apex of distal arm of male sternum IX without distinct group of stout

bristles 26

(25') Aedeagal fender present; dorsal margin "of fixed process of clasper sin-

uate; spermatheca with cribose bulga, its ventral margin interrupted

at bulga-hilla junction (with probable exception of P. hopkinsi , n. sp.) . . 27

Aedeagal fender absent (Fig. 30C); dorsal margin of fixed process of

clasper usually slightly convex; spermatheca not cribose, its ventral

margin continuous at bulga-hilla junction (Fig. 30E)

Polygenis pradoi (Wagner), p. 140

(26 ) Aedeagal side piece above basal part of inner tube; aedeagal ribs not

numerous (Fig. 25B) Polygenis delpontei, new species, p. 138

Aedeagal side piece absent or below basal part of inner tube; aedeagal

ribs very numerous 28

(27') Aedeagal lateral lobes very reticulate; subapical ridge of median dorsal

lobes of aedeagus present (Fig. 2 1C)

Polygenis caucensis, new species, p. 137

Aedeagal lateral lobes not reticulate or faintly reticulate; subapical ridge

of median dorsal lobes of aedeagus absent 29

(28') Labial palpus extended to trochanter I (Fig. 26 A); several distal arm

bristles of sternum IX very long, approximately four times maximum width

of distal arm (Fig. 26B) . . . Polygenis dunni (Jordan and Rothschild), p. 139

Labial palpus not extended to trochanter I; distal arm bristles of sternum

IX short or as moderate length, not very long 30

(29') Aedeagal fender well developed, half-moon shaped; distal arm of sternum

IX distinctly broad medially (Fig. 3 1 B)

Polygenis roberti beebei (I. Fox), p. 140

Aedeagal fender reduced to slender, arched, inconspicuous structure;

distal arm of sternum IX not distinctly broad medially (Fig. 27C)

Polygenis hopkinsi, new species, p. 139

(16') Genal comb present 32

Genal comb absent 33

(31 ) First spine of genal comb almost overlapped by second (Fig. 8A);

trabecula centralis present; labial palpus 5-segmented; both sexes with

three antesensilial bristles. . . . Neotyphloceras rosenbergi (Rothschild), p. 117

First spine of genal comb not overlapped by second (Fig. 9A); trabecula

centralis absent; labial palpus 4-segmented; both sexes with two antesen-

silial bristles Adoratopsylla intermedia copha Jordan, p. 117

(31') Pronotal comb with more than 24 spines; bristles of antennal segment 2

long in both sexes (Fig. 17A)

Dasypsyllus gallinulae perpinnatus ( Baker), p. 131

Mammalian-Siphonapteran Associations

117

33' Pronotal comb of most specimens with less than 24 spines; bristles of

antennal segment 2 short in male, not extended to apex of club in

female 34

34 (33') Protibia with seven dorsal notches with paired bristles; meso- and meta-

tibia with six dorsal notches with paired bristles proximal to only single

dorsal bristle Pleochaetis smiti Johnson, p. 131

34' Protibia with five or six dorsal notches with paired bristles; meso- and

metatibia with five dorsal notches with paired bristles proximal to only

single dorsal bristle Pleochaetis equato ris equatoris (Jordan), p. 131

SUPERFAMILY CERATOPHYLLOIDEA

FAMILY HY STRICHOPS YLLIDAE

SUBFAMILY CTENOPHTALMINAE

TRIBE NEOTYPHLOCERATINI

Neotyphloceras rosenbergi (Rothschild)

(Figure 8)

Typhloceras rosenbergi Rothschild, 1904, Novit. Zool., 11: 639, PI. 13, Fig. 68-69; PI. 14, Fig. 71, 74.

Material examined. — Ex Didelphis marsupialis. Depto. del Valle, Municipio de Cali - 26, Pichindd, 1600m, VIII.

Ex Didelphis azarae. Depto. del Valle - 6 , Finca Holanda (nr. Paramo de Chinche), 2700m, X.

Ex Oryzomys caliginosus. Depto. del Valle, Municipio de Cali - 3 6, 4? Quebrada Honda, nr. Pichindd, 1800m, I, III,

VIII, IX.

Ex Oryzomys alfaroi. Depto. del Valle, Municipio de Cali - 2? Quebrada Honda, nr. Pichinde', 1800 m, III, IX; 9 ,

Florida, 8 km. S.E. “La Diana”, 1700m., X.

Ex Oryzomys albigularis. Depto. del Valle, Municipio de Cali - 4(3, 4? , Valle del Rib Pichinde', 1700 - 1900 m., I, X,

XII; 2(3, Cerro Munchique, 60 km. by road W. Popayan, Pena del Perro, 2160m., V; 9, Finca La Flora, Quebrada Norte,

Pichindd, 1900m., VIII; 5(3, Saladito, Km. 12, 2000m., Ill; 4<3, 59, Finca Holanda (nr. Pdramo de Chinche), 2700 m., X.

Depto. del Cauca - 9, Cerro Munchique, Finca El Retiro, 2200m., V.

Ex Oryzomys (Oligoryzomys) species. Depto. del Valle, Municipio del Cali - 9, Quebrada Honda, nr. Pichindd, 1800m.,

X.

Ex Rhipidomys latimanus. Depto. del Valle, Municipio de Cali - 9, Quebrada Honda, nr. Pichindd, 1800m., X; <3, 29,

Valle del Ri'o Pichindd, 1700 - 1900 m., VII, XII.

Ex Rhipidomys similis. Depto. del Cauca — 9, Cerro Munchique (60 km. by road W. Popayan, sitio No. 1), 2000m., V.

Ex Thomasomys aureus. Depto. del Cauca — 9 > Pilimbala, 3100m., V.

Ex Thomasomys cineriventer. Depto. del Cauca, Cerro Munchique -9,60 km. by road W. Popayan, Sitio. No. 1, 2500m.,

V; (3, sitto No. 3, V; 6, 60 km. by road W. Popayrfn, Pena del Perro, 2160 m., V. Depto. del Narino - 6(3, 9 , Laguna de La

Cocha, 2700 m., V; <3, Km 38, between Pasto & Sibundoy, Comisario de Putumayo, 3100 m., V; <3, Km. 33, between Pasto

& Sibundoy, 2900 m., V.

Ex Thomasomys fuscatus. Depto. del Valle, Municipio de Cali - 3 <3, Valle del Rib Pichindd, 1700 - 1900 m., I, III, XI;

2(3, 29, Pichindd, 1900 m., V; 2(3, 9, Pichindd, Rincon del Yarumal, V; <3, Pichindd, Finca La Flora, VII; 6(3, 8 9, Saladito,

Km. 12, 2000 m., II. Depto. del Narino - 9, Laguna de La Cocha, 2700 m.

Remarks. - In addition to Colombia, this species is distributed in Venezuela, Ecuador and

Peru. Specimens are regularly found on a variety of rodents and less commonly on marsupials.

Our material contains specimens from 1 2 species of mammals, indicated above. Other hosts

in the literature for this species are the following: Philander opossum, Marmosa, Akodon, Chil-

omys, Rheomys, Stictomys, and Sigmodon. The variety of hosts on which fleas of this species

are found indicates a low degree of host specificity.

TRIBE ADORATOPSYLLINI

Adoratopsylla (Tritopsylla) intermedia copha Jordan

(Figure 9)

Stenopsylla intermedia copha Jordan, 1926, Novit. Zool., 33: 391, Fig. 15.

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Fig. 6. General appearance of female of Ctenocephalides felis (Bouche). AN. - Antenna, A.S. - Antesensilial seta, A.S.T. - Anal stylet of female, B.C. - Bursa copulatrix, C. - Coxa, CL. - Claw of tar-

sus, D.A.L. - Dorsal anal lobe, E. - Eye, EP. - Epipharynx, F. - Femur, FR. - Frons, G.CT. - Genal ctenidium, L. - Lacinia, L.P. - Labial palp, L.P.B. - Lateral plantar bristles, M.L. - Maxillary lobe,

M.P. - Maxillary palp, MPM. - Mesepimere, MPS. - Mesepisternum, MSN. - Mesonotum, MTM. - Metepimere, MTN. - Metanotum, MTS. - Metasternum, O. - Occiput, O.B. - Ocular bristle, P. - Prono-

tum, P.CT. - Pronotal ctenidium, PRM. - Proepimere, S. - Sensilium, SP. - Spermatheca, ST. - Sternum, T. - Tergum, TA. - Tarsus, TI. - Tibia, V.A.L. - Ventral anal lobe.

Mammalian-Siphonapteran Associations

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DLL

Fig. 7. Structures of apex of aedeagus of Scolopsyllus colombianus Mdndez. A.M.S. - Apico-median sclerite, AP.S. - Apodemal

strut, CR. - Crochet, Cr.P. - Crochet processes, C.S. - Crescent sclerite, DL.L. - Distolateral lobes, F.M. - Fluted membrane,

H.L. - Heel at base of aedeagal pouch, I.T.-A - Apical portion of sclerotized inner tube, I.T.-B - Basal portion of inner tube,

L.L. - Lateral lobes, M.D.L. - Median dorsal lobes, P.R. - Penis rods, PS.T. - Pseudotube.

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Fig. 8. Neotyphloceras rosenbergi (Rothschild). Female. A. Head, prothorax and procoxa; B. Modified abdominal segments;

C. Spermatheca.

Mammalian-Siphonapteran Associations

121

Material examined. — Ex Didelphis azarae. Depto. del Valle - 3(5, 3?, vie. Cali, X.

Ex Didelphis marsupialis. Depto. del Valle, Municipio de la Cumbre - (5, 9 , La Maria, 1400 m, XI; Municipio de Cali -

7(5, 79, Pichindd, 1600 m., VIII; 10(5, 159, Lago Calima, 1450 m., II.

Ex Philander opossum. Depto. del Valle - Pichindd, 1600 m., VIII; 39, Alto Anchicaya', 650 m., II.

Ex Oryzomys caliginosus. Depto. del Valle - (5, Pichindd, 1600 m., I.

Ex Thomasomys fuscatus. Depto. del Valle - 2(5, 39, Saladito, (Km. 12), 2000 m., II.

Remarks. — This subspecies has been reported from Colombia, Panama, Ecuador and Peru,

from sea level to over 3000 meters. It is a common parasite of marsupials and displays a high

degree of infestation on some single host animals (Tipton and Mendez, 1966). In southwestern

Colombia specimens have been obtained from the hosts indicated above. Elsewhere specimens

have been recorded from Oryzomys caliginosus and Proechimys semispinosus.

FAMILY PYGIOPSYLLIDAE

SUBFAMILY PYGIOPSILLINAE

Ctenidiosomus rex Johnson

Ctenidiosomus rex Johnson, 1957, Mem. Ent. Soc. Wash. 5:50, PI. 20.

Remarks. — Type material (2 males and 2 females) from San Agustin, Departamento de

Huila, Colombia, represents the only specimens known of this taxon. These specimens were

collected from Thomasomys, Oryzomys and Rhipidomys. It seems probable that a species of

Thomasomys (probably T. laniger ), is the natural host of this flea species.

Ctenidiosomus traubi Johnson

(Figure 10)

Ctenidiosomus traubi Johnson, 1957, Mem. Ent. Soc. Wash. 5:49-50, PI. 17; PI. 18, Fig. 5; PI. 19, Fig. 1, 2.

The original description is based on the holotype female, ex Caenolestes obscurus, Colom-

bia, Depto. de Antioquia, Sanson, 7 km. E. of Paramo, 3,160 m., 18 Oct. 1950, P. Hershkovitz

collector. Males are described below.

Material examined. — Ex Caenolestes obscurus. Depto. del Cauca - 9 , Puracd Park, 3520 m., V.

Ex Thomasomys aureus. Depto. del Cauca - (5, Pilimbala, 3100 m., V.

Ex Thomasomys cinereiv enter. Depto. del Narino - (5, Comisam Putumayo, Km. 77, between Sibundoy & Mocoa, 2200

m., V.

Description of male. — Head (Fig. 10A). Strongly fracticipit, with frons evenly rounded, without clypeal tub-

ercle. Preantennal region with 3 distinct discs and abundant micropores; principal bristles of preantennal region arranged in

2 rows, secondary bristles very short, scattered mainly on preocular area. Eye deeply excised ventrally, weakly pigmented.

Postocular area with 2 pits just below eye. Genal area bilobed, its anterior lobe or genal process broadly rounded, not accum-

inate. Posterior lobe of gena evenly rounded, moderately broad. Postantennal region with anterior micropores, several pits

profusely distributed, 2 rows of bristles, in addition to short bristles on dorsal margin and in front of antenna, mainly

on antennal fossa. Pedicellus of antenna covered with short bristles.

Thorax. Very setose. Pronotum with 2 rows of bristles preceding comb of about 26 spines. Mesonotum at least with 3

well defined rows of bristles, remaining bristles short, concentrated on anterior pronotal region. Mesepisternum with few

non-prominent bristles on anterodorsal area near pleural ridge of mesothorax. Mesepimere with bristles of different sizes.

Metanotum with about 3 or 4 defined rows of bristles, posteriormost row of long bristles and short intercalaries. Other

bristles not arranged in rows in anterior metanotal area. Lateral metanotal area apparently with no more than 1 bristle. Met-

episternum with oval outline interrupted by anterior projection of metasternum, provided with few bristles. Metepimere

with 3 rows of uneven bristles.

Abdomen. Combs on terga II- V, and with various number of spines (in the two specimens examined), respectively, 14-15;

14-15; 12-13; 14-15. Upper antesensilial birstle about 1/2 as long as lower bristle.

Modified abdominal segments. Tergum VIII reduced to subtriangular plate provided with broad spiracle, with group of

short bristles near antesensilial bristles. Sternum VIII large, ensheating principal structures of genitalia, provided with num-

erous marginal and inner bristles, caudal margin entire, without sinus. Clasper large, somewhat pyriform, projected anterior-

ly into short manubrium curved upward. Process with subrounded apical expansion, largely squamose, with group of bristles

on outer and inner surfaces. Posterocaudal margin of process with four long bristles. Movable process of clasper broadest

near basal area, narrowed apically, with bristles of varied size. Sternum IX (Fig. 10B) like those of other males of genus.

Distal arm club-shaped, with broad and rounded apex with 4 stout bristles on caudal margin; remaining bristles smaller,

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Fig. 9. Adoratopsylla intermedia copha Jordan. Male. A. Head, prothorax and procoxa; B. Genitalia; C. Apex of aedeagus,

A.I.T. - Armature of inner tube, CR. - Crochet, C.S. - Crescent sclerite, L.L. - lateral lobes, L.S.I. - Lateral sclerotization of

inner tube, S.I.T. - Sclerotized inner tube. Female. D. Spermatheca and 7th abdominal segment. From “The Fleas (Siphonap-

tera) of Panama” by Tipton and Mdndez, in “Ectoparasites of Panama”, Field Museum of Natural History, Chicago (1966).

Mammalian-Siphonapteran Associations

123

Fig. 10. Ctenidiosomus traubi Johnson. Male. A. Head, prothorax and procoxa; B. Ninth sternum; C. Clasper; D. Apex of

aedeagus.

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Mendez

scattered over most of arm. Proximal arm of sternum IX narrow at base, broad at apex. Aedeagus like that of Ctenidiosomus

perplexus Tipton and Machado-Allison. Median dorsal lobe broad, with rounded dorsal margin, produced as caudal subacum-

inate blade. Lateral lobes narrower than distal lobe, slightly arched. Crochets crescent-shaped. Aedeagal apodemal rod arched

apically. Penis rods strongly coiled, extensively fimbriate on apical portion.

Remarks. — Presently, Ctenidiosomus traubi is known only from Colombia, where it has

been collected in localities with elevation ranging from 2200 m to 3500 m. The scant material

available has been collected from one marsupial and two cricetine rodent species. As Lewis

(1974) pointed out, it is likely that the preferred hosts of C. traubi are rodents.

FAMILY STEPHANOCIRCIDAE

SUBFAMILY CRANEOPSYLLINAE

TRIBE CRANEOPSYLLINI

Cleopsylla monticola Smit

(Figure 1 1)

Cleopsylla monticola Smit, 1953, Bull. Brit. Mus. (Nat. Hist.) Entomol., 3(5): 193, Fig. 13, 15, 17, 19, 20.

Material examined. — Ex Oryzomys albigularis. Depto. del Cauca - <5, 9 , Cerro Munchique, 60 km. by road

W. Popaydn, sitio No. 1, 2500 m., V. Depto. del Valle - 13 <5, 69, Finca Holanda (nr. Paramo de Chinche), 2700 m., X.

Ex Rhipidomys similis. Depto. del Valle - 2(5, 9 , Finca Holanda, 2700 m., V.

Ex Thomasomys cinereiv enter. Depto. del Narino — 2c5, 9 , Laguna de La Cocha, 2700 m., V.

Ex Thomasomys fuscatus. Depto. del Valle - (5, Pichindd, Finca La Flora, 1900 m., V.

Remarks. — Reports of this helmet flea are from Ecuador, Colombia and Venezuela. In

Colombia, specimens have not been collected above 2700 meters elevation. The range of ver-

tical distribution for this species in Venezuela is from 120 meters to 1443 meters (Tipton and

Machado-Allison, 1971). Five species of cricitines harbor C. monticola in southwestern Colom-

bia. (see above for details) Other hosts recorded in the literature are Caenolestes fuliginosus,

Didelphis marsupialis, Marmosa fuscata, M. dryas, Oryzomys minutus, Rhipidomys venustus,

Rhipidomys sp., Thomasomys hylophilus, T. laniger, T. vestibus, Chilomys ins tans, and birds.

Sphinctopsylla diomedes Johnson

(Figure 12)

Sphinctopsylla diomedes Johnson, 1957, Mem. Ent. Soc. Wash. 5:68, PI. 32.

This species was originally described from two male specimens ex Caenolestes obscurus,

Colombia: Depto. of Huila, San Agustin, San Antonio, left bank of Rio Magdalena (Cordillera

Central), 2200 m, 24 Aug. 1950. P. Hershkovitz collector.

Material examined. — Ex Caenolestes obscurus. Depto. del Cauca - 8(5, 79, PuracdPark, 3500 m., IV- VI. Depto.

de Cundinamarca, Municipio de Soacha - 9, Soche, 2700 m., IX.

Description of female. — Head (Fig. 12A). Frons margin moderately rounded. Helmet comb of 13 spines. Genal

comb of 5 spines.

Thorax. Pronotum (Fig. 12 A) with 2 rows of bristles and conspicuous comb of 9 spines per side. Remaining thoracic

structures and legs as in male.

Abdomen. Terga I-IV with apical spineletes. All terga with two rows of bristles but anteriormost row more reduced. Ter-

gum VII with 2 subequal antesensilial bristles.

Modified abdominal segments (Fig. 12B). Posterior margin of tergum VIII sinuous, with 2 groups of large, stout spini-

form bristles, upper group of 4-6 bristles, lower group of 1-3. Both groups preceded by scattered bristles of various size and

location, in addition to short and moderate size marginal bristles. Spiracle of tergum VIII with very broad basal portion.

Sterna II- VI with 1 row of 6 bristles. Sternum VII with group of 3-4 stout spiniform bristles on each side, in addition to sev-

eral inconspicuous bristles. Posterior margin of this sternum almost straight, not indented. Sensillum with about 11 sensory

pits per side. Dorsal anal lobe with several bristles. Ventral anal lobe with only 1 or 2 bristles. Anal stylet short and stout,

dorsally and ventrally convex, its apical bristle about twice the length of stylet body, with minute ventral bristle and mesal

bristle of moderate size. Spermatheca (Fig. 12B, 12C) with divided bulga, anterior section globular, fairly reticulate, and pos-

terior section not reticulate, followed by short, upturned unpigmented hilla. Main body of bursa copulatrix short, sinuous,

weakly sclerotized.

Mammalian-Siphonapteran Associations

125

B

Fig. 11. Cleopsylla monticola Smit. Male. A. Head, prothorax and procoxa; B. Mesothorax, metathorax and tergum I.

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Mendez

B

Fig. 12. Sphinctopsylla diomedes Johnson. Female. A. Head, prothorax and procoxa; B. Modified abdominal segments; C

Spermatheca.

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127

Remarks. — This species is endemic to southwestern Colombia where specimens have been

found in areas between 2200 and 3500 meters. All specimens presently existing in collections

are from Caenolestes obscurus individuals, which seem to be the natural host.

Sphinctopsylla tolmera (Jordan)

Craneopsylla tolmera Jordan, 1931, Novit. Zool., 36:314, Fig. 5.

Material examined. — Ex Thomasomys cinereiventer. Depto. del Narino -(5,9, Laguna de La Cocha, 2700 m.,

V. Depto. del Valle - <5, Finca Holanda (nr. Paramo de Chinche), 2700 m., X.

Remarks. - The geographical range of S. tolmera involves Colombia, Ecuador and Venezuela,

in some areas exceeding 2000 meters elevation. Rodents of the genus Thomasomys probably

are the preferred hosts in Colombia and Ecuador. In Venezuela this flea species seems to be

more associated with Oryzomys minutus. Of 36 males and 76 females recorded by Tipton

and Machado- Allison (1972), 32 males and 63 females were recovered from 46 specimens of

this rodent species.

Plocopsylla phyllisae Smit

(Figure 13)

Plocopsylla phyllisae Smit, 1953, Bull. Brit. Mus. (Nat. Hist.) Entomol., 3:197, Fig. 25, 26, 28, 30.

Material examined. — Ex Caenolestes obscurus. Depto. del Cauca — 7(5, ll9, PuracdPark, 3500 m., IV, V.

Depto. de Cundinamarca, Municipio de Soache — 29, El Soche, 2700 m., IX.

Remarks. — Some populations of this species live in territories over 3000 meters elevation

in Ecuador and Colombia. The holotype male was secured from Oryzomys sp; nevertheless,

the material (11 <5c5 and 2199) available to us for study came from six specimens of Caenoles-

tes obscurus, which is probably the preferred host. Individuals of this terrestrial marsupial

live in dark damp forests of paramos and are crepuscular or nocturnal. They seem to be pri-

marily insectivorous (Osgood, 1921 ; Tate, 1931).

Plocopsylla thor Johnson

(Figure 14)

Plocopsylla thor Johnson, 1957, Mem. Ent. Soc. Wash., 5:73-74, PI. 38 (Fig. 1 , 2, 3, 6, 7), PI. 39 (Fig. 1, 2, 3).

Material examined. — Ex Thomasomys cinereiventer. Depto. del Cauca - 9, Cerro Munchique, 60 km. by road

W. Popaya'n, sitio No. 3, V. Depto. de Narino - Laguna de La Cocha, 2700 m., V.

Remarks. - Plocopsylla thor seems to be restricted to areas of high elevation (particularly

between 2000 and 3000 meters) in Colombia. Specimens have been found associated with

cricetine rodents of the species Oryzomys albigularis and Thomasomys spp. The typical host

for this flea is probably Thomasomys cinereiventer.

FAMILY ISCHNOPSYLLIDAE

SUBFAMILY ISCHNOPSYLLINAE

Sternopsylla distincta speciosa Johnson

(Figure 15)

Sternopsylla distincta speciosa Johnson, 1957, Mem. Ent. Soc. Wash. 5: 100; PI. 48, Fig. 3, 4; PI. 50, Fig. 3, 8.

Remarks. - The description of this subspecies is based on a holotype male, an allotype fe-

male, and three female paratypes ex Tadarida brasiliensis, Peru: Dept, of Cuzco, Quince Mil,

19 June 1950, C. Kalinowski collector. One male and three female paratypes ex Tadarida sp.,

Colombia: Dept, of Huila, Pitalico, 1350 m, 28 Nov. 1951, P. Hershkovitz collector. No other

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Mendez

Fig. 13. Plocopsylla phyllisae Smit. Male. A. Head, prothorax and procoxa; B. Mesothorax, metathorax and tergum I; C

Modified abdominal segments; D. Spermatheca.

Mammalian-Siphonapteran Associations

129

Fig. 14. Plocopsylla thor Johnson. Male. Head, prothorax and procoxa.

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Mendez

Fig. 15. Sternopsylla distincta speciosa Johnson. Male. A. Head, prothorax and procoxa; B. Eight sternum; C. Process and

movable finger of clasper; D. Distal arm of ninth sternum. Female. E. Modified abdominal segments; F. Anal stylet and ven-

tral anal lobe; G. Spermatheca. From “The Fleas (Siphonaptera) of Panama” by Tipton and Mdndez, in “Ectoparasites of

Panama”, Field Museum of Natural History, Chicago (1966).

Mammalian-Siphonapteran Associations

131

Colombian records are mentioned in the literature.

FAMILY CERATOPHYLLIDAE

SUBFAMILY LEPTOPSYLLINAE

Leptopsylla segnis (Schonherr)

(Figure 16)

Pulex segnis Schonherr, 1811, K. svenska Ventenskakad. Handl. (2) 32:98, PI. 5, Fig. A. B.

Material examined. — Ex Rhipidomys latimanus. Depto. del Valle, Municipio de Cali - 9, Quebrada Honda, nr.

Pichindd, 1800 m., IX.

Ex Rattus rattus. (3, same locality as above.

Remarks. — Leptopsylla segnis is a cosmopolitan species which has been introduced with

commensal rodent hosts to many parts of the world. In South America it seems to be confined

to areas of high elevation in Colombia, Ecuador, Peru, Venezuela, Brazil, Chile and Argentina.

Typical hosts are various species of Muridae; however, the true host is the house mouse, Mus

mus cuius.

SUBFAMILY CERATOPHYLLINAE

Dasypsyllus gallinulae perpinnatus (Baker)

(Figure 17)

Ceratophyllus perpinnatus Baker, 1904, Proc. U.S. Nat. Mus., 27: 386, 391, 445, Fig. 1-6.

Material examined. — Ex Thomasomys cineriventer. Departo. del Cauca, Cerro Munchique - (3, 60 km. by road

W. Popay^n, sitio No. 1, 2500 m., V; <3, sitio No. 3, 2500 m., V; 9, Pena del Perro, 2160 m., V.

Remarks. - Dasypsyllus gallinulae perpinnatus is a widespread bird flea species, recorded

from several countries in the New World. Our specimens represent the first report of this tax-

on for Colombia. It is also known from Canada, United States of America, Panama, Venezuela

and Argentina.

Our specimens were obtained from different sites and dates on three Thomasomys cinerei-

venter specimens. These rodents were trapped on the ground; however, the fact that they were

parasitized by this bird flea suggests that T. cinereiventer is perhaps partly arboreal. The fleas

were probably obtained from bird nests located on trees visited by the rodents. At the present

time little is known about the habits of T. cinereiventer.

Pleochaetis equatoris equatoris (Jordan)

Ceratophyllus equatoris Jordan, 1933, Novit. Zool., 38: 344, Fig. 63, (partim ).

Material examined. — Ex Thomasomys cinereiventer. Depto. del Narino 9, Laguna de La Cocha, 2700 m., V;

9, Km. 77, between Sibundoy & Mocoa, Comisaria de Putumayo, 2200 m., V.

Remarks. — Pleochaetis equatoris equatoris has been reported from Peru, Ecuador and Col-

ombia. According to Johnson (1957), it is likely that specimens from Peru assigned by Macch-

iavello (1948) to P. equatoris equatoris are P. dolens quitanus.

Pleochaetis smiti Johnson

(Figure 18)

Pleochaetis smiti Johnson, 1954, Jour. Wash. Acad. Sci., 44(9): 291, 295, Fig. 1, 3, 6-8, 10, 12, 13, 16, 21, 25, 26, 31.

Material examined. — Ex Caenolestes obscurus. Depto. del Cauca — 9, Puracd Park, 3500 m., IV.

Ex Oryzomys albigularis. Depto. del Cauca - <3, Cerro Munchique, 60 km. by road W. Popayan, sitio No. 3, 2500 m., V.

Depto del Valle - 2(3, 29, Finca Holanda (nr. Paramo de Chinche), 2700 m., X.

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Mendez

Ex Thomasomys cinereiventer. Depto. de Narino - <3, Laguna de La Cocha, 2700 m., V; Comisaria de Putumayo - 9,

Km. 38, between Pasto & Sibundoy, 3100 m, V; <3, Km. 77, between Sibundoy and Mocoa, 2200 m., V.

Remarks. - P. smiti is to date known from Colombia, Ecuador and Venezuela. Judging from

our collection and the data available in the literature, the vertical distribution of this species ex-

tends from 1980 to 3810 meters. Tipton and Machado- Allison (1972) report abundant mat-

erial (203 males and 208 females) from Venezuela. Evidence is presented by these authors that

the characteristic host of P. smiti in Venezuela is Oryzomys minutus. They recovered 364 spe-

cimens of P. smiti from 158 specimens of this rodent.

I suspect that in Colombia, P. smiti is probably more specific on Oryzomys albigularis\

however, flea specimens collected in this country are few and do not allow final interpretation

as to preferred host species.

SUPERFAMILY RHOPALOPSYLLOIDEA

FAMILY RHOPALOPSYLLIDAE

SUBFAMILY RHOPALOPSYLLINAE

TRIBE PARAPSYLLINI

Tetrapsyllus comis Jordan

(Figure 19)

Tetrapsyllus comis Jordan, 1931, Novit. Zool., 37:135, Fig. 1.

Material examined. — Ex Caenolestes obscurus. Depto. del Cauca - 2?Purace Park, 3500 m., V.

Remarks. — Tetrapsyllus comis was hitherto known only from Ecuador. Our record is the

first for the Republic of Colombia. The scant information about this flea does not allow for

determination of host preference. We have two females from Caenolestes obscurus while the

Ecuadorian holotype female was taken on Sigmodon sp. Apparently, T. comis is a typical

member of the Andean fauna of the northwest portion of South America, perhaps limited in

its distribution to Colombia and Ecuador. The male of this species remains unknown.

TRIBE RHOPALOPSYLLINI

Polygenis bohlsi bohlsi (Wagner)

(Figure 20)

Pulex bohlsi Wagner, 1901, Hor. Soc. Ent. Ross., 35:21, PL 1, Fig. 6.

Material examined. — Ex Oryzomys albigularis. Depto. del Valle - <3, Valle del Rio Pichindd, 1700 - 1900 m.,

Ill; Saladito (Km. 12), 2000 m., II.

Ex Oryzomys alfaroi. Depto. del Valle, Municipio de Buga - 5(3, 79 , Sonso, 1000 m., V.

Ex Oryzomys caliginosus. Depto. del Valle, Municipio de Buga - 14(3, 149, Sonso, 1000 m., Ill, VI. Depto. del Valle,

Municipio de Cali - 4(3, 79, Valle del Rio Pichindd, 1700 - 1900 m., Ill, IV, VI, VII, 6, 59 , Pichindd, 1800 - 1880 m., I,

III, IV, VII. Depto. del Valle - 39 , La Buitrera, 1000 m., II; 35(3, 479 , Lago Calima, 1450 m., II, III; (3, Florida, 8 km.

S.E. “La Diana”, 1700 m., XII.

Ex Thomasomys fuscatus. Depto. del Valle, Municipio de Cali - 9 , Pichindef, IV.

Remarks. — Our records of P. bohlsi bohlsi extend the range of this taxon which is now

known from Colombia, Ecuador, Venezuela, Trinidad, Brazil, Argentina and Paraguay. The

majority of our southwestern Colombia specimens are from oryzomine rodents obtained from

1000 to 2000 meters elevation. Tipton and Machado-Allison (1972) suggest that the optimum

habitat of this flea species is at elevations between 1000 and 1500 meters and the preferred

hosts are cricetine rodents and perhaps more specifically of akodont stock.

Mammalian-Siphonapteran Associations

133

Fig. 16. Leptopsylla segnis (Schonherr). Male. A. Head, prothorax and procoxa; B. Mesothorax, metathorax and tergum I;

C. Process and movable finger of clasper; D. Hind femur and hind tibia. Female. E. Modified abdominal segments; F. Sperma-

theca.

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134

Mendez

Fig. 17. Dasypsyllus gallinulae perpinnatus (Baker). Male. A. Head, prothorax and procoxa; B. Genitalia. Female. C. Sperma-

theca and 7th abdominal segment. From “The Fleas (Siphonaptera) of Panama” by Tipton and Mdndez, in “Ectoparasites of

Panama”, Field Museum of Natural History, Chicago (1966).

Mammalian-Siphonapteran Associations

135

B

D

Fig. 18. Pleochaetis smiti Johnson. Male. A. Head, prothorax and procoxa; B. Process and movable finger of clasper; C. Distal

arm of 9th sternum. Female. D. Modified abdominal segments; E. Spermatheca.

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136

Mendez

Fig. 19. Tetrapsyllus comis Jordan. Female. A. Head, prothorax and procoxa; B. Modified abdominal segments; C. Spermathe-

Mammalian-Siphonapteran Associations

137

Polygenis caucensis, new species

(Fig. 21,22)

Type Material. — Holotype 6 ex Oryzomys caliginosus, (00036), Alto Anchicaya', 650 m, Depto. del Valle, Colom-

bia, 11.11.1974, E. Mdndez & L. Vela'squez; allotype 9, (00032), same locality, date and collector; 9paratype, (00042), same

locality, date and collector; <3 paratype (Acc. No. B-571), ex Didelphis marsupialis, Curiche River, Depto. del Choco, Colom-

bia, 19. VI. 1967, D.G. Young

Holotype and allotype are in the National Museum of Natural History, Washington, D.C.;

I paratype is in the British Museum (Natural History) and 1 paratype is in the Gorgas Memor-

ial Laboratory’s collection.

Diagnosis. - This species appears to be closest to Polygenis roberti beebei I. Fox. Males are

differentiated by the more reticulate aedeagal lateral lobes. In addition, P. caucensis males have

an aedeagal median dorsal lobe with a subapical ridge, a structure apparently absent from males

of P. r. beebei.

Description. — Male. Head (Fig. 21A). Frons fairly rounded, interrupted by short, angular tubercle protruded up-

ward. Preantennal region with moderate number of micropores and 2 pits in front of antennal scape. Preocular row of 2 bri-

stles inserted near eye. Post-ocular row with few minute bristles on lower and upper portions of preocular region. Arch of

tentorium not conspicuous. Oral angle well defined. Genal lobe semiangular. Eyes subovate, not incised, large and well pig-

mented. Maxilla with acuminate tip extended to last segment of maxillary palp. Post-antennal region with 3 rows of bristles

behind antennal scape. Antennae densely covered with minute and prominent bristles irregularly distributed.

Thorax. Pronotum and mesonotum with 2 rows of bristles. Mesepisternum with 2 large bristles per side. Mesepimeron

apparently with 2 or 3 bristles per side. Metanotum with 3 rows of bristles. Metanotal flange with about 6 spinelets. Metep-

isternum with single bristle. Metepimeron with 9 bristles in 2 rows.

Legs. Posteromarginal notches of metatibia with strong bristles with following distribution: 2-2-2-3-2-3.

Abdomen. Terga I-V with spinelets and 2 rows of bristles. Unmodified sterna with single row of bristles, those of sterna

II and III preceded by very few marginal and submarginal bristles.

Modified abdominal segments (Fig. 21C). Sternum VIII somewhat reduced, with several unequal bristles. Fixed process

of clasper (Fig. 21B) broad, with its total length in excess of maximum width, and with barely projected apical lobe, dorsal

margin shallowly sinuate, posterior and ventral margins strongly sinuate, last one moderately indented. Chaetotaxy as illus-

trated. Movable finger of clasper (Fig. 21B) not extended to apical lobe, with several marginal and inconspicuous inner bris-

tles. Proximal arm of sternum IX shorter than distal arm, of irregular shape, narrowed at basal portion, then considerably ex-

panded and sinuate, terminated in subangular projection. Distal arm of sternum IX (Fig. 22A) curved cephalad, more dilated

medially but gradually tapered toward its subrounded tip. Chaetotaxy of this arm limited to some apical bristles and postero-

marginal bristles of various sizes. Heel of sternum IX prominent, with slender, straight terminal tendon. Sternum VIII with

wide subrounded caudal lobe and single row of unequal bristles. Apodeme of aedeagus (Fig. 22B) broad, shorter than termi-

nal portion of aedeagus, with subrounded apex. Terminal portion of aedeagus (Fig. 22B) with semirounded distolateral lobes.

Median dorsal lobe large, almost reaching posterior portion of distolateral lobes, with upper margin shallowly convex. Crochet

not apparent. Lateral lobe gently reticulated, broadly curved, except for short basal angular prominence. Side piece prominent,

almost triangular, its longest side fairly convex. Fender well sclerotized, arched and conspicuous. Apical portion of inner tube

larger than basal portion, strongly convoluted, with final portion slender. Basal portion of inner tube of moderate proportions,

with narrow foramen on anterior half. Crescent sclerite short, indistinct. Side piece broad, of irregular shape, with anterior

margin convex, posterior margin angulate. Fulcral latero-ventral lobe represented by short, slightly curved knob-like structure.

Pseudo tube long, sinuate, well sclerotized. Lateral thickening of end chamber very long, sinuous, reaching heel at base of ae-

deagal pouch. Heel conspicuous, slightly curved upward, with apex subrounded. Fluted membrane prominent.

Female. Head, thorax, legs and unmodified abdominal somites essentially as in male.

Modified abdominal segments. Tergum VII ventrally expanded beyond longitudinal axis of abdomen, with 2 rows of bri-

stles. Tergum VIII large, with sinuous posterior margin and bristles of different length. Sternum VII with posterior margin

subtruncate, with several uneven bristles apparently in single row. Sternum VII broad but not conspicuous, with subangular

caudal margin. Dorsal anal lobe and ventral anal lobe both with subtruncate apex and several bristles. Anal stylet about two

times as long as maximum width, with 2 minute ventral bristles preceding long apical bristle. Spermatheca (Fig. 22C) with

humped bulga well delimited from short, upturned hilla. Dorsal margin of bulga and apex of hilla respectively with short pro-

jection.

Length. Holotype, 2.96 mm; allotype, 3.31 mm.

Remarks. — The trivial epithet of this taxon has been adopted from the Depto. del Cauca,

where part of the type material was obtained.

Quaest. Ent., 1977 13 (2)

138

Mendez

Polygenis delpontei, new species

(Fig. 23, 24, 25)

Type material. — Holotype 6 and allotype 9(HTC-315) ex Oryzomys caliginosus, Colombia, Depto. del Valle,

Municipio de Cali, Quebrada Honda near Pichinde', elevation 1800 m, 7. X. 1967, H. Trapido. Paratypes: 3 9 with same data

as holotype; 1 (5(HTC-146) with same host, locality and collector as holotype but 13. VIII. 1965; 1<3, 1 9(HTC-212) with

same data as holotype but 8.IX.1965; 1 9(HTC-213) ex Reithrodontomys mexicanus, other data as HTC-212; 1 <3(HTC-

243) with same data as HTC-212 but 16.IX.1965; 1 9(HTC-302 ) ex Oryzomys (Oligoryzomys) sp. (?) with same data as

HTC-212 but 4.X.1965. Following paratypes with same data as holotype except date: 2 5 (HTC-317) 11. X. 1963; l9(HTC-

330) 19.X.1965; l9 (HTC-393) 17. XI. 1965; 2 6 (HTC-397) 18.XI.1965; l5(HTC-429) with same data as holotype but

29.XI.1965; 1 9(HTC-454) with same data as holotype but 14.XII.1965; 1 5(HTC-1307) with same data as holotype but

13.XII.1966, 2 (5(00007) same host, Rincon del Yarumal, Pichinde', Depto. del Valle, 21-25. I. 1974, M. Thomas & L. Vel-

a'squez; 15(00010), La Buitrera, Depto. del Valle, 7. II. 1974, E. Mendez & L. Velasquez; ex Thomasomys fuscatus, 1(5,

(HTC-2955), Rincdn del Yarumal, Pichinde', Depto. del Valle, 8.V.1969, H. Trapido, id, with same host and locality but

28.V.1969; l9 (00064), Saladito (Km 12), 2000 m, Depto. del Valle, 17.11.1974, E. Mdndez & L. Vela'squez; l5(HTC-805)

with same data as holotype but elevation 1700 - 1900 m, 5. II. 1966; l5(HTC-1317) with same data as HTC-605 but 13. XII.

1966.

Holotype and allotype are in the U.S. National Museum of Natural History. Paratypes are

in the collections of the following institutions and specialists; British Museum (Natural History),

Universidad del Valle, Colombia, Gorgas Memorial Laboratory, Robert Traub and Phyllis T.

Johnson.

Diagnosis. — Polygenis delpontei is closely related to P. brachinus Jordan. It differs from

this species primarily in that it has the terminal portion of the aedeagal sclerotized inner tube

rod-like and upturned. In P. brachinus the terminal portion of the sclerotized inner tube is

flat and bent downward.

Description. — Male. Head (Fig. 23A). Frons evenly rounded. Frontal tubercle barely projected out of margin, sub-

rounded by wide sclerotized area. Preantennal region profusely covered with micropores. Preantennal row of 7 unequal bri-

stles evenly spaced. Preocular row of 3 medium size bristles and 2 or 3 secondary short bristles. Premarginal bristles of anten-

nal fossa weak, inconspicuous. Oral angle acute but not prominent. Genal lobe subangular. Eye oval, with small ventral in-

dentation and moderate pigmentation. Post antennal region essentially with 3 rows but posteriormost bristle somewhat dis-

placed, thus suggesting 4th row.

Thorax. Pronotum and mesonotum each with 2 rows of bristles, last row with intercalaries. Mesepisternum somewhat re-

ticulated, with 2 bristles. Mesepimeron with 3 bristles. Metanotum with 4 rows of bristles; anterior row reduced to about 3

or 4 bristles. Flange of metanotum with 9 to 11 spinelets.

Legs. Typical of genus. Metatibia with posteromarginal notches with strong bristles arranged from upper to lower as fol-

lows: 2-2-2- 3-2-3.

Modified abdominal segments (Fig. 23C). Tergum VIII small, with single row of bristles. Sternum VIII relatively large,

caudally in form of subangular lobe, with 1 row of 5 to 7 unequal bristles. Caudal flaps of sternum VIII opened at 0.33 of

ventral margin. Dorsal and anal lobes of proctiger as in other Polygenis. Fixed process of clasper (Fig. 23B) with subangular

apical lobe. Dorsal margin barely sinuate, ventral and posterior margins strongly sinuate but without true indentations. Dor-

sal area of fixed process very setose; rest of bristles of this structure scantly distributed over ventral, posterior and inner ar-

eas. Movable process of clasper (Fig. 23B) slightly shorter than distal arm, with broad apex, usually of 3 lobes. Distal arm of

sternum IX (Fig. 23A) curved cephalad, with broad base, but with about equal width throughout most of its length, numer-

ous bristles oriented posteriorly. Heel of sternum IX small, with short slender terminal tendon. Apodeme of aedeagus (Fig.

24B) with very broad portion before subrounded apex. Terminal portion of aedeagus (Fig. 24B) with prominent rounded dis-

tolateral lobes. Median dorsal lobe with posterior semiangular projection. Crochet indistinct. Lateral lobe broadly convex.

Dorsal area of terminal portion of aedeagus with distinct striated section facing apical portion of sclerotized inner tube. Lat-

eral thickening of end chamber sinuous, ended almost at level with basal segment of sclerotized inner tube. Fender long, sin-

uous and slender. Basal portion of sclerotized inner tube much shorter than apical portion of this tube, with large foramen

on posterior half. Ribs scant, with irregular distribution. Crescent sclerite short, sinuous. F ulcral latero-ventral lobe curved

caudad, with rounded apex. Heel at base of aedeagal pouch subacute, connected with apodemal rod. Vesicle ventrally ex-

panded into thick ridge of irregular shape. Final section of this ridge apparently receiving anterior portion of fluted mem-

brane. Penis rod coiled, ending on thick nob-like portion. Fluted membrane moderately developed.

Female. Head (Fig. 25A). Frons more evenly rounded than in male. Thoracic structures and legs similar to those of male.

Tergum I with 2 or 3 rows of bristles and posteromarginal row of spinelets. Other unmodified abdominal somites as in male.

Modified abdominal segments (Fig. 25B). Sternum VIII moderately developed, extended beyond longitudinal axis of ab-

domen, with 2 rows of bristles and single antepygidial bristles. Tergum VIII long, with posterior margin somewhat angular,

with several posteromarginal and submarginal bristles preceded by long irregular row of bristles. Sternum VII with postero-

caudal margin abruptly truncate, not incised, with combination of long and short bristles. Sternum VIII reduced, caudally

Mammalian-Siphonapteran Associations

139

truncate. Sternum IX short, with posterior margin straight or barely arched. Dorsal anal lobe of proctiger (Fig. 25C) with

truncate apex and prominent apical and subapical bristles. Anal stylet with 2 minute ventral bristles before long apical bris-

tle. Spermatheca (Fig. 25 B, D) with cribose bulga clearly separated from short hilla with terminal portion upturned, dilated,

semi-globular.

Length. Holotype, 2.04 mm; allotype, 2.01 mm.

Remarks. — This species is dedicated to the memory of the late Dr. Eduardo Del Ponte,

whose excellent work on Siphonaptera and other blood-sucking insects contributed to the

foundation of medical entomology in South America.

Polygenis dunni (Jordan & Rothschild)

(Figure 26)

Rhopalopsyllus dunni Jordan & Rothschild, 1922, Ectop., 1:269, Fig. 261, 262.

Remarks. - Although this species has not been yet recorded from Colombia, there is a

strong possibility that it occurs in this country. It is presently known from Panama, Venezuela

and Trinidad, and parasitizes an assemblage of hosts (see Tipton and Mendez, 1966 and Tip-

ton and Machado- Allison, 1972).

Polygenis hopkinsi, new species

(Fig. 27, 28)

Type material. — Holotype male ex Oryzomys albigularis (HTC-1838), Cerro Munchique (60 kms by road west

of Popayin); Pena del Cerro, elevation 2160 m, Departamento del Cauca, Colombia, 11.V.1967, H. Trapido.

The holotype is in the U.S. National Museum of Natural History.

Diagnosis. — Males of Polygenis hopkinsi are similar to those of P. litargus Jordan & Roths-

child, from which they are readily differentiated by having the terminal portion of the scler-

otized inner tube bent upward. In P. litargus males, this structure ends barely sinuate and or-

iented cephalad.

Description. — Male. Head (Fig. 27A). Frons rounded, its contour not interrupted by unpronounced clypeal tubercle.

Preantennal area above clypeal tubercle with micropores. Preantennal row of bristles of 5 bristles, one near antenna exceeds

others in size. Preocular row of 3 principal long bristles and short secondary bristles separated from displaced minute bristle

inserted near ventral margin. Eye subovate, moderately pigmented, with deep ventral indentation. Tentorium with arched

anterior arm preceding principal stem. Oral angle short, acuminate. Tip of maxilla extended to anterior portion of last seg-

ment of maxillary palpus. Genal angle subacuminate. Postantennal region with 3 rows of gradually increased number of bri-

stles. Micropores limited to space between antennal base and first row of bristles. Group of short and thin bristles located

near scape of antenna.

Thorax. Mesonotum with 3 rows of bristles, first row reduced, of 2 or 3 bristles. Mesepisternum with 2 bristles. Mesepi-

meron with 3 bristles. Lateral metanotal area with 2 large bristles preceded by 2 short bristles. Metepisternum with single

long bristle and group of short bristles on anteromarginal projection.

Legs. As in other Polygenis. Metatibia with strong bristles inserted in notches as follows: 2-2-2-3-2-3.

Abdomen. Tergum I with 3 rows of bristles, first row reduced to about 4 bristles. Remaining unmodified terga with 2

rows of bristles. Sternum I with scattered ventral, subventral and inner bristles. Other unmodified sterna with single row of

bristles.

Modified abdominal segments (Fig. 27C). Tergum VIII short but extended to abdominal axis. Sternum VIII large, with

truncate caudomarginal expansion, its ventral division clearly posterior to row of bristles. Fixed process of clasper (Fig. 27B)

with dorsal margin moderately sinuate, gradually elevated toward apical portion. Posterior margin of fixed process very un-

dulate, with short semiangular protrusion proximad to fovea of fixed process. Ventral margin deeply indented. Bristles of

fixed process as illustrated. Movable finger (Fig. 27B) slightly bent upwards, not extended to apex of fixed process of clas-

per, clothed with several marginal, submarginal and inner bristles. Distal arm of sternum IX (Fig. 28A) semi-falcate, almost

as long as proximal arm, with anterior margin devoid of bristles and posterior margin with bristles of moderate size distri-

buted over half its length. Heel of sternum IX heavily sclerotized, with relatively short tendon. Distolateral lobes of aedeagus

(Fig. 28B) with anterior portion subrounded and posterior portion obtuse, prolonged into large ventromarginal extension.

Median dorsal lobe with subangular apical portion. Apicomedian sclerite of aedeagus striated. Crochet imperceptible. Later-

al lobe undulate, with distinct superior subangular projection. Dorsal area of terminal portion of aedeagus with small stria-

ted section facing junction of apical and basal portions of sclerotized inner tube. Lateral thickening of end chamber sinuous,

ended at level of vesicle. Lateroventral sclerite sickle-shaped but with rounded tip. Heel at base of aedeagal pouch semi-acum-

inate, attached to apodemal rod. Fender semi-arched, slender and elongate. Apical portion of sclerotized inner tube with 2

Quaest. Ent., 1977 13 (2)

140

Mendez

loops, its terminal section wide, upturned, oriented cephalad, with semiglobular apex. Basal portion of sclerotized inner tube

with mesal foramen of moderate size. Ventral nodular section of basal portion large and prominent. Ribs distributed from

area near vesicle to fender. Crescent sclerite arched, not prominent. Side pieces semi-triangular, with opposite ends acuminate.

Length. Holotype, 2.31 mm.

Remarks. — This species is named for the late G.H.E. Hopkins in recognition of the outstand-

ing contributions he made to the systematics of Siphonaptera and Anoplura.

Polygenis klagesi (Rothschild)

(Figure 29)

Pulex klagesi Rothschild, 1904, Novit. Zool., 11:620, PI. 9, Fig. 28; PI. 10, Fig. 35, 39.

Material examined. — Ex Proechimys semispinosus. Departo. del Valle - <5, Rio Raposo, XI.

Ex Hoplomys gymnurus. Depto. del Valle - 11(5, 11? , Alto Anchicaya', 650 m., II.

In addition to material from the southwest part of Colombia, I have examined 1 1 males and

17 females from Carimagua, Depto. del Meta, 8 males and 1 1 females from the Depto. de Antio-

quia and large series of males and females from Depto. del Choco.

Remarks. - Our specimens of Polygenis klagesi taken on Hoplomys gymnurus from Alto

Anchicaya, Depto. del Valle, and Curiche, Depto. del Choco, do not agree in certain features

with specimens taken on Proechimys semispinosus which we are tentatively interpreting as typ-

ical Polygenis klagesi samuelis. In view of the considerable amount of variation displayed by

P. klagesi , it is advisable not to try to segregate Colombian populations into subspecies until

an adequate study of material obtained throughout the geographical range of these fleas is

done.

Polygenis klagesi is presently known from Brazil, Panama, Costa Rica, Colombia, Venezuela,

Trinidad and Ecuador, from sea level to altitudes below 900 meters.

The P. klagesi complex occurs on a spectrum of hosts; however, these fleas are more natur-

ally associated with the spiny rat family Echimyidae, particularly with Proechimys.

Polygenis pradoi (Wagner)

(Figure 30)

Rhopalopsyllus pradoi Wagner, 1937, Zeits. Parasit., 9:420, Fig. 4.

Material examined. — Ex Oryzomys albigularis. Depto. del Valle, Municipio de Cali — 2(5, 29, Pichindd(La

Esperanza), 1900 m., I, VIII.

Ex Oryzomys caliginosus. Depto. del Valle, Municipio de Cali - 39, Quebrada Honda, nr. Pichinde', 1800 m., IX, X; 9(5,

189, Valle del Rio Pichindd, 1700- 1900 m„ I, III, VI-XI;4(5, 1 1 9 , Pichindd, 1780-1900 m„ X.

Ex Rhipidomys latimanus. Depto. del Valle, Municipio de Cali — (5, Pichindd, 1900 m., XI.

Ex Thomasomys fuscatus. Depto. del Valle, Municipio de Cali - 9, Valle del Rio Pichindd, 1700 - 1900 m., XI.

Remarks. — Polygenis pradoi has been reported from Brazil. Our specimens from southwes-

tern Colombia represent the first records of this flea species for the country. Oryzomys (Mel-

anomys) caliginosus , probably the commonest rodent in southwestern Colombia, stands out

as the more favored host in this territory. Of 48 specimens of P. pradoi obtained, 42 were from

that host while four were taken on Oryzomys albigularis , one on Rhipidomys latimanus and

one on Thomasomys fuscatus. The reports of this species from Brazil concern the following

hosts: Nasua socialis, Didelphis cancrivora, Oryzomys physodes, Akodon sp. (possibly cursor ),

a wild rat and wild mouse (Johnson, 1957).

Polygenis roberti beebei (I. Fox)

(Figure 31)

Rhopalopsyllus beebei I. Fox, 1947, Zool: N.Y. Zool. Soc., 32:117 , Fig. 2.