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Quaestiones
Entomologicde
A periodical record of entomological investigations/
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.
VOLUME 15
1979
CONTENTS
ii
Book Review — Traugott-Olsen and E. Schmidt Neilsen. 1977. The Elachistidae
(Eepidoptera) of Fennoscandia and Denmark 1
Dosdall and Lehmkuhi — The Stoneflies (Plecoptera) of Saskatchewan 3
Corrigenda — Upper Oligocene fossil pupae and larvae of Chaoborus tertiarius
(von Hey den) (Chaoboridae, Diptera) from Western Germany 117
Editorial — To Compute or Not Compute 119
Borkent — Systematics and bionomics of the species of the subgenus Schadonophasma
Dyar and Shannon Chaoboridae, Diptera) 122
Jones and Heming — Effects of temperature and relative humidity on embryogenesis in
eggs of (Walker) (Eepidoptera: Noctuidae) 257
Guest Editorial — New Tools in Publishing 295
Kuster — Comparative structure of compound eyes of Cicindelidae and Carabidae
(Coleoptera): Evolution and scotopy and photopy 297
Wong — Insect damage to old oak beams at Lower Fort Garry, Manitoba 335
Shorthouse — Observations on the snow scorpionfly Boreus brumalis Fitch (Boreidae:
Mecoptera) in Sudbury, Ontario 341
Straneo — Notes about classification of the South American Pterostichini with a key for
determination of subtribes, genera and subgenera (Coleoptera: Carabidae) 345
Kurtak — Food of black fly larvae (Diptera: Simulidae); seasonal changes in gut contents
and suspended material at several sites in a single watershed 357
Editor’s Aknowledgements 375
Bell and Bell — Rhysodini of the world Part II. Revisions of smaller genera
(Coleoptera: Carabidae or Rhysodidae) 377
Wheeler — Revision and cladistics of the Middle American genus CreagropHoms Matthews
(Coleoptera: Leiodidae) 447
Corrigenda — Volume 15(3) 481
INDEX
iii
Abbot, W.S., 270,292
abbreviata, Isocapnia, 32, 33
abnormis, Acroneuria, 3,13,46,49,50,65,
70,79,80,89,109
abnormis. Per la, 49
Acanthametropodinae, 75
Acipenser fulvescens, 7, 75
acrea, Estigmene, 265, 278, 290
Acroneuria, 13, 16, 18, 48, 77
abnormis, 3, 13, 46, 49, 50, 65, 70,
79, 80, 89, 109
lycorias, 3, 13, 46, 47, 49, 50, 51, 65,
69, 79, 89, 109
Okanagan, 5 1
pad fie a, 51,72
perbranchiata, 50
pumila, 51
salvelini, 46
(Hesperoperla) pacifica, 51
Acroneuriinae, 46
Adephaga, 313
Agonini, 345
Agrotis orthogonia, 290
Aitken, T.H.G., 135, 205
Akehurst, S.C., 191, 195, 205
AlRawy, A., 285,286,292
albatus, Chaoborus, 126, 127, 135, 161, 198
alevins, 7
algae, 55, 357, 358, 362, 363, 371
filamentous, 363
Alhonen, P., 199, 205
Allan, J.D., 127, 205
Allee, W.C., 285, 292
Allen, J.L., (see W.H. Miller), 298, 309, 316,
317
Allocapnia recta, 1
Allonarcys, 19
Alloperla, 7, 43, 45, 76
lineosa, 44
signata, 45
(Suwallia) lineosa, 44
(Triznaka) signata, 45
alpinus, Chaoborus, 125, 213
alticolor, Geocoris, 292
Amblycheila, 307,309,31 1,317
cylindriformis, 315
schwarzi, 297, 298, 299, 300, 301, 302,
305, 307, 309, 310, 31 1,313, 316,
321, 322, 323, 324, 325
americana, Arcynopteryx, 53
am eric ana, Perlodes, 54
americanus, Chaoborus, 126, 127, 158, 176,
177, 183, 185, 189, 190, 202, 206, 207
americanus, Elaphrus, 291 , 302, 305, 307,
310, 332, 333
Amphinemura, 12, 15, 18
linda, 3, 12, 37, 38, 39, 40, 66, 61, 69,
80, 86, 87, 105
Analetris eximia, 70, 75
Anderson, D., 162, 205
Anderson, J.M., (see D. Ludwick), 286, 293
Anderson, J.R., 358, 372
Anderson, R.S., 127, 205
angulata, Paracapnia, 3, 12, 25, 26, 66, 81,
82, 84, 102
annulatus, Chaoborus, 135
Anobiidae, 336, 338
anomalus, Chaoborus, 126, 127, 135, 183,
198
Anoplognathus, 317
pallidicollis, 309
Anoplura, 209
Antarctiina, 345, 347
aphids, 187
Apidae, 309
A pis mellifera, 311, 317
mellifica, 309,310
appendiculata, Corethrella, 187
apricots, 7
aquatic insects, 5, 72, 73, 75
aquatic mites, 363
arctica, Claassenia, 48
arctica, Nemoura, 68
arcticus, Thyrnallus, 1
Arcynopteryx, 13, 16, 17, 19, 53, 54
americana, 53
compacta, 3, 13, 53, 54, 66, 68, 69, 90,
111
IV
Arcynopteryx ignota, 53
inornata, 53
lineata, 53
minor, 53
(Skwala) parallela, 54
armyworm, 287, 293, 294
Arnett, R.H., 311,314
arthropods, 199,343,363,
arthropods, terrestrial, 214
Ashlock, P.D., 194, 205
aspen, 8
assimilis, Dineutes, 317
astictopus, Chaoborus, 135, 157, 197, 198
Atchley, W.R., 162, 205
Atmar, G., (see K.W. Stewart), 53, 78
atricolour, Geocoris, 286
auratus, Carabus, 309
aureum, Simulium, 373
Australomochlonyx, 199
Autrum, H., 310,316
auxiliaris, Chorizagrotis, 290
Bacon, O.G., (see D.M. Dunbar), 286, 292
bacteria, 362, 372
bacteria, coliform, 372
bacteria, rheophilic, 372
badia, Pteronarcella, 3, 12, 21, 22, 66, 69,
80, 83, 103
badia, Pteronarcys, 21
Bailey, C.G., 258, 260, 265, 286, 288, 292
Baker, W.L., 336, 337
Ball, G.E., 346,354
balsamifera, Populus, 183
Balvay, G., 126, 205
Banks, N., 4, 36, 43, 44, 45, 48, 51, 61, 72
Barathra configurata, 293
Bates, D.H., 261, 263, 264, 270, 278
bathana, Chagasia, 158
Bauer, T., 310,314
Baumann, R.W., 19, 20, 23, 33, 34, 35, 36,
37, 40, 45, 51, 72
Beal, R.S., Jr., 337
Beck, A.P., 183, 205
beetle, 297, 299, 300, 301, 305, 307, 31 1, 315,
316, 317
adephagan, 313
anobiid, 336
carabid, 203
cicindelid, 322, 323, 325, 326
death watch, 335, 336
ground, 3 1 1
larder, 336, 337
predatory, 317
tiger, 398, 311, 315, 317
Beirne, B.P., 258, 265, 292
Belehradek, J., 285, 292
belfragei, Cicindela, 297, 301, 302, 305, 310,
312, 313, 328, 329
Belkin, J.N., 136, 205, 208
Bell, J.R., 375
Bell, R.T., 375
Bell, W., (see R.Macy), 7,75
Bellamy, R.E., (see H.G. James), 127, 209
Belton, P., (see H.G. James), 127, 209
Berg, K., 135,206
Bernard, F., 309, 310, 311, 316
Bernard, G.D., 298, 315, 317
Bernhard, C.G., 300, 315
Bertha armyworm, 258
Betney, E., (see J. Green), 126, 208
bicaudata, Phryganea, 55
bicaudata, Diura, 3, 13, 55, 66, 67, 68, 69,
81, 90, 112
Bidlingmayer, W.L., 129, 206
biguttatus, Notiophilus, 307
bilineata, Isoperla, 3, 13, 58, 59, 60, 61, 66
61, 70, 82, 91, 92, 1 13
bilineata, Sialis, 60
birch, white, 342
Bird,C.D., 69,72
birds, 7, 203
birds, passerine, , 7, 73
black fly, 295, 357, 358, 363, 371, 372, 373,
374
Blackwelder, R.E., 345, 346, 354
V
Blanchard, R., 135, 206
Bliss, C.L, 270,292
blood-sucking Corethella, 2 1 1
Bodnaryk, R.P., 258, 265, 292
bolls, 258
Bolyphanthes index, 344
Bombyx mori, 288, 289, 293
Bonomi, G., 126, 206
boreal forest, 8, 69
borealis, Chaoborus, 213
Boreidae, 295, 341, 344
borer, corn, 28 6,293
Boreus, 341, 343, 344
brumalis, 295, 341, 342, 343, 344
calif ornicus, 344
hyemalis, 341, 344
nivoriundus, 341
Borkent, A., 122, 127, 207
Bott, H.R., 310, 314
Brachyptera fosketti, 23
zelona, 23
Brachypterinae, 22, 77
Bradley, J.D., 311,315
Bradshaw, W.E., 126, 206
Braitenberg, V., 311,315
brakeleyi, Corethrella, 187
brasiliensis, Chaoborus, 196, 197, 198
Bras sic a campestris, 258
napus, 258
brevis, Chloroperla, 43
brevis, Hastaperla, 3, 13, 43, 44, 66, 67, 69
82, 88, 108
Brinck, P., 24, 53, 54, 72
brook trout, 7
Broscus cephalotes, 310
Brous, H.A., 311, 316
Brown, B.W., 209
Brown, W.J., 133,206
brumalis, Boreus, 295, 341, 342, 343, 344
Brundin, L., 194,206
Brunetti, E., 146, 206
brunskilli, Chaoborus, 123, 127, 140, 146,
155, 157, 159, 175, 176, 177, 199, 225
Brydon, H.W., 126, 206
bryophytes, 69, 72
Bugnion, E., 309,315
bull-dogfish, 210
Burghause, E., 310,315
Bursell, E., 264, 285, 286, 287, 289, 290, 291,
292
Burton, G.J., 358, 372
Bush, G.L., 203, 206
Buxton, P.A., 264, 287, 292
Byers, G.W., 341, 344
cabbage looper, 389,392
Cable, R.M., (see D. Ludwick), 290, 293
caddisflies, 10, 77
calcarea, Hastaperla, 43
californica, Malenka, 3, 7, 12, 39, 40, 66,
67, 69, 71, 86, 87, 94, 105
californica, Nemoura, 39
californica, Nemoura (Malenka), 39
californica, Pteronarcys, 19, 73
californicus californicus, Omus, 297, 298, 313
californicus, Boreus, 344
californicus, Omus, 299, 300, 302, 305, 307,
309, 310, 321, 322, 325
Callosamia promethea, 293
campestris, Brassica, 258
campestris, Cicindela, 310
campestris, Gryllus, 309,315
Capnia, 9,12,15,18,25,26,30,75
coloradensis, 3, 12, 26, 27, 28, 29, 66, 69,
71, 84, 85, 93, 100
Columbiana, 30, 75
confusa, 3,12, 26, 27, 28, 29, 66, 69, 71
81, 82, 84, 85, 93, 102
crinitia, 32
gracilaria, 3, 12, 26, 27, 29, 66, 70, 71, 80,
81, 84, 85, 93, 100
nearctica, 68
nivalis, 28
opis, 25
trava, 3 1
vernalis, 3, 12, 25, 27, 28, 30, 66, 69, 84,
85, 93, 101
(Utacapnia) trava, 32
Capniidae, 5, 14, 18, 33, 72, 74, 75, 84, 85, 93
Capniids, 25
Capniinae, 25
VI
Carabidae, 295, 298, 305, 307, 308, 310, 31 1, Chaoborus ceratopogones, 183, 198
313, 314, 315, 316, 375
Carabus auratus, 309
glabmtus, 310
carinatus, Hadrobregmus, 338
Carlson, S.D., 309,315
Carlsson, M., 358, 372
Carolina, Megacephala, 299, 300, 301, 302,
305, 307, 309, 310, 321, 322, 323, 325
Carter, J.C.H., (see G.E. Pope), 126, 127, 212,
Casey, T.L., 310, 311, 315
Castle, G., 36, 72
Catapiesini, 345
caterpillar, 278
saltmarsh, 290
cecropia, Platysamia, 293
cephalotes, Broscus, 310
ceratopogones, Chaoborus, 183, 198
Chaetogenyina, 345, 347
Chagasia bathana, 158
chalcid parasite, 289
Chance, Mary M., 358, 373
Chaoboridae, 117, 1 19, 129, 187, 196, 205,
206, 207, 208, 211, 212, 213, 214, 215
Chaoborinae, 187,206,211,213
Chaoborus, 119, 125, 126, 127, 128, 129,
131, 134, 135, 136, 137, 155, 156, 157,
158, 160, 161, 175, 180, 183, 184, 185,
189, 190, 191, 192, 193, 194, 195, 196,
197, 198, 199, 200, 201, 202, 203, 205,
206, 207, 209, 210, 21 1, 212, 213, 214,
215, 252
albatus, 126,127,135,161,198
alpinus, 125, 213
americanus, 126, 127, 158, 176, 177,
183, 185, 189, 190, 202, 206, 207
annulatus, 135
anomalus, 126, 127, 135, 183, 198
astictopus, 135, 157, 197, 198
borealis, 213
brasiliensis, 196, 197, 198,
brunskilli, 123, 127, 140, 146, 155, 157,
159, 175, 176, 177, 199, 225
cooki, 123, 127, 129, 130, 131, 133, 134,
137, 139, 140, 146, 147, 151, 154, 155,
156, 158, 159, 160, 161, 162, 165, 166,
170, 173, 175, 176, 177, 178, 179, 180,
181, 184, 185, 187, 188, 189, 190, 191,
192, 193, 194, 195, 198, 199, 200, 201,
203, 217, 219, 220, 222, 223, 227, 229,
233, 234, 236, 237, 238, 239, 240, 241,
242, 243, 244, 247, 248
cooki-nyblaei, 201, 203, 255
crystallinus, 126, 127, 156, 187, 189,
190, 198, 199, 205, 209, 213
edulis, 136, 156, 214
flavicans, 125, 126, 127, 155, 156, 157,
158, 159, 160, 161, 176, 179, 183, 187,
189, 190, 198, 199, 205, 208, 212, 213,
214
fuliginosus, 128
knabi, 123
maculipes, 135
magnificus, 196, 198
nyblaei, 123, 127, 134, 136, 137, 138, 139
140, 145, 147, 151, 152, 154, 155, 161,
162, 165, 173, 175, 176, 177, 178, 181,
184, 185, 187, 192, 195, 199, 200, 201,
203, 223, 226, 230, 235, 244
obscuripes. 111, 189
pallidipes, 127, 156, 198, 208
pallidus, 154, 191, 196, 198
pilipes, 152
punctipennis, 125, 126, 127, 135, 146, 183,
197, 198, 206, 209, 210
tertiarius, 117,206
trivattus, 140
trivittatus, 123, 127, 128, 129, 130, 133,
134, 137, 138, 139, 140, 144, 146, 147,
154, 155, 156, 157, 158, 159, 160, 161,
162, 165, 166, 170, 173, 175, 176, 177,
178, 179, 180, 181, 183, 184, 185, 187,
188, 189, 190, 191, 192, 193, 194, 195,
199, 200, 201, 202, 207, 214, 219, 220
222, 225, 228, 231, 232, 236, 237, 239,
240, 242, 243, 244, 245, 246, 248, 249,
250, 255
vu
Chaoborus underwoodi, 127
velutinus, 127
(Sayomia) edulis, 21 1
Chapman, J. A., 341,344
Chapman, R.F., 289, 292
Chaston, I., 127, 206
Chi, C., 309,315
Chionea, 344
Chipley, J.R., 372,373
Chironomidae, 48, 49, 51, 52, 53, 55, 56,
129, 130, 160, 187, 191, 210, 213, 123,
343, 344
Chironomus riparius, 130, 208
Chloroperla, 43
brevis, 43
si^nata, 45
transmarina, 65
Chloroperlidae, 5, 16, 19, 42, 43, 53, 67, 73,
76, 88, 97
Chloroperlinae, 43
Chorizagrotis auxiliaris, 290
Choudhuri, D.K., 286, 292
Christenson, R.O., (see J. Needham), 64, 76
cichlids, endemic, 208
Cicindela, 298, 301, 302, 307, 312, 317
befragei, 297, 301, 302, 305, 310, 312,
313, 328, 329
campestris, 310
duodecimguttata, 207
hybrida, 310,317
lepida, 297, 298, 301, 305, 309, 310, 312^
313, 327, 328
limbalis, 297, 305, 307, 312
limbata, 312
limbata nympha, 297
longilabris, 297,312
maritima, 207
repanda, 312
repanda repanda, 297
silvatica, 310
tranquebarica, 291 , 298, 299, 300, 301,
302, 305, 307, 310, 312, 320, 321,322,
323, 325
Cicindelida lemniscata, 312
pilatei, 312
Cicindelidae, 295, 298,299, 302, 305, 307,
311, 313, 314, 334
Cicindelini, 311,312
Cimex lectularius, 293
Cinclus mexicanus, 75
cinctipes, Mochlonyx, 187, 21.1
cinctipes, Nemoura, 36
cinctipes, Nemoura (Zapada), 36
cinctipes, Zapada, 3, 12, 36, 66, 69, 70, 80,
81, 86, 87, 106
Claassen, P., 5, 1 1, 14,20,21,22,27,28,29,
30, 32, 33, 35, 36, 38, 39, 40, 41, 42, 43, 44,
45^ 46,47, 48, 49, 50, 51, 52, 54, 56, 57, 60,
60, 62, 63, 72
Claassenia, 13, 16, 18,47,48
arctica, 48
languida, 48
sabulosa, 3, 13, 46, 47, 48, 65, 69, 78, 80
89, 110
Cladocera, 130,191,205,207,208
Claire, E., 7, 72
clarki, Salmo, 1
clay, 362
Clearwater, J.R., 258, 292
Cleugh, T.R., 127, 144, 206
Clifford, H.F., 36, 66, 72
Cnephia mutata, 360,369,371,372
Coccinellidae, 293
Cockle, J.W., 343,344
codling moth, 294
, Coleman, M.J., 23,73
Coleoptera, 75, 286, 292, 293, 295, 298, 314,
315, 316, 317, 337, 375
coliform bacteria, 372
Collembola, 212, 286, 292, 343
colloids, 362, 372
coloradensis, Capnia, 3, 12, 26, 27, 28, 29, 66,
69, 71, 84, 85, 93, 100
colubrinus, Isogenoides, 3, 13, 56, 57, 66, 71
81, 82, 90, 112
colubrinus, Isogenus, 56
colubrinus. Isogenus (Isogenoides) frontalis, 56
Columbiana, Capnia, 30, 75
Common Merganser, 7
compacta, Arcynopteryx, 3, 13, 53, 54, 66, 68,
69,90,111
viii
com pacta, Dictyopteryx, 53
Comstock, J.H., 11,73
configurata, Barathra, 293
configurata, Mamestra, 1 19, 257, 258, 259,
260, 261, 262, 263, 264, 265, 270, 277,
278, 286, 287, 288, 289, 290, 291, 292,
293, 294
confusa, Capnia, 3, 12, 26, 27, 28, 29, 66, 69,
71, 82, 84, 85, 93, 102
confusum, Tribolium, 286
Cook, E.F., 125, 126, 127, 140, 206, 209
Cook, W.C., 290, 292.
cooki, Chaoborus, 123, 127, 129, 130, 131,
133, 134, 137, 138, 140, 146, 147, 151,
154, 155, 156, 158, 159, 160, 161, 162,
165, 166, 170, 173, 175, 176, 177, 178,
179, 180, 181, 184, 185, 187, 188, 189,
190, 191, 192, 193, 194, 195, 198, 200,
201, 202, 203, 217, 219, 220, 222, 223,
227, 229, 233, 234, 236, 237, 238, 239,
240, 241, 242, 243, 244, 247, 248
cooki-nyblaei, Chaoborus, 201, 203, 255
Cooper, K.W., 341, 343, 344
copepods, 130, 191
Corbet, P.S., 177, 206
Corbet, S.A. (see J. Green), 126, 208
Corethra, 206, 207, 211
nyblaei, 152
pilipes, 152
plumicornis, 205, 215
trivittata, 136, 139
Corethrella, 126, 187
appendiculata, 187
brakeleyi, 187
japonica, 187
wirthi, 187
Corethrinae, 206,212
coriaceus, Procrustes, 310
corn borer, 286, 293
corn earworm, 286, 293
corrupta, Epilachna, 286, 293
cotton, 292
Coulianos, C.-C., 343, 344
Coupland, R.T., 7, 73
Craighead, F.C., 336, 337
Crampton, G.C., 5,73,341,343,344
Cratocerina, 345, 346, 352
crayfish, 47
Creagrophorus, 375
Creagrophorus, revision and cladistics, 447-479
Cremer, R., (see H.F.J. Swilste), 126, 214
Creophilus erythrocephalus, 309, 316
cricket, 8,315
crin i tia, Capn ia, 3 2
crinitia, Isocapnia, 3, 12, 32, 33, 66, 70, 81,
82, 85, 102
Cross, H.F., (see C.R. Twinn), 127, 214
Crozier, W.K., 285, 287, 292
Cryophila, 199
crystallinus, Chaoborus, 126, 127, 156, 187,
189, 190, 198, 199, 205, 209, 213
Csiki, F., 345, 346, 354
culcid, 158, 160, 191
culcid larvae, 1 3 1
Culicidae, 158, 205, 206, 207, 213, 214
culiciformis, Mochlonyx, 21 1
Culicinae, 140,213
Culicites tertiarius, 118
Culiseta inornata, 130
Cummins, K.W., 75,371,373
cupreus, Elaphrus, 310,314
Cushing, C.F., 5,21,50,73
cut-throat trout, 7
cutworm, 292
Cy bister, 309,318
Cyclorrhapha, 208
cylindriformis, Amblycheila, 3 1 5
Cymbella, 363, 369
Dahlbominus fuscipennis, 289
damselflies, 77
Darlington, P.J. Jr., 194, 206
Davidson, J., 287, 292
Davies, D.M., 358,360,373
Davies, R.G., 341,344
death watch beetle, 335,336
decepta, Leuctra, 41
Deckart, M., 135,206
Decker, G.C., (see Matteson, J.W.), 286, 373
decolorata, Isoperla, 3, 13, 58, 59, 61, 66,
68, 69, 71, 91, 92, 98, 115
decolorata. Per la, 60
IX
decorum, Simulium, 360
Deevy, E.S., 127, 206
delicatula, Nemoura, 35
delicatula, Nemoura (Podmosta), 35
delicatula, Podmosta, 3, 12, 35, 66, 70, 71, 86,
87, 94, 105
Deonier, C.C., 126, 135, 138, 156, 157, 206,
210
Dermestes lardarius, 337,339,
dermestid, 337
diatoms, 54, 357, 362, 363, 369, 371,
372,
Dicke, R.J., 358, 372
Dickenson, W.E., 127, 206
Dictyopterygella postica, 55
Dictyopteryx compacta, 53
Diesendorf, M.O., 309,315,316
dilatatus, Nitrocris, 73
Dineutes assimilis, 310, 317
Diptera, 48, 1 17, 1 19, 205, 206, 207, 208,
209, 211, 212, 213, 215, 286, 292, 295,
373
Diura, 13, 17, 19, 53, 55
bicaudata, 3, 13, 55, 66, 67, 68, 69, 81,
90, 112
Dixa, 212
Dixidae, 212
Dixinae, 207
Dodson, S.I., 127, 206
domestica, Musca, 309, 311
dorsata, Pteronarcys, 3, 7, 12, 20, 21, 65,
70, 76, 80, 83, 103
dorsata, Sialis, 20
Doving, K.B., 298,309,315
Downes, J. A., 187,206
downesi, Plecopteracoluthus, 47, 51, 77,
Drosophila, 289
melanogaster, 290, 293
Dunbar, D.M., 286, 292
duodecimguttata, Cicindela, 207
Dyar, H.G., 126, 127, 207
Dytiscidae, 75, 309, 317
Dytiscus, 316
marginalis, 3 1 0
earworm, corn, 286, 293
Edman, J.D., 126, 215
edulis, Chaoborus, 136, 156,214
edulis, Chaoborus (Sayomyia), 211
Edwards, F.W., 126, 127, 207
Edwards, J.L., (see M.K. Hecht), 127, 208
Edwardsops, 126
eggs, 7, 119, 123, 124, 127, 128, 130, 134,
136, 137, 152, 178, 183, 185, 187, 189,
190, 192, 193, 201, 202, 208, 259, 260,
262, 263, 277, 278, 288, 293, 337, 338
Eggleton, E.E., 183,207
Elachistidae, 1, 2
Elaphrus, 313
americanus, 297, 302, 305, 307, 313,
332, 333
cupreus, 310,313
riparius, 310, 314
Elder, J.A., 20, 73
elephant snout fish, 210
Elson, J.A., 69, 73
Embioptera, 6
embryos, 3, 259, 260, 262, 263
embryos, salmon, 76
embryos, salmonid, 72
Emery, W.T., 358,373
endemic cichlids, 208
Ephemeroptera, 47, 48, 52, 72, 75
Epilachna corrupta, 286, 293
Erickson, A.M., 299, 315
Erioptera nyblaei, 151
Erwin, T.L., 346, 354
erythrocephalus, Creophilus, 309, 316
Estigmene acrea, 265, 278, 316
Euchroina, 345, 347, 352
Eucorethra, 127, 196, 200
underwoodi, 196
Eutrichomerus terricola, 309
eximia, Analectris, 70, 75
Exner, S., 298, 309, 315
exuviae, 127, 129, 161, 162, 187, 188, 192
falsa, Pseudohadrotoma, 337
fasciatus, Oncopeltus, 211 , 286, 290, 294
Fedorenko, A.Y., 126, 127, 207, 214
Felt, E.P., 127, 207
ferruginea, Lecutra, 3, 13, 40, 41, 66. 69, 81,
82, 86, 87, 107
X
ferruginea, Nemoura, 41
filamentous algae, 363
Filipalpia, 6,
firefly, 316
fish, bull-dog, 210
fish, elephant-snout, 210
fishes, 7, 48, 75, 126, 210
Fitch, A., 22, 73
Fjellberg, A., 342, 344
flavicans, Chaoborus, 125, 126, 127, 155, 156,
157, 152, 159, 160, 161, 176, 179, 183, 187,
189, 190, 198, 199, 205, 208, 212, 213, 214
flies, muscoid, 293
flies, fruit, 290, 293,
flies, tephritid, 203
Flitters, N.E., 290,293
flowers, pistillate, 187
flowers, wild 187
fontanum, Prosimulium, 360, 369, 371
fontium, Iso per la, 63
forest, boreal, 8, 69
forest, savanna, 2 1 2
fosketti, Brachyptera, 23
fosketti, Oemopteryx, 3, 12, 22, 23, 24, 66, 70,
71, 80, 82, 94, 98, 103
fossil, 123, 124, 127, 199, 201, 206, 207, 209
Fragilaria, 363
Frankenberg, G. von, 126,207
Fraser, F.C., 341, 342, 344
Fredeen, F.J.H., 372, 373
Freitag, R., 133, 207, 312, 315
Frey, D.G. 156,207
Friedrichs, H., 298, 310, 315
Frison, T.H., 11, 22, 25, 32, 39, 40, 41, 43, 46,
47, 48, 49, 50, 51, 52, 54, 57, 58, 60, 61, 62,
63, 64
frogs, tree, 187,211
frontalis, Isogenoides, 3, 13, 56, 57, 66, 69, 72,
89, 111
frontalis. Isogenus, 56, 57
frontalis. Isogenus (Isogenoides) frontalis, 57
frontinalis, Salvenius, 1
fruit, 7, 258
fruit flies, 290, 293
Fry, R.E., 265, 278, 290, 292
fuliginosus, Chaoborus, 127
fulveseens, Acipenser, 7, 75
fumosa, Isoperla, 64
fungi, 362
furciferus, Onychiurus, 286
furs, 336
fuscipennis, Dahlbominus, 289
fuscum, Prosimulium, 360
gairdneri, Salmo, 1
Gambusia georgei, 209
Gammarus, 205
Gaskins, C.T., (see W.R. Atchley), 162, 205
Gaufin, A.R., 6, 7, 11, 14, 20, 21, 22, 27, 28,
29, 30, 31, 32, 33, 35, 36, 37, 39, 40, 42,
43, 44, 45, 48, 49, 51, 52, 54, 56, 58, 62,
63, 64, 72, 73, 75, 76
Gelechioidea, 1
Geocoris alticolor, 292
atricolour, 286
pallens, 292
punctipes, 292
georgei, Gambusia, 209
Giddings, C., 309, 310, 316
Giles, G.M., 127, 207
Gimmerthal, B.A., 152,207
Gissler, C.E., 307, 315
glabratus, Carabus, 310
Glenn, C.L., 7, 73
Gnathonemus macrolepidotus, 210
gnats, 207, 208, 210
Goff, A.M., 130, 208
Goldring, P., 335, 337
Goldsmith, T. A., 298,315
Goldspink, C.R., 126,208
Gomphonema, 363
gorbushca, Oncorhynchus, 76
Goulden, C.E., 199, 208
gracilaria, Capnia, 3, 12, 26, 27, 29, 66, 70,
72, 80, 81, 84, 85, 93, 100
Grantham, B.J., 126,215
grass, trailing, 371
grasses, 8
grassland, 8
gravel, 360
grayling, 7, 77
Green, J., 126,208
Grenacher, El., 300, 315
XI
Greve, H., (see E.T. Nielson), 189, 211
Greve, L., 342, 344
Griffiths, G.C.D., 194, 208
Groves, A.E., 7, 73
Gryllidae, 309
Gryllus campes tris, 309, 315
Guppy, J.C., 287, 292,
Gyrinidae, 310,313,315,317
Gyrinus, 313
natator, 310,315
nator subtriatus, 310,315
subtriatus, 310,317
Haavisto, M., 199,205
Hadrobregmus carinatus, 338
Hagvar, S., 343, 344
Hagen, H.A., 20, 52, 56, 73
Hales, D.C., 64, 73
Hall, J.E., 7, 73
Hamilton, A.L., 127, 208
Hamilton, W., 6, 73
Hancey, J.E., (see A.W. Lindquist), 136, 210
Hanson, J., 10, 19, 25, 26, 27, 28, 29, 32, 40,
42, 53, 54, 56, 57, 74
Harden, P., 19, 20, 22, 30, 38, 41, 43, 46, 49,
50, 56, 60, 61, 62, 64, 74
hargreavesi, Simulium, 372
Harper, P., 7, 22, 25, 26, 30, 37, 38, 41, 44,
47, 50, 51, 65, 66, 67, 74
Hasler, A.D., (see K.W. Malueg), 146, 210
Hasselmann, E-M., 309,315
Hastaperla, 9, 13, 16, 19, 43
brevis, 2, 12, 43, 44, 66, 67, 69, 82, 88,
108
cal car ea, 43
orpha, 43
Hauser, B.W., (see T.R. Cleugh), 127, 206
Hays, R.A., 27, 28, 29, 33, 36, 39, 45, 48,
51, 62, 63, 73
Hazama, N., (see D. Miyadi), 126,211
Hearle, E., 127,208
Hecht, M.K., 126, 208
Heiman, D.R., 47, 65, 74
Heinemann, S.J., 136, 205, 208
Heleomyzidae, 343
Heliothis zea, 286
Hemiptera, 286
Henneberry, T.J., (see A.M. tCishabo), 296, 293
Hennig, W., 156, 201, 208
herbaceous plants, 129
Herms, W.B., 126,208
Hesperoboreus, 341
hesperomydis, Hoplopleura, 209
Hesperoperla, 13, 15, 51
pacifica, 3,7, 13, 46, 51, 65, 70, 78, 89,
109
Heteroptera, 292
Heyden, C. von, 127, 208
hickory, 336
hides, 336
Hilsenhoff, W.L., 20, 74
Hinton, H.E., 208, 336, 337
Hiodon tergisus, 7, 73
Hirvenoja, M., 126, 127, 208
Hitchcock, S.W., 6, 1 1,13, 20, 25, 30, 40,
41, 43, 44, 46, 49, 50, 52, 58, 62, 64,
74, 126, 209
Hocking, B., (see C.R. Twinn), 127, 214
Hodson, A.C., 285, 286, 292
Hofmann, W., 199, 209
Hollenberg, M.J., 299, 315
holometabolous insects, 73
Home, E.M., 308, 310, 315
Honegger, H.W., 309,315
honey, 187, 259
honey bee, 3 1 1
honey-water, 187
honeybee, 317
honeydew, 187
Hongve, D., 157, 209
Hope, F.W., 147, 208
Hopkins, D.M., 167, 202, 209
Hoplopleura, hesperomydis, 209
Hoplopleuridae, 209
Horn, G.H., 298, 311, 312, 315
Horridge, G.A., 298, 309, 310, 315, 316,
317
Howe, P.A., 287, 293
Howe, R.W., 285, 287, 290, 292
Hubbs, C., 133,209
hudsonicus. Isogenus, 57
Hull, D.L., 127,209
Humason, G.L., 261, 293, 299, 316
xii
hyalita, 31
hybrida, Cicindela, 310,318
Hydroperla parallela, 54
hyemalis, Boreus, 341, 344
Hyla, 187, 211
Hynes, 13, 22, 25, 26, 30, 37, 41, 65,
66, 67, 71, 72, 74
ignota, Arcynopteryx, 53
lilies, J., 11, 19, 33, 35, 36, 37, 38, 39, 40,
41, 42, 43, 44, 45, 51, 53, 54, 56, 57, 75
imperfectus, Onychiurus, 292
index, Bolyphanthes, 344
inornata, Arcynopteryx, 53
inornata, Culiseta, 130
Insecta, 44, 77
insects, 5, 11, 20, 61, 72, 73, 75, 77, 206, 208,
209, 210, 21 1, 213, 258, 259, 262, 264, 277,
287, 289, 294, 295, 317, 337
insects, aquatic, 4, 72, 73, 75
insects, holometabolous, 73
insects, Neuropteroid, 72
invertebrates, 65, 74, 287
loannides, A.C., (see G.A. Horridge), 298, 309,
316
Isocapnia, 12, 15, 18, 25, 31, 32, 77
crinitia, 3, 12, 32, 33, 66, 70, 80, 81, 84, 101
missourii, 3, 12, 32, 33, 66, 70, 85, 101
Isogeninae, 15, 53, 55, 89
Isogenoides, 13, 16, 17, 19, 53, 55, 56, 57,
74
colubrinus, 3, 13, 56, 57, 66, 71, 81, 82,
90, 112
frontalis, 3, 13, 56, 57, 66, 69, 71, 90, 1 12
Isogenus colubrinus, 56
frontalis, 56, 57
hudsonicus, 57
Isogenus (Isogenoides) frontalis colubrinus, 56
Isogenus (Isogenoides) frontalis frontalis, 57
Isoperla, 13, 17, 19, 52, 53, 57, 61, 63,
bilineata, 3, 13, 58, 59, 60, 61, 66, 67, 70,
82, 91, 92, 113
decoLorata, 3, 13, 58, 59, 61, 66, 68, 69, 71,
91, 92, 98, 115
fontium, 63
fumosa, 64
longiseta, 3, 13, 58, 59, 61, 66, 67, 70, 91,
91, 114
Isoperla marly nia, 3, 11, 13, 58, 59, 62, 66,
69, 92, 1 14
patricia, 3, 13, 58,59, 62, 63, 66, 70,
91, 92, 113
petersoni, 3, 13, 58, 59, 63, 64, 66, 69,
91. 92, 115
transmarina, 3, 13, 58, 59, 60, 64, 66, 67,
69.91.92, 1 15
Isoperlinae, 17,53,90,91
jack pines, 77
James, H.G., 127, 209
Jamnback, H.A., 360, 373
Janisch, E., 285, 287, 293
Japanese beetle, 290
japonica, Corethrella, 187
japonica, Popillia, 293
Jenkins, D.W., 127, 209
je'nningsi, Simulium, 363, 369
Jewett, S.G., 20, 21, 22, 28, 34, 35, 36, 39,
40, 42,44,48,54,62,63,74,75
Johannsen, O.A., 127, 209
Johnels, A.G., 343, 344
Johnson, C.G., 262, 277,285 , 286, 291, 293
Johnson, D.L., 167, 209
Jones, M.P., 126, 288, 293
Juday, C., 126,209
kannume, Mormyrus, 210
Kathroperla, 43
Kavanaugh, D.H., 194, 209
keta, Oncorhynchus, 76
Kim, K.C., 133,209
King, K.M., 258, 263, 264, 293
Kirchoffer, O., 209, 310, 316
Kishabo, A.M., 289, 293
Klapalek, F., 33, 75
kPabi, Chaoborus, 123
knabi, Sayomyia, 139, 146
Knight, A.W., 47, 74, 75
Knight, K.L., 131,209
Kolb, G., 310,316
Krogh, A., 286, 293
Kruger, E.J., 126, 210
Krumholz, L.A., 57, 65, 77
Kunze, P., 309, 316
Kurtak, D.C., 360,371,372,373
Kuster, J.E., 299, 302, 305, 308, 313, 316
xm
Lacordaire, M.T., 311,312,316
lucustrine plain, 8
Laffoon, J.L., 131,210
lake strugeon, 7
Lane, J., 126,210
LaRow, E.J., 157,192,210
lardarius, Dermestes, 331 , 339
larder beetle, 336,337
Larsen, J.R., 310,317
Larson, D.J., 68, 75
larvae, 1, 43, 48, 64, 77, 117, 123, 126,
127, 129, 130, 131, 132, 133, 134, 135, 140,
144, 146, 147, 151, 154, 155, 156, 157, 158,
159, 160, 161, 162, 165, 166, 173, 175, 178,
179, 180, 183, 184, 185, 190, 191, 192, 194,
195, 196, 197, 198, 200, 202, 206, 207, 210,
211, 212, 213, 214, 228, 229, 230, 241, 245,
246, 247, 249, 252, 258, 259, 260, 263, 270,
285, 286, 288, 290, 291, 336, 337, 357, 358,
360, 362, 363, 369, 371, 372, 373, 374
larvae, culcid, 131
lead, 336
lectularius, Cimex, 293
Lehmkuhl, D.M., 20, 26, 70, 75
Leiodidae, 375
Lemke, A.E., (see A.V. Nebecker), 23, 76
lemniscata, Cicindelida, 312
Leng, C.W., 307, 311, 316
lepida, Cicindela, 297, 298, 301, 305, 309, 310,
312, 313, 326, 327
Lepidoptera, 1,119, 258, 286, 288, 292, 293,
294
Leuctra, 1 1, 13, 17, 40, 73
decepta, 41
ferruginea, 3, 13, 40, 41, 66, 69, 81, 82, 86,
87, 107
occidentalis, 42
Leuctridae, 4, 13, 17, 33, 40, 42, 74, 77, 85,
86
Lewis, W.M., Jr. 157, 210
lichen-woodland, 8
Lienk, S.E., 126, 213
limbalis, Cicindela, 297, 305, 307, 312
limbata, Cicindela, 312
limbata nympha, Cicindela, 297
lime, 335, 336
limestone, 335, 339
Lin., S., 277, 285, 286, 289, 290, 293
linda, Amphinemura, 3, 12, 37, 38, 39, 40,
66, 67, 69, 80, 86, 87, 105
linda, Nemoura (Amphinemura), 31
Lindquist, A. W., 126,135,210
Lindroth, C.H., 346, 354
Lindsey, C.C., 68, 69, 75
lineata, Arcynopteryx, 53
lineosa, Alloperla, 44
lineosa, Alloperla (Suwallia), 44
lineosa, Suwallia, 3, 13, 44, 45, 66, 70, 71,
88, 97, 108
Linnaeus, C., 75
Linyphiidae, 344
lobata, Nemoura, 39
Loew, H., 127,210
loewii, 23
longilabris, Cicindela, 291, 312
longiseta, Isoperla, 3, 13, 58, 59, 62, 66, 67,
70, 91, 92, 114
looper, cabbage, 289, 293
Loricera pilicornis, 307
Love, D., 69, 75
Luckmann, W.H., 286, 293
Ludwick, D., 264,286,287,288,293
lycorias, Acroneuria, 3, 13, 46, 47, 49, 50,
51, 65, 69, 78, 89, 109
lycorias. Per la, 50
Lygaeidae, 292
MacDonald, W.W., 126, 210
macdunnoughi, Podmosta, 35
Maciolek, J.A., 363, 373
MacLachlan, R., 53, 75
macrolepidotus, Gnathonemus, 210
maculipes, Chaoborus, 135
Macy, R., 7, 75
madidus, Steropus, 309,311
magnificus, Chaoborus, 196, 198
Magnin, E., 7, 44, 74, 75
magnum, Prosimulium, 360, 371, 372
Main, R.A., 146, 210
Malenka, 12, 15, 18
californica, 3, 7, 12, 39, 40, 66, 67, 69,
71, 86, 87, 94, 105
XIV
Malueg, K.W., 146,210
Mamestra configurata, 1 19, 257, 258, 259, 260,
262, 262, 263, 264, 265, 270, 277, 278, 286,
287, 288, 289, 290, 291, 292, 293, 294
mammals, 187, 203, 344
manicatus, Repsimus, 309
maple, red, 342
marginalis, Dytiscus, 3 1 0
mariana, Picea, 183, 184, 185
maritima, Cicindela, 207
Marland, F.C., 199, 210
Marlin, J.C., (see D.W. Webb), 341, 342, 344
marlynia, Isoperla, 3, 11, 13, 58, 59, 62, 66,
69, 92, 114
Martini, E., 126, 210
Marzolf, G.R., 157, 210
Maslin, T.P., 196, 210
Matheson, R., 127, 210
Matteson, J.W., 286, 293
maura, Taeniopteryx, 22
Maw, M., (see H.G. James), 127,209
mayflies, 10, 54, 70
Mayr, E., 133,211
McCloy, J.S., 190, 211
McDuffie, W.D., (see C.R. Twinn), 127, 214
McGowan, L.M., 126,211
McKeever, S., 126,211
McPhail, J.D., 68, 69, 75
Mecoptera, 295, 344
media, Paragnetina, 3, 13, 46, 47, 65, 69, 74, 7'
78, 89, 1 10
media, Per la, 46
media, Togoperla, 46
Magacephala, 3 1 1
Carolina, 299, 300, 301, 302, 305, 307.
309, 310, 320, 321, 322, 323, 325
Carolina mexicana, 291 , 298, 313
Megacephalina, 312
Megacephalini, 311, 312
Megaleuctra, 25, 40
Meinert, E., 158,211
melanarius, Pterostichus, 291 , 299, 302, 305,
307, 310, 330, 331
melanogaster. Drosophila, 290, 293
mellifera. Apis, 309, 31 1, 317
mellifica, Apis, 309,310
Melon tha volgaris, 309
Melvin, R., 286, 293
Menzel, R., 310,316
Merganser, Common, 7
Meridion, 363
meromixus, 207
Merritt, R.W., 23, 56, 75
Me so Cyclops, 215
Messenger, P.S., 290, 293
Metoliophilus uraticus, 75
Meunier, F., 127,211
mexicana, Megacephala Carolina, 291 , 298,
313
mexicanus, Cinclus, 15
Meyer-Rochow, V.B., 309, 317
Magdefrau, K., 200,210
Mickel, C., 19, 20, 22, 30, 38, 41, 43, 46, 49,
50, 52, 56, 60, 61, 62, 64, 74
Microcephalina, 345, 347, 352
Microcephalina (=Tichoniina), 346
microfossils, cladoceran, 208
midges, 205,207,214
Miliam, P., 27, 28, 29, 33, 36, 39, 45, 48, 51,
62, 63, 73
milkweed bug, 286
Miller, W.H., 298, 309, 3 1 5, 3 1 7
Miner, M., 27, 28, 29, 33, 36, 39, 45, 48, 51,
62, 63, 73
mineral, 357, 362, 363, 369
, minor, Arcynopteryx, 53
minor, Perlodes, 53
missourii, Isocapnia, 3, 12, 32, 33, 66, 70,
85, 101
mites, aquatic, 363
Miyadi, D., 126, 211
Miyagi, F., 126, 211
Mochlonyx, 127,158,196,198,199,200,215
cinctipes, 187,211
culiciformis, 211
velutinus, 187, 196, 209
mogila, 32
Montchadsky, A.S., 126,211
mooneye, 7, 73
mori, Bombyx, 288, 289, 293
Morionina, 345, 346, 352
XV
Mormyridae, 210
Mormyrus kannume, 210
mosquito, non-anopheline, 2 1 3
mosquitoes, 126, 129, 187, 206, 209, 21 1,
213
mosses, 341, 342
moth, 258
moth, codling, 294
moth, noctuid, 292
moth, silkworm, 293
moths, saturniid, 293
Munroe, E.G., 68, 75
Musca domestica, 309, 3 1 1
Muscidae, 309,311
muscoid flies, 293
mutata, Cnephia, 360, 369, 371, 372,
Muttkowski, R.A., 126, 211
Mykelbust, R.J., 211
napus, Brassica, 258
Narf, R.P., 20, 74
natator, Gyrinus, 310, 315
nator subtriatus, Gyrinus, 310, 315
Naumann, E., 358, 373
nearctica, Capnia, 68
Neave, F., 5, 28, 50, 63, 64, 75
Nebeker, A.V., 21, 22, 27, 28, 29, 30, 31, 36,
40, 62, 73, 75
Nechols, J.R., (see C.A. Tauber), 203, 214
Needham, J., 5, 1 1, 14, 20, 22, 29, 32, 33, 35,
36, 38, 40, 41, 42, 43, 44, 45, 46, 48, 49,
50, 51, 52, 54, 57, 60, 61, 62
Needham, J.G., 10, 63, 72
Neeley, J.A., 358, 373, 481
Nelson, C., 20, 76
Nematocera, 211,215
Nemoura, 12, 15, 18, 33, 34,
arctica, 68
californica, 39
cinctipes, 36
delicatula, 35
ferruginea, 41
lobata, 39
nivalis, 22
rickeri, 3, 12, 34, 66, 69, 71, 85, 87, 94, 106
rotunda, 38
sahlbergi, 34
Nemoura (Amphinemura) linda, 37
(Malenka) californica, 39
(Podmosta) delicatula, 35
(Shipsa) rotunda, 38
(Zapada) cinctipes, 36
Nemouridae, 5, 10, 1 1, 14, 17, 33, 72, 74,
86, 87, 94
Neochaoborus, 126
Nesse, V., 309,317
Neuroptera, 73, 76, 77
Neuropteroid insects, 72, 77, 309, 310
Newcomer, E.J., 7, 76
Newman, E., 33, 49, 50, 57, 64, 76
Newport, G., 30, 76
ni, Trichoplusia, 289
Nicola, S., 7, 76
Nielson, E.T., 189, 211
Nilssen, L.M., (see M. Carlsson), 358, 372
Ninham, B.W., (see G.A. Horridge), 298,
315
Nitrocris dilatatus, 73
nivalis, Capnia, 28
nivalis, Nemoura, 22
nivalis, Taeniopteryx, 3, 12, 22, 69, 79, 83,
104
nivoriundus, Boreus, 341
Noctuidae, 119, 258, 290, 292, 293, 294,
non-anopheline mosquito, 213
Northcote, T.G., 146, 155, 157, 214
Notemourinae, 25
Notiophilus biguttatus, 307
Notonomus, 309
nubilalis, Ostrinia, 286
Nutting, W.L., 6, 76
nyblaei, Chaoborus, 123, 127, 134, 136, 137,
138, 139, 140, 146, 147, 151, 152, 154,
155, 161, 162, 165, 173, 175, 176, 177,
178, 181, 184, 185, 187, 192, 195, 199,
200, 201, 203, 223, 226, 230, 235, 244
nyblaei, Corethra, 152
nyblaei, Erioptera, 151
O’Conner, C.T., 200, 211
oak, 337, 339
oak, red, 342
obscuripes, Chaoborus, 127, 189
XVI
occidentalis, Lecutra, 42
occidentalis, Paraleuctra, 42
occidentalis auct., Paraleuctra, 42
Odonata, 77
Oedegonium, 372
Oemopteryx, 9, 12, 14, 17, 23
Oemopteryx fosketti, 3, 12, 22, 23, 24, 66,
70, 71, 81, 83, 95, 99, 104
Okanagan, Acroneuria, 51
Omina, 309,311
Omus, 307, 31 1, 312, 313
californicus, 299, 300, 302, 305, 307,
309, 310, 320, 321, 322, 325
californicus californicus, 291 , 298, 313
Oncopeltus, 293
fasciatus, 211 , 286, 290, 294
Oncorhynchus gorbuscha, 76
Oncorhynchus keta, 76
Onychiurus furciferus, 286
Onychiurus imperfectus, 292
op is, Capnia, 25
opis, Paracapnia, 25, 26
orchard insects, 76
oregona, Cicindela, 317
orioles, 2 1 2
orpha, Hastaperla, 43
orthogonia, Agrotis, 290
orthopteroid insects, 5, 73, 76
Oryctes rhinoceros, 309
Ostbye, E., 343, 344
ostracods, 160, 191
Ostrinia nubilalis, 286
Ostryakova-Varshaver, U.P., 288, 293
Pacaud, A., 358, 373
pacifica, Acroneuria, 51, 72
pacifica, Acroneuria (Hesperoperla), 51
pacifica, Hesperoperla, 3, 7, 13, 46, 51, 65,
70, 78, 89, 109
pacifica, Taenionema, 1
Page, W.A., (see J.N. Belkin), 136, 206
pallens, Geocoris, 292
pallidicollis, Anoplognathus, 309
pallidipes, Chaoborus, 127, 156, 198, 208
pallidus, Chaoborus, 154, 191, 196, 198
Panday, R.S., 183,212
Panorpatae, 344
Pantin, C.F.A., 299, 317
Pappas, C.D., 310,317
Paracapnia, 9, 12, 15, 18, 25, 74
angulata, 3. 12, 25, 26, 66, 80, 82, 84,
102
opis, 25, 26
Paragnetina, 13, 16, 18, 46, 76
media, 3, 13, 46, 47, 65, 69, 74, 77, 78,
79, 110
Paraleuctra, 13, 14, 18, 40, 42, 74
occidentalis, 42
occidentalis auct., 42
vershina, 3, 13, 42, 66, 70, 82, 86, 87,
107
parallela, Arcynopteryx (Skwala), 54
parallela, Hydroperla, 54
parallela, Skwala, 3, 13, 54, 66, 70, 80,
82, 90, 111
Paraperla, 43
parasites, 7, 206, 260
parasite, chalcid, 289
Parma, S., 126, 212, 214
parnassum, Simulium, 360
passerine birds, 74
patricia, Isoperla, 3, 13, 58, 59, 62, 63, 66,
70, 91, 92, 113
peaches, 7,
Peaks, L.M., 261, 285, 286, 293
Peden, A.C.,(seeC. Hubbs), 133,209
Penny, N.D., 341, 344
perbranchiata, Acroneuria, 50
Perimegatoma, 337
Perla, 49
abnormis, 49
decolorata, 60
lycorias, 50
media, 46
placida, 52
sabulosa, 48
Perlesta, 13, 16, 18, 52
placida, 3, 11, 13, 46, 52, 66, 69, 77, 110
Perlidae, 5, 10, 18, 46, 47, 51, 53, 72, 76,
77, 78, 89
Perlinae, 46
Perlites, 75, 76
Perlodes americana, 54
xvu
Period es minor, 53
Perlodidae, 5, 16, 19, 53, 55, 61, 74
Perlodinae, 17, 53, 55, 89
Perlodini, 77
Perlomyia, 40
Peteronarcys californica, 73
Peterson, A., 260, 293
Peterson, B.V., 360, 373
petersoni, Isoperla, 3, 13, 58, 59, 63, 64, 66,
69, 91, 92, 115
Petr, T., 126, 212
Pens, F., 126, 127, 183, 212
Peusomyia, 126
Phalaenidae, 294
Phasganophora, 47
pheromones, 258
Phillips, R., 7, 72
Phormia regina, 286
Phryganea bicaudata, 55
Picea mariana, 183, 184, 185
pictipes, Simulium, 360, 363, 369, 371, 372
pilatei, Cicindelida, 312
pilcornis, Loricera, 307
pilipes, Chaoborus, 152
pilipes, Corethra, 152
Pilon, J.-G., 57, 50, 51, 74
pines, 8, 69
pines, jack, 77
Pinkovsky, D.D., 358, 360, 372. 373
Pinus strobus, 363
Pisces, 210
pistillate flowers, 187
placida, Perla, 52
placida, Perlesta, 3, 1 1, 13, 46, 52, 66, 69, 77,
109
plain, lacustrine, 8
plants, 61, 187, 258, 357, 362, 363, 372
plants, herbaceous, 129
plants, rape, 259
Platysamia cecropia, 293
Plecoptera, 3, 5, 6, 9, 10, 1 1, 31, 33, 67, 70, 71,
72, 73, 74, 75, 16, 82, 1 16. 343
Plecoptera setipalpian, 74
Plecopteracoluthus downesi, 47, 51, 77
plumicornis, Corethra, 205,215
plums, 7
Podmosta, 12, 15, 18, 35
delicatula, 3, 12, 35, 66, 70, 71, 86, 87, 94,
105
macdunnoughi, 35
Poje, G.V., 183,205
polyphemus, Telea, 286, 293
Pope, G.E., 126, 127, 212
Popillia japonica, 290, 293
poplar, 8
Popoff, M., 309, 315
Populus balsamifera, 183
tremuloides, 185
postica, Dictyopterygella, 55
Power, G., (see G.E. Pope), 126, 127, 212
prairies, 8, 293
Prashad, B., 203,212
Precambrian rock, 8
predaceous water beetles, 75
predatory beetles, 317
Prest, V.K., 68, 76, 212
Priobium sericeum, 336, 337, 339
Procladius, 211
Procrustes coriaceus, 3 1 0
Prokesova, V., 126,212
promethea, Callosamia, 293
Promochlonyx, 199
Prosimulium, 313
fontanum, 360,369,371
fuscum, 360
magnum, 360,371,372
prunes, 187
Pseudaletia separata, 292
unipuncta, 287, 292
Pseudo-neuroptera, 73
Pseudohadrotoma, 331 , 339
falsa, 331
Pteronarcella, 14,17,21
badia, 3 12,21,22,66,69,80,83,103
triloba, 21
Pteronarcidae, 5, 12, 14, 19, 76, 82
Pteronarcinae, 77
Pteronarcys, 12, 14, 19, 20, 21
badia, 21
californica, 20
dorsata, 3, 1, 12, 20, 21, 65, 69, 75, 80,
83, 103
regalis, 1 6
xvm
Pterostichina, 345, 347
Pterostichini, 295
Pterostichini, Notes about classification of
the South American, 345 — 354
Pterostichus melanarius, 297,299, 302, 305,
307, 310, 331, 332
Ptinus tectiis, 286
Ptychopteridae, 212
Pucat, A.M., 188, 212
pumila, Acroneuria, 51
punctipennis, Chaoborus, 125, 125, 127, 135,
146, 183, 197, 198, 206, 209, 210
punctipes, Geocoris, 292
pupa, U7, 125, 126, 127, 129, 130, 133, 134,
138, 146, 155, 175, 176, 179, 180, 188, 192,
194, 196, 198, 206, 213, 258, 259, 260, 336
Puri, I.M., 358, 373
Putnam, L.G., 265, 293
Pyenson, L., 286, 293
Quercus, 337
Raasveldt, L.G., 127, 205
Raby, S., 8, 76
rainbow trout, 7
raisins, 187
rape plants, 258, 259, 260
Rapoport, E.H., 212
Rawson, D.S., 7, 76
recta, Allocapnia, 7
red maple, 342
red oak, 342
Reeves, B.O.K., 167, 212
regalis, Pteronarcys, 11
re gin a, Phormia, 286
Reichardt, H., 345, 346, 354
reindeer moss, 8
Reisen, W.K., 183,212
Rempel, J.C., 258
Rempel, J.G., 260, 294
Repsimus manicatus, 309
repanda, Cicindela, 312
repanda, Cicindela repanda, 297
rheophilic bacteria, 372
rhinoceros, Oryctes, 309
RhyacophUidae, 64
Rhysodidae, 375
Rhysodini, 375
Rhysodini of the world. Part II, 377-446
Ribi, W.A., 311, 317
Richards, A.G., 286, 289, 294
Richards, J.H., 8, 76
Richards, O.W., 341, 344
Richards, T., 8, 76
Richardson, J.W., 22, 48, 52, 54, 76
Ricker, W.E., 5, 11, 19, 20, 21, 22, 23, 24, 27,
28, 29, 30, 31, 32, 33 35, 36, 37, 38, 39, 40,
42, 43, 44, 45, 46, 47, 48, 49, 50, 51, 52,
53, 54, 55, 56, 57, 60, 61, 62, 63, 64, 68,
60, 73, 76, 77
rickeri, Nemoura, 3, 12, 34, 66, 69, 71, 85,
87, 94, 106
riparius, Chironomus, 130, 208
riparius, Elaphrus, 310,315
Rising, J.D., 212,
Rivalier, E., 311, 312, 317
river snails, 7
Roback, S.S., 7,77,212
rock, Cretaceous, 8
rock, Precambrian, 8
rock, sedimentary, 8
Roff, D., 126,212
Rohlf, FJ., 214, 299, 317
Ross, H.H., 11, 21, 22, 23, 40, 77, 212
Roth, J.C, 126,212
rotunda, Nemoura, 38
rotunda, Nemoura (Shipsa), 38
rotunda, Shipsa, 3, 12, 38, 66, 67, 69, 86, 87,
106
Rowe, J.S., 8, 73
rufovillosum, Xestobium, 335, 336
Russell, L.S., 69, 77
sabulosa, Claassenia, 3, 13, 46, 47, 48, 65,
69, 79, 80, 89, 110
sabulosa, Perla, 48
Saether, O.A., 125, 126, 127, 129, 138, 213,
213
sahib ergi, Nemoura, 34
Sailer, R.L, 126, 213
Salix, 184
Salmo clarki, 1
Salmo gairdneri, 1
salmon, 7
saimonid embryos, 72
XIX
saltmarsh caterpillar, 290
salvelini, Acroneuria, 46
Salvelinus fontinalis, 1
Sarnia walkeri, 293
sand, 360, 362
Sanderson, E.D., 285, 287, 294
Sartallus signatus, 309,316
saturniid moths, 293
savanna forest, 212
Sawchyn, W.W., 10, 77
Say, T., 60, 77
Sayomyia, 126, 135, 197
knabi, 139, 146
trivittata, 139
Scarabaeidae, 309,317
scavenger, 337
Schadonophasma, 119, 123, 125, 126, 127,
128, 129, 131, 133, 134, 135, 136, 137, 140,
147, 154, 155, 156, 173, 175, 176, 179, 191,
192, 193, 194, 195, 196, 198, 199, 200, 201,
202, 203, 204, 206, 221, 222, 253, 254
Schaupp, F.G., 311, 317
Schlee, D., 129,213
Schroer, W-D., 310, 317
schwarzi, Arnblycheila, 291 , 298, 299, 300,
301, 302, 305, 307, 309, 310, 31 1, 313, 317,
320, 321, 322, 323, 324, 326
scorpion, 341
scorpionflies, 295, 343, 344
Scott, D.B.C., 126,208
Scudder, S.H., 127, 213
sedimentary rock, 8
seed pods, 258
Senior-White, R., 127, 213
separata, Pseudaletia, 292
sericeum, Priobium, 292
Serres, M. de, 199,213
Service, M.W., (see Hinton, H.E.), 208, 336, 337
Sessions, J., 27, 28, 29, 62, 73
Setipalpia, 6
setipalpian Plecoptera, 74
Seguy, E., 153,213
Shannon, R.C., 126, 127, 207
Shelford, V.E., 286, 294
Shemanchuk, J.A., 126,213
Shipsa, 12, 15, 18, 38
rotunda, 3, 12, 38, 66, 67, 69, 86, 87,
106
shrubs, 8, 342
Sialis biline at a, 60
dorsata, 20
signata, Alloperla, 45
signata, Alloperla (Triznaka), 45
signata, Chloroperla, 45
signata, Triznaka, 3, 13, 45, 66, 69, 71, 88,
96, 97, 108
signatus, Scartallus, 309, 316
signifer, Thymallus, 76
Sikorowa, A., 125,126,213
silkworm, moth, 293
silvatica, Cicindela, 310
Simeone, J.B., 336, 338
Simuliidae, 48, 295, 373
Simulium, 371,373,374
aureum, 373
decorum, 360
hargreavesi, 372
jenningsi, 363, 369
parnassurn, 360
pictipes, 360,363,369,371,372
tuberosum, 360, 363, 369, 372
underhilli, 373
venustum, 360
verecundum, 360
vittatum, 360, 373
Siphloneuridae, 75
Skierska, B., 126, 213
Skwala, 13, 17, 19, 53
parallela, 3, 13, 54, 66, 70, 81, 82, 90,
111
Sladeckova, A., 369, 373
Smith, B.C., 126, 127, 209, 213
Smith, D.H., 52, 77
Smith, L.W., 9, 20, 21, 53, 77
snails, 69, 77
Sneath, P.H.A., 194, 214
Snoddy, E.L., 372, 373
Snodgrass, R.E., 10, 77
Snyder, A.W., 310, 317
soil, 258, 358
XX
Sokal, R.R., 214, 299, 317
Solon, B., (see K.W. Stewart), 53, 78
Southwood, T.R.E., 190,214
spiders, 343, 344
spruce, 8, 336
Stahl, J.B., 199,214
Stanford, J.A., 32, 33,77
Staphylinidae, 309,316
Stark, B.P., 49, 51, 77
Steffan, A., 47, 49, 50, 51, 77
Steropus madidus, 309, 31 1
Stewart, K.W., 52, 77
Stinner, R.E., 287, 294
Stizostedion vitreum, 7, 77
Stone^ A., 360,373
stoneflies, 3, 4, 5, 6, 7, 8, 9, 10, 1 1, 14, 19, 25,
26, 42, 47, 49, 59, 54, 60, 65, 67, 69, 70, 71,
72, 73, 74, 75, 76,
stoneflies, winter, 7, 74
Straneo, S.E., 345, 354,
strobus, Firms, 363
Struble, D.L., 258, 294
Suannaksa, S., 286
subtriatus, Gyrinus, 310, 317
Surber, D.E., 265, 278, 290
Surdick, R.F., 20, 33, 36, 37, 45, 51, 72
Suwallia, 9, 13, 16, 19, 43, 44
lineosa, 3, 13, 44, 45, 66, 70, 71, 88, 97, 108
Svensson, B.J., (see M. Carlsson), 358, 373
Sweetman, H.L., 286, 293
Swiecimski, J., 300,310,317
Swift, M.C., 126, 127, 207, 214
Swilste, H.FJ., 126, 214
Syme, P.D., 358, 360, 373
Taenionema pacifica, 7
Taeniopterygidae, 5, 14, 21, 22, 33, 74, 82
Taeniopteryginae, 21
Taeniopteryx, 9, 12, 14, 17, 22, 77
rnaura, 23
nivalis, 3, 12, 22, 69, 80, 81, 83, 104
Tarter, D.C., 47, 65, 77
Tauber, C.A., 214
Tauber, M.J., (see C.A. Tauber), 203, 214
tectus, Ptinus, 286
Telea polyphemus, 286, 293
Tendipedidae, 212
Tennessen, K.J., 74
tephritid flies, 203
Teraguchi, M., 214
tergisus, Hiodon, 7, 73
terricola, Eutrichomerus, 309
tertiarius, Chaoborus, 117,206
Thaumaleidae, 373
Theobold, F.V., 127, 214
Thompson, M.J., 311,312,317
Thymallus arcticus, 1
signifer, 16
tiger beetles, 8,298,31 1,315,317
till, 8
Tipulidae, 343
Tj0nneland, A., 183, 214
Togoperla media, 46
Totani, K., 289, 294
trailing grass, 371
tranquebarica, Cicindela, 291 , 298, 299, 300,
301, 302, 305, 307, 310, 312, 321, 322, 323
323, 324, 326
transmarina, Chloroperla, 64
transm.arina, Isoperla, 3, 13, 58, 59, 60, 64,
66, 61, 69, 91, 92, 1 15
trava, Capnia, 31
trava, Capnia (Utacapnia), 31
trava, Utacapnia, 3, 12, 31, 66, 69, 70, 84,
85, 101
tree frogs, 187,211
trees, 128, 129, 185, 342
trees, deciduous, 8
tremuloides, Populus, 185
Tribolium, 293
confusum, 286
Trichoceridae, 343
Trichoplusia ni, 289
Trichoptera, 47, 52, 77
triloba, Pteronarcella, 21
trivattus, Chaoborus, 140
trivittata, Corethra, 136, 139
triv itta ta. Say o my ia, 139
trivittatus, Chaoborus, 123, 127, 128, 129,
130, 133, 134, 137, 138, 139, 140, 144,
146, 147, 154, 156, 157, 159, 160, 161,
XXI
trivattatus, Chaoborus, 162, 165, 166, 170, 173,
175, 176, 177, 178, 179, 180, 181, 183, 184,
185, 187, 188, 189, 190, 191, 192, 193, 194,
195, 199, 200, 201, 202, 207, 214, 219, 229,
222, 225, 228, 231, 232, 236, 237, 238, 240,
242, 243, 244, 245, 246, 248, 249, 250, 255
Triznaka, 9, 13, 16, 19, 43, 45
signata, 3, 13, 45, 66, 69, 71, 88, 96, 97,
108
trout, 47
trout, brook, 7
trout, cut-throat, 7
trout, rainbow, 7
tuberosum, Simulium, 360, 363, 369, 372
Tullock, G.S., 127,214
Twinn, C.R., 127, 214
Typha, 128,185,190
Ulfstrund, S., (see M. Carlsson), 358, 373
underhilli, Simulium, 373
underwoodi, Chaoborus, 127
underwoodi, Eucorethra, 196
unipuncta, Pseudaletia, 287, 292
United States Department of Agriculture, 362,
United States Geological Survey, 358, 373
uraticus, Metoliophilus, 75
Utacapnia, 9, 12, 15, 18, 25, 30
trava, 3, 12, 31, 66, 69, 70, 84, 85, 101
Utaperla, 43
Uvarov, B.P., 285, 187, 290, 294
variduzeea, 23
Varela, F.G., 199, 211, 219
Vaurie, P., 312,319
vegetational zones, 8
velutinus, Chaoborus, 127
velutinus, Mochlonyx, 187, 196, 209
venustum, Simulium, 360
Verbeke, J., 127, 215
verecundum, Simulium, 360
vernalis, Capnia, 3, 12, 26, 27, 28, 30, 66, 69,
84, 85, 93, 101
Vernberg, F.J., 143, 215
vershina, Paraleucira,, 3, 13, 42, 66, 70, 82, 86,
107
vitreium, Stizostedion, 7, 76
vittatum, Simulium, 360, 373
volaris, Melontha, 309
Wachmann, E., 310, 317
Walcott, B., (see G.A. Horridge), 309, 315,
Walker, F., 41, 46, 60, 78
walkeri, Sarnia, 293
Waller, W.T., 183, 205
walleye, yellow, 16
Wallis, J.B., 311, 317,
Walton, C.P., (see W.L. Hilsenhoff), 2, 74
water beetles, predaceous, 75
water-striders, 48
Webb, D.W., 341, 342, 344
Weber, H., 311, 317
Wehner, R., 310, 317,
Weismann, A., 215
Welch, P.S., 362, 374
Wesenberg-Lund, C., 143, 215
Westgate, J.A., 68, 69, 78
Wheeler, Q.D., 375
White, R.E., 336, 338
white birch, 342
Whitehead, D.R., 346, 354
Wichard, W., 131, 215
Wigglesworth, V.B., 215, 288, 290, 294
Wiitanen, W., 298,310,318
wild cherry, 336
wild flowers, 187
Williams, J.A., 126, 215
Williams, T.R., 358, 374
Wniis, H.L., 300, 311, 317, 318
willows, 128, 187
Winston, P.W., 261, 263, 264, 270, 278,
294
winter stoneflies, 7, 74
wirthi, Corethrella, 187
Withycombe, C.L., 341, 342, 343, 344
Wojtusiak, H., 342, 343, 344
woodlands, deciduous, 341
Woodmansee, R.A., 126,215
Wotton, R.S., 358, 372, 374
Xestobium rufovillosum, 335, 336
Xiphidiocercariae, 73
Yeatman, C.W., 69, 77
yellow walleye, 76
Young, J.O., 126, 208
xxii
Zapada, 12, 13, 15, 17, 36
cinctipes, 3, 12, 36, 66, 69, 70, 80, 81, 86,
87, 106
zea, Heliothis, 286
Zealeuctra, 40, 77
zelona, Brachyptera, 23
Zetterstedt, J.W., 127, 215
zones, grassland, 8
zones, vegetational, 8
zooplankton, 205
Zwick, P., 11, 22, 30, 31, 78
Zygoptera, 77
Enf.
Quaestiones
Entomologicae
A periodical record of entomological investigations,
published at the Department of Entomology, '
University of Alberta, Edmonton, Canado.
JANUARY 1979
VOLUME 15
NUMBER 1
QUAESTIONES ENTOMOLOGICAE
ISSN 0033-5037
A periodical record of entomological investigation published at the Department of Entomology,
University of Alberta, Edmonton, Alberta.
Volume 15 Number 1 January 1979
CONTENTS
Book Review -Traugott-Olsen, E. and E. Schmidt Neilsen. 1977 The Elachistidae (Lepidoptera) of
Fennoscandia and Denmark 1
Dosdall and Lehmkuhl - The Stoneflies (Plecoptera) of Saskatchewan 3
Corrigenda - Upper Oligocene fossil pupae and larvae of Chaoborus tertiarius (von Heyden)
(Chaoboridae, Diptera) from Western Germany 1 17
Editorial 119
BOOK REVIEW
TRAUGOTT-OLSEN, E. and E. SCHMIDT NEILSEN. 1977. The Elachistidae (Lepidoptera) of
Fennoscandia and Denmark’. Scandinavian Science Press Ltd. Klampenborg, Denmark. 299 pages.
This is a frustrating text because it contains more information than is usually presented in a general
handbook, but less than one might expect in a revision.
However, the nine introductory sections provide an excellent basis for understanding the treatment
of elachistine genera and species found in northern Europe. Structure of adults and immatures is
described and all characters used in the text are described and figured. The position of the family
Elachistidae within the Gelechioidea is discussed and the family diagnosed . A generic phylogeny is
presented, and status of questionable genera is exlained. There are brief sections on bionomics,
zoogeography, and nomenclature.
The remaining seven sections of text consist of keys to genera and species, and descriptions of taxa
known to occur in the area covered. This part of the book is well organized, as are the keys, which
contain very few ambiguous couplets. Where possible, keys for larvae are also presented. Descriptions
are detailed and uniform in presentation, with short sections covering biology, distribution, and major
synonymies. Unfortunately, the authors chose not to point out diagnostic characters in species
descriptions, so that while keys are sufficient for identification purposes, differences between closely
related species are not readly appreciated from the descriptions, and even less so in the keys. This could
be confusing if the descriptions are to be used in conjunction with the keys, and it is certainly confusing
to the reader.
The figures are generally clear and sharp, although the presentation of larval mining patterns is less
than satisfactory. These are presented as half-tone pencil drawings which makes details in many of them
difficult to perceive. Simple line drawings would have presented these patterns much more clearly.
Scales are not given on the plates, but Figures 1 through 152 are stated to be approximately 5.5 times
2
magnification. Figures 153 through 536 are without indication of magnification, as are all text figures.
Distributional data are not well presented, and are dealt with only superficially. There are no
distribution maps, although, with considerable effort, they could be constructed from data presented.
This is an area where treatment is weak and where the synthesis falls short of that which might be
expected in a revision of the group. In defense of the authors, a catalogue arranged by species and
province is presented in which records are shown by a full or half dot. However, no indication is given
of the significance of the two symbols used. A half dot appears to be used where two provinces are
combined in the catalogue and appear to represent presence in one, but not in the other; there is a 50
percent chance of guessing correctly. No precise locality data are presented, and type localities are not
mentioned. Biogeography is treated very superficially.
Although a generic phytogeny is presented, species relationships are not discussed beyond broad
species groups in the largest genus. Little attempt is made to discuss trends in structure or bionomics.
The text is generally free of errors.
If you are interested in Fennoscandian Elachistidae, this is a reasonably good text; if you are not,
there is little here for you. If this review seems harsh, it is simply because the authors have missed a
number of opportunities to synthesize their data more completely, and thus make the work more
valuable to those outside a rather restricted field of interest.
E.M. Pike
Department of Entomology
University of Alberta
Edmonton, Alberta, Canada
T6G 2E3
STONEFLIES (PLECOPTERA) OF SASKATCHEWAN
LLOYD M. DOSDALV
Biology Department
University of Saskatchewan
Saskatoon, Saskatchewan
S7N0W0
Quaestiones Entomologicae
15:3-116 1979
D.M. LEHMKUHL
Biology Department
University of Saskatchewan
Saskatoon, Saskatchewan
S7N0W0
Forty-one species, twenty-nine genera and eight families of Plecoptera are recorded from
Saskatchewan. Distinguishing characters are given and keys are provided. Species recorded are:
Pteronarcys dorsata (Say), Pteronarcella badia (Hagen), Taeniopteryx nivalis (Fitch), Oemopteryx
fosketti (Ricker), Capnia coloradensis Claassen, Capnia confusa Claassen, Capnia gracilaria Claassen,
Capnia vernalis Newport, Paracapnia angulata Hanson, Isocapnia crinita (Needham and Claassen),
Isocapnia missourii Ricker, Utacapnia trava (Nebeker and Gaufin), Nemoura rickeri Jewett, Shipsa
rotunda (Claassen), Amphinemura linda (Ricker), Zapada cinctipes (Banks), Malenka californica
(Claassen), Podmosta delicatula (Claassen, Paraleuctra vershina Gaufin and Ricker, Leuctra ferruginea
(Walker), Acroneuria abnormis (Newman), Acroneuria lycorias (Newman), Hesperoperla pacifica
(Banks), Claassenia sabulosa (Banks), Paragnetina media (Walker), Perlesta placida (Hagen), Isoperla
bilineata (Say), Isoperla longiseta Banks, Isoperla transmarina (Newman), Isoperla patricia Frison,
Isoperla decolorata (Walker), Isoperla marlynia Needham and Claassen, Isoperla petersoni Needham
and Christensen, Arcynopteryx compacta (Mac Lachlan), Skwala parallela (Frison), Isogenoides
colubrinus (Hagen), Isogenoides frontalis (Newman), Diura bicaudata (Linnaeus), Triznaka signata
(Banks), Suwallia lineosa (Banks) and Hastaperla brevis (Banks). Nymphs of the following ten species
are described for the first time: Oemopteryx fosketti, Triznaka signata, Suwallia lineosa, Isoperla
decolorata, Nemoura rickeri, Malenka californica, Podmosta delicatula, Capnia coloradensis, Capnia
confusa, and Capnia gracilaria. Keys to species are provided for mature nymphs except that nymphs of
Isocapnia crinita and Isocapnia missourii are unknown, and nymphs o/Malenka californica and
Amphinemura linda are inseparable, as are nymphs of Capnia coloradensis from Capnia confusa and
Isogenoides frontalis/rom Isogenoides colubrinus.
The following information is presented for each species: selected literature references, diagnostic
information, bionomics, and distribution which consists of a brief summary of the species range and a
map showing Saskatchewan collection localities. Diagnostic illustrations are provided.
Life history patterns and seasonal succession of Saskatchewan stoneflies are discussed. Life cycles
range from more than one year to univoltine. Some species with a one-year life history diapause as
embryos, others do not. The post-glacial dispersal of stoneflies to Saskatchewan is discussed.
'Present address: P.O. Box 7415, Saskatoon, Saskatchewan, S7N 4J3
4
Dosdall& Lehmkuhl
Integration of the geographical and ecological distributions of extant species with the post-glacial
history of Saskatchewan leads to the inference that the Saskatchewan stonefly fauna is derived mainly
from post-glacial dispersal from refugia to the south and northwest of the Wisconsin ice sheets.
Quarante et-une especes de Plecopteres se rencontrent en Saskatchewan. Ces esp^ces se distribuent
en huit families et 29 genres. L’adulte de chaque especes est caracterise, et nous pourvoyons une clef
permettant leur identification. Les especes de cette region sont: Pteronarcys dorsata (Say),
Pteronarcella badia (Hagen), Taeniopteryx nivalis (Fitch), Oemopteryx fosketti (Ricker), Capnia
coloradensis (Claassen), Capnia confusa (Claassen), Capnia gracilaria Claassen, Capnia vernalis
Newport, Paracapnia angulata Hanson, Isocapnia crinita (Needham et Claassen), Isocapnia missourii
Ricker, Utacapnia trava (Nebeker et Gaujin), Nemoura rickeri Jewett, Shipsa rotunda (Claassen),
Amphinemura linda (Ricker), Zapada cinctipes (Banks), Malenka cdi\\iovn\c2i(Claassen), Podmosta
delicatula (Claassen), Paraleuctra vershina Gaufin et Ricker, Leuctra ferruginea (Walker), Acroneuria
abnormis (Newman), Acroneuria lycorias (Newman), Hesperoperla pacifica (Banks), Claassenia
sabulosa (Banks), Paragnetina media (Walker), Perlesta placida (Hagen), Isoperla bilineata (Say),
Isoperla longiseta Banks, Isoperla transmarina (Newman), Isoperla patricia Prison, Isoperla decolorata
(Walker), Isoperla marylnia Needham et Claassen, Isoperla petersoni Needham et Christensen,
Arcynopteryx compacta (Mac Lachlan), Skwala parallela (Prison), Isogenoides colubrinus (Hagen),
Isogenoides frontalis (Newman), Diura bicaudata (Linnaeus), Triznaka signata (Banks), Suwallia
lineosa (Banks) et Hastaperla brevis (Banks). Nous decrivons pour la premise fois les stades nymph
aux des especes suivantes: O. fosketti, T. signata, S. lineosa, I, decolorata, N. rickeri, M. californica, P.
delicatula, C. coloradensis, C. confusa et C. gracilaria. Nous pourvoyons une clef des especes pour les
nymphes au dernier stade, excepte pour cells d'\. crinita et d’l. missourii qui ne nous sont pas connues.
De plus les nymphes des especes paires suivantes ne peuvent etres separees: le M. californica, d’N. linda,
le C. coloradensis de C. confusa, et /’I. frontalis d’ I. colubrinus.
Pour chaque especes nous presentons les donnees suivantes: une serie de references choisies, un
diagnostic des stades, des notes sur la bionomique, et une distribution d^crite brievement dans le texte
et illustree en detail sur une carte de la Saskatchewan. Nous avons illustre tous characteres structuraux
dhmportance.
Nous discutons pour les Plecopteres de la Saskatchewans les patrons d’histoire naturelle et de
succession saisonniere. Le cycle vital s’etend sur une annee (univoltine) our plus (multivoltine).
Quelques unes des especes univoltines ont une periode de diapause au stade d'embryo, les autres especes
ne I’ont pas. Nous discutons la dispersion post-glaciale des Plecopteres de cette region. L’integration des
donnees ecologiques et geograhiques des especes presentes avec I’histoire post-glaciale de la
Saskatchewan suggere que la faune des Plecopteres s'est dispersee apres les temps glacaux a partir de
refuges au sud au nord-ouest des glaciers partir de refuges au sud au nord-ouest des glaciers de la
periode du Wisconsin.
Stoneflies of Saskatchewan
5
TABLE OF CONTENTS
Introduction 5
Materials and Methods 8
Table I 12
Systematics 14
Pteronarcidae 19
Taeniopterygidae 21
Capniidae 25
Nemouridae 33
Leuctridae 40
Chloroperlidae 43
Perlidae 46
Perlodidae 53
Discussion 65
Summary and Conclusions 7 1
Acknowledgements 71
References 72
INTRODUCTION
Introductory background and objectives of the study
Plecoptera, or stoneflies, are a small order of aquatic insects with approximately 2,000 species
known in the world, and 470 species recognized from America north of Mexico. Stoneflies comprise a
significant part of many lotic ecosystems, and to a lesser extent some lentic ecosystems. The nymphal
stage forms an important link in the aquatic food web between the algal-detrital and vertebrate trophic
levels. Also, nymphs have potential importance as indicators of water quality. Specific identification as
well as knowledge of the distribution and way of life of stoneflies is a preliminary requirement to their
use as indicators in assessment of water quality and to understanding dynamics of aquatic communities
of which they are a component.
Studies of Plecoptera in Western Canada have largely excluded Saskatchewan. Banks (1907)
reported species known from Alberta and British Columbia and Neave (1929, 1933, 1934) published
records for several species from Alberta, British Columbia and Manitoba. Needham and Claassen
(1925) listed one species from Saskatchewan. Ricker (1943) published a thorough study of the stonefly
fauna of southwestern British Columbia and later (1944, 1946, 1965), published records of prairie and
northern stoneflies including nine species from Saskatchewan. Cushing (1961) reported two species,
from Montreal River, not previously known in the province. Records for these 1 1 species were from
fewer than 10 Saskatchewan localities. It was evident, with the tremendous diversity and number of
habitats represented in Saskatchewan, that a varied and extensive fauna should occur in the province.
Lotic systems are well represented, especially in the boreal part of the province and, to a lesser extent, in
the prairies and Cypress Hills regions.
Quaest.Ent., 1979, 15(1)
6
Dosdall& Lehmkuhl
This study of Saskatchewan Plecoptera had three main objectives. The first objective was to
document nymphal as well as adult stages. Although adults of North American stoneflies are well
known, less attention has been given to taxonomic characters of the nymphs. For many species, only
mature nymphs with underlying adult characters can be definitely recognized at present, though the
nymph is the stage of life cycle most commonly encountered in the field. Keys presented in this study,
especially for immature stages of Plecoptera, will be useful for future ecological studies.
The second objective was to infer possible sources for the Saskatchewan stonefly fauna by
examining their known North American distributions. Until this study, little could be stated of the
post-glacial stonefly dispersal to Saskatchewan. Also, distributional information recorded here will be
useful in assessing the effects of man's future alterations of Saskatchewan's rivers and streams.
The third objective was to summarize published information on life-cycles and to synthesize this
with observations made on the way of life of Saskatchewan stoneflies.
A total of 41 species are known from Saskatchewan, including 30 new records for the province and
four new Canadian records. New descriptions are presented for nymphs of ten species.
Morphology
The following text describes basic morphological characters of Plecoptera and is summarized from
Hitchcock (1974) Gaufinc/ al. (1972), Merritt and Cummins (1978) and others.
Plecoptera are hemimetabolous, belonging to the orthopteroid group of insects. They are most
closely related to Embioptera(Crampton, 1932, Nutting, 1951).
The eyes are generally posterior on the head, but in some species they are placed far forward. A few
species have only two lateral ocelli, the majority have three: two lateral and one median with an
imaginary or “ocellar triangle” connecting the three ocelli. The Y-shaped ecdysial line forks just
posterior to the lateral ocelli. The posterior part of the head, or occipital region, may bear short bristles
or long setae forming an occipital ridge.
The mouthparts are of the chewing type, though they are reduced in some adults. Nymphal
mandibles may have grinding surfaces or be sharply toothed, depending on feeding habits (Fig. 166).
Maxillary appendages include the 5-articled palpus, galea, and laciniae(Fig. 165). Laciniae may have
one or two sharp teeth and a row or tuft of bristles. The labium bears a 3-articled palpus and two pairs
of lobes distally, the outer paraglossae and inner glossae. Glossae and paraglossae are approximately of
equal length in Filipalpia (Fig. 21), but in Setipalpia the glossae are much reduced and in some genera
appear only as small projections on the inner side of the paraglossae (Fig. 20). There is a small mentum
proximal to the labium, and beyond it a larger submentum (Fig. 20).
The pronotum may have a median longitudinal stripe or a reticulate pattern. The basisternum is the
largest thoracic sternal sclerite and this is preceded by the small presternum and followed by the
furcastrum (Fig. 18, 19). On each side of the furcasternum are the furcal pits and in certain species they
are joined by a transverse ridge with another ridge posterior to each pit forming a Y-shaped line when
they join and run together (Fig. 24, 25).
Nymphal fore and hindwing pads may be aligned at an angle (Fig. 14), or they may be subparallel
with forewing and hindwing pads in line with each other (Fig. 15).
Adult wing venation shows little variation from the basic form. Except for three North American
species, hindwings always bear an enlarged lobe in the anal region. Forewings can have up to three anal
veins, with the second anal vein forked in some species. Generally, there is only one anal cell although
several anal cells occur in some species. The number and position of crossveins, the point of origin and
the degree of upturn of the first branch of the radial vein are important systematic characters.
Stoneflies of Saskatchewan
7
Brachypterous individuals show distorted venation.
Many nymphs have gills on various parts of the body. Submental gills are small and finger-like
arising from the posterior corners of the submentum (Fig. 20). Cervical gills on the neck may be simple
(Fig. 1 1) or branched (Fig. 12). Thoracic gills may also be simple or branched. Some species have
simple coxal gills occurring singly on the ventral surface of each coxa (Fig. 13). Abdominal gills occur
laterally on basal abdominal segments (Fig. 6, 7), and subanal gills occur between the cerci. Except for
coxal gills, larval gill remnants are visible on the adult.
Males bear various genitalic processes, used in mating, and which are important taxonomically.
These projections range from small, hardly perceptible processes, to very elongate projections. The
male genitalia generally consist of a median epiproct or supra-anal process (an unpaired structure
attached to the tenth tergum) which can be simple (Fig. 93), complex (Fig. 50, 51), or not readily visible;
there are sclerotized paragenital plates at its base. In some species additional processes occur at the base
of the epiproct. Paraprocts are located on each side of the epiproct and can range in structure from
being simple and fingerlike (Fig. 117, 1 18) to complex structures (Fig. 120). The seventh, eighth or ninth
sterna have vesicles or basal lobes in some species (Fig. 62, 64) and the ninth sternite may have a heavy
knob or “hammer” (Fig. 9). Recurved pointed hooks or “genital hooks” can emerge as paraprocts
(Fig. 102) or as part of the tenth tergite (Fig. 8).
The relatively simple female abdomen has the seventh or eighth sternum produced to form a
subgenital plate (Fig. 1 16).
Single-, or many-articled cerci arise from the tip of the abdomen, and may bear characteristic
bristles or hairs.
Economic importance of Plecoptera
Stoneflies are economicaly important mainly as natural fish food. A few fish species known to eat
stoneflies include grayling ( arcticus), (Rawson, 1950), lake sturgeon {Acipenser fulvescens),
(Magnin and Harper, 1970), yellow walleye {Stizostedion vitreum), (Rawson, 1956) and mooneye
{Hiodon tergisus) (Glenn, 1975). In Saskatchewan, adults of Malenka calif ornica were found in
stomachs of brook trout {Salvenius frontinalis), (D. Larson, pers. comm.).
Stoneflies also provide a significant food source for some birds. Hamilton (1933) noted that the
winter stonefly Allocapnia recta constituted an important food source for five species of passerine
birds. In Saskatchewan, Pteronarcys dorsata nymphs have been found in crops of the Common
Merganser (H. Stelfox, pers. comm.).
Generally stoneflies are restricted to well-oxygenated, unaltered streams and so have been used as
indicators of water purity. Roback (1974) examined 23 species of stoneflies and found that none
occurred in dissolved oxygen concentrations of less than four parts per million (ppm.), alkalinity
greater than 1000 ppm., or chloride greater than 1000 ppm. Gaufin (1958) found that fewer stonefly
species occurred in the zone of degradation than in the zone of clean water in a polluted stream, and
none were found in extremely polluted water.
Taenionema pacifica is a minor pest in orchards in the Pacific Northwest. It has been reported
feeding on foliage, buds and fruit of apricots, peaches and plums (Newcomer , 1918, 1933).
Claire and Phillips (1968) observed nymphs of Hesperoperla pacifica attacking and killing eggs and
alevins of rainbow trout {Salmo gairdneri) and cut-throat trout {Salmo clarki). However, Nicola (1968)
found that scavenging by stonefly nymphs {Alloperla sp.) on dead salmon eggs was beneficial since it
prevented the spread of fungal infestation to living eggs.
Macy and Bell (1968) found that Hesperoperla pacifica was an alternate intermediate host to an
internal parasite of birds. Hall and Groves (1963) noted that cercariae of seven of nine trematode
Quaest.Ent., 1979, 15(1)
8
Dosdall& Lehmkuhl
species harboured in river snails entered nymphs of Hesperoperla pacifica.
The study area
The limits of the study area were the political boundaries of Saskatchewan. Battle Creek in Cypress
Hills was most easily accessible for collecting near Reesor Lake, Alberta, about one-half mile west of
the Saskatchewan border. Battle Creek flows east into Saskatchewan, and since stoneflies collected here
undoubtedly occurred in Saskachewan also, species collected near Reesor Lake were considered part of
the Saskatchewan fauna.
Saskatchewan encompasses a wide range of vegetational, climatic and geological zones. Two of the
major North American vegetational zones are represented in the province: Grassland in the semi-arid
climate of the southwest, and Boreal Forest in the subhumid north (Coupland and Rowe, 1969).
The southern Grassland zone is mainly higher than 600 meters above sea level and composed of
extensive till and lacustrine plain overlaying Cretaceous rock (Richards, 1969). The major drainage
basin in the Grassland region is the extensive Saskatchewan River System composed of the North and
South Saskachewan Rivers which unite near the edge of the Boreal Forest zone to form the
Saskatchewan River. The flow is to the northeast, eventually into Hudson Bay. The Saskatchewan
River System is fed both by prairie surface runoff and by snowmelt near its source in the Rocky
Mountains resulting in a definite water flow regime (Raby and Richards, 1969). Relatively low winter
flows are followed by a rise in early spring when there is snowmelt in the prairies and foothills. After a
spring peak, flow tends to decrease but rises again in May-June because of meltwater runoff from the
Rocky Mountains. Declining flow leads to low flows in August and September with an even base-flow
maintained in winter months.
In the southwestern corner of Saskatchewan, rolling plains gradually rise to the Cypress Hills
uplands which reach a maximum elevation of 1393 m, the highest in the province. The highest parts of
Cypress Hills are believed to have been unglaciated (Richards, 1969). Battle Creek, which flows through
some of the highest Cypress Hills elevations, and Frenchman River which flows down the eastern slope
of Cypress Hills, are the two prominent rivers of this area.
The Boreal Forest zone, mostly between 365 and 550 m above sea level, is an area of lakes, bogs,
coniferous and mixedwood forest and rock outcrop. In the far north, east of Lake Athabasca, is
subarctic lichen-woodland of open coniferous forest and reindeer moss. Extending southward to the
Churchill River System is the northern coniferous forest of spruce and pine underlain mainly by
Precambrian rock. To the south of the Canadian Shield, soils deepen over sedimentary rock and
spruce, pine and poplar are abundant. The southern edge of the Boreal Forest is the aspen belt,
transitional between Boreal Forest and Grassland. It is composed mainly of deciduous trees and a
variety of shrubs and grasses on rich grey-black soils (Coupland and Rowe, 1969).
On the Shield much of the drainage flows southward to the Churchill River System or to Lake
Athabasca in the northwest. Northeastern drainage is primarily to Reindeer Lake and Wollaston Lake.
In the Boreal Forest zone, practically any depression in the bedrock is occupied by a lake, bog, or
marsh. Streams and rivers are numerous and because of climate and other factors, water levels fluctuate
far less than in the south.
MATERIALS AND METHODS
Collection sites
Many areas of the province were collected extensively. These included Cypress Hills, the
Saskatchewan River System and much of the boreal forest region. Collecting trips to rivers and creeks
Stoneflies of Saskatchewan
9
in southcentral and southeastern Saskatchewan produced no specimens so this area was sampled only
superficially. In addition, access roads are poor or nonexistent in the northeastern (north of Southend,
Sask.) and northwestern (north of Buffalo Narrows, Sask.) parts of the province, so collections are
meagre from these areas. Extensive collecting in this far northern region may lead to new provincial
records and certainly will modify some of the distributions presented here.
Collection and preservation of specimens
Stonefly nymphs were most commonly collected with a sweep net where one stirs up the river
substrate with the feet while walking backward against the current. Stonefly nymphs which had been
dislodged from the rocks were carried by the current into the net. The net content was examined for
stoneflies by placing it in a white pan with water in the bottom.
Another collection method was simply to examine such submerged objects as rocks or logs by
hand-picking and to remove any stoneflies present. Mature final-instar nymphs were collected from
rocks close to shore where they had crawled to emerge. The latter method was ideal for streams too
small to use the sweep net in, but was less productive than sweeping in larger, more swiftly flowing
rivers. Cast skins were found under bridges, on rocks near streams, or on vegetation which borders
streams. Nymphs and cast skins were preserved in 70-90 percent alcohol.
In order to rear nymphs for associations with known adults, they were transported to the laboratory
in glass bottles or Nalgene plastic bottles containing river water. Since the survival rate of Plecoptera
was higher when kept cool, these containers were placed in an ice chest during the period of transit.
Periodic oxygenation with a battery operated airpump or replacement of the water in the container
with fresh water was necessary on long trips.
Adults were collected by rearing mature nymphs in laboratory aquaria, by sweeping vegetation near
streams with an aerial net or by examining bridges, posts, or tree trunks near flowing water. Some
adults hide under rocks near the shore and were collected by turning over the rocks. Winter stoneflies
{Capnia, Oemopteryx, Taeniopteryx, Utacapnia) were collected in large numbers on the snow and ice
of river banks, or on bridges. Adults of Paracapnia were collected by throwing panfuls of water onto
the riverbank. In a few minutes the adults were flooded out of cavities in the ground and were collected
with forceps. Although a few species are known to come to light traps, such equipment was not used in
this study. Since stoneflies are weak fliers, finding adults was often more difficult than capturing them.
Adults were preserved in 70-90 percent alcohol. Specimens of certain species changed color in
alcohol. Adults of Hastaperla and Suwallia which were bright yellow in life, and Triznaka which were
green, faded to white when preserved.
Rearing
Rearing nymphs to adults was necessary in order to associate the immature and adult stages of the
life cycle. Two methods were used: the first was similar to that described by Smith (1975). An aquarium
with dimensions of 1 .7 x 0.3 x 0.3 m was filled with cold tap water and dechlorinated using two drops
per gallon of a sodium thiosulfate solution (700 grams sodium thiosulfate per liter of water). The water
was vigorously aerated, and cooled somewhat by running tap water through glass tubing in the bottom
of the aquarium.
Plastic sheets of dimensions 29 cm x 45 cm were suspended from the edge of the aquarium just
above the surface of the water. These sheets contained several holes, into which the rearing containers
were fitted (Fig. 16). Rearing containers were made from sections of Nitex screening, with a 0.33 mm
mesh size glued together to form a tube. A circular piece of screening glued to the lower end of the Nitex
tube formed the bottom and a piece of plastic tubing covered with Nitex screening formed the lid
Quaest.Ent., 1979, 15(1)
10
Dosdall & Lehmkuhl
(Fig. 17). The rearing container was then fitted through a hole in the plastic sheet but the upper portion
rested on top of the sheet. A mature nymph placed in the container was able to crawl up the screening
and out of the water when emerging. The lid prevented escape of the adult. In instances where it was
necessary to rear nymphs which were not mature, the appropriate food could be placed in the container
with the nymph. Detrivorous species were given dead leaves which had soaked for months in water;
carnivorous species were supplied with larval mayflies, caddisflies or other stoneflies.
Rearing of stonefly nymphs which emerge in early spring required temperatures in aquaria of about
1 C to 5 C, much colder than were possible in the apparatus just described, which was kept in a
laboratory at room temperature. The second rearing method used was similar to the one described by
Sawchyn (1971). Containers with nymphs in them were kept in a Sherer-Gillet controlled-environment
cabinet with temperature and photoperiod controls. Temperatures could be maintained within ±1 C of
the desired level and the fluorescent lights were controlled by timers. The rearing chambers each
consisted of an outside plexiglass vessel measuring 28 x 18 x 10 cm, each holding 15 containers. The
cylindrical containers had a frame of plexiglass and were covered with a plastic screening of 8 meshes
per cm. The cylinders were suspended in the vessel by wires so that there was sufficient room for mature
nymphs to crawl out of the water to emerge. Plastic petri dishes (^1007), placed over the chamber tops,
prevented escape of adults. Dechlorinated tap water was used, and aeration was provided by a Model
120 Silent Giant pump.
Clearing of genitalia
It was unnecessary to clear genitalia of Plecoptera in order to observe taxonomic details.
Abdominal sclerites of some specimens were contracted when preserved in alcohol, but after placing the
abdomen in KOH at room temperature for 5-6 hours, the contracted sclerites were gently pulled apart.
This enabled more efficient examination of the genitalia.
Material examined and its disposition
Material examined in this study was based primarily on collections made during the spring and
summer of 1974 and 1975 with a few specimens collected in 1976.
In addition, I have examined Saskatchewan material from the personal collections of Drs.
D.M. Lehmkuhl and D.J. Larson, Mr. D.H. Smith, and Mr. Ron Demaray. I have also examined some
material collected from Stoney Rapids and Black Lake by Mr. L. Kratt and from the Churchill River
and Cluff Creek in the collections of the Saskatchewan Research Council.
I was unable to collect either males or females of certain species, but I was usually able to borrow
missing specimens from the personal collection of W.E. Ricker for drawings.
Small representative collections of Saskatchewan Nemouridae and Perlidae were sent to R.W.
Baumann (Brigham Young University, Provo, Utah) and to B.P. Stark (Mississippi College, Clinton,
Miss.), respectively. The Churchill River and Cluff Creek collections remain with the Saskatchewan
Research Council. Most of the remaining specimens from other sources have been returned to their
collectors. The remainder of the material is in my collection.
Illustrations
Line drawings, prepared using the camera lucida, and photographs using the Zeiss Tessovar
photographic unit, are presented to illustrate taxonomic keys and augment descriptions. Illustrations of
homologous structures in related species are from the same aspect.
Stoneflies of Saskatchewan
11
Terms
Most terms for insect structures are based on the usage of Snodgrass ( 1935). Those specific to
Plecoptera are from Needham and Claassen( 1925), Prison (1935, 1942), Ricker (1943, 1949, 1952, 1959,
1965), Hanson (1946) and Ricker and Ross (1969). The nomenclature for wing venation follows the
system proposed by Comstock and Needham (1898-1899). The section on Morphology (page six)
provides a discussion of some of the taxonomically useful characters and includes some explanation of
the most important terms.
Verification of species identifications
With the exceptions of Perlesta placida (Hagen) and Isoperla marlynia Needham and Claassen, all
species described in this study have been verified. However, since these two species are particularly
distinctive, there is little doubt that my identifications are correct. R.W. Baumann of Brigham Young
University, Provo, Utah verified the Nemouridae, and W.E. Ricker of the Fisheries Research Board of
Canada, Nanaimo, B.C. confirmed the remainder of the species identifications.
Organization of family, genus and species accounts
The systematic arrangement used for all taxa is that of lilies (1966) and Zwick (1973). A systematic
list of Saskatchewan stoneflies is presented in Table 1. Descriptions of each family and genus
represented in Saskatchewan are presented after the keys. Information for these descriptions was
obtained from Needham and Claassen (1925), Claassen (1931), Prison (1935, 1942a), Ricker (1943,
1952), Gaufin et al. ( 1972) and Hitchcock (1974) as well as from my collections.
For genera with more than one species known from Saskatchewan, a key to adults is provided, and
keys to nymphs are given where possible.
Each species heading is followed by a list of references pertaining to the species. No attempt was
made to include complete synonomies but the references indicate the various names proposed for each
species and may include important taxonomic studies, faunal works, catalogue listings and biological
studies.
The North American range of species is outlined in very general terms from published distributional
records. Saskatchewan locality records for each species are listed and are plotted on a map.
Measurements are presented at the beginning of species descriptions; length is measured from the
most anterior portion of the head to the tip of the folded wings and for brachypterous specimens is from
the anterior margin of the head the end of the abdomen.
The species description is a diagnostic statement presenting the principal characters which separate
males, females, and nymphs of the species from related forms, even though the related types may not
occur in Saskatchewan.
Illustrations augment keys and descriptions. Drawings of male and female genitalia are given for
nearly every species. For species where representatives were not collected in this study, and were not
borrowed from W.E. Ricker for examination, drawings are not presented.
Quaest.Ent., 1979, 15(1)
12
Dosdall& Lehmkuhl
Table 1 . Systematic List of the Stoneflies of Saskatchewan.
ORDER PLECOPTERA
Family Pteronarcidae
Genus Pteronarcys Newman 1838
Pteronarcys dorsata (Say) 1823
Genus Pteronacella Banks 1900
Pteronarcella badia (Hagen) 1873
Family Taeniopterygidae
Genus Taeniopteryx Pictet 1841
Taeniopteryx nivalis (Fitch) 1847
Genus Oemopteryx Klapalek 1902
Oemopteryxfosketti{Kic\iQv) 1965
Family Capniidae
Genus Capnia Pictet 1841
Capnia confusa Claassen 1936
Capnia coloradensis Claassen 1937
Capnia gracilaria C\aa^?,Qn 1924
Capnia vernalis Newport 1848
Genus Utacapnia Nebeker and Gaufin 1971
Utacapnia trava (Nebeker and Gaufin) 1965
Genus Isocapnia Banks 1938
Isocapnia crinita (Needham and Claassen) 1925
Isocapnia missourii Ricker 1959
Genus Paracapnia Hanson 1946
Par acapnia angulata Hanson
Family Nemouridae
Genus Nemoura Latreille 1976
Nemoura rickeri Jewett 1971
Genus Podmosta Ricker 1952
Podmosta delicatula (Claassen) 1923
Genus Zapada Ricker 1952
Zapada cinctipes (Banks) 1897
Genus Amphinemura Ris 1902
Amphinemura linda (Ricker) 1952
Genus Shipsa Ricker 1952
Shipsa rotunda {CXaa^^tn) 1923
Genus Malenka Ricker 1952
Malenka calif ornica (Claassen) 1923
Family Leuctridae
Genus Leuctra Stephens 1835
Stoneflies of Saskatchewan
13
Leuctra ferruginea (Walker) 1852
Genus Paraleuctra Hanson 1941
Paraleuctra vershina Gaufin and Ricker 1975
Family Chloropterlidae
Genus Hastaperla Ricker 1935
Hastaperla brevis (Banks) 1895
Genus Suwallia Ricker 1943
Suwallia lineosa (Banks) 1918
Genus Triznaka Ricker 1952
Triznaka signata (Banks) 1895
Family Perlidae
Genus Paragnetina Klapalek 1907
Paragnetina (Walker) 1852
Genus Claassenia Wu 1934
Claassenia sabulosa {Banks) 1900
Genus Acroneuria Pictet 1841
Acroneuria lycorias (Newman) 1839
Acroneuria abnormis (Newman) 1838
Genus Hesperoperla Banks 1938
Hesperoperla pacifica (Banks) 1900
Genus Per lest a Banks 1906
Perlesta placida {HagQn) 1861
Family Perlodidae
G^nns ArcynopteryxYAapaltk 1904
Arcynopteryx compacta (MacLachlan) 1872
Genus Skwala Ricker 1943
Skwala parallela (Frison) 1936
Genus Diura Billberg 1820
Diura bicaudata {VmnaQUs) 1758
Genus Isogenoides Klapalek 1758
Isogenoides colubrinus (Hagen) 1874
Isogenoides frontalis (Newman) 1838
Genus Isoperla Banks 1906
Isoperla bilineata (Say) 1823
Isoperla decolorata (Walker) 1852
Isoperla longiseta Banks 1906
Isoperla marlynia Needham and Claassen 1925
Isoperla Patricia Frison 1942
Isoperla peter soni Needham and Christensen 1927
Isoperla transmarina (Newman) 1838
Quaest.Ent., 1979, 15(1)
14
Dosdall & Lehmkuhl
SYSTEMATICS
The following keys to families and genera apply only to stoneflies known from Saskatchewan, and
are modified from Needham and Claassen (1925), Harper and Hynes (1971b, 197 Id), Gaufin et al.
(1972) and Hitchcock (1974). Keys for nymphs will not separate the very early instars.
Pteronarcella, p.
.. Pteronarcys,p.
Key to families and genera of adult Plecoptera known from Saskatchewan
1 Paraglossae and glossae subequal in length (Fig. 21)
r Paraglossae much longer than glossae (Fig. 20)
2 ( 1 ) Remnants of branched gills on abdominal segments 1 and 2 (Fig. 6, 7); anal area of
forewing with two or more rows of crossveins
Pteronarcidae
2' No gill remnants on abdominal segments 1 and 2; anal area of forewing without
crossveins or with only one row of them (Fig. 33, 34, 36, 37, 39)
3 ( 2 ) Gill remnants on abdominal segment 3 (Fig. 7)
3' Gill remnants absent from abdominal segment 3 (Fig. 6)
4 ( 2') Tarsal article 2 much shorter than article 1 in lateral view (Fig. 23)
4' Article 2 at least as long as article 1 in lateral view (Fig. 22)
Taeniopterygidae
5 (4') Each coxa with small, round membranous areas on its ventral surface; male cercus
with one article (Fig. 40, 43); female sternum 9 without long projection extended across
sternum 10 (Fig. 44) Taeniopteryx, p.
5' Coxae without membranous areas on ventral surfaces; male cercus with at least 3
articles (Fig. 42); female sternum 9 with long projection extended across sternum 10
(Fig. 45) Oemopteryx, p.
6 ( 4 ) Cercus of more than one article (Fig. 50-72); A2 of forewing unbranched(Fig. 37)
Capniidae
6' Cercus of only one article (Fig. 73-92); A2 of forewing branched(Fig. 34, 36)
7 (6') Wings flat at rest; last article of labial palpus in ventral view subcircular, larger than
subterminal article (Fig. 29) Nemouridae
7' Wings rolled around body at rest; last article of labial palpus in ventral view longer
than wide, equal in length to subterminal article (Fig. 21)
Leuctridae
8 (7') m-cu of hindwing connected to Cui before it divides (Fig. 36); male cercus normal,
without point or projections (Fig. 84) Leuctra, p.
8' m-cu crossvein of hindwing connected to Cui after it divides (Fig. 34); male cercus with
sharp projections or pointed (Fig. 8 1 , 83) Paraleuctra, p.
9 ( 7 ) Males: Supra-anal process present (Fig. 73-77, 79); flaplike ventral lobe arising from
base of 8th abdominal sternum (Fig. 78, 80)
9' Females: no supra-anal process; no ventral lobe; 8th abdominal sternum modified as
subgenital plate (Fig. 85-90)
10 ( 9') Gill remnants under neck or head (Fig. 10)
10' Gill remnants absent..
11 ( 10 ) Gills of most specimens of five branches Zapada, p.
1 1 ' Gills with six or more branches
2
23
3
4
21
19
6
5
22
23
20
7
9
8
41
42
15
10
11
13
36
12
Stoneflies of Saskatchewan
15
12 (in
12'
13 (10')
13'
21 (20)
21'
22 (21')
22'
23 (!')
23'
24 (23)
24'
7th abdominal sternum with large, sclerotized posterior projection which extends onto
8th sternum (Fig. 88) Amphinemura, p.
7th abdominal sternum with small pointed posterior projection not extended onto 8th
sternum (Fig. 85) Malenka, p.
Sternum 7 sclerotized, produced over full length of 8, its hind margin straight to
broadly rounded (Fig. 89) Nemoura, p.
Sternum 7 only slightly or not at all produced over 8; though the narrowly rounded
margin of the subgenital plate of the sternum 8 in Shipsa may be mistaken for
sternum 7
Sternum 8 with subgenital plate terminated anterior to its well-developed hind margin
(Fig. 87) Shipsa, p.
Sternum 8 with narrow median sclerotized band contrasted sharply with unsclerotized
field at either side (Fig. 90) Podmosta, p.
Gills present, cervical (Fig. 10)
Gills absent
Gills of most specimens with five branches Zapada, p.
Gills with six or more branches
Cerci with mesobasal lobe (Fig. 73) Malenka, p.
Cerci without mesobasal lobe (Fig. 77) Amphinemura, p.
Cerci elongate, heavily sclerotized distally, inner surface membranous, tip sharp or
with two or more processes (Fig. 79, 80) Nemoura, p.
Cerci membranous or weakly sclerotized, tip blunt, without spines or processes
(Fig. 75,76)
Sides of 10th tergum produced into erect in-curved spiny processes (Fig. 15) ..Shipsa, p.
Sides of 10th tergum not as above (Fig. 76) Podmosta, p.
Male supra-anal process simple, comprised of one element; female subgenital plate
without notch or conspicuous pattern
Male supra-anal process of two distinct elements, one dorsal one ventral (Fig. 50, 51);
female subgenital plate slightly notched, sternum 8 with conspicuous pattern (Fig. 69) ..
Utacapnia, p.
Ri of forewing strongly bent upward at origin, Ai bent abruptly caudad at junction of
cu-a, then curved laterad (Fig. 37) Capnia, p.
Ri and/or Ai of forewing straight (Fig. 33,39)
Prothoracic and mesothoracic presterna broadly united with basisterna (Fig. 18);
forewing Ri straight at origin (Fig. 39); male with lobe on 9th sternum (Fig. 62)
Iso capnia, p.
Prothoracic and mesothoracic presterna separated from basisterna (Fig. 19); Ri of
forewing slightly curved upward at origin (Fig. 33); male 9th sternum without lobe
Paracapnia, p.
remains of branched gills at lower angles of thorax; cu-a of forewing in most specimens
either in anal cell or distant from it by no more than its own length
Remains of branched gills absent from thorax; cu-a, if present, generally distant from
anal cell by more than its own length (Fig. 32, 35, 38)
Males with raised knob or “hammer” on 9th sternum (Fig. 9); female subgenital plate
without deep notch (Fig. 99, 100, 104, 106)
Males without “hammer” on 9th sternum, female subgenital plate deeply notched
37
39
34
14
38
35
16
18
36
17
39
37
34
19
38
35
21
31
26
22
32
25
24
28
26
25
Quaest.Ent., 1979, 15(1)
16
Dosdall& Lehmkuhl
Distinct Y-shaped mesosternal ridge pattern (Fig. 24, 25) hind margin of male 10th
tergum deeply cleft, with dorsal projections (genital hooks) extended forward from
sides of the cleft; female subgenital plate not hairy (Fig. 105) Paragnetina, p.
Mesosternal ridge pattern not in distinct, dark Y-shape; hind margin of male 10th
tergum not cleft, no dorsal projections on 10th tergum, paraprocts joined; female
subgenital plate covered with long hairs Perlesta, p.
Male tergum 10 with hooks arising from lateral angles; paraprocts normal (Fig. 8)
Claassenia, p.
Male tergum 10 unmodified; paraprocts formed as recurved hooks (Fig. 101-103)
Subanal lobes of male very broadly triangular, terga 9, 10 not covered with spinules
(Fig. 101); female subgenital plate considerably produced (Fig. 104) ... Hesperoperla, p.
Subanal lobes of male slender hooks, terga 9 and 10 with many spinules (Fig. 102, 103);
female subgenital plate little if any produced(Fig. 99, 100) Acroneuria, p.
A2 of forewing either not forked or forked beyond anal cell (i.e., two main anal veins,
and 2nd of which is forked in some species)(Fig. 32)
Chloroperlidae
Fork of A2 of forewing included in anal cell, so that two branches leave cell separately
(i.e., there are three main anal veins)(Fig. 35, 38) Perlodidae
Anal area of hindwing apparently absent (Fig. 32) Hastaperla, p
Anal area of hind wing present
Head unmarked except for ocellar rings (Fig. 163); males with finger-like process
directed inward from basal article of each cercus (Fig. 95); female subgenital plate
emarginate (Fig. 98) Suwallia, p.
Head with conspicuous markings of black on yellow, ocellar triangle dark with
anterior mark on head as long as broad (Fig. 164); no process at base of male cerci
(Fig. 94); female subgenital plate broadly rounded (Fig. 97) Triznaka, p.
Males: paraprocts modified as recurved hooks, or considerably produced posterad, or
genitalia complicated by various stylets and sclerotized areas
Females: paraprocts unmodified; genitalia simple, 8th sternite generally produced as
genital plate
10th tergum completely cleft; genitalia complicated by stylets or various unsclerotized
structures (Fig. 110, 111) Isogeninae
10th tergum entire, at most slightly notched; genitalia simple (Fig. 107)
Wings with four-to-many crossveins beyond cord, generally arranged in irregular
network (Fig. 35); 7th sternum without lobe
Wings with no more than two crossveins beyond cord (Fig. 38); 7th sternum generally
with lobe Isogenoides , p.
Supra-anal process very long, needle-like; lateral stylets absent (Fig. 113)
Arcynopteryx, p.
Supra-anal process blunt, not unusually long; lateral stylets present (Fig. 112)
Skwala, p.
Paraprocts produced postero-mesad, meeting along their inner faces (Fig. 107); no lobe
on 8th sternum Perlodinae Diura, p.
Paraprocts forked as hooks or only slightly modified (Fig. 1 17-123); 8th sternum
always with a lobe
46
52
48
27
51
49
29
31
43
30
44
45
32
36
33
35
34
55
53
54
55
Isoperlinae
Isoperla, p.
57
Stoneflies of Saskatchewan
17
Numerous irregular crossveins between Rs and R (Fig. 35) 37
Apical crossveins few or absent 38
Subgenital plate generally produced more than halfway across ninth sternum; caudal
margin slightly emarginate (Fig. 115) Arcynopteryx, p. 53
Subgenital plate generally produced less than halfway across ninth sternum; caudal
margin straight (Fig. 1 14) Skwala, p. 54
Submental gills present (Fig. 20) Isogenoides, p. 55
Submental gills absent 39
General color dark brown with yellow median stripe on pronotum; hairs on margin of
groove in femur of proleg not noticeably longer than other hairs on femur; subgenital
plate produced halfway or more across 9th sternum (Fig. 108, 109) Diura, p. 55
Not as above Isoperla,p. 57
Key
1
1'
2
2'
2
3'
4
4'
5
5'
6
9
9'
10
10'
11
IF
12
to families and genera of Plecoptera nymphs known from Saskatchewan
Glossae and paraglossae subequal in length (Fig. 21) 2
Paraglossae extended greatly beyond glossae (Fig. 20) 16
( 1 ) Branched gill tufts on abdominal segments 1 and 2 (Fig. 6,1)
Pteronarcidae 3
Branched gill tufts absent from abdominal segments 1 and 2 4
( 2 ) Gill tufts on first two abdominal segments (Fig. 6) Pteronacys, p. 19
Gill tufts on first three abdominal segments (Fig. 7) Pteronarcella, p. 21
( 2') Tarsi in lateral view with 2nd article much shorter than 1st (Fig. 23) 6
Tarsi in lateral view with 2nd article at least as long as 1 st (Fig. 22) 5
( 4') A single retractile gill on each coxa (Fig. 13) Taeniopteryx, p. 22
No coxal gills Oemopteryx, p. 23
( 4 ) Extended hind legs exceed end of abdomen; hindwing pads divergent from axis of
body (Fig. 14); cervical gills present in some species(Fig. 11,12)
Nemouridae 7
Extended hind legs not exceeding end of abdomen; hindwing pads subparallel to axis
of body (Fig. 15); gills absent 11
( 6 ) Species with four prosternal gills (Fig. 11, 12) 8
Species without gills 9
( 7 ) Prosternal gills commonly five-branched (Fig. 1 1) Zapada, p. 36
Prosternal gills with six or more branches(Fig. 12)
Malenka, p. 39, Amphinemura, p. 37
( 7') Pronotum with well-defined lateral fringe of hairs (Fig. 146) Nemoura, p. 34
Pronotum without definite fringe of hairs; bristles on lateral margins of pronotum may
sometimes be longer than dorsal bristles, but are never set in a distinct line 10
(9') Femora with continuous fringe on long silky hairs; legs banded Shipsa, p. 38
Femora without fringe; legs not banded Podmosta, p. 35
( 6') First eight abdominal segments divided to tergum and sternum by membranous fold
(Fig. 138-140) Capniidae 13
First six or less abdominal segments divided to tergum and sternum by membranous
fold (Fig. 27) Leuctridae 12
(IF) First four abdominal segments divided laterally (Fig. 27); labial palpi extended well
beyond paraglossae (Fig. 21) Leuctra, p. 41
Quaest.Ent., 1979, 15(1)
18
Dosdall & Lehmkuhl
First six abdominal segments divided laterally; labial palpi extended approximately to
tip of paraglossae Paraleuctra, p.
Long swimming hairs along cerci (Fig. 26) Isocapnia, p.
Cerci without long swimming hairs
Body with numerous conspicous bristles; head capsule with reticulate purplish pattern .
Par acapnia, p.
Bristles inconspicuous; head capsule without elaborate pattern
Erect bristles on posterior margin of abdominal terga very long, nearly as long as
mid-dorsal length of segment Utacapnia, p.
Erect bristles shorter, half (or less) mid-dorsal length of tergum (Fig. 138-140)
Capnia, p.
Tufts of filamentous gills on thorax ..Perlidae
Thoracic gills absent
Spinules or long hairs set in row across back of head forming occipital ridge (Fig. 4, 5) .
No spinules in row across back of head except near hind margin of eye (i.e., no
occipital ridge)(Fig. 2)
Subanal gills absent (Fig. 5) Paragnetina, p.
Subanal gills present (Fig. 4)
Abdomen with numerous “freckles” Perlesta, p.
Abdomen without numerous “freckles”, but uniformly brown above (Fig. 4)
Claassenia, p.
Head with central light spot anterad of median ocellus (Fig. 3) Hesperperla, p.
Head without central light spot; with a light M-pattern in front of anterior ocellus
(Fig. 1,2) Acroneuria, p.
Hindwing pads at angle to axis of body (Fig. 168); cerci as long as, or longer than,
abdomen; body commonly patterned; particles of maxillary palpus evenly tapered
from 1st to 5th (Fig. 170) Perlodidae....
Hindwing pads subparallel to axis of body (Fig. 159); cerci three-quarters length of
abdomen; body usually uniformly brown; last article of maxillary palpus abruptly
thinner than previous articles (Fig. 162) Chloroperlidae
Inner margin of hindwing pads straight; body of mature nymph 7 mm or
less Has taper la, p.
Inner margin of hindwing pads sinuate or notched (Fig. 159); body of mature nymph
larger than 7 mm
Abdominal terga each with two light colored spots laterally, and central median stripe
(Fig. 159) Triznaka, p.
Abdomen uniformly brown above Suwallia, p.
Arms of mesosternal Y-ridge approach or meet anterior corners of furcal pits (Fig. 25) .
Arms of mesosternal Y-ridge meet posterior corners of furcal pits (Fig. 24)
Denticles numerous along both sides of outer cusps of both mandibles (Fig. 28)
Skwala, p.
Denticles absent from cusps of nymphal mandible, or a few present on outer left cusp
only Arcynopteryx, p.
Single fingerlike gills at each basal corner of submentum (Fig. 20) Isogenoides, p.
Submental gills absent
Abdominal terga dark, with two dorsolateral spots and a few small lateral spots;
42
32
14
25
15
31
26
17
21
18
20
46
19
52
48
51
49
24
22
43
23
45
44
25
26
54
53
55
27
Stoneflies of Saskatchewan
19
lacinia of maxilla with sharp angle below small tooth (Fig. 30) Diura, p. 55
27' Abdominal terga with longitudinal or transverse markings or dark dots (Fig. 168);
lacinia without sharp angle low small tooth, rounded or, more commonly, tapered
from tooth to base (Fig. 170) Isoperla, p. 57
Family Pteronacidae
Pteronacids are large, primitive stoneflies occurring in North America and eastern Asia. They are
characterized by many-articled cerci, long antennae and wings with numerous cross-veins. The adult
male has no vesicle on the ninth sternum and has a conspicuous epiproct and paraprocts on the cleft
10th tergum. Nymphs have gill tufts on the thorax and first two abdominal segments. There are two
North American genera in this family; both occur in Saskatchewan.
Genus Pteronarcys Newman
Ricker (1925) proposed two subgenera of Pteronarcys, Allonarcys and Pteronarcys sensu strictu,
which were later ranked as genera by lilies (1966). Allonarcys occurs in eastern North America and
adults are characterized by an upright epiproct, cupped paraprocts, the ninth sternum unnotched and
without peglike setae at the tip, divided hemitergal lobes on the male tenth tergum and produced female
subgenital plates. Nymphs have paired lateral projections on abdominal segments. Pternarcys adults
differ by having massive epiprocts, fleshy, rounded paraprocts, the ninth sternite notched and with
peglike setae near the tip, hemitergal lobes on the tenth tergite projected posterad (Fig. 46), and
unproduced female subgenital plates (Fig. 47, 48). Nymphs lack paired lateral projections of the
abdominal segments. Six species of Pteronarcys are known in North America and the range of one
species extends into Saskatchewan.
Pteronarcys dor sat a Say
(Fig. 6, 46-48, 180)
Stalls dorsata {Secy) 1823: 164.
Pteronarcys dorsata, Frison 1942a: 242.- Harden and Mickel 1952: 9. - Ricker 1964: 68. - Hitchcock
1974: 234. - Baumann, Gaufin, and Surdick 1977: 1 16.
Pteronarcys dorsata, the one species of this genus found in Saskatchewan, has a transcontinental
distribution which extends from Laborador to Alaska, south in the Rockies to Wyoming and in the east
to the Great Lakes and south to Georgia. Previous Saskatchewan records include Cushing’s (1961)
report from Montreal River, Ricker’s (1944) record from Wapus River in the Reindeer Lake region, a
record from Saskatchewan River (Smith, 1917) and a report from the South Saskatchewan River at
Saskatoon (Ricker, 1946). New Saskatchewan records (Fig. 180) include: North Saskatchewan River at
Jet. Hwy. 5 (Borden Bridge) and at the ferry 10 mi. E. of Prince Albert, Sask.; South Saskatchewan
River at ferry N of Birch Hills, Sask. and at the ferry N. of Lemsford, Sask.; Nipekamew River, Jet.
Hwy. 165; Weyakwin River, Jet. Hwy. 2; Waskesiu River, Jet. Hwy. 2; Torch River, Jet. Hwy. 106;
Arsenault River, Jet. Hwy. 104; Nemeiben River, Jet. Hwy. 2; Puskwakau River, Jet. Hwy. 2;
Puskwakau River, Jet. Hwy. 106; river at mile 34, Jet. Hwy. 105; creek at mile 120, Jet. Hwy. 105;
stream 85 mi. N. of La Ronge, Sask., Jet. Hwy. 102; Battle River, 4 mi. S. of Lashburn, Sask.;
Overflowing River, Jet. Hwy. 109; Jackfish Creek, Jet. Hwy. 8; Meeyomoot River, Jet. Hwy. 165;
Quaest.Ent., 1979, 15(1)
20
Dosdall & Lehmkuhl
McDougal Creek, Jet. Hwy. 120; Bear River, Jet. Hwy. 106; Martineau River, Jet. Hwy. 55; Maekay
Creek, Jet. Hwy. 2; Maekenzie Creek, near Hwy. 165; Bow River, Jet. Hwy. 165, Swan River, Jet. Hwy.
8; Ballantyne River, Jet. Hwy. 106; Churehill River, Jet. Hwy. 2, Wintego Lake Rapids, and Iskwatam
lake; Fond du Lae River at outflow of Blaek Lake; stream at mile 30, Wollaston Lake Road; Caribou
Creek, Jet. Hwy. 106; Red Deer River, 2 mi. S. of Hudson Bay, Sask.; and Toreh River, Jet. Hwy. 106.
Rieker (1964) presented a North Ameriean distribution map for the speeies.
Diagnostic Characters. - Average length of males, 40.0 mm; females, 60.0 mm (from anterior
margin of head to tip of folded wings). Male genitalia with ninth sternum straight near tip; epiproet
large, flattened, eomplex in shape (Fig. 46). Female subgenital plate straight, or with two small
projeetions (Fig. 47, 48).
Nymphal females with projection on tenth tergum projected postero-dorsad to point; nymphal
males with projection extended postero-ventrad with peg on caudal surface. Mature male nymphs with
a nearly rectangular projection posterad on ninth abdominal sternum.
Needham and Claassen (1925), Nelson and Hanson (1971) and Hitchcock (1974) figured the adult
genitalia, and Claassen (1931) figured the nymph.
Bionomics. - Nymphs of this species are common in streams and large rivers. Nymphs occur in
rapids (Smith, 1917) and in trash from eddies below stony rapids (Gaufin et al., 1972).
Hilsenhoff and Narf (1972) stated that the life cycle of Pteronarcys dorsata is at least two years in
Wisconsin; the aquatic stage of Pteronarcys calif ornica Newport lasts three years (Elder and Gaufin,
1973). In Saskatchewan, three size classes of Pteronarcys dorsata nymphs have been collected at one
time, suggesting a three year life cycle. Eight mature nymphs with fully developed wing pads collected at
Weyakwin River on June 9, 1975 averaged 34.2 mm long; seven immature nymphs with no wing pad
development averaged 17.0 mm long and ten very immature nymphs averaged 12.3 mm in length. Eggs
appear to hatch directly and nymphs are likely the overwintering stage since ten nymphs averaging
5.1 mm long were collected at the same river on July 16, 1975, about two months after adult emergence.
In Saskatchewan, adult emergence occurs in early June and lasts approximately three weeks.
Pteronarcys dorsata nymphs are detrivores, eating leaves and other vegetable matter that falls into
the water. They can be maintained in an aquarium on a diet of dead leaves which are skeletonized
(Harden and Mickel, 1952).
Nebeker (1971a) found the longevity of adults was related inversely to the temperature at which the
nymphs were maintained. At 10 C the mean life span was 36 days; it was 31 days at 15 C and 17.5 days
at 20 C. It was also found that the stonefly could live at higher temperatures than levels where good
development and successful emergence occurs. Highest feeding rates were observed at 20 C (no feeding
occurred at 1 C and 35 C); the best temperature for emergence was 15 C. Females reared at 15 C
produced, on average, 475 eggs, but at 20 C only one female oviposited and then laid only 175 eggs.
Water temperature had a great influence on development: at 5 C nymphs did not develop for a nine
month testing period (Nebeker, 1971a), but at a constant 20 C adults emerged five months earlier than
they normally do in the field, and the separate emergence of the sexes was far more pronounced than is
normal (Nebeker, 1971b).
Cushing (1961) reported collecting Pteronarcys dorsata in rapids of Montreal River above four
associated lakes, but not in rapids below the lakes. A possible explanation for this absence was
proposed by Lehmkuhl (1972), who pointed out that one major effect of a large reservoir is to alter the
thermal regime of an outflowing river, making it possible for some aquatic insects to complete their life
cycles in the river near the reservoir.
Stoneflies of Saskatchewan
21
Genus Pteronarcella Banks
Adults of Pteronarcella resemble those of Pteronarcys but are only about half the size. Crossveins
are fewer, and they are entirely absent from radial areas of the wing. The male ninth abdominal segment
is elevated in a broad, tranversely recurved scoop-like lobe. Some of the segments above this lobe bear
paired dorsal humps at the sides. At rest, the large U-shaped supra-anal process is concealed between
the subanals and the divided halves of the tenth segment. Female subgenital plates are not produced
over the ninth sternite (Fig. 44). Nymphs have gills on the first three abdominal segments, with the tenth
segment produced into a triangular, pointed, conical process. Two species are known in North
America; one is found in Saskatchewan.
Pteronarcella badia (Hagen)
(Fig. 49, 180)
Pteronarcys badia Hagen 1874: 573.
Pteronarcella triloba. Smith 1917: 461, 462.
Pteronarcella badia, Claassen 1940: 23. - Jewett 1956: 57. - Ricker 1964: 55. - Baumann, Gaufin and
Surdickl977: 112.
Pteronarcella badia, the only species of Pteronarcella in Saskatchewan, is common in the Rocky
Mountain region and extends eastward in Montana, Utah and Arizona. New Saskatchewan records,
(Fig. 180) include: Broad Creek, Jet. Hwy. 104 and Mistohay Creek, Jet. Hwy. 226.
Diagnostic Characters. - Average length, males 15.0 mm (from anterior portion of head to tips of
folded wings); females, 20.0 mm. Male genitalia with appendage on ninth dorsal abdominal segment
broadly rounded at apex, side margins sinuous. Hind margin of female subgenital plate rounded,
truncate and slightly trilobate but not acutely notched (Fig. 49).
Nymphal gill filaments at least twice as long as basal conical process of gill tufts.
Needham and Claassen (1925), Jewett (1956), Gaufin et al. (1966) and Gaufin et al. (1972) figured
the adult genitalia, and Claassen (1931) figured the nymph.
Bionomics. - Richardson and Gaufin (1971) report that nymphs generally occur in slow areas of
streams in Utah and Colorado where debris collects. However, in Saskatchewan, nymphs were
commonly collected from swiftly flowing areas of the streams.
Gaufin et al. (1972) state that the life cycle probably lasts two years. In Saskatchewan, adult
emergence was during the first week of June in 1975. It appears that the eggs hatch in a short period
since 1 1 nymphs averaging 3.9 mm long were collected in mid-July. The nymphs would be the
overwintering stage.
Nymphs of Pteronarcella badia are predominantly herbivorous though some individuals feed on
animal material when plant matter is unavailable or scarce. (Richardson and Gaufin, 1971).
Family Taeniopterygidae
Taeniopterygids have tarsal articles subequal in length; epiprocts of males are prominent and
paraprocts are generally greatly modified. Adults are also characterized by cu-m intercubital crossveins.
The range includes North America, Eurasia, and northern Africa.
Zwick (1973) recognized two subfamilies. Taeniopteryginae includes the single genus Taeniopteryx,
and 13 genera are recognized in the Brachypterinae. Ricker and Ross (1975) discussed the
distinguishing characters of the two subfamilies. In Brachypterinae, the male ninth sternite is scoop-like
Quaest.Ent., 1979, 15(1)
22
Dosdall & Lehmkuhl
and greatly produced beyond the tenth segment. Inner members of male subanal lobes are complex,
asymmetrical and partly membranous. Male cerci have at least two articles. The female ninth sternite is
produced well beyond the base of the subanal lobes and coxal gills are absent.
In Taeniopteryginae the male ninth sternite is not scoop-shaped and extends very little beyond the
tenth segment. Males have simple, symmetrical and sclerotized subanal lobes, and lack a basicercal
process. The female subgenital plate is somewhat produced over the tenth segment and rounded but
does not extend to the subanal lobes. Male cerci have a single article and coxal gills are present. Both
subfamilies of Taeniopterygidae are represented in Saskatchewan by Taeniopteryx nivalis of the
Taeniopteryginae, and Oemopteryx fosketti of the Brachypterinae.
Genus Taeniopteryx Pictet
Nymphs of this genus have coxal gills (Fig. 13). Adults have membranous circular areas on each
coxa, which represent coxal gill scars. Male paraprocts are concealed within the ninth sternite and there
are usually two membranous lobes behind the female subgenital plate (Fig. 44). Eight Taeniopteryx
species are known in North America; one species occurs in Saskatchewan.
Taeniopteryx nivalis (Fitch)
(Fig. 13,40,41,43,44, 181)
Nemoura nivalis Fitch 1847: 274.
Taeniopteryx nivalis, (in part) Needham and Claassen 1925: 240. - Harden and Mickel 1952: 12. -
Ricker and Ross 1968: 1434.
Taeniopteryx maura, {indent) Frison 1942a: 248. - Jewett 1959: 55, and 1960: 151.
Taeniopteryx nivalis ranges from Laborador through Quebec and Ontario to Minnesota and south
to Pennsylvania, northern Indiana and Illinois. Western records include Oregon and Alberta with this
as the first Saskatchewan record. The Saskatchewan distribution (Fig. 181) includes the following
localities: Waskesiu River, Jet. Hwy. 2; Crean River, Jet. Hwy. 2; Weyakwin River, Jet. Hwy. 2;
Montreal River, Jet. Hwy. 2 (mile 65); Little Red River near Prince Albert, Sask.; Jackfish Creek, Jet.
Hwy. 8; Torch River, Jet. Hwy. 106; McDougal Creek, Jet. Hwy. 120, Nipekamew River, Jet. Hwy.
165; and Caribou Creek, Jet. Hwy. 120.
Diagnostic Characters. - Average length males, 11.0 mm (from anterior margin of head to tip of
folded wings); females, 17.0 mm. Males without spurs or other projections on hind femur; with vesicle
or ventral lobe on ninth sternite, 2-3 times long as wide. Hairs on hind margin of the ninth sternite
directed downward, and forward generally much shorter than those situated more anteriorly on sternite
(Fig. 40). Aedeagus without brown sclerotized band between the two lateral lobes (Fig. 41). Females
with strongly sclerotized V-shaped notch on eighth sternite, produced sides of the notch in contact at
acute angle (Fig. 44).
Abdomen of nymph with mid-dorsal stripe or several light dots on the posterior margin of each
segment. Abdominal tergites with short, thick bristles and a few long hairs on hind margins.
Needham and Claassen (1925) figured the female; Claassen (1931) and Harper and Hynes (1971c)
illustrated the nymphs. Figures of the male genitalia are presented in Fig. 40, 41, and 43.
Bionomics. - Harper and Hynes (1970) determined that the life cycle of Taeniopteryx nivalis is
univoltine; eggs are deposited in April and May, hatch directly, and nymphs diapause in an
undetermined instar (4 or 5). In diapausing nymphs, fat globules accumulate, all bristles are lost, cerci
are cast off, legs are folded and head and antennae are reflexed under the thorax. Diapausing nymphs
Stoneflies of Saskatchewan
23
have not been found in streams probably because they burrow deep into the substratum. Summer
diapause enables early instar nymphs to survive high water temperatures; later instars are extremely
heat sensitive (Nebeker and Lemke, 1968). Diapause ends in late September (Harper and Hynes, 1972)
and most growth occurs in winter (Harper and Hynes, 1970).
Coleman and Hynes (1970) and Harper and Hynes (1972) reported rapid growth from October
through the winter months. Emergence of adults begins in mid-March and lasts about one week.
Mating occurred soon after emergence. Harper and Hynes (1972) noted that oviposition began about
one week after the maximum period of emergence and adults lived for about one month in the field.
Under simulated stream conditions in the laboratory, egg hatching occurred about 40 days after
oviposition.
In Saskatchewan, adults emerged during the end of March of 1976. Adults remained in ice cavities
under bridges in early May when all other ice had melted. If removed, adults immediately crawled back
to the same cavity or entered another cavity.
Harper and Hynes (1972) found that the species was primarily detrivorous, but occasionally
ingested animal matter.
Genus Oemopteryx Klapalek
Wings of adult Oemopteryx lack costal crossveins and Rs and Cui each have two branches. Males
have two epiproct prongs which are usually set together. Outer members of the paraprocts are
elongated and heavily sclerotized (Fig. 42). Female subgenital plates are variable, and nymphal
characters have been little studied.
Ricker and Ross ( 1975) noted that there are four well-marked species groups of Oemopteryx, the
loewii group from central Europe, the contorta group from the Appalachian Mountains, the vanduzeea
group from California and Xhtglacialis group from northeastern North America and the plains region.
The one species of Oemopteryx known from Saskatchewan belongs to iht glacialis group.
Oemopteryx fosketti (Ricker)
(Fig. 42,45, 154-158, 171-176, 181)
Brachyptera fosketti Ricker 1965: 475.
Brachyptera zelona, Ricker 1965: All.
Oemopteryx fosketti, Baumann 1973: 95. - Ricker and Ross 1975: 140
Oemopteryx fosketti is known from the North and South Saskatchewan Rivers, and has also been
reported from Utah. The type locality is the South Saskatchewan River at Clarksboro, Sask. The
Saskatchewan distribution (Fig. 181) includes the following localities: South Saskatchewan River at
Lemsford Ferry, Saskatoon, Sask., and Ferry near Clarksboro, Sask.; and the North Saskatchewan
River at Hwy. 5 (Borden Bridge).
Diagnostic Characters. - Average length, females, 10.5 mm (from anterior portion of head to tip of
folded wings); males average 7.5 mm (from anterior margin of head to end of abdomen). Males
brachypterous, with forewings greatly reduced (about 2.5 mm long) and upturned near the tip;
hindwings narrowed and of less than normal length. Male genitalia with supra-anal process divided into
four parts: basal bulb brown and smoothly rounded with low, dark, flat, conical apex next to groove
separating it from anterior erect member; anterior erect member slender and curved forward with low
swellings laterally near tip; posterior erect member near anterior erect member but broader and
terminated in two hemispherical membranous bulges separated by angular groove; hind surface of
Quaest.Ent., 1979, 15(1)
24
Dosdall& Lehmkuhl
posterior erect member opened to membranous sleeve or posterior portion of epiproct (Fig. 42).
Females with distinct subgenital plate, with hind margin anterior to the hind margin of eighth sternite;
subgenital plate rounded and shallowly excavated medially (Fig. 45). The previously unknown nymph
is described below.
Description. - Total length of mature nymphs: 10.5 mm (males); 12.0 mm (females).
General color dark brown to nearly black, light yellow ventrally. Head uniformly dark with few
dark mottlings near antennae; antennae almost as long as body. Antennal scape and pedicel patterned
dorsally as in Fig. 155. Mouthparts as in Fig. 156, 157.
Thoracic nota dark brown; mesonotum with two subrectangular patches of sclerotization at its
anterolateral corners. Metanotum with anterior subrectangular sclerotized area separated by
membranous area from large metathoracic sclerotized block. Wing pads of mature male nymph pointed
and small (the adult is brachypterous); female wing pads normal. Legs yellowish brown, darker at
joints; fringes of long hairs along femora, tibiae and tarsi.
Each abdominal tergite with anterior half dark brown, and posterior half light brown. Abdominal
terga covered with long clothing hairs; the posterior tergal margin beset with very short bristles and few
long hairs (Fig. 158). Sterna 1-7 unsclerotized. Cerci longer than body; cereal articles 1-8 with long
bristles, about twice as long as the cereal article; the other articles with whorls of very short bristles,
about one-fifth the length of the segment.
Until comparative studies are made of closely related species of this genus, distinguishing characters
of the nymph of Oemopteryx fosketti cannot be specified.
Bionomics. - Adults emerged on April 16, 17, 18, and 19, 1975 and April 1-4, 1976, at the North
Saskatchewan River near Hwy. 5 (Borden Bridge). Water temperature was 0.8 C for the 1975 dates and
1.0 C for the 1976 dates.
Males emerge earlier than females. On April 17, 1975, 121 adults were collected from a small section
of the river bank; 106 were males and 15 were females. From a random sample of 79 nymphs collected
the same day and reared to adults, 59 were females and 20 were males. This indicated that a significant
number of females had yet to emerge. By April 19, 1975, males and females were equally abundant in
collections along the river bank: of 77 adults, 41 were males and 36 were females. Earlier male
emergence is not uncommon in stoneflies and has been documented for many species. Brinck (1949)
suggested a definite advantage to this discrepancy between emergence of the sexes: males are fully
developed and ready to mate as soon as females emerge.
Time of ice break-up is probably a critical factor in determining onset of adult emergence. On
April 16, 1975, a main channel of water had just started to form but no adults had emerged. By April 17
a main water channel had formed (Fig. 176) and adults were abundant. On April 1, 1976, runoff from
highway ditches had melted the river ice under Borden Bridge causing a small opening and adults were
present only in this small area. When a large water channel had opened the river adults were numerous
along the entire river bank.
Mating occurred immediately after emergence for Oemopteryx foskettv. in fact, some males
attached to emerging females. After the female had completed eedysis, they mated immediately.
Males are very active and walk about on snow searching out females (Fig. 171). Many females often
remain in small cavities or pockets within the snowbanks (Fig. 172). When discovered by a male,
mating usually occurs in the cavity. In one instance, a male encountered a mating pair and also
attempted to mate with the female. A struggle ensued (Fig. 173) until one male was driven off. The
remaining male mated with the female (Fig. 174).
Abdomens of several Oemopteryx fosketti were examined to ascertain degree of egg maturation.
Abdomens used were taken from a mature female nymph, an emerging nymph in which the wings had
Stoneflies of Saskatchewan
25
just begun to leave the nymphal wing pads, a newly emerged female, a fully emerged female collected in
the field and a fully emerged female reared in the laboratory.
Abdomens were soaked in 100 percent ethanol for two periods of one hour each, benzene for one
hour, one-half benzene and one-half paraffin for one hour and finally were embedded in paraffin.
Abdomens were sectioned at 8 microns and placed on glass slides. The slides were dried, the paraffin
removed with xylene and stained using Harris’ Hematoxylin and Eosin. The tissue was then mounted
using Permount. All slides were treated alike.
The eggs appeared similar in all slides studied. In a mature females the eggs stained purple, with a
resistant outer covering. No difference was found in a mature female nymph - eggs were in the
abdominal cavity and appeared to be as mature as those of a fully developed female. In all abdomens,
the eggs appeared identical to eggs which had been deposited on the ice.
Mating soon after adult emergence and the occurrence of mature ova in newly-emerged females is
probably an adaptation to the type of habitat in which Oemopteryx fosketti lives. After ice break-up
occurs, excessive runoff from the surrounding terrain often causes flooding in the North Saskatchewan
river. Since emergence occurs as soon as the ice begins to break up, ability to mate and lay eggs
immediately ensures that the eggs are fertilized and deposited before the adults may be killed by a flood.
Males are brachypterous and could not likely escape a high flood.
The life cycle of this species is probably similar to that described for several other winter stoneflies
(Harper and Hynes, 1970). Eggs of winter stoneflies hatch within a month and early-instar nymphs are
thought to diapause throughout the summer. Diapause was proposed as an adaptation to survival of
high water temperatures in summer. Nymphs in a diapause-like state burrowed deep into the
substratum, and this probably explains the absence of members of O. fosketti from summer benthic
samples. When collected in September, nymphs did not seem to have made any significant growth.
Most growth and development occurs during winter.
Family Capniidae
Capniids are small, dark stoneflies characterized by many-articled cerci, wings at rest folded flat on
the back, the second tarsal article short, and forewings with only one or two median crossveins and one
cubital crossvein. The species are arranged in two subfamilies - the Capniinae of North America,
Eurasia and northern Africa, and the Notemourinae of Africa, South America, and Australia and with
the single genus Megaleuctra in northwestern North America. Four genera of Capniinae Par acapnia,
Capnia, Utacapnia and Isocapnia occur in Saskatchewan, but the Notemourinae are not known in the
province.
Genus Par acapnia Hanson
Paracapnia members are distinct in having the mesothoracic postfurcasternal plate united with the
furcasternum and the spinasternum (Fig. 19); the meso- and metafurcasternum are transverse and
almost rectangular. Ri of the forewing is bent caudally beyond its base and Cui of the hindwing
generally has its apical portion missing (Fig. 33). Two species of this genus are known, and one occurs
in Saskatchewan.
Quaest.Ent., 1979, 15(1)
26
Dosdall& Lehmkuhl
Paracapnia angulata Hanson
(Fig. 19,33, 60,61, 179)
Paracapnia angulata Hanson 1961: 29. - Hitchcock, 1974: 69.
Capnia opis, Prison 1942a: 264.
Paracapnia angulata and Paracapnia opis (Newman) are very closely related and have only recently
been recognized as distinct. The geographic range of Paracapnia angulata cannot be delimited until
previously identified specimens have been checked. This is the first Saskatchewan record of the species;
specimens have been collected from the Montreal and Nipekamew Rivers (Fig. 179).
Diagnostic Characters. - Average length, females 8.5 mm (from anterior part of head to tip of
folded wings); brachypterous males 4.0 mm (from anterior part of head to end of abdomen). Males of
Paracapnia angulata are distinguishable from those of Paracapnia opis only by the male supra-anal
process. In lateral view, Paracapnia angulata males have both inner and outer margins of the epiproct
angled at the base (Fig. 61). The inner margin is acutely angled or may approach a right angle, but it is
not evenly curved as in Paracapnia opis. In dorsal view, the epiproct of Paracapnia angulata is
broadened once and then tapers evenly to the tip (Fig. 60), but the epiproct of male Paracapnia opis
shows two slight enlargements in dorsal view. Females of these two species are indistinguishable;
members of both have the posterior margin of the subgenital plate broadly rounded and with a small,
mesal light-colored or membranous spot (Fig. 68).
Nymphs of Paracapnia angulata have a long intermediate bristle at the ventral base of the middle
and distal cereal articles and short bristles on the inner surface of the tibiae about half as long as the
width of the tibia. In Paracapnia opis intermediate bristles are absent from cereal articles and the tibial
bristles are as long as the width of the tibia.
Hanson (1961) and Harper and Hynes (197 lb) figured the adult genitalia; figures of the nymph were
published by Harper and Hynes (1971b).
Bionomics. - Harper and Hynes (1972) noted that the emergence pattern of Paracapnia angulata
was relatively short and synchronous, with the main peak lasting only eight days. Oviposition occurred
about one week after emergence and the first nymphs were collected about two months after the
appearance of ovipositing females. Growth proceeded rapidly through summer and autumn and by
mid-November nearly all specimens had reached the penultimate instar. The last instar was reached by
December. Unlike other winter stoneflies, members of Paracapnia angulata do not undergo summer
diapause, but show significant summer growth.
Harper and Hynes (1970) state that this species is restricted to cold spring-fed streams and thus
nymphs are not exposed to high summer water temperatures. Diapause, an adaptation to survival of
high summer water temperatures, may then be superfluous since individuals are exposed to low summer
water temperatures. In Saskatchewan, Paracapnia angulata has been collected in Montreal River and
Nipekamew River. Neither of these rivers are spring-fed, but both drain lakes. Lake drainage may affect
the temperature scale of out-flowing rivers (Lehmkuhl, 1972), resulting in summer water temperatures
which are low enough for Paracapnia angulata nymphs to withstand.
Genus Capnia Pictet
Adults of this genus are characterized by having vein Ri of the forewing bent upward toward the
costal margin at its origin, and Ai of the forewing bent caudad at its junction with cu-a and then curved
outward (Fig. 37). Thoracic spinasterna are fused to the basisterna, and metathoracic postfurcasternal
plates and thoracic presterna are free. Nymphs lack a fringe of cereal bristles. In North America, most
Stoneflies of Saskatchewan
27
Capnia species are known only from the western part of the continent. Fifty-nine species are known in
North America, and four occur in Saskatchewan.
Key to Saskatchewan species of Capnia
Adults
1 With supra-anal process recurved forward and above ninth and tenth terga (males)
1 ' Supra-anal process absent (females)
2 (1) Conspicuous hump on tergum 8; supra-anal process with conspicuous downturned
lobe at apex (Fig. 58, 59) Capnia coloradensis Claassen p.
2' No hump on tergum 8; hump on tergum 7 or absent from all terga
3 (2') Supra-anal process extended to tergum 7; a low hump on tergum 7 near anterior edge
(Fig. 52, 53) Capnia gracilaria Claassen, p.
3' Supra-anal process extended at most to middle of tergum 8, no hump on tergum 7
4 (3') Supra-anal process expanded at the basal 0.66 of length and narrowed near tip
(Fig. 56, 57) Capnia vernalis Newport, p.
4' Supra-anal process tapered evenly from base to tip (Fig. 54, 55)
Capnia confusa Claassen, p.
5 ( 1 ') Sterna 7 and 8 united by a median sclerotized connection of various widths
5' Sterna 7 and 8 not connected, but separated by membranous area
6 (5') Posterior edge of sternum 7 membranous with narrow median sclerotized projection
extended to or overlapping part of sternum 8 (Fig. 70) Capnia confusa Claassen, p.
6' Posterior edge of sternum 7 sclerotized, uniform, unmodified, medially; posterior edge
of subgenital plate projected posteriorly usually with low median and lateral
projections (Fig. 72) Capnia gracilaria Claassen, p.
7 (5 ) Conspicuous subcuticular oval object visible in center of subgenital plate; subgenital
plate separated from lateral sclerotized patches by narrow membranous area (Fig. 71) ..
Capnia coloradensis Claassen , p.
7' Subgenital plate without median oval object, subgenital plate united to lateral
sclerotized patches (Fig. 67) Capnia, vernalis Newport, p.
Nymphs
1 Tip of galea expanded, with fringe of long hairs (Fig. \<X\).. Capnia vernalis Newport, p.
1 ' Tip of galea evenly tapered or pointed (Fig. 142-144)
2 ( 1') Long setae on head about equal to diameter of eye, male supraanal lobe long (Fig. 138)
Capnia gracilaria Claassen, p.
2' Long setae on head about equal to half diameter of eye, male supra-anal lobe short
(Fig. 139, 140) Capnia confusa Claassen, p. 28; Capnia coloradensis Claassen, p.
2
5
27
3
29
4
30
28
7
6
28
29
27
30
30
2
29
27
Capnia coloradensis Claassen
(Fig. 58,59,71, 140, 144, 177)
Capnia coloradensis Claassen 1937: 79. - Claassen 1940: 92. - Hanson 1946: 238. - Ricker 1965: 487. -
Gaufin, Nebeker and Sessions 1966: 48. -Gaufin, Ricker, Miner, Milam and Hays 1972: 66.
Capnia coloradensis is known from the Rocky Mountain areas of Montana, Idaho, Wyoming, and
Colorado in the United States, and from the Yukon, Canada. This is the first Saskatchewan record. The
species has been collected (Fig. 177) from: Scarth River, Jet. Hwy. 120; Mackenzie Creek near Bow
Quaest. Ent., 1979, 15(1)
28
Dosdall & Lehmkuhl
River, Jet. Hwy. 165; Trapper Cabin Creek, Jet. Hwy. 120; Cub Creek, Jet. Hwy. 106; and MeDougal
Creek, Jet. Hwy. 120.
Diagnostic Characters. - Average length, males, 5.0 mm; females, 7.0 mm (from anterior margin of
head to tip of folded wings). Males with narrow heavily selerotized eurved band on anterior margin of
tergum 7 and median shining area behind it; latter flat in most speeimens, but in Saskatehewan
speeimens with barely pereeptible dark knob rising from it. Narrow dark band extended along anterior
margin of terga 2-6, interrupted medially. Prominent hump on tergum 8 and supra-anal process long
with conspicuous downturned lobe at apex. Sternum 9 without ventral lobe. Female sternum 7
moderately selerotized, with many long, stout pale hairs on posterior half, sclerotizat-
ion continuous across intersegmental groove. Subgenital plate with posterior margin small, rounded
and with conspicuous subcuticular oval area visible in center; subgenital plate separated from lateral
selerotized areas of sternum 8 by narrow membranous area. The previously unknown nymph is
described below.
Description. - Total length of mature nymph: 3. 5-4.5 mm.
General color light brown, lighter ventrally; darker on borders of pronotum, anterior corners of
meso- and metanota and anteriorly to lateral ocelli. Appendages and wing pads light yellow; no
conspicuous color pattern. Maxillae as in Fig. 144.
Head mostly glabrous with few long setae on top of the head, longest about equal to half diameter
of eye (Fig. 140).
Pronotum with few long bristles on anterior and posterior borders, with shorter bristles between.
Wing pads with many short hairs. Legs with usual fringe of hairs on tibiae and tarsi; tibiae with short
stout bristles below fringe. Outer surface of femur with many short bristles and a few interspersed
longer ones.
Abdomen covered with short clothing hairs, and a few stout bristles on posterior tergal margins
(Fig. 140). Cerci of usual type.
Ricker (1965), Gaufin et al. (1966) and Gaufin et al. (1972) figured the adult genitalia; drawings of
the nymph are presented in Fig. 140 and 144.
Bionomics. - Gaufin et al. (1972) noted that adults of Capnia coloradensis emerge in March and
April. In Saskatchewan, adults were collected on April 18, 1975 at Trapper Cabin Creek, Jet. Hwy. 120;
May 29, 1975 at Cub Creek, Jet. Hwy. 106; April 12, 1976 at Mackenzie Creek, Jet. Hwy. 165 and on
May 30, 1975 at MeDougal Creek, Jet. Hwy. 120. Nymphs were absent from benthic samples after the
spring adult emergence indicating a univoltine life history.
Capnia confusa Claassen
(Fig. 37,54, 55,70, 139, 142, 179)
Capnia nivalis 1929: 163.
Capnia confusa Claassen 1936: 623. - Hanson 1946: 238. - Jewett 1959: 43. - Gaufin, Nebeker and
Sessions 1966: 47. - Gaufin, Ricker, Miner, Milam and Hays 1972: 67.
Capnia confusa is known from Alaska, Alberta and British Columbia, south to Utah, and east to
Wyoming and Montana. This is the first Saskatchewan record, and the species has been collected
(Fig. 179) from: North Fork of Scarth River, Jet. Hwy. 120; the stream at mile 83, Jet. Hwy. 106;
Mackenzie Creek, near Bow River, Jet. Hwy. 109; River 40 mi. N. of of Hudson Bay, Sask., Jet. Hwy.
109; and Waskwei River, Jet. Hwy. 109.
Ricker (1964) presented a North American distribution map of the species.
Stoneflies of Saskatchewan
29
Diagnostic Characters. - Average length, males 5.5 mm; females, 7.5 mm (from anterior part of
head to tip of folded wings). Males of this species closely resemble those of Capnia vernalis, but females
of Capnia confusa and Capnia vernalis are markedly different. Male supra-anal process long, about 10
times longer than wide (Fig. 54), fairly uniform in width throughout its length and tip not down-turned
in side view (Fig. 55). Humps absent from both terga 7 and 8. Sterna 7 and 8 of females not united by
median sclerotized connection; posterior edge of sternum 7 membranous with narrow median
sclerotized projection extended to or overlapping part of sternum 8 (Fig. 70). The first description of a
nymph of this secies is given below.
Description. - Total length of mature nymph: 4.5 -6 mm.
General color light brown, lighter ventrally; darker on borders of pronotum, anterior corners of
meso- and metanota and anteriorly to lateral ocelli. Appendages and wing pads yellow. Maxillae as in
Fig. 142.
Head mostly glabrous with few long setae, longest about half diameter of eye (Fig. 139).
Pronotum with a few long bristles on anterior and posterior borders, and shorter bristles between
meso- and metanota with a few long hairs extended anteriorly and shorter bristles on the disc; wing
pads with many short hairs. Legs with usual fringe of hairs on tibiae and tarsi; tibiae with short stout
bristles below fringe. Outer surface of femora with many short bristles and few interspersed longer ones.
Abdomen covered with short clothing hairs, and few stout bristles on posterior tergal margins
(Fig. 39). Supra-anal lobe of male as in Fig. 139; cerci of the usual type.
I have been unable to distinguish the nymph of this species from the nymph of Capnia coloradensis .
Gaufin et al. (1966) figured the male and female genitalia; first figures of the nymph are presented in
Fig. 139 and 142.
Bionomics. - Gaufin et al. (1972) noted that Capnia confusa nymphs are common in creeks and
adults emerge in February to July. In Saskatchewan, adults were collected from the North Fork of
Scarth River, Jet. Hwy. 120 on May 15, 1975, stream at mile 83, Jet. Hwy. 106 on April 26, 1976,
Mackenzie Creek near Bow River, Jet. Hwy. 165 on April 16, 1976, and May 7, 1976, McDougal Creek,
Jet. Hwy. 120, on May. 30, 1975, and from Waskwei River, Jet. Hwy 109 on June 11, 1975. Nymphs
were absent from benthic samples after the adults emerged in spring, indicating a univoltine life history.
Capnia gracilaria Claassen
(Fig. 52, 53,72, 138, 143, 177)
Capnia gracilaria Claassen 1924: 57. - Needham and Claassen 1925: 258. - Claassen 1940: 93. - Ricker
1943: 99. - Hanson 1946: 239. - Gaufin, Nebeker and Sessions 1966: 46. - Gaufin, Ricker, Miner,
Milam and Hays 1972: 67.
Capnia gracilaria is known from British Columbia and Manitoba south to Oregon, Montana and
Utah. This first Saskatchewan record is from Cypress Hills at Battle Creek near Reesor Lake (Fig. 177).
Diagnostic Characters. - Average length, males, 4.0 mm (from anterior portion of head to tip of
folded wings); females, 7.5 mm. Males without conspicuous hump on tergum 8 but most specimens with
low hump near anterior edge of tergum 7. Supra-anal process long (about 10 times longer than wide)
and round in cross-section, extended to posterior edge of tergum 7 (Fig. 52). In lateral view, supra-anal
process gently S-shaped and of same width throughout its length (Fig. 53). Female abdomen with very
broad dorsal membranous stripe on segments 1 to 8. Subgenital plate of most specimens slightly
anterior to membranous border of eighth sternum and set off laterally by weakly sclerotized areas;
posterior edge of subgenital plate is usually straight or slightly rounded often with small median blunt
protuberance, plate with 3-toothed appearance (Fig. 72). The nymph is described below for the first
Quaest.Ent., 1979, 15(1)
30
Dosdall& Lehmkuhl
time.
Description. - Total length of mature nymph: 5.0-6. 5 mm.
General color light brown, reddish brown in very mature specimens, lighter ventrally, appendages
and wing pads light yellow; no conspicuous color pattern.
Head mostly glabrous with a few long setae on top of head, longest about equal to diameter of the
eye (Fig. 138). Galea of maxilla without definite fringe of hairs (Fig. 143).
Pronotum with few long bristles at anterior and posterior borders, and few short hairs between.
Meso- and meta-nota with few long bristles anteriorly, and few stout bristles along notal discs; wing
pads glabrous. Legs with usual fringe of hairs on tibiae and tarsi; tibiae with short stout bristles below
fringe. Outer surface of femora with many short bristles and a few interspersed longer ones.
Abdomen with a few stout bristles on posterior tergal margins and a few short hairs along rest of
each tergum(Fig. 138). Supra-anal lobe of male as in Fig. 138; cerci of usual type.
Needham and Claassen (1925) figured the male genitalia; Ricker (1943) illustrated the female
genitalia. Gaufin et al. (1966) and Gaufin et al. (1972) figured both male and female genitalia. First
nymphal figures are presented in Figs. 138 and 141.
Bionomics. - Gaufin (1972) noted that nymphs of this species are common in creeks and small
rivers, with adult emergence occurring from January through May. In Saskatchewan, adults were
collected at Battle Creek, Cypress Hills on April 20, 1975, March 28, 1976 and May 19, 1976. Nymphs
were absent from benthic samples after the emergence of adults in spring indicating a univoltine life
cycle.
Nebeker( 1971c) studied the adult emergence of Capnia graclaria at altitudes of 2590 meters down to
1530 meters in the Wasatch Mountains, Utah. Emergence was found to be spread up to five months
apart depending on altitude (i.e. temperature) for nymphs which received the same photoperiod.
Capnia vernalis Newport
(Fig. 56, 57,67, 141, 178)
Capnia vernalis Newport 1848: 451. - Claassen 1931: 109. - Ricker 1938: 135. - Claassen 1940: 95. -
Harden and Mickel 1952: 24. - Harper and Hynes 1971b: 938. - Hitchcock 1974: 45.
Published records of Capnia vernalis are from Alberta, Manitoba, Ontario, Labrador, Minnesota
and Ohio. Although specimens were not collected in the present study and there are no published
records of the species from Saskatchewan, W.E. Ricker (pers. comm.) has examined a female Capnia
vernalis from this province. Saskatoon, Sask.; May 19, 1940; A.R. Brooks, coll. An additional female is
deposited in the Biology Department Entomology Museum, University of Saskatchewan, Saskatoon,
Saskatchewan, with the following collection data: Saskatoon, Sask.; April 24, 1918. The genitalia are
shrivelled since this specimen has been pinned, but it appears very close to Capnia vernalis. The
Saskatchewan distribution of this species is indicated in Fig. 178.
Diagnostic Characters. - Average length, males 4.5 mm; females, 6.5 mm (from anterior part of
head to tip of folded wings). Male sternum 9 produced posteriorly into bluntly pointed subanal plate
with elongate process at tip lying between subanal lobes; elongate process with second acute anterior
projection dorsal to sternum. Subanal lobes each subacute and marked off from sternum by deep
notch. Supra-anal process extended forward to hind margin of tergum 8 (Fig. 56); its distal one-third
narrower in side view and the tip pointed (Fig. 57). Female subgenital plate with strongly sclerotized
posterior lip about one-third width of sternum, and slightly anterior to its hind margin. Sclerotized
floor of genital tract visible through exoskeleton (Fig. 67) anterad to posterior lip of subgenital plate.
Sterna 7 and 8 connected by narrow sclerotized bridge.
Stoneflies of Saskatchewan
31
Nymph with tip of galea of maxilla not evenly pointed, but expanded and covered with fringe of
long hairs (Fig. 141).
Ricker (1938) figured the type specimens; Harper and Hynes (1971b) illustrated the adult genitalia,
wings and nymph.
Bionomics. - There is no available biological information on this species.
Genus Utacapnia Nebeker and Gaufin
Nebeker and Gaufin (1965) studied the Capnia columbiana complex and reported 10 species in
western and arctic America. Further study of this group and other North American Capnia resulted in
recognition of subgeneric status for this complex under the name Utacapnia (Nebeker and Gaufin,
1967), and it was later assigned generic status by Zwick (1973).
Males vary from long-winged to apterous. There is no lobe on sternum 9. With the exception of
wingless adults, postfurcasternal plates are partially fused to the spinasternum. The prothoracic
spinasternum is fused at lateral angles to the mesothoracic basisternum by a connection four times
longer than wide. The supra-anal process is bipartite with the upper part furcate and enlarged at its tip
(Fig. 50, 51). Females have the subgenital plate bounded on either side by a well-defined lateral
sclerotized plate with various median and anterior sclerotization (Fig. 69). Nymphs are poorly known
and inseparable at present. One species of Utacapnia is known from Saskatchewan.
Utacapnia trava (Nebeker and Gaufin)
(Fig. 50,51,69, 178)
Capnia trava Nebeker and Gaufin 1965: 479.
Capnia (Utacapnia) trava, Nebeker and Gaufin 1967: 236.
Utacapnia trava, Zwick 1973: 392.
Utacapnia trava was previously reported from Montana and Idaho. This is the first Saskatchewan
and Canadian record (Fig. 178). Specimens have been collected at Battle Creek, near Reesor Lake,
Cypress Hills; Scarth River, Jet. Hwy. 120; Trapper Cabin Creek, Jet. Hwy. 120; and McDougal Creek,
Jet. Hwy. 120.
Diagnostic Characters. - Average length: males, 7.5 mm (from anterior margin of head to end of
abdomen); females, 8.5 mm (from anterior margin of head to tip of folded wings). Male genitalia with
fan-shaped enlargement at tip of upper supra-anal process; lower process massive and greatly enlarged,
about as wide as tips of upper process; posterior end of supra-anal process enlarged and notched in
dorsal view (Fig. 50, 51). Females with anterior sclerotization of subgenital plate joined by continuous
pigmentation from anterior sclerotization extended posteriorly to pigmented tip of subgenital plate;
posterior margin notched(Fig. 69). Nymphs of this species are unknown.
Nebeker and Gaufin (1965) and Gaufin et al. (1972) illustrated the adult genitalia.
Bionomics. - Adults of this species are usually collected from February to April, but in one
collection from a glacial lake emergence was July 11, 1964 as a result of the extremely cold water in
which the nymphs lived (Nebeker and Gaufin, 1967). In Saskatchewan, adults were collected from
McDougal Creek and Trapper Cabin Creek on April 18, 1975, and from Battle Creek, on April 2, 1975
and March 28, 1976.
Quaest. Ent., 1979, 15(1)
32
Dosdall& Lehmkuhl
Genus Isocapnia Banks
Compared to most capniines, specimens of Isocapnia are unique because of their large size and the
rarity of their collection. Adults are characterized by basally straight veins Ri and Ai and the forewing,
two or more crossveins in the costal area beyond the cord (Fig. 39), the mesothoracic furcasternum
faintly demarked from the postfurcasternal plates with which it is united, metathoracic furcasternum
united with and faintly demarked from the first abdominal sternum, and the prothoracic presternum
broadly fused with the basisternum (Fig. 18). The known nymphs of Isocapnia differ from those of
other stonefly species by long swimming hairs along the cerci (Fig 26). Dwarf males occur in some
Isocapnia species in w'hich wings are very reduced (1-2 mm long) and the body is generally small.
Though brachyptery is not rare in Plecoptera, Ricker (1959) noted that the unusual aspect of
brachyptery in Isocapnia is the occurrence of extremely short-winged and completely long-winged
individuals in the same population without intergradation.
Ricker (1959) established three well-defined North American species groups of Isocapnia for the
eleven species known. These are ihtgrandis, hyalita, and abbreviata species groups. One member of the
grandis group and one member of the hyalita group occur in Saskatchewan.
Key to Saskatchewan species of Isocapnia
Adults
1 Costal crossveins one to four before end of Sc, and none to two beyond it; male 9th
tergum with prominent posterior notched or bilobed process; female subgenital plate
without median recessed tongue set off by unsclerotized notch on either side (Fig. 66)
Isocapnia missourii Ricker p.
1 ' Costal crossveins four to eight before end of Sc, two to four beyond it (Fig. 39); no
raised process on male 9th tergum (Fig. 63, 64); female subgenital plate with median
recessed tongue set off by unsclerotized notch on either side (Fig. 65)
Isocapnia crinita (Needham and Claassen), p.
Nymphs
Since the nymph of Isocapnia missourii is unknown, a nymphal species key is not provided.
33
32
Isocapnia crinita (Needham and Claassen)
(Fig. 26, 39, 62-65, 179)
Capnia crinita Needham and Claassen 1925: 269.
Isocapnia crinita, Claassen 1940: 96. -Frison 1942b: 69. - Hanson 1946: 239. - Ricker 1959: 643.
Isocapnia crinita has been reported from Colorado, Utah and Montana with this as the first
Saskatchewan and Canadian record. It has been collected from Battle Creek, Cypress Hills (Fig. 179).
Diagnostic Characters . - Average length, males, 1 1.5 mm (from most anterior margin of head to tip
of the folded wings); females, 15.0 mm. Male tenth tergum cleft dorsally, with posterior angle of cleft
filled by broad, triangular, basal portion of supra-anal process; latter curved gently anterad and
upward, with very short blunt tip extended abruptly anterad (Fig. 62-64). Females with dorsal
unsclerotized stripe on terga 2-8; all sterna with heavily sclerotized areas anteriorly, broadly
interrupted at midline. Sterna 1-6 with series of irregular dots on either side of midline, sternum 7 with
two nonsclerotized areas posterlaterally. Subgenital plate situated slightly anteriorly to level of lateral
margins of sternum; central sclerotized portion with margins more markedly sclerotized but pigment
not extended to hind margin of plate (Fig. 65).
Stoneflies of Saskatchewan
33
Male nymph with sheath of supra-anal process curved forward as in adult.
It is interesting that a single dwarf male of this species was collected at Battle Creek on
May 19, 1976. Long-winged specimens were not collected. Isocapnia crinita can now be added to
Ricker's (1959) list of Isocapnia species in which dwarfing is known to occur, leaving three species
abbreviata, grandis and mogila in which only normal males are known.
Prison (1942a), Ricker (1959), and Gaufinc/ al. (1972) figured the male and female genitalia, Ricker
(1959) figured the terminalia of the male exuviae.
Bionomics. - Specimens of Isocapnia crinita are rarely collected. Gaufin et al. (1972) state that in
Montana adults emerge from March to May. A single female was found on June 3, 1975 at Battle Creek
near Reesor Lake, Cypress Hills, and 15 females and one male were collected on May 19, 1976 at the
same locality. Stanford and Gaufin (1974) reported nymphs of Isocapnia crinita in subterranean waters
about four meters below and 50 meters laterally from the Tobacco and Flathead River channels in
Montana. It appears that most of the two-year nymphal life cycle is spent in these hyporheic
communities and only when nymphs are mature do they migrate to the surface waters and molt to
adults. If a similar hyporheic community exists below Battle Creek, it would explain the absence of
nymphs from benthic samples taken a month before adults were collected.
Isocapnia missourii Ricker
(Figs. 66, 178)
Isocapnia missourii Ricker 1959; 651. - Gaufin, Ricker, Miner, Milam and Hays 1972: 82. - Baumann,
Gaufin and Surdick 1977: 80.
Isocapnia missourii is known from western United States in Montana and Utah. This first
Saskatchewan (and Canadian) record is from Battle Creek near Reesor Lake (Fig. 178).
Diagnostic Characters. - Average length, females, 14.5 mm (from anterior margin of head to tip of
folded wings), dwarf males, 9.5 mm (from anterior margin of head to end of abdomen). Wings with one
to four costal crossveins before end of subcosta, and none to two (usually one) costal crossveins beyond
it. Male genitalia with tergum 9 with prominent notched or bilobed process; supra-anal process long
and uniformly slender from base to apex. Female subgenital plate entirely sclerotized, without median
recessed sclerotized tongue set off by unsclerotized notch on either side (Fig. 66).
The nymph is unknown.
Ricker (1959) and Gaufin c/ al. (1972) figured the adult genitalia.
Bionomics. - Gaufin c/ al. (1972) reported emergence of adults of Isocapnia missourii from March
to May in Montana. I collected four females at Battle Creek near Reesor Lake on May 19, 1976.
Nymphs were not collected in sweep net samples on this date, nor on March 28, 1976.
Stanford and Gaufin (1974) reported collecting nymphs of Isocapnia missourii as well as of
Isocapnia crinita in subterranean waters below two Montana rivers. Apparently most of the two-year
nymphal life cycle is spent below the main river channel and mature nymphs migrate to the surface
stream to molt to adults. A similar community probably exists below Battle Creek; this would explain
the absence of nymphs from benthic collections.
Family Nemouridae
Nemouridae is the largest family of Plecoptera comprising 373 species occurring in North America,
Eurasia and northern Africa. The family was established by Newman (1853) but Klapalek (1905) later
arranged this group in four families: Nemouridae, Capniidae, Leuctridae, and Taeniopterygidae.
Quaest.Ent., 1979, 15(1)
34
Dosdall& Lehmkuhl
Needham and Claassen(1925) described one subgenus of Nemouridae. Ricker (1952) recognized 12
subgenera in North America, which were later ranked as genera by lilies (1966). Baumann (1975)
revised the family for the world adding three new genera and a new subfamily.
Useful structures in generic definitions include clear, pigmented or banded wings, presence or
absence of cervical gills, membranous or partly sclerotized cerci, and presence or absence of a lobe at
the base of the male ninth sternum. Adults of this family have a slanting crossvein between the costa
and vein Ri of the forewing, the wings at rest folded flat over the body and the last article of the labial
palp subcircular in lateral view. The male supra-anal process is fully sclerotized and anteriorly recurved
in members of most taxa. Either the seventh or eighth sterna of female are produced to form the
subgenital plate. Nymphs are distinguished by number, size and arrangement of body setae. Six genera
of Nemouridae live in Saskatchewan.
Genus Nemoura Latreille
Baumann (1975) observed that species belonging to Nemoura are mainly in more northern regions.
Males are distinguished by terminal hooks on sclerotized cerci (Fig. 79, 80) and females have a large
pregenital plate (on the seventh sternum) (Fig. 89), with lightly sclerotized truncate cerci. Nymphs lack
cervical gills. Four species occur in North America; one lives in Saskatchewan.
Nemoura richer i Jewett
(Fig. 79,80,89,145,146,151,183)
Nemoura rickeri 1971: 190. - Baumann 1975: 21.
Nemoura rickeri has previously been reported from two Alaskan localities only. This first
Saskatchewan and Canadian report (Fig. 183) is from the following localities: Puskwakau River, Jet.
Hwy. 106; Mackay Creek, Jet. Hwy. 2; stream 40 mi. N. of Hudson Bay, Sask., Jet. Hwy. 109; stream at
Promontory Campground, 15 mi. N. of La Ronge, Sask., Jet. Hwy. 2; and the stream at mile 5, Jet.
Hwy. 165.
There is some question whether Nemoura rickeri is conspecific with the Palearctic Nemoura
sahlbergi Morton, but it sould be considered a valid species until a detailed study of all northern species
is completed (R.W. Baumann, pers. comm.).
Diagnostic Characters. - Average length, males, 7.5 mm (from anterior portion of head to tip of
folded wings); females, 9.0 mm. Male cerci strongly sclerotized on outer surface, directed upward;
cereal tips each with outwardly directed, sharply pointed hairy tooth. Male genitalia with tenth tergum
deeply incised medially; epiproct recurved, massive, and rectangular, in dorsal view tip blunt and twice
as long as wide, but in lateral view tip is bluntly pointed; paraprocts simple and broad (Fig. 79, 80).
Female seventh sternum produced over most of sternum eight in form of broad rounded plate, hairy
and heavily sclerotized along border; ninth sternum with small median plate extended anteriorly to
margin of extended seventh sternum (Fig. 89). Since the nymph of this species has not previously been
known, a detailed description follows.
Description. - Total length of mature nymph: 5. 5-8.0 mm.
General color medium to light brown, underside very light brown; thin white eedysial line the only
marking. Head with dark brown subtriangular spots just behind antennae. Antennae light brown, first
three articles dark brown; femora light brown, tibiae and tarsi slightly darker; cerci light brown, first
five articles darker; each cereal article with dark band at base, width about one-fifth length of article.
Head with numerous short stout bristles slightly longer behind eyes and with a few long hairs on
Stoneflies of Saskatchewan
35
anterior region of head.
Pronotum trapezoidal, wider anteriorly, and with short dorsal bristles and pronotal fringe
(Fig. 146); meso- and meta-nota with numerous short stout bristles longer at anterior corners.
Legs with numerous short stout bristles; no tibial fringe but several long hairs along length of tibial
margin; femora in side view about four times as long as wide (Fig. 145).
Abdomen with numerous short stout bristles with marginal bristles longer, up to one-third tergal
length. Tenth abdominal tergum of mature male nymph with distinct pattern (Fig. 148). Cereal bristles
in regular whorls, bristle length between one-quarter and one-half that of cereal article (Fig. 151).
Until comparative study is made of nymphs of Nemoura, it is impossible to identify distinguishing
characters of Nemoura rickeri nymphs.
Bionomics. - In Alaska, Nemoura rickeri specimens were collected on June 30, 1968. Saskatchewan
collection dates for adults are: Puskwakau River, Jet. Hwy. 106, June 10, 1975, and May 30, 1976;
stream 40 mi. N. of Hudson Bay, Sask., June 11, 1975; stream at mile 5, Hwy. 165, May 29, 1975; and
stream at Promontory Campground, 15 mi. N. La Ronge, Sask., June 21, 1976. The secies appears to
have an extended emergence because a large series of nymphs collected at the stream 15 mi. N. of of La
Ronge, Sask. showed a wide variation in wing pad development from very mature to having wing pads
just beginning to form.
Genus Podmosta Ricker
This genus is characterized by absence of cervical gills, simple male cerci and a male supra-anal
process which is short, thick, slightly curved, complex in structure and mostly membranous. The male
tenth tergum has a deep median depression (Fig. 76). The female seventh sternum is unmodified, but
the eighth sternum is produced into a subgenital plate which is straight or excavated posteriorly and
usually slightly notched. There is a distinct darkly sclerotized stripe along the midline of the subgenital
plate (Fig. 90). Five North American species of Podmosta are known and one is found in
Saskatchewan.
Podmosta delicatula (Claassen)
(Fig. 76, 90, 150, 152, 182)
Nemoura delicatula Claassen 1923: 285. - Claassen 1940: 54.
Nemoura (Podmosta) delicatula, Ricker 1952: 43. - Jewett 1959: 33.
Podmosta delicatula, lilies 1966: 219. - Baumann 1973: 92.
Podmosta delicatula is known from the Rocky Mountain areas of North America including British
Columbia, California, Colorado, and Utah. First Saskatchewan records (Fig. 182) include Battle
Creek, Cypress Hills near Reesor Lake; Shuard Creek, 1 1 mi. S. of Piapot, Sask.; and Bear Creek 10
mi. S. of Piapot, Sask.
Diagnostic Characters. - Average length, males,5.5 mm (from anterior margin of head to tip of
folded wings); females, 7.0 mm. Male genitalia with supra-anal process divided into long narrow ventral
portion and short broad dorsal portion, with two thin, forked, sclerotized processes at apex of dorsal
process (Fig. 76). Females with seventh sternum unmodified, and with median sclerotized stripe of
eighth sternum three to four times as long as its greatest width and of fairly uniform width throughout
its length; margin of eighth sternum with tiny median notch (Fig. 90). Since the nymph of this species
has not previously been known, a detailed description follows.
Quaest.Ent., 1979, 15(1)
36
Dosdall& Lehmkuhl
Description. - Total length of mature nymph: 4.0-5. 5 mm.
Color light brown; head darker, especially anterior to lateral ocelli, white markings extend from
ecdysial line to tip of abdominal terga. Head with central light spot between lateral ocelli extending
anteriorly about halfway between lateral ocelli and median ocellus. Antennae golden-brown; scape and
pedicel darker. Cerci very light brown; lower half of each cereal article slightly darker.
Head covered with very short setae; ocelli form an equilateral triangle.
Pronotum narrower than head, trapezoidal, narrowed posteriorly with numerous short bristles;
short setae extended over mesonotum and metanotum. Legs with short bristles; a few long hairs on
tibiae, longest about equal to greatest width of tibiae. No tibial fringe.
Abdomen with numerous very short setae, largest bristles about one-quarter mid-dorsal tergum
length. Each dorsal abdominal segment with distinct row of bristles along its posterior margin
(Fig. 150). Whorls of cereal bristles at base of each cereal article, longest about one-third article length;
dorsal and ventral bristles longer on distal cereal articles, where they may be 0.50-0.75 article length
(Fig. 152).
The distinguishing characters of the nymph of this species cannot be specified now.
Needham and Claassen (1925) and Gaufin et al. (1972) figured the male and female genitalia. The
first illustrations of nymphs are presented in Fig. 150 and 152. Podmosta delicatula is closely related to
the eastern species Podmosta macdunnoughi (Ricker) but differs by not having the male dorsal process
bent near its middle.
Bionomics. - Gaufin e/ al. (1972) state that the species is common in creeks and rivers throughout its
range with adult emergence occurring from April to August. In Saskatchewan, adults were collected
from Battle Creek, Cypress Hills near Reesor Lake on June 3, 1975, June 23, 1975, and July 13, 1975;
from Bear Creek, 10 mi. S. of of Piapot, Sask., on May 17, 1975; and from Shuard Creek, 1 1 mi. S. of
of Piapot, Sask. on June 3, 1975, June 23, 1975 and May 19, 1976. Nymphs were absent from benthic
samples after the spring adult emergence, suggesting a univoltine life history.
Genus Zapada Ricker
Specimens of Zapada are commonly encountered, especially in western North America. Adults
generally emerge in early spring. Males have large angular outer paraproctal lobes and a short broad
epiproct with a well developed dorsal sclerite (Fig. 74). The female seventh sternum is slightly produced
over an unsclerotized eighth sternum (Fig. 86). The four cervical gills are unbranched except in Zapada
cinctipes whose specimens have four to five branches.
Nymphs have whorls of large spines on all femora. Seven North American species are known; one in
Saskatchewan.
Zapada cinctipes (Banks)
(Fig. 10, 11, 14, 74, 86, 183)
Nemoura cinctipes Banks 1897: 21. - Needham and Claassen 1925: 212. - Claassen 1940: 53.
Nemoura (Zapada) cinctipes, Ricker 1952: 57. - Jewett 1959: 35. - Gaufin, Ricker, Miner, Milam and
Hays 1972:41.
Zapada cinctipes, lilies 1966: 250. - Baumann 1975: 31. - Baumann, Gaufin and Surdick 1977: 42.
Zapada cinctipes is common in western Canada and United States from Alaska to California and
Utah and east to Manitoba, South Dakota, Montana and Ohio. This first Saskatchewan report
(Fig. 183) is based on material from the following localities: stream 80 mi. N. of La Ronge, Sask., Jet.
Stoneflies of Saskatchewan
37
Hwy. 102; stream 39 mi. N. of Hudson Bay, Sask., Jet. Hwy. 109; Broad Creek, Jet. Hwy. 104; stream
at Otter Lake, Missinipi, Jet. Hwy. 2; Mackay Creek, Jet. Hwy. 2; Caribou Creek, Jet. Hwy. 106; north
fork of Scarth River, Jet. Hwy. 120; Mackenzie Creek near Bow River, Jet. Hwy. 165; Cub Creek, Jet.
Hwy. 106; Battle Creek, Cypress Hills, near Reesor Lake; and east block of Cypress Hills Provincial
Park, stream one-third mi. W. of Park Gates.
Diagnostic Characters. - Average length, males, 9.5 mm (from anterior part of head to tip of folded
wings); females, 13 mm. Adults with four groups of branched gill remnants, membranous cerci and
wings with dark, transverse bands. Male genitalia with subanal lobes broad with short spine-like
processes on inner margins; supra-anal process recurved, largely membranous and spinulose on outer
halves (Fig. 74). Female seventh sternum produced over entire eighth sternum as a broadly rounded
subgenital plate (Fig. 86).
Nymphs with distinct transverse line of bristles on femora; cervical gills with 4-5 branches.
Needham and Claassen (1925) and Jewett (1959, 1960) figured the adult genitalia. Castle (1939)
described the nymph.
Bionomics. - Emergence of Zapada cinctipes is regulated by an initial photoperiodic response and a
temperature stimulus (Nebeker, 1971c).
Clifford (1969) found that while most adults emerged in early spring after ice started breaking up in
the Bigoray River, Alberta, some nymphs had molted to adults while the stream was still completely
ice-covered. These were found in air spaces between the ice and water. Early-instar nymphs were first
collected in samples in late June, and growth was steady throughout fall and winter. The life history was
univoltine.
At Battle Creek, near Reesor Lake, Cypress Hills, adults and mature nymphs were collected in this
study on March 28, 1976, and adults were still abundant, though no nymphs were found, on
May 19, 1976.
Genus Amphinemura Ris
Adults of this genus are characterized mainly by the presence of cervical gill remnants and
unmodified cerci. Male subanal lobes are divided into two narrow parts which may be recurved forward
alongside the supra-anal process. Usually both inner and outer parts bear few to many heavy spinules
(Fig. 77, 78). The female seventh sternum is produced about halfway over sternum 8 which bears a
median notch and may be produced (Fig. 88). Nymphs have branched prosternal gills (Fig. 12). Ten
North American species are known, and one occurs in Saskatchewan.
Amphinemura /mt/a (Ricker)
(Fig. 12, 77, 78,88, 182)
Nemoura (Amphinemura) linda Ricker 1952: 22.
Amphinemura linda, lilies, 1966: 181. - Baumann, Gaufin and Surdick 1977: 26.
Amphinemura linda is a common species in Canada and northern United States. Previous records
include British Columbia, Alberta, Manitoba, Quebec, Ontario, Labrador, and Michigan. This first
Saskatchewan record fills in the central Canadian distributional gap for the species. Specimens have
been collected in the following localities (Fig. 183): Mistohay Creek, Jet. Hwy. 226; Nemeiben River,
Jet. Hwy. 2; stream 10 mi. E. of Squaw Rapids Power Station; stream 80 mi. N. of of La Ronge, Sask.,
Jet. Hwy. 102; stream 85 mi. N. of of La Ronge, Sask., Jet. Hwy. 102, Mackay Creek, Jet. Hwy. 2;
Nipekamew River, Jet. Hwy. 165, stream 37 mi. N. of Green Lake Sask., Jet. Hwy. 155; stream at mile
Quaest. Ent., 1979, 15(1)
38
Dosdall& Lehmkuhl
98, Jet. Hwy. 155; creek between Bedard Creek and Bisset Creek, Jet. Hwy. 106; Puskwakau River, Jet.
Hwy. 106; creek at mile 135, Jet. Hwy. 106; Swan River, Jet. Hwy. 8; and the stream at mile 34 on the
Wollaston Lake Road.
Diagnostic Characters. - Average length, males 6.5 mm (from anterior margin of head to tip of
folded wings); females, 8.5 mm. Male genitalia with anterior lobe on sternum 9 and posterior
projection; subanal lobes double with both branches bearing spines, most specimens with two spines on
outer branch, and four to five on inner branch (Fig. 77); subanal lobes not recurved forward, and
usually seen only in ventral view (Fig. 78); supra-anal process mainly membranous with sclerotized
sides. Female seventh sternum produced halfway over sternum 8, and subgenital plate sclerotized and
sinuate (Fig. 88).
Nymphs with long and pointed pronotal bristles; cereal bristles at least three-quarters length of each
cereal article; and large femoral and tibial bristles darker in color than rest of leg.
Ricker (1952) figured the adult genitalia, and Harper and Hynes (197 Id) illustrated the nymph.
Bionomics. - Harper (1973b) noted that Amphinemura linda was an autumnal species in which the
first adults were found in late August and adult emergence was extended throughout September. The
emergence pattern of males and females was similar, but males emerged earlier than females. The
intensity of emergence was about the same throughout. Oviposition occurred mainly at midday and
early afternoon of late September and early October.
The life history of Amphinemura linda was shown to be univoltine (Harper, 1973b). Eggs hatched in
the laboratory under simulated stream conditions showed a short initial development in October,
embryonic diapause for five months in winter and resumed development in March. Hatching occurred
in April. Nymphal growth was continuous throughout the summer and ended just before emergence of
adults. In one river with warm winter water temperatures, embryonic diapause lasted only two months
although emergence occurred about the same time. This slower summer growth was due to a cooler
summer stream temperature.
In Saskatchewan, Amphinemura linda emerges in July and August. Adults were collected from the
creek between Bedard Creek and Bisset Creek, Jet. Hwy. 106 on July 2, 1975, from Puskwakau River,
Jet. Hwy. 106 on July 17, 1975, from the stream at mile 98 near Ile-a-la-Crosse, Sask. on July 10, 1975,
and from Mistohay Creek, Jet. Hwy. 226 on August 12, 1975.
Genus Shipsa Ricker
Adults of this genus have banded wings, unmodified cerci and lack cervical gills. The male tenth
tergum is produced into long terminal projections with one on each side of the epiproct. The epiproct is
modified by having the ventral sclerite extending to the dorsal surface and it terminates in a prominent
forcep-shaped structure (Fig. 75). The female seventh sternum is produced and laterally excavated with
the produced part of sternum 7 attached basally to sternum 8. The eighth sternum is not produced
(Fig. 87). Nymphs have a fringe of long hairs along the posterior margins of the tibiae.
Shipsa rotunda (Claassen)
(Fig. 75,87, 183)
Nemoura rotunda Claassen 1923: 290. - Needham and Claassen 1925: 219. - Harden and Mickel 1952:
17.
Nemoura (Shipsa) rotunda, Ricker 1952: 50.
Shipsa rotunda, lilies 1966: 247.
Stoneflies of Saskatchewan
39
Shipsa rotunda, the single known species of this genus, has been reported from Alaska,
Saskatchewan, Ontario, the Maritimes and eastern United States. Ricker (1944) reported the species
from Black Lake, Saskatchewan and additional collection records include (Fig. 183): Little Red River,
Prince Albert, Sask.; Nipekamew River, Jet. Hwy. 165; Mackay Creek, Jet. Hwy. 2; stream at south
end of Wollaston Lake, Jet. Hwy. 105; Torch River, Jet. Hwy. 106; North Saskatchewan River, Jet.
Hwy. 5; and North Saskatchewan River at Prince Albert, Sask.
Diagnostic Characters. - Average length, males, 7.5 mm (from anterior part of head to tip of folded
wings); females, 10.5 mm. Since this is the only known species of Shipsa, generic characters also define
specific characters.
Needham and Claassen (1925) figured the adult genitalia; Harden and Mickel (1952) illustrated
nymphal characters.
Bionomics. - Harper (1973b) reported collecting early-instar nymphs of Shipsa rotunda in
November in a southern Ontario stream. Growth was rapid and without arrest throughout the winter
even though the stream had a thick ice cover. Nymphal development was complete by mid-April, about
one week before adult emergence.
In Saskatchewan, Shipsa rotunda was most abundant at the North Saskatchewan River at Hwy. 5
(Borden Bridge). Mature nymphs were collected on April 25, 1974, May 2, 1974, and May 4, 1974.
Adults were collected on May 4, 1974.
Genus Malenka Ricker
This common western North American genus is characterized by the presence of mesobasal lobes on
male cerci (Fig. 73), a nipple-like ventral projection on the female seventh sternum and a notch on the
posterior margin of the female eighth sternum (Fig. 85). Nymphs have four branched cervical gills.
Eleven species are known in North America; one occurs in Saskatchewan.
Malenka californica (Claassen)
(Fig. 76,90, 147, 149, 153, 182)
Nemoura californica Claassen 1923: 284. -Frison 1942a: 261.
Nemoura lobata, Frison 1936: 260.
Nemoura (Malenka) californica, Ricker 1952: 33. - Jewett 1959: 32. -Gaufin, Ricker, Miner, Milam
and Hays 1972: 32.
Malenka californica,l\\iQS 1966: 191.
This species has been recorded mainly from the cordillera including New Mexico and California
north to British Columbia and western Alberta. This first Saskatchewan report (Fig. 183) is from the
following localities: Broad Creek, Jet. Hwy. 104; Mackenzie Creek, Jet. Hwy. 165; Waskwei River, Jet.
Hwy. 109; and Scarth River, Jet. Hwy. 120.
Diagnostic Characters. - Average length, males, 8.0 mm (from anterior margin of head to tip of
folded wings); females 8.5 mm. Male cercus with mesobasal lobe sclerotized, sharply pointed, and
directed inward and backward. Male genitalia with outer part of subanal lobes broad and with notch on
mesal margin (Fig. 73). Female seventh sternum with erect nipple-like protuberance on produced
portion with its base anterior to hind margin of sternum; sternum 8 notched about halfway across and
with no margin of extra sclerotization (Fig. 85). The nymph is described below for the first time.
Description. -Total length of mature nymph: 5.0-6. 5 mm.
Color medium brown; antennae a little darker; legs brown; bristles on legs slightly darker than
Quaest.Ent., 1979, 15(1)
40
Dosdall & Lehmkuhl
ground color of leg; cerci pale.
Short bristles covering head capsule, those behind eyes longer and stout. Antennal bristles large
near base but very small and short near tip.
Pronotum rectangular, covered with short bristles and hairs. Pronotal fringe well-defined consisting
of long pointed bristles which are long on corners but very short along medial margin (Fig. 147). Meso-
and metanota covered with medium-sized bristles especially on anterior corners. Legs with long stout
bristles, the longest femoral bristles equal to about two-thirds of greatest femoral width; no tibial fringe
of hairs but profuse tibial bristles.
Prosternal gills in four tufts; each tuft of 6-8 filaments forming a whorl about a central axis.
Abdomen with numerous long bristles, longest about 1.3 times as long as mid-dorsal length of
corresponding tergum(Fig. 149). Whorls of cereal bristles regular; length of bristles equal to total
length of cereal article (Fig. 153).
The nymph of this species is almost identical to that of Amphinemura linda, but if the two are
compared there are several distinguishing features. Amphinemura linda has an overall orange-red
color, while Malenka californica is brown. The body bristles of Malenka californica are much darker
than the body; in Amphinemura linda, the bristles, especially the leg bristles, are only slightly darker
than the body. The two species are also temporally separated. Amphinemura linda is a mid- to
latesummer species with mature nymphs occurring in mid-July. However, Malenka californica is a fall
species with mature nymphs occurring in late August.
Needham and Claassen(1925), Jewett (1959) and Gaufin et al. (1972) figured the male and female
genitalia. Nymphal characters are illustrated in Fig. 147, 149 and 153.
Bionomics. - Malenka californica is found in small streams, creeks and small springs. Adults
emerge in late summer or fall (Baumann, 1975), as soon as there is a sharp decrease of stream
temperature (Nebeker, 1971c).
In Saskatchewan, adults have been collected on August 8, 1975 at Mackenzie Creek, Jet. Hwy. 165,
September 1, 1975 at Waskwei River, Jet. Hwy. 109, and on September 25, 1975 at Scarth River, Jet.
Hwy. 120. Adults of this species have been found in stomachs of brook trout at Scarth River
(D. Larson, pers. comm.).
Nymphs were not present in samples taken at Mackenzie Creek, Jet. Hwy. 165 in April or May of
1976. However, early instar nymphs were collected at the same locality on June 25, 1975. There is
probably an embryonic diapause over the winter and early spring, with the main nymphal development
occurring throughout June, July, and early August.
Family Leuctridae
Leuctridae are characterized by having wings rolled around the body at rest, cu-m and intercubital
crossveins in the forewings, a forked cubitus and simple media in the hindwings of most genera, single
articled cerci (simple in most genera), an inconspicuous male epiproct and absence of gills. Male
leuctrids are unique by having a long median process or subanal probe between the paraprocts (Ricker
and Ross, 1969). Nymphs have the first tarsal article longer than the second, hindwing pads subparallel
to the body axis, and the first six or less abdominal segments divided into tergum and sternum by a
membranous fold. The family occurs in North America, Eurasia and northern Africa.
At present, the nomenclature of Leuctridae is somewhat confused. Hanson (1941) proposed the
genus name Paraleuctra for seven North American species previously in the genus Leuctra. Frison
(1942a), Ricker (1943, 1952), Jewett (1956, 1959, 1960) and Gaufin (1972) considered Paraleuctra
a subgenus of Leuctra, but lilies (1966) maintained the generic status of Paraleuctra, this was accepted
Stoneflies of Saskatchewan
41
by Ricker and Ross (1969) and Hitchcock (1974). I follow the nomenclature of lilies (1966). Both
Leuctra and Paraleuctra occur in the province but the other North American genera of this family,
Perlomyia, Megaleuctra and Zealeuctra, were not found.
Genus Leuctra Stephens
Adults of Leuctra differ from the closely related genera Paraleuctra and Zealeuctra by having the
m-cu crossvein in the hindwing proximal to the dichotomy of Cui, and Rs and M originating from
different points on the radius (Fig. 36). Male cerci are unmodified. The number and shape of processes
on the male seventh and eighth terga are often used for species determinations. Nymphs differ from
other genera because the first four abdominal segments are divided into tergum and sternum by a
membranous fold (Fig. 27), and labial palpi extend well beyond the paraglossae (Fig. 21).
Members of this genus seem to prefer smaller streams (Hitchcock, 1974). The adults are small,
brownish and inconspicuous. Of 22 North American species, one species of Leuctra was collected in
Saskatchewan, Leuctra ferruginea.
Leuctra ferruginea (Walker)
(Fig. 21, 27, 36, 84, 92, 184)
Nemoura ferruginea Walker 1852: 183.
Leuctra decepta, Claassen 1923: 260. -Claassen 1940: 77. -Frison 1942a: 257. - Harden and Mickel
1952:21.
Leuctra ferruginea, lilies 1966: 87. - Hitchcock 1974: 76.
Leuctra ferruginea ranges from eastern Canada (Ontario, Quebec, Newfoundland, Nova Scotia,
and New Brunswick), eastern United States (Maine to Florida) to Illinois and Minnesota in the west.
The new Saskatchewan record is the most western extension of the species range. Specimens were
collected from the following localities (Fig. 1 83): stream 85 mi. N. of La Ronge, Sask., Jet. Hwy. 102;
Broad Creek, Jet. Hwy. 104; stream at mile 30 on the Wollaston Lake Road; Mackay Creek, Jet. Hwy.
2; Low Creek, Jet. Hwy. 104 and Waddy River, Jet. Hwy. 102.
Diagnostic Characters. - Average length, males, 6 mm (from anterior margin of head to tip of
folded wings); females, 8.5 mm. Male genitalia with subanal lobes slender and considerably longer than
specilla; supra-anal lobe broadly rounded and cerci unmodified, small raised process on eighth
abdominal tergum in most specimens, but in some individuals missing (Fig. 84). Some specimens with
dark sclerotization on tergum 7 which resembles a process but not raised. Female sternum 8 produced
into broad and deeply notched subgenital plate which reaches nearly halfway across sternum 9, two
rounded lobes of subgenital plate separated by membranous tissue (Fig. 92).
Nymphs with all abdominal terga covered with short stout bristles but no stout bristles on
abdominal sterna.
Needham and Claassen (1925), Frison (1942a) and Hitchcock (1974) figured the adult genitalia;
Claassen (1931) and Harper and Hynes (197 la) figured the nymph.
Bionomics. - Harper and Hynes (1971a) noted that Leuctra ferruginea occurs predominant-
ly in small cold streams although it may be found in a wide variety of streams and even small warm
rivers. Harper (1973b) found that adult emergence in southern Ontario was extended, lasting from early
May to September of 1968, although the majority of individuals emerged in late May and early June.
Males emerged slightly earlier than females. Adult feeding began soon after emergence but mating did
not occur for a few days. Oviposition was observed in the field from mid-June until mid-October. Each
Quaest.Ent., 1979, 15(1)
42
Dosdall & Lehmkuhl
egg batch contained an average of 204 eggs. Females produced an average of 26 egg batches.
Eggs maintained at a constant temperature of 10 C and photoperiod of 12 hours light: 12 hours
darkness hatched in 49 days (Harper, 1973b). There was no evidence of embryonic diapause. Cooler
temperatures lengthened embryonic development; therefore, eggs laid in mid-October would not hatch
until spring.
Nymphal growth and emergence appeared to depend upon water temperatures. Only newly hatched
nymphs were collected from October to May in streams coldest in summer and warmest in winter.
Nymphs hatching in October grew rapidly in fall but little growth occurred during winter. Some of
these nymphs emerged in late summer, but the majority of the population was only half grown by the
end of the summer and did not emerge until the following year. Growth continued until October,
slowed for winter, and resumed in March. Most nymphs matured in May and June. The number of
individuals which completed their nymphal growth in one year was very small; most required two years.
A semivoltine growth pattern was still apparent in the warmest stream where Leuctra ferruginea was
common, but in this case a large part of the population was univoltine. Adult emergence was short and
late (most mature nymphs were collected in July).
It is not known whether the life history of this species is univoltine or semivoltine in Saskatchewan.
Adults were collected on July 4, 14, and August 7 of 1974 from the stream 85 mi. N. of La Ronge, Sask.,
Jet. Hwy. 102. This was the only Saskatchewan stream sampled where the species was abundant.
Genus Paraleuctra Hanson
Paraleuctra differs from other leuctrid genera in having the m-cu crossvein reach Cui after it has
divided (Fig. 34), heavily sclerotized male cerci (Fig. 83), the male ninth tergum entire and the female
dorsum with a longitudinal sclerotized stripe on either side. Nymphs have labial palpi extending to the
tip of the paraglossae, the first six abdominal segments divided into tergum and sternum by a
membranous fold, and the subanal lobes not partly fused but separate. Eight species are known in
North America and one occurs in Saskatchewan.
Paraleuctra vershina (Gaufin and Ricker)
(Fig. 34,81-83,91, 184)
Leuctra occidentalis, Needham and Claassen 1925: 231.
Paraleuctra occidentalis, Hanson 1941: 57. - Jewett 1959: 39 and 1960: 141. -(in part) lilies 1966: 1 14.
Paraleuctra occidentalis auct,, Hanson 1962: 135.
Paraleuctra vershina Gaufin and Ricker 1974: 285.
Paraleuctra vershina ranges from the cordilleran regions of Canada and the U.S. (British Columbia,
Alberta, California and Washington) and east to Colorado, Montana and Utah. This, the first
Saskatchewan record, is from Battle Creek near Reesor Lake (Fig. 184).
Diagnostic Characters. - Average length, males 6.0 mm (from anterior margin of head to tip of
folded wings); females, 8.0 mm. Male genitalia with tergum 10 partly cleft; supra-anal lobe triangular
and bearing slender, recurved, whip-like process at tip (Fig. 81, 82); cerci modified into chitinous armed
processes, bulbous at base and each bearing sharp tooth above and below (Fig. 83); subanal lobes
modified into probe with each lobe receding into ninth segment. Female subgenital plate produced into
two lobes lined with long hairs and separated by a broad notch (Fig. 91). Nymphs of this species are
unknown.
Needham and Claassen (1925) illustrated the adult genitalia. Hanson (1962) figured the male cerci
Stoneflies of Saskatchewan
43
showing the characters which distinguish Paraleuctra vershina from other Paraleuctra.
Bionomics. - Little is known of the biology of this species. In Battle Creek adult specimens were
collected on June 23, July 13, and July 20, 1975.
Family Chloroperlidae
Chloroperlidae are small to medium size stoneflies charcterized by having paraglossae much longer
than glossae, gills absent, three ocelli, reduced wing venation (only two anal veins) and a small or absent
anal area in the hind wing. The family is found in North America, eastern and western Asia and Europe.
Male genitalia have unmodified paraprocts and a well developed epiproct. The male tenth tergum is
cleft and lobes are absent from all male sterna. Female subgenital plates are variable.
Nymphs lack anal gills, have cerci about three-quarters the abdominal length, and have hindwing
pads subparallel to the axis of the body. The last article of each nymphal maxillary palp is abruptly
thinner than previous segments.
In North America, Chloroperlidae presently consists of two subfamilies, ten genera, and about 56
species. The subfamily Paraperlinae of western North America is comprised of the genera Utaperla
Ricker and Kathroperla Banks which are monotypic and the genus Paraperla Banks with two species.
The subfamily Chloroperlinae, of eastern and western North America, originally consisted of three
genera: Hastaperla Ricker, Chloroperla Newman and Alloperla Banks. The five subgenera of Alloperla
have recently been assigned full generic status (lilies, 1966).
Three genera of Chloroperlinae are known in Saskatchewan: Hastaperla, Suwallia and Triznaka.
Genus Hastaperla Ricker
Adults of Hastaperla are distinguished from other chloroperlid genera by the lack of a fold
separating the anal area from the remainder of the hindwing. The anal cell of the forewing gives rise to a
single unbranched vein (Fig. 32). Nymphs of Hastaperla cannot be distinguished from Chloroperla;
both have the inner margins of the hindwing pads straight and mature larvae are 7 mm in length or less.
Of the three species of Hastaperla known in North America, one lives in Saskatchewan.
Hastaperla brevis (Banks)
(Fig. 32, 93,96, 185)
Chloroperla brevis Banks 1895: 314.
Hastaperla calcarea, Ricker 1935: 200.
Hastaperla brevis, Claassen 1940: 197. - Frison 1942a: 340. - Harden and Mickel 1952: 62. - Ricker
1964: 54. - Hitchcock 1974: 168.
Hastaperla brevis is the most widespread species of Chloroperlidae (Gaufin, 1964), ranging from
Nova Scotia through New England, south to Georgia, and west to Oklahoma and Manitoba. Though
not previously reported in Saskatchewan new collection records (Fig. 185) for the province include:
South Saskatchewan River at the ferry North of Birch Hills, Sask.; Weyakwin River, Jet. Hwy. 2;
Ballantyne River, Jet. Hwy. 106; Mackenzie Creek, Jet. Hwy. 165; Bow River, Jet. Hwy. 165;
Nipekamew River, Jet. Hwy. 165; Red Deer River, Jet. Hwy. 8; Waskwei River, Jet. Hwy. 109;
McDougal Creek, Jet. Hwy. 120; Puskwakau River, Jet. Hwy. 106; Torch River, Jet. Hwy. 106; stream
at mile 83, Jet. Hwy. 106, Waddy River, Jet. Hwy. 102; Overflowing River, Jet. Hwy. 109; Crean River,
Jet. Hwy. 2; Pine Creek, Jet. Hwy. 165; Cluff Creek near Cluff Lake; Churchill River at Wintego Lake
Quaest.Ent., 1979, 15(1)
44
Dosdall & Lehmkuhl
rapids; Mackay Creek, Jet. Hwy. 2; Fond du Lac River at Black Lake; and Montreal River Jet. Hwy. 2.
Diagnostic Characters. - Average length, males, 6.5 mm (from anterior margin of head to tip of
folded wings); females, 9.0 mm. Male genitalia with ninth abdominal segment produced ventrad,
rounded and hairy behind, with penis lying beneath; curved chitinous process on each side of penis
joined at posterior end to form blunt, prong-like tip (may be conspicuous only in cleared-mount); male
epiproct rounded (Fig. 93). Female subgenital plate produced over most of ninth sternum, triangular
and rounded at tip (Fig. 96).
Nymphal hindwing pads subparallel to axis of body and inner wing pad margins straight; cerci
three-quarters abdominal length; mature larvae 7 mm in length or less (from anterior tip of head to end
of abdomen).
Adult genitalia have been figured by Needham and Claassen (1925), Ricker (1935), Frison (1935,
1942a) and Gaufin^/ (1966); nymphs were illustrated by Frison (1935). Adults differ from the closely
related species, Hastaperla orpha (Frison) by lacking a dorsal abdominal stripe.
Bionomics. - Harper and Magnin(1969) studied the life cycle of Hastaperla brevis in Quebec. Adult
emergence began in June, with the period of time for egg laying and hatching being relatively short.
Nymphs were first collected in October, and growth was rapid during fall and early winter but slow in
January. In May growth continued and maximum size was reached. It was concluded that growth was
not closely synchronized since nymphs in various stages of maturity were collected in each sample.
Though Hitchcock (1974) stated that the nymphs were herbivorous. Harper and Magnin (1969)
contended that nymphs were at least partly carnivorous.
Genus Suwallia Ricker
Males of this genus are characterized by having slender, curved finger-like processes developed from
the basal article of the cerci. The male supra-anal body is a membranous lobe with a small hairy process
at its tip (Fig. 95). Adults have a small anal lobe on the hindwing, a dark dorsal stripe on the abdomen,
and dark U-marks on the meso- and metanota. Few nymphs of Suwallia have been described, so
nymphal generic characters are not defined.
Suwallia lineosa (Banks)
(Fig. 95,98, 163, 165-167)
Alloperla lineosa Banks 1918: 7. - Claassen 1940: 186. - Ricker 1964: 68.
Alloperla (Suwallia) lineosa, Ricker 1943: 139. - Jewett 1955: 151. -Gaufin 1964: 42.
Suwallia lineosa, lilies 1966: 449.
Suwallia lineosa occurs in the Rocky Mountains of western Canada (British Columiba and Alberta)
and westward to the Cascade and Wallowa Mountains (Colorado, Montana, Oregon, Washington,
Wyoming). Though not previously reported in Saskatchewan, specimens were collected in Cypress
Hills at Battle Creek near Reesor Lake (Fig. 185). It was not collected further downstream in Battle
Creek near Consul, Sask.
Diagnostic Characters. - Average length, males, 7.8 mm (from anterior margin of head to tip of
folded wings); females, 8.9 mm. Adult without head or pronotal markings except for ocellar rings and
faint stripe in centre of pronotum (Fig. 163). Male genitalia with aedeagus with single V-shaped patch
of sclerotized spinules; cerci with slender, curved, finger-like processes developed from basal article;
supra-anal body membranous, hairy and weakly sclerotized (Fig. 95). Female subgenital plate
emarginate; produced completely over sternum 9 and nearly all of sternum 10 (Fig. 98). A first
Stoneflies of Saskatchewan
45
description of the mature nymph is given below.
Description. - Length from apex of head to end of abdomen in mature nymphs, 7.8 mm.
General color of head, thorax and abdomen pale brown; head and abdomen without conspicuous
bands, spots or stripes of contrasting colors. Abdomen pale ventrally. Prothorax suboval, at least twice
as long as broad, and dark brown on margin but may be lighter laterally. Mesothorax margined with
dark brown at anterior end.
Wing pads with lateral margins broadly rounded.
Head with three ocelli forming nearly equilateral triangle; distance between ocelli about same as
distance to inner margin of compound eye. Labium, maxillae and mandibles as Fig. 165-167. No
occipital ridge.
Jewett (1955) figured the male genitalia, and Needham and Claassen(1925) and Gaufinc? a/. (1972)
figured the female genitalia. Nymphal mouthparts are figured for the first time in Fig. 165-167.
Bionomics. - The biology of Suwallia lineosa is poorly known. In Battle Creek, adult emergence
began early in July, 1975 and lasted at least three weeks. Several stages of maturity of nymphs from one
sample were observed, indicating little synchrony of emergence. By the end of July nymphs were absent
from benthic collections, suggesting a one-year life cycle since all had emerged.
Genus Triznaka Ricker
This genus was originally proposed as a subgenus of AUoperla (Ricker, 1952). Species of Triznaka
have the male supra-anal process short and lying along the surface of and fused to the tenth tergum. It is
usually in a slight depression but never in a deep groove. The tip of the process is directed upward and
may be curved anterad, but there is no apparatus of erection (Fig. 94). Adults also have a black stripe
on the abdominal terga, and a small anal lobe on the hindwing. Nymphal generic characters are not
defined since few of the nymphs have been described.
Triznaka signal a (Banks)
(Fig. 94, 97, 159-162, 164, 185)
Chloroperla signata Banks 1895: 314.
AUoperla signata,C\2i?i^^Qn 1940: 188.
AUoperla (Triznaka) signata, Ricker 1952: 186. -Gaufin 1964: 47. -Gaufin, Ricker, Miner, Milam and
Hays 1972: 146.
Triznaka signata, IWics 1966: 154. -Baumann, Gaufin and Surdick 1977: 184.
Triznaka signata ranges from Wasington and Montana south to Colorado and Utah. New
collection records include Broad Creek and Mistohay Creek in northeastern Saskatchewan (Fig. 185).
Diagnostic Characters. - Average length, males 7.5 mm (from anterior margin of head to tip of
folded wings); females, 9.8 mm. Adult head with anterior mark as long as broad and produced in
midline to median ocellus in most specimens; ocellar triangle dark; pronotal rugosities only lightly
pigmented; median pronotal mark broadly T-shaped, little produced laterally rearward (Fig. 164).
Female subgenital plate extended over ninth sternum and broadly rounded posteriorly (Fig 97). A first
description of the nymph is given below.
Description. - Length from apex of head to end of abdomen of mature nymphs, 8.0 mm.
General color of head, thorax and abdomen pale brown. Head with small light spot in front of
anterior ocellus, and small light area at base of ocellar triangle extended posteriorly to end of head.
Each compound eye connected to lateral ocelli by a light area. Prothorax suboval, at least twice as long
Quaest.Ent., 1979, 15(1)
46
Dosdall & Lehmkuhl
as wide with a lighter subcircular area in center, margin of dark brown, but broken laterally and at
midline. Meso- and metathoracic regions patterned. Central median stripe on dorsal abdomen with two
light spots on either side of stripe per segment. Abdomen pale ventrally; lateral margins of wing pads
broadly rounded (Fig. 159).
Head with three ocelli forming a nearly equilateral triangle; distance between ocelli about same as
distance to inner margin of compound eye. Labium, maxillae and mandibles as in Fig. 160-162. No
occipital ridge.
Needham and Claassen (1925) and Gaufine/ a?/. (1972) illustrated the male and female genitalia; first
figures of the nymph are presented in Fig. 159-162.
Bionomics. - In Saskatchewan, mature nymphs were collected near the end of May and adults
emerged in June of 1975 at Broad Creek. Nymphs of various stages of maturity could be collected in
one sample indicating little synchrony of emergence. By July, nymphs were not present, indicating that
all had emerged and a univoltine life history is probable.
Family Perlidae
Perlidae have profusely branched gills at lower angles of the thorax, the male epiproct reduced and
inconspicuous and the Cu-A crossvein of the forewing located either in the anal cell or very close to it.
Both subfamilies of Perlidae, Acroneuriinae and Perlinae occur in Saskatchewan. The genital hooks
of male acroneuriines are modified paraprocts. The subfamily occurs in the Americas and in
southeastern Asia. Genital hooks of perlines are outgrowths of a cleft tenth tergum and paraprocts are
little modified. This subfamily occurs in Africa, Eurasia and North America. Saskatchewan Perlidae,
Paragnetina media and Claassenia sabulosa are in the subfamily Perlinae: Acroneuria abnormis,
Acroneuria lycorias, Hesperoperla pacifica and Perlesta placida belong to the subfamily Acroneuriinae.
Genus Paragnetina Klapalek
Paragnetina members are characterized by the absence of anal gills, three ocelli, and a Y-shaped
median ridge on the prosternum and mesosternum. Males are distinguished from those of other genera
by having genital hooks arising from the side of the cleft tenth tergum and extending to the hind margin
of the ninth tergum. The female subgenital plate is slightly produced past the hind margin of the eighth
sternum (Fig 105). Nymphs have a transverse line of spinules on the occiput (Fig. 5). Five species of
Paragnetina are presently known in North America and one occurs in Saskatchewan.
Paragnetina media (Walker)
(Fig. 5, 105, 187)
Per la media Walker 1852: 145.
Acroneuria salve lini, Ricker 1935: 26.
Togoper la media, Claassen 1940: 150.
Paragnetina media, Ricker 1949: 287, and 1964: 59. - Harden and Mickel 1952: 51.
Paragnetina media occurs in the Maritimes to northern Ontario, central Quebec, Manitoba,
northern Saskatchewan, northern Illinois, southern Michigan, central Pennsylvania and southern New
England. A North American distribution map for the species was given by Ricker (1964) and the
Saskatchewan distribution is illustrated in Fig. 187. Ricker (1944) reported the species from Wapus
River in the Reindeer Lake region, and additional Saskatchewan collection localities include Mackay
Stoneflies of Saskatchewan
47
Creek, Jet. Hwy. 2; Weyakwin River, Jet. Hwy. 2; Churchill River, Jet. Hwy. 2; stream 85 mi. N. of La
Ronge, Sask., Jet. Hwy. 102; Bow River Jet. Hwy. 165; and Caribou Creek, Jet. Hwy. 106.
Diagnostic Characters. - Average length, males, 18 mm (from most anterior portion of head to tip
of folded wings); females, 32 mm. Male fifth tergum produced very shallowly, broady excavated or
practically straight. Male genitalia with hooks rounded at tip; male sternum 9 without raised knob or
“hammer”. Female subgenital plate little produced with base of median notch in line with sides of hind
margin of sternum 8 (Fig. 105). Adult pronotum with black margins and black median stripe.
Nymphal abdominal terga uniformly brown, some specimens with faint median line (Fig. 5).
Needham and Claassen ( 1925), Frison (1935) and Hitchcock (1974) illustrated the adult genitalia;
Claassen (1931) and Frison (1935) figured the nymph.
Bionomics. - Ricker (1949) noted that Paragnetina media was absent from colder trout streams and
rivers of that region. Nymphs are usually found under stones in the larger streams but can also be
collected in lakes and ponds (Claassen, 1931).
Though Claassen (1931) stated that nymphs of Paragnetina media probably require three years to
complete their life cycle. Tarter and Krumholz (1971) found that in Kentucky Paragnetina media
required only two years to complete the life cycle. In Saskatchewan, two size classes can be collected at
one time, also suggesting a two year life cycle. For example, at Mackay Creek on June 5, 1974 seven
mature nymphs with well-developed wing pads ranged from 20-23 mm in length (average length,
20.5 mm) and were near the end of their life cycle since the species emerged in the first week of July in
both 1974 and 1975. Four specimens of the immature size class of nymphs, characterized by having
poorly developed wing pads, ranged from 8. 6-9. 8 mm in length (average length, 9.5 mm). No very small
nymphs were collected in mid-August and either the eggs from adults laid in July had not yet hatched or
nymphs were yet too small to be collected in a sweep net. A significant growth of both hatchling
nymphs and year-old nymphs must occur between fall and spring.
Harper and Pilon (1970) noted that the adult emergence period of Paragnetina media lasted from 5
to 25 days.
Based on dissected nymphs, the females produce an average of 802 eggs (Tarter and Krumholz,
1971) and the egg incubation period is 30 days at room temperature (Heiman and Knight, 1970).
Steffan (1965) described larvae of the new genus and species of Chironomidae (Diptera),
Plecopteracoluthus downesi, which live phoretically on nymphal Paragnetina media in a gelatinous
case and feed on detritus caught in the long hairs of the stonefly. Of the Perlidae carrying chironomids,
six percent were Paragnetina media. In Saskatchewan, phoretic Chironomidae have not been found on
Paragnetina media though they are commonly encountered on nymphs of Acroneuria lycorias.
Adults of Paragnetina media are rarely encountered in the field. Sweeping the vegetation along
stream banks and searching under rocks and under bridges yielded no specimens. This was also
observed by Tarter and Krumholz (1971). The only adults examined in this study were from reared
specimens. Females reared in laboratory aquarium cages where no males were present were found to
produce egg masses even though copulation could not have occurred. Harper (1973a) incubated eggs
laid by a virgin female and found that 13 percent hatched. This ability of eggs to develop
parthenogenetically was thought to act as a safeguard in large carnivorous species, which are rarely
abundant in their habitat, because the chances of meeting mates may be small. Frison (1935) noted that
adults of Paragnetina media are diurnal and that mating occurs during the day.
Nymphs are carnivorous, feeding primarily on Ephemeroptera, Trichoptera and Diptera, but adults
do not feed. Larvae are food for fish and crayfish (Tarter and Krumholz, 1971).
Quaest . Ent ., 1979, 15(1)
48
Dosdall& Lehmkuhl
Genus Claassenia Wu
Males of Claassenia are distinct from other genera of Perlidae by having genital hooks darkly
sclerotized at their tips and arising from lateral angles of tergum 10 (Fig. 8). Males also have a raised
knob or “hammer” on sternum 9. The female subgenital plate is broadly rounded, usually with a
shallow median recession and is little produced over sternum 9. It has with a row of spinules along its
posterior margin (Fig 106). Nymphs resemble the eastern genus Phasganophora by having anal gills
and an occipital ridge composed of closely set spinules, but are distinguished by having dorsal
abdominal segments almost wholly brown and a banded pattern on the legs (Fig. 4).
Claassenia sabulosa, the single North American species of this genus, occurs in Saskatchewan.
Claassenia sabulosa (Banks)
(Fig. 4,8, 106, 187)
Per la sabulosa Banks 1900: 242.
Claassenia languida, Needham and Claassen 1925: 100. -Claassen 1940: 181.
Claassenia arctica, Frison 1942a: 285.
Claassenia sabulosa, Ricker 1952: 190. - Jewett 1959: 90. - Gaufin, Ricker, Miner, Milam and Hays
1972: 156.
Claassenia sabulosa ranges from the cordilleran region of British Columbia and Alberta through the
Cascade and Rocky Mountains to New Mexico and east to northern Manitoba and northern Ontario.
Ricker (1964) provided a North American distribution map for the species. Claassenia sabulosa has not
previously been reported from Saskatchewan, and new collection records include McDougal Creek,
Jet. Hwy. 120; Creek at Mile 145, Jet. Hwy. 105; Overflowing River, Jet. Hwy. 109; Cluff Creek near
Cluff Lake; and the South Saskatchewan River at the ferry N. of Lemsford, Sask. (Fig. 187). Though
the species is abundant at McDougal Creek, it has rarely been collected in the South Saskatchewan
River.
Diagnostic Characters. - Average length, brachypterous males, 19.6 mm (from anterior margin of
head to end of abdomen); females, 31.4 mm (from anterior margin of head to wing tips). Since
Claassenia is monotypic, generic characters also define specific characters.
Ricker (1938), Gaufin et al. (1966) and Gaufin et al. (1972) figured the adult genitalia; Claassen
(1931) and Frison (1942a) figured the nymph.
Bionomics. - The habitat of Claassenia sabulosa is under stones in swift riffle areas where food is
abundant (Richardson and Gaufin, 1971).
Three size classes of nymphs present in the same sample suggest a three-year life cycle. For example,
on May 28, 1975 at McDougal Creek three nearly mature male nymphs with well developed wing pads
averaged 23.0 mm and two females averaged 32.5 mm in length. Fourteen specimens in the next size
class, with little wing pad development, averaged 13 mm in length and four specimens in the youngest
age class, with no wing pad development, averaged 7.4 mm long. In 1975 adults emerged in the second
week of July.
Richardson and Gaufin (1971) found that the species is carnivorous and Ephemeroptera make up
the largest percentage of food ingested (37 per cent). Simuliid and chironomid larvae also comprised a
large percentage of ingested material.
Frison (1942a) noted that the adults are nocturnal, emerging at nightfall and when active at dusk or
night have the ability to move on the surface of the water somewhat like water-striders. Mating can take
place as soon as a female emerges from its nymphal skin.
Stoneflies of Saskatchewan
49
Genus Acroneuria Pictet
Sternal ridge patterns of nymphs and adult Acroneuria lack a distinct Y-shape, but three ocelli are
present. MdAt Acroneuria have an undivided tenth tergum, paraprocts that are recurved into genital
hooks and a hammer on the ninth sternum (Fig. 9, 102, 103). The female subgenital plate is little to
moderately produced over the ninth sternum (Fig. 99, 100). Nymphs of most species lack an occipital
ridge of closely set spinules, but if one is present it is broken at the midline. Seventeen species are known
in North America; two occur in Saskatchewan.
Key to Saskatchewan species of Acroneuria
Adults
1 Remnants of subanal gills visible either dorsally or ventrally on tenth abdominal
segment; head with ocellar triangle enclosed with dark brown to black sclerotization
and extended anteriorly; female subgenital plate slightly emarginate (Fig. 100)
Acroneuria lycorias (Newman), p. 50
r No remnants of subanal gills; head with ocellar rings as only dark markings; female
subgenital plate broadly rounded (Fig. 99) Acroneuria abnormis (Newman), p. 49
Nymphs
1 With subanal gills; general color dark brown and yellow dorsally (Fig. 2)
Acroneuria lycorias (Newman), p. 50
1 ' Without subanal gills; general color light brown and yellow dorsally (Fig. 1)
Acroneuria abnormis (Newman), p. 49
Acroneuria abnormis (Newman)
(Fig. 1,99, 103, 186)
Perla abnormis 1838: 177.
Acroneuria abnormis, Needham and Claassen 1925: 178. - Claassen 1940: 171. - Harden and Mickel
1952: 54. -Ricker 1964: 54. -Frison 1935: 391. -Hitchcock 1974: 149.
Acroneuria abnormis, a common prairie stonefly (Ricker, 1946), ranges from northern Quebec and
the Maritimes to New England and south to Florida. It occurs west to Manitoba, Illinois and
Minnesota. Stark and Gaufin (1976) reported Acroneuria abnormis from the South Saskatchewan
River at Saskatoon, Sask., and additional collection localities include the North Saskatchewan River at
Hwy. 5 (Borden Bridge), and 10 mi. E. of Prince Albert, Sask.; the South Saskatchewan River at the
ferry N. of Lemsford, Sask.; and the Saskatchewan River 2 mi. S.W. of Nipawin, Sask. (Fig. 186).
Diagnotic Characters. - Average length, males, 25.0 mm (from anterior margin of head to tip of
folded wings); females, 42.0 mm. Male genitalia with paraprocts broad, triangular and sharp; spinules
present on terga 9 and 10 (Fig. 103). Female subgenital plate slightly produced and broadly rounded
(Fig. 99).
Nymphs without anal gills; with light M-pattern in front of median ocellus (Fig. 1).
Needham and Claassen (1925) and Frison (1935) figured the adult genitalia and the nymph was
illustrated by Claassen (1931) and Frison and Hitchcock (1974). Hitchcock (1974) noted that there was
a wide range of variability in the abdominal color pattern of Connecticut nymphs, and a similar
variation occurs in Saskatchewan specimens. It may vary from being all dark to having two broad,
paired, semicircular dark patches on the basal part of most abdominal terga.
Quaest.Ent., 1979, 15(1)
50
Dosdall& Lehmkuhl
Bionomics. - Most nymphs of Acroneuria abnormis are found under larger rocks in water about
one meter deep. Steffan (1967) noted that the species occurs in rivers where it is exposed to the strongest
current.
The species probably has a three-year life cycle since three nymphal size classes can be collected at
one time. For example, the average length of six near-mature nymphs with well-developed wing pads
collected at the South Saskatchewan River (Lemsford Ferry) on May 23, 1974 was 25.5 mm. Six
nymphs with little wing pad development averaged 17.6 mm in length, and two nymphs with no wing
pad development averaged 8.2 mm long.
In Saskatchewan, adult emergence began in the first week of July of 1974 and 1975 and lasted
approximately two weeks. Eggs evidently hatch soon after they are laid because a nymph measuring
2.66 mm long was collected at the ferry north of Lemsford, Sask. on August 9, 1974. The nymph would
be the overwintering stage. Harper and Pilon (1970) graphed the emergence of the species in Quebec
and suggested that emergence variation from year to year was dependent on water temperature. Males
were found to emerge earlier than females. Hitchcock (1974) stated that nymphs are carnivorous and
after a drought in which streams had dried up reported collecting small nymphs from the streams when
they had again started to flow. Steffan (1967) observed that Acroneuria abnormis was active when the
water temperature was -0.5 C, presumably in the absence of ice.
Steffan (1967) found that larval phoresis of a chironomid on Acroneuria abnormis was common in
a Quebec stream. Eighty-three percent of chironomids on stonefly nymphs were on specimens of
Acroneuria abnormis, evidently the preferred host. This type of phoresis is known for Acroneuria
lycorias in Saskatchewan, but has not been observed for Acroneuria abnormis.
Acroneuria lycorias (Newman)
(Fig. 2, 100, 102, 186)
Perla lycorias Newman 1839: 35.
Acroneuria perbranchiata,'^QSL\Q 1933: 236. -Claassen 1940: 175.
Acroneuria lycorias, Claassen 1940: 174. - Frison 1942a: 283. - Harden and Mickel 1952: 55. - Ricker
1964: 54. - Hitchcock 1974: 155.
Acroneuria lycorias occurs in northern Quebec and south through New England to Florida, and
west to Ontario, Manitoba, Saskatchewan and Tennessee. It is common throughout its range. Cushing
(1961) reported the species from Montreal River, and additional Saskatchewan collection records
include (Fig. 186): Weyakwin River, Jet. Hwy. 2; stream 85 mi. N. of La Ronge, Sask., Jet. Hwy. 102;
Mackay Creek, Jet. Hwy. 2; Nipekamew River, Jet. Hwy. 165; Jackfish Creek, Jet. Hwy. 8; Cub Creek,
Jet. Hwy. 106; Nemeiben River, Jet. Hwy. 2; Torch River, Jet. Hwy. 106; Cole Creek, Jet. Hwy. 104;
Puskwakau River, Jet. Hwy. 106; Arsenault River, Jet. Hwy. 104; Overflowing River, Jet. Hwy. 109;
Red Deer River, Jet. Hwy. 23, and Jet. Hwy. 109; Pine Creek, Jet. Hwy. 165; Montreal River, Jet.
Hwy. 2; Waterhen River, Jet. Hwy. 226; stream 87 mi. N. of Southend, Sask.; Ballantyne River, Jet.
Hwy. 165; Churchill River, Jet. Hwy. 2, Pita Lake Rapids and Wapumon Lake Rapids; Caribou Creek
near Caribou Lake, Jet. Hwy. 106; Bear River, Jet. Hwy. 106; Martineau River, Jet. Hwy. 55; Little
Red River, near Prince Albert, Sask.; Bow River, Jet. Hwy. 165; Waskesiu River, Jet. Hwy. 2; and
Beaver River, Jet. Hwy. 155.
Diagnostic Characters. - Average length, males, 18.0 mm (from anterior margin of head to tip of
folded wings); females, 35.0 mm. Male genitalia with paraprocts flattened, triangular and sharply
pointed at anteriorly directed tips; spinules present on terga 9 and 10 (Fig. 102). Female subgenital plate
produced and flattened or emarginate apically (Fig. 100). Adult head with ocellar triangle dark. Adult
Stoneflies of Saskatchewan
51
with anal gill remnants.
Nymphs with anal gills; banded pattern on abdominal terga; light M-pattern in front of anterior
ocellus (Fig. 2).
Needham and Claassen (1925), Hitchcock (1974) and Prison (1942a) figured the adult genitalia, and
Prison (1942a) illustrated the nymph.
Bionomics. - In Saskatchewan, Acroneuria lycorias seems to have a three-year life cycle since three
size classes of nymphs can be collected at one time. For example, at the Nemeiben River on
June 5, 1974, the average length of seven mature nymphs with well-developed wing pads was 25.3 mm;
the average length of five immature nymphs which had little wing pad development was 20.0 mm, and
the average size of six very immature nymphs with no wing pad development was 10.1 mm.
In Saskatchewan, emergence occurs about the first two weeks of July. Harper and Pilon (1970)
noted that the emergence period is short.
Nymphs have most often been collected from under rocks in rapids, and are probably carnivorous.
Steffan (1967) noted that chironomid larvae of Plecopteracoluthus downesi were phoretic on
nymphs of Acroneuria lycorias. Of the Perlidae carrying chironomid larvae, three per cent were
Acroneuria lycorias. In Saskatchewan, phoresis of a chironomid on Acroneuria lycorias nymphs is very
common in some rivers.
Genus Hesperoperla Banks
In Hesperoperla male paraprocts are flattened, triangular and pointed; spinules are absent from
both the ninth and tenth abdominal terga (Fig. 101). The female subgenital plate is produced over half
of sternum 9; the hind margin is slightly angulate and usually has squarish shiny spot at its apex
(Fig. 104). The nymph has anal gills but lacks an occipital ridge. The nymphal head has a yellow mark
extending from labrum to the anterior ocellus, and the abdominal terga are brown (Fig. 3).
Hesperoperla, a monotypic genus, occurs in Saskatchewan.
Hesperoperla pacifica (Banks)
(Fig. 101, 104, 186)
Acroneuria pacifica Banks 1900: 242. -Frison 1942b: 72. - Jewett, S.G. 1959: 89. - Claassen 1940: 175.
Acroneuria pumila. Banks 1906: 335.
Acroneuria okanagan, Ricker 1935: 262. - Claassen 1940: 173.
Acroneuria (Hesperoperla) pacifica, Ricker 1943: 130. -Gaufin, Ricker, Miner, Milam and Hays 1972:
154.
Hesperoperla pacifica, lilies 1966: 336. - Baumann, Gaufin and Surdick 1977: 162.
Hesperoperla pacifica ranges from British Columbia to California and New Mexico and east to
Alberta and South Dakota. This first Saskatchewan collection record is from Cypress Hills at Battle
Creek near Reesor Lake (Fig. 186). It was abundant at this locality but was not collected further
downstream in the same river near Consul, Sask.
Diagnostic Characters. - Average length, males, 25 mm (from anterior margin of head to tips of
folded wings); females, 37 mm. Since this is the only known species of Hesperoperla, generic characters
also define specific characters.
Needham and Claassen (1925), Gaufin et al. (1972), Stark and Gaufin (1966) figured the adult
genitalia; Claassen (1931) figured the nymph.
Quaest. Ent., 1979, 15(1)
52
Dosdall & Lehmkuhl
Bionomics. - Three size classes of nymphs have been collected at one sampling time suggesting a
three-year life cycle. For example, four near-mature nymphs with well-developed wing pads were
collected at Battle Creek on June 3, 1975 which averaged 28.5 mm long. Nineteen nymphs with little
wing pad development averaged 14.9 mm long and seven very immature nymphs with no wing pad
development averaged 5.3 mm long.
In 1975 adults emerged about the last week of June and emergence lasted for approximately two
weeks.
Richardson and Gaufin (1971) found the species to be carnivorous, its diet consisting of 88.3 per
cent animal matter. The prey was chiefly Ephemeroptera nymphs and chironomid and Trichoptera
larvae.
Genus Perlesta Banks
Adults of this genus have remnants of nymphal subanal gills, a yellowish-white costal margin in the
forewings, and lack anal crossveins. Males have spinule patches only on the tenth abdominal tergum
and lack a hammer on sternum 9. Paraprocts are sclerotized and recurved over the tenth tergum. The
female subgenital plate is slightly produced, and distinctly notched mesally. Nymphs have a distinct
occipital ridge and dark freckled spots on abdominal terga.
Superficially, Perlesta adults resemble Isoperla but the gill remnants and light coloring in the costal
area of the wing are distinctive.
Two species are presently recognized in North America and one occurs in Saskatchewan.
Perlesta placida (Hagen)
(Fig. 187)
Perla placida Hagen 1861: 28.
Needham and Claassen 1925: 158. -Frison 1942a: 271. - Ricker 1964: 54. - Hitchcock
1974: 161.
Perlesta placida is a widely distributed species occurring from the Maritimes to Florida and west to
Manitoba, Wyoming and Texas. This first Saskatchewan record is the most northerly range extension
of the species. It has been collected from Bisset Creek, Jet. Hwy. 55, and from the Assiniboine River
nearTadmore, Sask., Jet. Hwy. 9 (Fig. 187).
Diagnostic Characters. - Average length, males, 8.5 mm (from anterior margin of head to tip of
folded wings); females, 14 mm. Adult head with central, longitudinal black stripe darkest between
ocelli. Male genitalia with genital hooks tapered from bases to tips. Female subgenial plate slightly
produced, notched in center, and covered with long hairs.
Nymphs with speckled appearance; with single light stripe extended down midline of abdomen;
abdominal terga lined with intercalary spinules.
Needham and Claassen (1925), Frison (1935, 1942a), and Hitchcock (1974) figured the adults;
Claassen(1931) and Frison (1935) figured the nymph.
Bionomics. - Stewart al. (1969) studied the mating behaviour of Perlesta placida. Males lived four
to six days and females lived seven to eight days; both could mate several times during this period.
Mating occurred the first day after adult emergence and began by the aggressive capture of females by
the males soon after contact. The male genital hooks attached to the lobes of the female subgenital plate
allowing a space between the male subanal lobes directly below the female genital opening. This space
served as a “channel” for the aedeagus which was everted from beneath the posterior portion of the
Stoneflies of Saskatchewan
53
subgenital plate, through the space, and into the female genital opening.
This species has been found in a wide variety of streams, even intermittent ones (Harden and
Mickel, 1952). In Saskatchewan, it emerges in mid-summer: mature nymphs were collected at Bisset
Creek on July 19, 1976; adults were found on July 22, 1976. Nymphs are carnivorous, feeding mainly on
chironomid larvae (Prison, 1935).
Family Perlodidae
Although few features characterize this family as a whole, perlodids all have slender maxillary palpi
and glossae reduced to small knobs fused to the side of the paraglossae. Perlodidae differ from Perlidae
by the absence of dissected thoracic gills, and pointed rather than rounded paraglossae. Although a few
exceptions occur, perlodids are distinct from chloroperlids by having the two branches of A2 in the
forewing leaving the anal cell separately and by having a flatter nymphal body.
This family occurs in North America, Eurasia and northern Africa and has been divided into three
subfamilies: the more primitive Isogeninae, and more highly advanced Perlodinae and Isoperlinae.
Isogninae have a cleft male tenth tergum, a well developed supra-anal process, and usually have
submental gills. Male paraprocts are generally produced into recurved hooks in Isoperlinae and project
upward or backward in Perlodinae. All three subfamilies are represented in Saskatchewan.
Isogenoides, Arcynopteryx and Skwala are in the subfamily Isogninae; Isoperla belongs to Isoperlinae
and Diura is a representative of Perlodinae.
Genus Arcynopteryx Klap^lek
Arcynopteryx is characterized by having a mesosternal Y-ridge attached to the anterior corners of
the furcal pits and the absence of a transverse ridge (Fig. 25). The male ninth tergum is emarginate and
bears many spinules on its posterior margin. The tenth tergum is cleft with the posterior section
modified into a pair of flat arms on dorsal lobes which are curved in a lateral direction. The supra-anal
process consists of a very long, thin spine projecting from an expansive membranous region (Fig. 1 13).
The female subgenital plate is broad, produced halfway or more over sternum 9, and is trilobed or
occasionally bilobed (Fig. 1 15). Nymphs lack abdominal and thoracic gills but have well-developed
submental gills. Nymphal mandibles usually lack denticles on their cusps but may have a few present on
the outer left cusp only. The major cusp of these mandibles is slightly serrate marginally.
This Holarctic genus is monotypic and occurs in Saskatchewan.
Arcynopteryx compacta (MacLachlan)
(Fig. 113, 115, 188)
Dictyopteryx compacta MacLachlan 1872: 53.
Arcynopteryx lineata. Smith 1917: 476.
Arcynopteryx ignota, Smith 1917: 479.
Arcynopteryx inornata. Smith 1917: 480.
Perlodes minor, Ricker 1938: 144.
Arcynopteryx minor , Hanson 1942: 396
Arcynopteryx americana, Ricker 1943: 114, and 1944: 183.
Arcynopteryx compacta, Brinck 1949: 58. - lilies 1966: 352.
Quaest.Ent., 1979, 15(1)
54
Dosdall& Lehmkuhl
In North America, Arcynopteryx compacta extends from the arctic southward to Colorado, Lake
Superior and New Hampshire. The single Saskatchewan record was given by Ricker (1944) from Tazin
River near Lake Athabasca. Ricker (1964) presented a North American distribution map of the species;
the Saskatchewan collection record is mapped in Fig. 188.
Diagnostic Characters. - Average length, brachypterous males, 1 1 mm (from anterior margin of
head to end of abdomen); females, 20 mm. (from anterior part of head to tip of folded wings). Since this
is the only known species of Arcynopteryx, generic characters also define specific characters.
Hanson (1942) figured the male and female genitalia; Brinck (1949) figured the nymph.
Bionomics. - Brinck (1949) found that adults of Arcynopteryx compacta emerge in mid-summer
during the night or early morning when humidity is highest. After mating, the female carries the
extruded egg mass on the abdomen and the eggs fall off as the female runs across the water. The eggs
then become attached to the substrate by adhesive projections. Nymphal growth is greatest in fall and
the following spring. Nymphs are mainly carnivorous feeding on mayfly and chironomid larvae.
Genus Skwala Ricker
This genus has the arms of the mesosternal Y-ridge terminating at the anterior corners of the furcal
pits; a transverse ridge is absent. The male tenth tergum is cleft with inward and forward directed lobes
which are fairly long and spinulose. The male lateral stylets are well-developed (Fig. 1 12). The female
subgenital plate is produced somewhat and truncate (Fig. 1 14). The nymphal mandibles have numerous
denticles along both sides of their outer cusps (Fig. 28).
This genus is represented by two North American species; one occurs in Saskatchewan.
Skwala parallela (Frison)
(Fig. 25,28,35, 112, 114, 177)
Perlodes americana, Needham and Claassen 1925: 61. -Claassen 1940: 111.
Hydroperla parallela, Frison 1936: 261, and 1942a: 298. -Claassen 1940: 104.
Arcynopteryx (Skwala) parallela, Ricker 1943: 113. - Jewett 1959: 64.
Skwala parallela, lilies 1966: 378.
Skwala parallela has been previously reported from western Canada and United States, including
British Columbia, Oregon, California and Utah. In Saskatchewan it occurs in the following localities
(Fig 188): McDougal Creek, Jet. Hwy. 120; Cub Creek, Jet. Hwy. 106; Mackenzie Creek, Jet. Hwy.
165; Shuard Creek, 1 1 mi. S. of Piapot, Sask.; and Battle Creek, Cypress Hills, near Reesor Lake.
Diagnostic Characters. - Average length, brachypterous males, 19.0 mm (from anterior margin of
head to end of abdomen); females, 25.0 mm (from anterior margin of head to tip of folded wings). Male
genitalia with tenth tergum cleft; subcylindrical lobes medially on tergum 10, 2-3 times as long as
breadth of their middle constricted portion (Fig. 112). Female subgenital plate extended only slightly
over sternum 9; posterior margin straight and corners angulate (Fig. 1 14).
Nymphs with serrations along major mandibular cusps (Fig. 28); partial occipital ridge, interrupted
in middle, borders hind margins of compound eyes.
Frison (1936, 1937) and Gaufin et al. (1972) figured the male and female genitalia; Frison (1942a)
figured the nymph.
Bionomics. - Mature nymphs of Skwala parallela were collected from Battle Creek near Reesor
Lake on March 28, 1976 and from Cub Creek, Jet. Hwy. 106 and McDougal Creek, Jet. Hwy. 120 on
April 12, 1976. Adults were collected from Battle Creek (Reesor Lake) on May 19, 1976.
Stoneflies of Saskatchewan
55
Richardson and Gaufin (1971) found that nymphs lived beneath rocks in swift, well-oxygenated
water and were principally carnivorous, feeding on mayfly and chironomid nymphs, and occasionally
on small stonefly nymphs. Filamentous algae, diatoms and detritus also formed significant parts of the
diet.
Genus Diura Billberg
Diura, the only North American Perlodinae, is characterized by having an uncleft male tenth tergum
and paraprocts which are extended backward to meet on the median margin of the tenth tergum
(Fig. 107). Females have the subgenital plate produced halfway or more over the ninth sternum
(Fig. 108, 109). Nymphs have dark abdominal terga with large paired spots laterally and a few smaller
lateral spots. The nymphal lacinia, or inner projecting process of the maxilla, has a sharp angle just
below the smaller tooth (Fig. 10) in contrast to other Saskatchewan perlodid genera in which the region
below the smaller tooth is rounded or, more commonly, tapering from the tooth to the base (Fig. 170).
This genus is represented by three species in North America: two are Holarctic {nanseni and
bicaudata) and onQ {knowltoni) occurs in western United States and Canada. Diura bicaudata occurs in
northern Saskatchewan.
Diura bicaudata (Linnaeus)
Phryganea bicaudata Linnaeus 1758: 548.
Dictyopterygella postica, Ricker 1944: 178.
Diura bicaudata, Brinck 1949: 61.
This Holarctic species has North American reports from Alaska, Yukon, Mackenzie, and Keewatin.
This first Saskatchewan record is based on a collection of a single female nymph from the shore of
Wollaston Lake (Fig. 189) by D.S. Rawson and identified by W.E. Ricker.
Diagnostic Characters. - Average length, brachypterous males, 12.0 mm (from anterior margin of
head to end of abdomen); females, 17.0 mm (from anterior margin of head to tip of folded wings). Male
genitalia with subanal lobes produced posterad, together subcylindrical, and glabrous with tips lighter
than the remainder (Fig. 107). Female subgenital plate variable. Ricker (1944) reported specimens with
subgenital plate produced about halfway over sternum 9, its sides broadly rounded and deeply
excavated. Brinck (1949) presented figures showing eight different forms of subgenital plate. Drawings
presented here (Fig. 108, 109) show two common forms of subgenital plate with posterior edge broadly
rounded (Fig. 108), and shallowly emarginate medially (Fig. 109).
Nymphs without underlying adult characters are presently indistinguishable from other species of
Diura.
Bionomics. - Brinck (1949) has reported females of Diura bicaudata ovipositing while walking
across the water surface or when flying. Egg incubation was found to last one to two months with
nymphs showing a high growth rate in autumn and spring, but very retarded winter growth.
Nymphs are mainly predatory, feeding on chironomid larvae but occasionally they ingest vegetable
matter (Brinck 1949).
Genus Isogenoides Klapalek
Isogenoides, the only Saskatchewan representative of Isogeninae, is characterized by having the
arms of the mesosternal Y-ridge meeting the posterior corners of the furcal pits in addition to having a
Quaest.Ent., 1979, 15(1)
56
Dosdall& Lehmkuhl
median ridge joining the transverse ridge to the fork of the Y (Fig. 24). Submental gills are long. The
male tenth tergum is completely cleft, and the genital lobes are directed medially or anteriorly rather
than posteriorly. The male supra-anal process is slender with a terminal or subterminal hook or hooks.
Lateral stylets are slender, acute or blunt, and the subanal lobes are quite expanded (Fig. 1 10, 1 1 1). The
female subgenital plate is moderately produced, and either rounded or notched (Fig. 1 16). Nymphal
mandibles usually have the ventral cusps serrate marginally but the denticles are minute.
Two of nine North American species of Isogenoides occur in Saskatchewan. These are the closely
related species Isogenoides colubrinus (Hagen) and Isogenoides frontalis (Newman) which were ranked
as subspecies of Isogenoides frontalis by Ricker (1952) but assigned full specific status by lilies (1966).
Isogenoides colubrinus and Isogenoides frontalis are separable taxonomically only by examination of
the male genital structures; Ricker (1952) was unable to establish characters to distinguish between
females. I have reared a large series of females of both species but also have been unable to find any
differences between the two which hold true for all or even most specimens. Nymphs of the two species
have shown a similar agreement in all characters examined.
Key to Saskatchewan species of Isogenoides
Males
1 Posterior inner angles of genital hooks (terminal sclerotized corners of 10th tergum)
sharp, practically a right angle (Fig. 1 10) Isogenoides colubrinus (Hagen), p. 56
1 ' Posterior inner angles of the genital hooks broadly obtuse or rounded (Fig. Ill)
Isogenoides frontalis (Newman), p. 57
Isogenoides colubrinus (Hagen)
(Fig. 20, 24, 38, no, 116, 189)
Isogenus colubrinus Hagen 1874: 576. -(in part) Claassen 1940: 106. - Ricker 1964: 55, 67.
Isogenoides frontalis , Hanson 1943: 660.
Isogenus frontalis. Harden and Mickel 1952: 33.
Isogenus (Isogenoides) frontalis colubrinus, Ricker 1952: 1 10.
Isogenoides colubrinus, lilies 1966: 363.
Isogenoides colubrinus is known in Alaska, the cordilleras to Utah, northern Alberta to the
Mackenzie River, Great Slave Lake, Lake Athabasca, central Saskatchewan and as far east as The Pas,
Manitoba. Ricker ( 1946) reported the species from St. Laurent, Sask. (South Saskatchewan River) and
additional collection records (Fig. 189) include the North Saskatchewan River, Jet. Hwy. 5 (Borden
Bridge); South Saskatchewan River at the ferry north of Lemsford, Sask., the ferry Fenton, Sask., and
the ferry north of Birch Hills, Sask.; and the Saskatchewan River 2 mi. S.W. ofNipawin, Sask. Ricker
(1964) mapped the North American distribution pattern of this species.
Diagnostic Characters. - Average length, males, 18.0 mm (from anterior part of head to tip of
folded wings); females, 24.0 mm. Adults with submental gills long; fork of the mesosternal Y-ridge
connected with transverse ridge (Fig. 24). Male genitalia with posterior angles of genital hooks slightly
acute and approaching right angles (Fig. 110); epiproct with hook at its tip. Female subgenital plate
only slightly produced with deep U-rounded notch (Fig. 1 16).
Nymphal mandibles in most specimens serrate marginally, with scarcely distinguishable denticles;
abdominal terga uniformly light brown.
Hanson (1943) and Gaufin et al. (1972) figured the male and female genitalia; Claassen (1931)
Stoneflies of Saskatchewan
57
figured the nymph.
Bionomics. - In Saskatchewan, mature nymphs were collected from Lemsford Ferry, Borden
Bridge, and Birch Hills Ferry in mid-May of 1974 and 1975. Adults were collected at the end of May of
1974. Mature nymphs were not found in collections from the North and South Saskatchewan Rivers
from mid-June onward in both 1974 and 1975 with the unusual exception of two mature nymphs from
Lemsford Ferry on July 7, 1975. They were reared and emerged in the laboratory on July 10, 1975. Eggs
apparently develop rapidly since six immature nymphs averaging 5 mm in length were collected from
Lemsford Ferry on August 10, 1974.
Isogenoides frontalis (Newman)
(Fig. Ill, 189)
Isogenus frontalis Newman 1838: 178. - (in part) Claassen 1940: 106. - Frison 1942a: 290. - Ricker
1964: 65,67.
Isogenus hudsonicus , Hanson 1943: 662.
Isogenus (Isogenoides) frontalis frontalis , Ricker 1952: 110.
Isogenoides frontalis, lilies 1966: 365.
Isogenoides frontalis is considered an eastern and northern species (Ricker, 1952) ranging from the
Maritimes to northeastern United States and west to Michigan, Minnesota and Churchill, Manitoba.
This first Saskatchewan report represents the most westerly range extension of the species. Specimens
have been collected from the following localities (Fig. 189): Weyakwin River, Jet. Hwy. 2; stream at
mile 83, Jet. Hwy. 106; stream at mile 85, Jet. Hwy. 102; stream 39 mi. N. of Hudson Bay, Sask., Jet.
Hwy. 109; Meeyomoot River, Jet. Hwy. 165; Torch River, Jet. Hwy. 106; Waskwei River, Jet. Hwy.
109; Puskwakau River, Jet. Hwy. 106; and Mistohay Creek, Jet. Hwy. 226.
Ricker (1964) presented a North American distribution map of this species. The species is probably
much more widespread than indicated on the Saskatchewan distribution map since I have collected
nymphs and females from several other northern rivers which key to Isogenoides, but without adult
males it is impossible to assign specific names to these specimens. I believe them to be Isogenoides
frontalis since Isogenoides colubrinus has been found only in the Saskatchewan River System.
Diagnostic Characters. - Isogenoides frontalis differs from Isogenoides colubrinus only in shape of
male genital hooks. In Isogenoides frontalis, posterior inner angles of genital hooks are broadly obtuse
or rounded (Fig. 1 1 1).
Needham and Claassen (1925), Frison (1942a) and Hanson (1943) figured the male and female
genitalia; Frison (1942a) figured the nymph.
Bionomics. - Isogenoides frontalis appears to have a one-year life cycle. Forty-three mature nymphs
with well-developed wing pads collected from Weyakwin River, Jet. Hwy. 2 on May 6, 1976 had an
average length of 20.2 mm. There were no nymphs of a younger age-class present (less than 17 mm in
length). Adult emergence occurred at the end of May and early June of 1974 and 1975. Nymphs were
absent from samples taken from the same river in late June and mid-July of 1974 and 1975. Egg
development and early nymphal growth appear rapid since twelve early-instar nymphs averaging
8.9 mm in length were collected from Weyakwin River on August 8, 1975.
Genus Isoperla Banks
Males of this genus are characterized by having an uncleft tenth tergum, the absence of both gills
and epiproct, and modified paraprocts. The male eighth sternum bears a lobe on its hind margin
Quaest.Ent., 1979, 15(1)
58
Dosdall & Lehmkuhl
(except in maxana Harden and Mickel which has lobes on both sterna 7 and 8 and in ebria (Hagen) and
trictura (Hoppe) in which lobes are absent). Females of Isoperla are difficult to separate from
Isogenoides, but in most Isoperla species females do not have subgenital plates produced as greatly as in
Isogenoides. Most nymphs have a longitudinal or transverse striped pattern on the abdomen.
This large, difficult genus is in great need of revision; many of the 50 known species probably should
be synonomized.
Because there is a great variation in both color patterns and genitalic structures within many species,
the keys, taken largely from Prison (1942a), Gaufin et al. (1972) and Hitchcock (1974) may be
inoperative for some specimens.
Key to Saskatchewan species of Isoperla
Adults
1
r
2 ( 1)
2'
3 ( 2')
3'
4 ( 3')
4'
5 ( 4')
5'
6 ( 5)
6'
7 ( 5')
7'
8 ( 1')
8'
9 ( 8')
Ninth ventral abdominal segment much produced posteriorly and recurved upward,
sternum 10 mostly or entirely concealed. Sternum 8 with small lobe in middle of
posterior portion (Fig. 1 18)( males)
Ninth ventral abominal segment poorly or not at all produced; tenth segment always
visible from below. Sternum 8 without small lobe in middle of posterior portion;
posterior margin either broadly rounded or strongly produced as subgenital plate
(females)
Cerci about twice as long as relaxed abdomen, length of 12th cereal article 6 to 8 times
its greatest width; two conspicuous semi-circular patches of spinules on ninth
abdominal tergum(Fig. 120) Isoperla longiseta Banks, p.
Cerci less than 1.5 times as long as relaxed abdomen, length of 12th cereal article 3 to 5
times its greatest width; if spinules present on abdominal tergum 9, spinules not
arranged in semi-circular pattern
Patches of short stout hairs on posterior margin of tergum 9; most specimens with red
tinge to some abdominal segments (Fig. 122) Isoperla patricia Prison, p.
No patches of stout hairs on tergum 9; most specimens without red tinge to abdominal
segments
Longitudinal dark stripes on abdominal terga(Fig. 1 19)
Isoperla transmarina (Newman), p.
Abdomen uniformly brown or yellow
Paraprocts projecting little if at all over tergum 10 (Fig. 1 17, 1 18, 121)
Paraprocts recurved upward and forward over tergum 10 (Fig. 123)
Paraprocts scarcely sclerotized (Fig. 117, 118) Isoperla bilineata (Say), p.
Paraprocts very heavily sclerotized (Fig. 121) Isoperla decolorata (Walker), p.
Aedeagus with slender process, forked at its base and visible as rod-shaped structure
through sternum 9 (Fig. 123, 124) Isoperla petersoni Needham and Christensen, p.
Aedeagus without slender sharp process
Isoperla marlynia Needham and Claassen, p.
Light spot in ocellar triangle broadly open caudally, forming a broad V; subgenital
plate subtriangular or slightly notched posteriorly (Fig. 125-128)
Isoperla bilineata (Say), p.
Ocellar triangle completely enclosed by dark sclerotization with central light spot
completely enclosed or narrowly open caudally; subgenital plate variable
Cerci about twice as long as relaxed abdomen, length of 12th cereal article 6 to 8 times
2
8
61
3
62
4
64
5
6
7
60
60
63
62
60
9
9'
10 ( 9')
10'
11 (10')
11'
12 (11')
12'
13 (12')
13'
Nymphs
1
r
2 (1)
2'
3
4 ( 3')
4'
5 (4')
5'
6 (5')
6'
Stoneflies of Saskatchewan
59
its greatest width; subgenital plate rounded in most specimens, slightly excavated, or
rarely slightly pointed (Fig. 129-132) Isoperla longiseta Banks, p. 61
Cerci less than 1 .5 times as long as relaxed abdomen, length of the 12th cereal article 3
to 5 times its greatest width; subgenital plates variable 10
Subgenital plate truncate (Fig. 135) Isoperla transmarina (Newman), p. 64
Subgenital plate not truncate 11
Subgenital plate subtriangular (Fig. 1 33) Isoperla decolorata (Walker), p. 60
Subgenital plate not subtriangular 12
Subgenital plate with deep wide excavation (Fig. 136); most specimens with red cast to
some abdominal segments Isoperla patricia Frison, p. 62
Subgenital plate without deep, wide excavation; without red tinge to abdominal
segments 13
Species with lateral light spots anterior to median ocellus and with crescent-shaped
light spot anterior to median ocellus (Fig. 137)
Isoperla marlynia Needham and Claassen, p. 62
Species without lateral light spots anterior to median ocellus and without subcircular
light spot anterior to median ocellus (Fig. 134)
Isoperla petersoni Needham and Christensen, p. 63
Abdominal terga with longitudinal stripes
Abdominal terga with transverse stripes with both posterior and anterior tergal
margins dark and middle tergal portion light
Isoperla marlynia Needham and Claassen, p.
Abdominal stripes indistinct; 6 to 8 dark dots on each abdominal terga
Isoperla bilineata (Say), p.
Abdominal stripes distinct, no dark dots on abominal terga
Cerci less than 1.5 times as long as relaxed abdomen
Abdominal terga with broad longitudinal yellow band in center bordered on each side
by dark band Isoperla petersoni Needham and Christensen, p.
Abdominal terga with narrow dark band in center bordered on either side by light
band
Light spot within ocellar triangle very small and inconspicuous (Fig. 168)
Isoperla decolorata (Walker), p.
Light spot within ocellar triangle at least as large in diameter as half the distance
between lateral ocelli
Abdominal terga with central dark band bordered on each side by a much broader
light band; light area within ocellar triangle equal in diameter to about half the distance
between lateral ocelli Isoperla patricia Frison, p.
Abdominal terga with central dark band bordered on each side by light band of about
the same width; light area within ocellar triangle equal in diameter to more than half
the distance between lateral ocelli Isoperla transmarina (Newman), p.
2
62
60
3
4
63
5
60
6
62
64
Quaest.Ent., 1979, 15(1)
60
Dosdall& Lehmkuhl
Isoperla biline at a (Say)
(Fig. 31, 117, 118, 125-128, 190)
Sialis bilineata Say 1823: 165.
Isoperla bilineata, Needham and Claassen 1925: 154. - Prison 1935: 436. - Claassen 1940: 198. -
Harden and Mickel 1952: 39. - Ricker 1964: 54.
Isoperla bilineata is a common species from central and eastern Canada and the United States. It
has been collected from Newfoundland, New York, New Jersey and North Carolina and west to
Saskatchewan, Minnesota and Colorado. Needham and Claassen (1925) reported the species from
“Saskatchewan”, and additional collection records (Fig. 190) include: Little Red River near Prince
Albert, Sask.; the North Saskatchewan River at Hwy. 5 (Borden Bridge), Hwy. 12, Prince Albert,
Sask., Cecil Ferry, and the ferry north of Maidstone, Sask.; the South Saskatchewan River at
Saskatoon, Sask., Fenton Ferry, ferry north of Lemsford, Sask., ferry east of Hague, Sask., ferry north
of Birch Hills, Sask.; Battle River, 4 mi. S. of Lashburn, Sask.; Fox Creek; and the river 87 mi. N. of
Southend, Sask. on the Wollaston Lake Road.
Diagnostic Characters. - Average length, males, 10.0 mm (from anterior margin of head to tip of
folded wings); females, 14.0 mm. Adults with head and body pale in color; ocelli in most specimens
connected by dark V-shaped mark. Male genitalia with paraprocts little modified, scarcely sclerotized
and somewhat recurved; lobe on sternum 8 broadly rounded (Fig. 117, 118). Female subgenital plate
sub-triangular and produced about halfway over sternum 9 (Fig. 126); or is slightly emarginate
posteriorly (Fig. 127, 128).
Nymphs with dark longitudinal stripes and six to eight dark dots on abdominal terga and light spot
in ocellar triangle.
Needham and Claassen (1925) and Prison (1935) figured the adult genitalia and wings; Prison (1935)
and Claassen (1931) figured the nymph.
Bionomics. - Prison (1935) noted that Isoperla bilineata was a member of the Illinois spring stonefly
fauna with adult emergence occurring at night and beginning by the end of March. Maximum
abundance of adults was reached in mid-May and adults were absent by the end of June. In
Saskatchewan, adults first appear by the end of May, and the species is very commonly collected
throughout June but practically absent by July. One peculiar collection was a female from the stream
87 mi. N. of Southend, Sask. on the Wollaston Lake Road on August 7, 1975. This record is interesting
because of the late collection date and northern location.
There is some confusion regarding the feeding habits of this species. Prison (1935) stated that the
species seemed to be herbivorous, but Harden and Mickel (1952) found the larval gut contents to be
principally Chironomidae. Eggs collected in June and kept in the laboratory did not hatch until
October (Harden and Mickel, 1952).
Isoperla decolor ata (Walker)
(Fig. 121, 133, 168-170, 192)
Per la decolorata Walker 1852: 170.
Isoperla decolorata, Claassen 1940: 199. - Ricker 1944: 183. - Ricker, 1955: 256. - Ricker 1964: 56.
Isoperla decolorata is a rare species known only from northern Canada (Great Bear Lake,
Keewatin, Fort Churchill, northern Ontario) and Alaska. The collection of a single male from the
North Saskatchewan River at Hwy. 5 (Borden Bridge) represents the most southerly known locality for
this species.
Stoneflies of Saskatchewan
61
Diagnostic Characters. - Average length, males, 10.5 mm (from anterior margin of head to tip of
wings); females, 11.5 mm. Adults with general dark brown color and yellow markings; ocellar spot
small and inconspicuous. Male sternum 8 with lobe broad at tip and constricted at base. Male genitalia
with darkly sclerotized paraprocts fairly broad at their bases, sharp at tips and recurved but not over
tergum 10 (Fig. 121). Female subgenital plate subtriangular with broad base extended about halfway
over sternum 9 (Fig. 1 13). A first description of the nymph is given below.
Description. - Total length of mature nymph about 10 mm.
General color yellow with fuscous areas on head, thorax and abdomen as in Fig. 168; light yellow
ventrally. Antennae light brown with first two articles darker; legs light yellow and darker at joints;
cerci light brown.
Head with ocelli forming nearly equilateral triangle; maxillae and mandibles as in Fig. 169, 170.
Abdominal terga with central longitudinal dark stripe and light subcircular areas on either side;
tergum 10 with central bilobed light area, as in Fig. 168. Abdominal terga covered with numerous
clothing hairs and longer bristles on posterior margins of each terga, longest about one-third the
mid-dorsal tergal length. Cerci with regular whorls of bristles near anterior margin of each article and a
few long hairs which become more numerous on distal cereal articles.
Ricker (1944) figured the male and female genitalia; the nymph is illustrated for the first time in
Fig. 168-170.
Bionomics. - Little is known of the biology of this species. From a sample of 205 Isoperla collected
and reared from the North Saskatchewan River at Hwy. 5 (Borden Bridge) on May 27, 1975 only one
was Isoperla decolorata; the remainder were Isoperla biline at a. The Isoperla decolor at a adult emerged
May 29, 1975 in the laboratory.
Isoperla longiseta Banks
(Fig. 120, 129-132, 191)
Isoperla longiseta Banks 1906: 337. - Needham and Claassen 1925: 156. - Claassen 1940: 203. -Frison
1942a: 318. -Ricker 1943: 124. -Harden and Mickel 1952: 41. -Ricker 1964: 56.
Isoperla longiseta is considered a typical prairie species (Ricker, 1964) and is abundant in large
rivers of the plains and inter-mountain regions from the Mississippi to the Great Basin and from
Colorado and Missouri northward. In Canada it occurs characteristically on the prairies but also as far
north as Norman Wells on the Mackenzie River. Ricker (1946) reported several collections of this
species from Saskatoon, Sask., and additional records (Fig. 191) include: the North Saskatchewan
River at Hwy. 5 (Borden Bridge), North Battleford, Sask., Cecil Ferry, ferry south of Maymont, Sask.,
Prince Alberta, Sask., and the ferry 20 mi. N. of Lioydminister, Sask.; the South Saskatchewan River
at the ferry north of Lemsford, Sask., the ferry north of BirchHills, Sask., and the ferry north of
Fenton, Sask.; the Saskatchewan River 2 mi. S.W. of Nipawin, Sask., and 10 mi. E. of the Squaw
Rapids Power Station; and a record from Melfort, Sask.
Ricker ( 1964) presented a North American distribution map for this species.
Diagnostic Characters. - Average length, males, 10.0 mm (from anterior margin of head to tip of
folded wings); females, 13.0 mm. Adults with yellowish color and darker markings on head; long cerci,
about twice abdominal length. Male genitalia with paraprocts sharp, slender and recurved over tergum
10; tergum 10 with two brown patches of setae and a bilobed brown patch on tergum 9 (Fig. 120).
Female subgenital plate produced about halfway across sternum 9; strongly sclerotized; posterior
margin of subgenital plate rounded in most specimens (Fig. 129), but slightly or greatly emarginate
(Fig. 130, 131) or even slightly pointed in some specimens (Fig. 132).
Quaest.Ent., 1979, 15(1)
62
Dosdall & Lehmkuhl
Nymphs with light color and faint longitudinal stripes on abdominal terga; light area within ocellar
triangle not completely enclosed by darker area.
Needham and Claassen (1925), Prison (1942a), Gaufin et al. (1966) and Gaufin et al. (1972) figured
the adult genitalia; Prison (1942a) figured the nymph.
Bionomics. - This species is apparently restricted to fairly large rivers. It is abundant in the
Saskatchewan River System with adult emergence beginning in mid-June and lasting until about the
end of July. The life-cycle appears to be univoltine since nymphs were absent from benthic collections
for a time following the summer adult emergence.
Isoperla marlynia Needham and Claassen
(Pig. 137, 191)
Isoperla marylnia Needham and Claassen 1925: 148. - Claassen 1940: 203. - Prison 1942a: 330. -
Harden and Mickel 1952: 203. - Hitchcock 1974: 202.
Isoperla marlynia is a common species of eastern Canada and United States. It has been reported
from Virginia to New Brunswick, and west to Manitoba and Minnesota. This first Saskatchewan
collection record (Pig. 191) is from Torch River, Jet. Hwy. 106, and Red Deer River near Chelan,
Sask., Jet. Hwy. 23.
Diagnostic Characters. - Average length, males, 12.0 mm (from anterior margin of head to tip of
folded wings); females, 13.0 mm. Adults with light color and brown markings; head with
crescent-shaped light spot and two lateral light spots anterior to median ocellus. Male genitalia with
paraprocts fairly long and slender, pointed at tips and recurved over tergum 10. Pemale subgenital plate
produced less than halfway over sternum 9; with shallow median emargination (Pig. 137).
Nymphs with three possible distinct color phases. Light phase: mainly yellow with black lateral
stripes on both posterior and anterior margins of each abdominal tergum. Intermediate phase: light
markings laterally on each abdominal tergum with few light markings on head and pronotum. Dark
phase: dark without yellow markings.
Needham and Claassen (1925) and Prison (1942a) figured the adult genitalia; Prison (1942a) figured
the nymph. The three possible color phases of nymphs are apparently equally common, and may occur
in the same population (Prison, 1942a). Only the light phase has been found in Saskatchewan
specimens.
Bionomics. - A female nymph collected from Torch River, Jet. Hwy. 106 on April 12, 1976 emerged
under laboratory conditions on June 15, 1976.
Isoperla patricia Prison
(Pig. 122, 136, 190)
Isoperla patricia Prison 1942a: 313. - Ricker 1943: 126. - Jewett 1959: 74. - Ricker 1964: 55. - Gaufin,
Nebeker and Sessions 1966: 70. - Gaufin, Ricker, Miner, Milam and Hays 1972: 118.
Isoperla patricia, a common species in western Canada and United States, has been collected from
British Columbia to California and Utah and east to Idaho, Montana and South Dakota. This first
Saskatchewan record (Pig. 190) is from the South Saskatchewan River at Lemsford Perry and the
following Cypress Hills localities: Bear Creek, 10 mi S. of Piapot, Sask.; Shuard Creek, 11 mi S. of
Piapot, Sask.; and Conglomerate Creek at Ravenscrag, Sask.
Diagnostic Characters. - Average length, males, 10.0 mm (from anterior margin of head to tip of
folded wings); females, 12.5 mm. Most adult specimens with red pigmentation on abdominal segments.
Stoneflies of Saskatchewan
63
Male genitalia with subanal lobes recurved upwards over tergum 10 and with slender, sharply pointed
tips; posterior margin of sternum 9 with patches of short stout hairs (Fig. 122). Female subgenital plate
produced about halfway over sternum 9 and deeply excavated in most specimens (Fig. 136).
Nymph with dark narrow band in center of abdominal terga bordered on each side by broader light
band.
Frison (1942a), Gaufin et al. (1966) and Gaufine/ al. (1972) figured the male and female genitalia;
Frison (1942a) figured the nymph.
Bionomics. - This species appears to have a two-year life cycle in Saskatchewan. From a sample of
45 nymphs collected on June 3, 1975 at Shuard Creek, 11 mi S. of Piapot, Sask., 21 nymphs ranging in
length from 4.0 mm to 9.5 mm (with an average length of 5.9 mm) had very little wing pad development
and could not have emerged until the following year. Twenty-four mature nymphs ranging in length
from 8.5 mm to 12.0 mm (with an average length of 10.3 mm) had full-grown wing pads and would have
emerged the same year. There were several stages of nymphal development present at one time among
the nearly mature specimens indicating little synchrony of emergence.
Isoperla petersoni Needham and Christensen
(Fig. 123, 124, 143, 192)
Isoperla petersoni Needham and Christensen 1927: 19. - Jewett 1959: 74. - Ricker 1964: 55. - Gaufin,
Ricker, Miner, Milam and Hays 1972: 118.
Isoperlafontium, Neave 1929: 161. - Ricker 1943: 122.
Isoperla petersoni is found in western Canada and United States including Alberta, British
Columbia, Utah and Montana. This first Saskatchewan record (Fig. 192) is from Mackenzie Creek near
Bow River, Jet. Hwy. 165 and the stream at mile 83, Jet. Hwy. 106.
Diagnostic Characters. - Average length, males 12.5 mm (from anterior margin of head to tip of
folded wings); females, 14.5 mm. Adults with yellow color and dark brown markings on head and
thorax. Male genitalia with paraprocts slender, recurved over tenth tergum and darkly sclerotized
(Fig. 123); aedeagus with slender sharp process forked at its base (Fig. 124), aedeagal process visible
through sternum 9 of most specimens as dark rod-shaped structure; when not visible, aedeagal process
may be exposed by dissecting away sternum 9. Female subgenital plate produced nearly halfway over
sternum 9, and slightly emarginate at tip (Fig. 134).
Nymphs with longitudinal striped pattern on abdomen, but differ from other Saskatchewan
Isoperla species by having broad central light band bordered on each side by dark band.
Neave (1929), Gaufin et al. (1966) and Gaufin et al. (1972) figured the female genitalia; Claassen
(1931) figured the nymph.
Brachyptery has been reported in males of this species in Montana (Gaufin c/ al., 1972) and in Utah
(Gaufin et al., 1966), but Saskatchewan specimens all have wings of normal length. The Isoperla species
key for Montana nymphs given by Gaufin et al. (1972) states that nymphs of Isoperla petersoni lack a
fringe of long hairs on the legs. Saskatchewan nymphs of this species all have a definite fringe of hairs
on the legs though the hairs are shorter than in other species of this genus.
Bionomics. - Hales and Gaufin (1971) stated that this species is restricted to spring-fed streams or
streams largely influenced by springs. Adult emergence was found to be relatively short, lasting about
two weeks. In Saskatchewan, adults emerged in the end of June and early July of 1974.
Mackenzie Creek empties into the Bow River near Hwy. 165, and it is interesting that this species
was found only in Mackenzie Creek. Samples from Bow River, even at the point of entry of Mackenzie
Creek, showed numbers of Isoperla transmarina, but not Isoperla petersoni. In both rivers where
Quaest. Ent., 1979, 15(1)
64
Dosdall& Lehmkuhl
Isoperla petersoni was collected, larvae of Rhyacophilidae (Trichoptera) were also abundant though
they are fairly uncommon in other Saskatchewan rivers.
Isoperla transmarina (Newman)
(Fig. 119, 135, 192)
Chloroperla transmarina Newman 1838: 499.
Isoperla fumosa, Neave 1933: 235.
Isoperla transmarina. Prison 1942a: 316. - Harden and Mickel 1952: 46. - Hitchcock 1974: 210.
Isoperla transmarina is a common North American species ranging from the Maritimes south to
New Jersey and west to Minnesota, Manitoba and British Columbia. This first Saskatchewan report is
from collections made at the following localities (Fig. 192): Little Red River, near Prince Albert, Sask.;
stream 80 mi. N. of La Ronge, Sask. on Hwy. 102; Martineau River near Cold Lake; Cole Creek, Jet.
Hwy. 104; Weyakwin River, Jet. Hwy. 2; Waskwei River, Jet. Hwy. 109; Mackay Creek, Jet. Hwy. 2;
Nemeiben River, Jet. Hwy. 2; Bear River, Jet. Hwy. 106; Puskwakau River, Jet. Hwy. 106; Ballantyne
River, Jet. Hwy. 106; Crean River, Jet. Hwy. 2; Churchill River at Wintego Lake Rapids, Otter Rapids,
Iskwatam Lake, and Pita Lake; Montreal River, Jet. Hwy. 165 and at La Ronge, Sask.; Waskesiu
River, Jet. Hwy. 2; Torch River, Jet. Hwy. 106; Overflowing River, Jet. Hwy. 109; Waterhen River, Jet.
Hwy. 226; Broad Creek, Jet. Hwy. 104; McDougal Creek, Jet. Hwy. 120; North Saskatchwan River at
Hwy. 5 (Borden Bridge); Bow River, Jet. Hwy. 165; Waddy River, Jet. Hwy. 102; Meeyomoot River,
Jet. Hwy. 165; Arsenault River, Jet. Hwy. 104; Mistohay Creek, Jet. Hwy. 226; Nipekamew River, Jet.
Hwy. 165; Caribou Creek, Jet. Hwy. 106; stream entering south end of Wollaston Lake, Jet. Hwy. 105;
Fond du Lac River at Black Lake; Cluff Creek near Cluff Lake; Cub Creek, Jet. Hwy. 106; and Green
Lake, Sask.
Diagnostic Characters. - Average length, males, 10.0 mm (from anterior margin of head to tip of
folded wings); females, 14.0 mm. Adult with light brown and yellow coloration; light spot of ocellar
triangle completely enclosed by darker areas in some specimens or may be slightly open posteriorly.
Male genitalia with paraprocts sharply pointed, darkly sclerotized and recurved over tergum 10. Male
abdominal terga with striped pattern, most easily seen on posterior segments (Fig. 119). Female
subgenital plate truncate and produced about halfway over sternum 9.
Nymph with large light spot within ocellar triangle; longitudinal striped pattern on abdominal terga
with central dark band bordered on each side by light stripes of about same width.
Ricker (1938) figured the male genitalia, and Neave (1933) figured the female genitalia (under the
neeoaQ fumosa). Frison( 1942a) figured the nymph.
Bionomics. ~ Gaufin (1958) studied the effects of pollution on the Mad River in Ohio and found
that the occurrence of Isoperla transmarina was limited entirely to the cleanest sections of the stream.
Harden (1942) stated that nymphs seemed to prefer a habitat of matted leaves and vegetation trapped
by submerged objects.
Harper (1973a) provided life cycle data for this species from southern Ontario. Adult emergence was
fairly synchronous, beginning in early May and lasting 40 days. Emergence occurred in early morning
and males emerged earlier than females. Oviposition was first observed in late May and early June. Egg
hatching lasted from 34 to 43 days under simulated stream conditions. Early-instar nymphs were
collected in the field in July which closely matched the laboratory incubation period. Rapid growth
ensued until January when growth was minimal up to April. Growth was then continued and completed
a few weeks before the May emergence.
Stoneflies of Saskatchewan
65
DISCUSSION
Types of life histories and seasonal succession of Saskatchewan Plecoptera
The invertebrate benthos of flowing waters in temperate climates shows a clear succession of events
as species appear and disappear from collections and one species after another completes its life cycle
(Hynes, 1970). A life cycle classification will be employed to categorize how and when Saskatchewan
stonefly species initiate and complete their development.
The system of classification proposed by Hynes (1961) can be applied to Saskatchewan stonefly life
cycles. Basically a distinction is made between univoltine and seasonal species and non-univoltine or
non-seasonal species. Non-seasonal species have individuals of all sizes present at all times while
seasonal cycles show a distinct change of size distribution with time. Univoltine species are then
separable into two groups. The “F” or fast type of cycle implies an embryonic diapause; species of “S”
or slow type do not diapause. Each of these types can be further arbitrarily separated into four groups
depending on the time of year when adults emerge. For example, group Fi contains species of the fast
seasonal type which emerge from mid-winter until April, adults of F2 species emerge in May and June,
F3 species emerge in July and early August and F4 species emerge in late August and in fall.
As can be seen from Table II, Saskatchewan Plecoptera have varied life histories and can contain, as
far as is presently known, species representing all types except S4 and F3. A large number of species
remain unclassified due to a paucity of life cycle data. Further life history studies on the uncategorized
species will certainly increase the variety of types depicted.
The simplest seasonal type is Si in which embryonic development is immediate. Harper and
Hynes (1972) further divided this category, separating species which undergo nymphal diapause
from those which do not. For the sake of simplicity this grouping has not been followed here.
Saskatchewan stoneflies with this type of life cycle are the so-called winter stoneflies.
Several Perlodidae and Chloroperlidae fall into the S2 type. There are, however, still differences
in the timing and deviation of adult emergence. For example, Isoperla bilineata emerges in late
May; Isoperla transmarina emerges in early to mid-June, and Hastaperla brevis emerges
throughout all of June.
Isoperla longiseta emerges in July and although it is not established for certain that an
embryonic diapause is absent, the species probably is of the S3 type.
Species with fast seasonal cycles undergo a long diapause at the onset of embryo-
genesis allowing them to survive the warm summer season as an underdeveloped egg.
Saskatchewan stoneflies with this type of development include Shipsa rotunda emerging in late
April (Fi), Diura bicaudata emerging in late spring (F2) and Amphinemura linda and probably
Malenka californica which emerge in late summer and autumn (F4). Amphinemura linda may
emerge as soon as early July in a few warm streams, but most often it does not emerge until
mid-August, and thus will be considered type F4.
Species which are listed in the uncertain section of Table 1 have inadequate life history
information available for placement in any of the other categories.
Figure 193 presents the seasonal distribution of adult Plecoptera in Saskatchewan. The late
winter and early spring fauna is evidently very rich, and this is succeeded by an equally diverse late
spring and early summer fauna. There are few summer species and only Amphinemura linda and
Malenka californica can be considered late summer and fall species. Several species, which have
not been indicated in Fig. 193, are adventitious and their seasonal distributions are poorly known.
Quaest.Ent., 1979, 15(1)
66
Dosdall& Lehmkuhl
Table II. Types of life histories known in Saskatchewan Plecoptera. Data are from the literature
and present study. The question marks (?) indicate that the life-cycle of the species is
incompletely investigated and that it is placed in the most likely category from data
available.
I. Non-seasonal species (life history longer than one year):
Pteronarcys dorsata (present study)
Claassenia sabulosa (present study)
Acroneuria lycorias (present study)
Acroneuria abnormis (present study)
Hesperoperla pacifica (present study)
Paragnetina media (Harper, 1973a; Heiman and Knight, 1970; Tarter and Krumholz, 1971;
present study)
II. Seasonal species (life history of one year duration):
A. Slow type (no embryonic diapause):
51 (adult emergence from mid-winter until April)
Zapada cinctipes {Clifford, 1969)
Paracapnia angulata (Harper and Hynes, 1970; Harper and Hynes, 1972)
? Oemopteryx fosketti
? Capnia confusa
? Capnia coloradensis
? Capnia gracilaria
? Utacapnia trava
52 (adult emergence in May and June):
Isoperla transmarina (Harper, 1973a)
? Isoperla biline at a
? Isoperla petersoni
? Isoperla patricia
? Skwala parallela
? Hastaperla brevis
? Triznaka signata
? Isogenoides frontalis
? Isogenoides colubrinus
53 (adult emergence in July and early August):
? Isoperla longiseta
B. Fast type (Embryonic diapause):
Fi (adult emergence from mid-winter until April):
Shipsa rotunda {R2irpQr, 1973b)
F2 (adult emergence in May and June):
Diura bicaudata {Brinck, 1949)
F4 (adult emergence in late August and fall):
Amphinemura linda (Harper, 1973b)
? Malenka calif ornica
Stoneflies of Saskatchewan
67
III. Species of uncertain life cycle.
Perlesta placida
Suwallia lineosa
Pteronarcella badia
Isoperla decolorata
Isoperla marlynia
Nemoura rickeri
Podmosta delicatula
Paraleuctra vershina
Leuctra ferruginea
Capnia vernalis
Isocapnia crinita
Isocapnia missourii
Arcynopteryx compacta
Although the period of adult occurrence overlaps for several species, the timing of their
maximum abundance differs.
The differences in stonefly life histories and adult seasonal successions are important in limiting
interspecific and intraspecific competition and allowing the coexistence of several stonefly species
in the same stream.
Winter stoneflies, which concentrate the largest proportion of their nymphal growth in fall and
winter, have utilized several ecological opportunities as Harper and Hynes (1972) point out. Food,
in the form of dead plant matter, is especially abundant in this time of year. Also, few other stream
insects are active during this season. Species which grow rapidly in early spring and undergo
nymphal diapause in summer are able to use an abundant spring food supply in form of dead
leaves which have soaked right through the winter.
Many carnivorous Perlodidae and Chloroperlidae show a wide range of size-frequency
distributions in a given sample indicating the simultaneous co-existence of specimens of different
sizes. Harper (1973a) argues that this has the result of lessening intraspecific competition because it
provides predatory individuals with a wider variety of prey organisms.
Some related species of the same genus or family show a distinct succession of adults, with the
obvious advantage of limiting possible mating encounters.
Saskatchewan stonefly species show a variety of life history patterns and adult seasonal
distributions. Life cycle differences enable the co-existence of several stonefly species in the same
stream, and a succession of adults permits maintenance of a higher level of diversity.
Origin and past dispersal of Saskatchewan Plecoptera
In order to reconstruct the faunal history of a region it is necessary to integrate information of
the geographical and ecological distribution of existent species with knowledge of the geological
and climatic history of the area. To determine past dispersals of species it is requisite to assume
that each species is now found in the same type of habitat which it occupied in the past and that
geographical changes in the distribution of a species’ habitat have also affected the possible range
of that species. Also, it is assumed that non-glaciated areas presently occupied by a species represent
refugia or regions where the species was able to pass the glacial maximum (Larson, 1975).
Quaest.Ent., 1979, 15(1)
68
Dosdall & Lehmkuhl
The destructive results of the Pleistocene glaciation had a major effect in shaping the
distributional history of the extant fauna. There were four major glaciations in the Pleistocene but
the last of the glacial advances, the Wisconsin, obliterated the effects of the previous glaciations
and virtually eliminated the fauna of Saskatchewan. The present-day faunal composition of areas
previously ice-covered originated with the retreat of Wisconsin ice.
In order to arrive at a reasonable explantion of post-glacial stonefly dispersal into
Saskatchewan, several events of the Wisconsin glaciation will be discussed including the full extent
of the ice sheets and locations of possible refugia, subsequent retreat of ice from the glacial
maximum and the important post-glacial lakes and river systems and their development to the
present.
The Wisconsin glaciation began about 50,000 years before present (B.P.) and extended about
1 1,000 years B.P. (McPhail and Lindsey, 1970). At its maximum extent (17,000 years B.P.) almost
all of Saskatchewan was ice-covered by the Laurentide ice sheet except Cypress Hills and a small
area in southcentral Saskatchewan on the present-day Canada-United States border (Prest, 1968)
By about 14,000 years B.P., Prest (1968) suggests that the glaciers had retreated from southwestern
Saskatchewan and by 13,000 years B.P. most of southcentral Saskatchewan was also ice-free.
Two-thirds of Saskatchewan was ice-free about 10,000 years B.P.
As the Laurentide ice sheet began its retreat. Glacial Lake Agassiz formed over roughly 200,000
square miles of the provinces of Ontario, Manitoba and Saskatchewan and the states of
Minnesota, North Dakota and South Dakota (Elson, 1967). The morphometry and the outlets of
Lake Agassiz changed as the glaciers advanced and retreated for short periods. Elson (1967) noted
that the lake originally drained south to the Mississippi River System and later to the Great Lakes,
and to the Athabasca River in the northeast. As the ice receded, the lake drained north to Hudson
Bay.
The Saskatchewan River System began to flow south, prior to 12,000 years B.P., into the Big
Muddy-Missouri System because the ice front prevented a northerly flow (Elson, 1967). Further
retreat of the ice sheet diverted its course to the Souris-Lake Agassiz drainage, and then to the
Qu’Appelle Valley-Assiniboine River drainage. Finally, the ice retreated sufficiently to allow a
northern drainage to Hudson Bay.
The present-day stonefly fauna of Saskatchewan is the result of post-glacial dispersal from
ice-free refugia. The Cypress Hills was unglaciated (Westgate, 1964), and could have served as a
refugium. Also, there were unglaciated areas in Alaska and Yukon which served as important
refugia for some organisms (McPhail and Lindsey, 1970; Munroe, 1956). Three important refugia
formed south of the continental ice sheet: the Pacific refugium west of the continental divide, the
Mississippi comprised of the Missouri River and Upper Mississippi River Systems and the Atlantic
refugium (McPhail and Lindsey, 1970).
Ricker (1964) proposed that the most likely species which survived glaciation in the
northwestern refuge of Alaska and Yukon are the present-day tundra species: Nemoura arctica,
Capnia nearctica, Isoperla decolorata, Diura bicaudata and Arcynopteryx compacta. These
species have since migrated as far east as Hudson Bay but as yet have been unable to pass around
its southern extremity. The latter three species have been found in Saskatchewan: Diura bicaudata
and Arcynopteryx compacta in the northern boreal region and a single specimen of Isoperla
decolorata collected in the North Saskatchewan River.
The collection of Diura bicaudata in northern Saskatchewan and a record from approximately
the same latitude near Hudson Bay in Manitoba represent the most southerly known point to
which this species has been able to penetrate the boreal forest. Arcynopteryx compacta, however.
Stoneflies of Saskatchewan
69
has not only been collected in northern Canada west of Hudson Bay, but also from the Great
Lakes, New Hampshire, southern Alberta and Wyoming. Ricker ( 1964) suggests that the species
must have lived in Glacial Lake Agassiz and dispersed eastward when the lake drained to the Great
Lakes. The common factor in the distribution of Arcynopteryx compacta is that it occurs
principally in cold lakes. Isoperla decolorata has crossed the tundra and boreal forest, reaching its
most southerly distribution on the Saskatchewan parkland. Nemoura rickeri has been collected
previously only in Alaska and also probably dispersed from a northwestern refugium.
A number of species reach their most westerly ranges in the boreal regions of Saskatchewan,
Alberta and northeastern British Columiba, and are abundant in nonglaciated areas of eastern
North America. These species probably originated in an eastern boreal refugium and dispersed to
the north and west. Saskatchewan species with this type of distribution include Acroneuria
lycorias, Perlesta placida, Paragnetina media, Pteronarcys dorsata, Isoperla transmarina, Isoperla
marlynia, Shipsa rotunda, Hastaperla brevis, Taeniopteryx nivalis, Amphinemura linda,
Isogenoides frontalis, and Leuctra ferruginea. The extent to which southeastern species have
invaded Saskatchewan varies from species to species. Paragnetina media, Taeniopteryx nivalis and
Isoperla marlynia are found in northeastern Saskatchewan but not in the northwest. Perlesta
placida occurs only in eastcentral Saskatchewan. Other species range entirely across the northern
part of the province and beyond: Hastaperla brevis extends to the Mackenzie River,
Amphinemura linda enters southern Yukon; Isoperla transmarina reaches northern British
Columbia and Shipsa rotunda extends to the Mackenzie River delta. Pteronarcys dorsata, an
exceptionally widespread species, occurs across Canada except in southern British Columbia and
on the tundra.
Several species are widespread in the mountains of western North America and also occur in
Saskatchewan boreal regions. They probably survived glaciation in the western portion of the
southern North American refugium and dispersed north and east from the eastern border of the
Rockies. Species with this type of distribution are: Pteronarcella badia, Isoperla petersoni, Capnia
confusa, Capnia coloradensis, Capnia vernalis, Utacapnia trava, Claassenia sabulosa, Zapada
cinctipes, Malenka californica and Triznaka signata. On reaching the boreal forest, these species
moved eastward in varying degrees: Pteronarcella badia and Triznaka signata occur only in
northwestern Saskatchewan and the rest of the species have reached northeastern Saskatchewan.
The Cypress Hills remained unglaciated in the Wisconsin (McPhail and Lindsey, 1970) and
there is no reason to believe that they could not have harbored stoneflies even though the area was
completely surrounded by ice. Westgate (1964) provided evidence in Cypress Hills of ponds with
vegetation and animals, not only at the ice edge, but actually on the ice surface where debris had
accumulated to provide a suitable substrate.
Bird (1962), studying bryophytes, Russell (1951), studying land snails and Yeatman(1967)
studying pines, demonstrated that the Cypress Hills flora and fauna has its principal affinities with
the Rocky Mountains. Russell (1951) contended that most or all of the land snails entered the
Cypress Hills postglacially across a “bridge” of suitable climatic and edaphic conditions connected
with the mountains. The bridge later disappeared as the glaciers retreated further and the climate
warmed. This proposal agrees with the statements of Love (1959) regarding the direction of floral
movement across the southern prairies.
Cypress Hills stoneflies are all montane species and their affinities are with present-day species
of the Rocky Mountains. Though not a likely possibility, these stoneflies could have been present
in the Cypress Hills prior to the Wisconsin and passed the glacial maximum in the Cypress Hills
refugia. This proposal assumes that the area was connected to the mountains by a similar type of
Quaest.Ent., 1979, 15(1)
70
Dosdall & Lehmkuhl
climatic and edaphic “bridge” sometime prior to the Wisconsin glaciation. Though little is known
of pre-Wisconsin geography, the formation of such a connection from the Cypress Hills to the
mountains could as easily have occurred after pre-Wisconsin glaciations just as it could following
the Wisconsin glaciation.
The second, more likely origin of Cypress Hills Plecoptera could be by post-Wisconsin
dispersal to the area from the Rockies by a connecting bridge of montane climatic and edaphic
conditions which is proposed to have formed as the ice retreated. As the climate warmed and the
bridge disappeared, the stoneflies were separated from their parent population.
Montane species occurring in the Cypress Hills include Hesperoperla pacifica, Podmosta
delicatula, Suwallia lineosa, Paraleuctra vershina, Capnia gracilaria, Isocapnia crinita, Isocapnia
missourii, Zapada cinctipes, Utacapnia trava, Skwala parallela and Isoperla patricia.
Cypress Hills stoneflies have been unsuccessful in dispersing to, or populating, surrounding
prairie rivers either because of the large expanses of prairie separating them from the rivers or
because of their ecological preference for cool water. The single exception is Isoperla patricia
which has been collected at Lemsford Ferry on the South Saskatchewan River, about 80 miles
north of Cypress Hills.
A group of species occurs in the Saskatchewan boreal forest region and in the Cypress Hills. All
are widespread in the western North American mountains including the western portion of the
southern North American refugium. These separated Saskatchewan populations probably have a
dual origin. The Cypress Hills fauna likely originated by a connection to the mountains either in
pre-Wisconsin or post-Wisconsin time. The western species occurring in the boreal forest probably
dispersed east and north from the eastern border of the Rockies. Species showing this distribution
are Zapada cinctipes, Utacapnia trava and Skwala parallela. It is unlikely that these species
originated from the Cypress Hills refugium and dispersed northward because they do not occur
either in the prairie rivers or in rivers on the southern margin of the boreal forest in Saskatchewan.
Some of the prairie and parkland fauna consists of a northern Great Plains component. These
species occur in the northern part of the Great Plains east of the Rockies and west of the
Mississippi. Species showing this form of distribution are Isoperla longiseta, Isoperla bilineata and
Acroneuria abnormis. These species probably entered the Saskatchewan River System when it
flowed south to the Big Muddy-Missouri Systems. The southeastern and southcentral prairie
species then had a direct aquatic dispersal route, and subsequent changes in drainage patterns have
isolated Saskatchewan River populations from their parent populations.
Oemopteryx fosketti occurs in the Saskatchewan River System and also in the Colorado
System. It probably dispersed northward in a manner similar to that proposed by Lehmkuhl (1976)
for the mayfly Analetris eximia Edmunds. A direct aquatic invasion route is not evident, but the
species could possibly have crossed from the Colorado to the Missouri System in Wyoming and
then entered the Saskatchewan System from tributaries of the Missouri in southern Alberta and
Saskatchewan.
The present-day stonefly fauna of Saskatchewan is of diverse origins, showing a variety of
distributional patterns and histories. Northern glacial refugia contributed to a small percentage of
the post-glacial stonefly re-colonization of Saskatchewan. The majority of the present fauna was
derived from refugia south of the main Wisconsin ice sheets.
Stoneflies of Saskatchewan
71
SUMMARY AND CONCLUSIONS
Saskatchewan has a diverse stonefly fauna which comprises at least 41 species. Nymphs were
reared in the laboratory in order to associate them with known adults and this has enabled the first
nymphal descriptions of the following species: Oemopteryx fosketti, Triznaka signata, Suwallia
lineosa, Isoperla decolorata, Nemoura rickeri, Malenka californica, Podmosta delicatula, Capnia
coloradensis, Capnia confusa and Capnia gracilaria . Keys are provided for most nymphs and all
adults except females of Isogenoides frontalis and Isogenoides colubrinus.
The life cycle classification devised by Hynes (1961) was used to categorize the life histories of
several Saskatchewan Plecoptera. Both univoltine and non-univoltine life cycles are evident in
Saskatchewan species with univoltine types ranging from slow with an early emergence (Si) to fast
with a late emergence (F4). Winter stoneflies all seem to be in type Si in which an embryonic
diapause is absent and adults emerge in late winter and early spring. The majority of non-winter
species appear to belong to type S2 in which embryonic diapause is also absent but adult emergence
is later in May and June. Few species have been classified as fast seasonal types with an embryonic
diapause. At present, life cycle types S4 and F3 are unknown in Saskatchewan species. Only 13 of
the 41 stonefly species known in the province have been definitely classified, indicating the large
amount of research required to determine all the life cycles of Saskatchewan Plecoptera.
All or most of the present-day Saskatchewan stonefly fauna originated with the retreat of the
Wisconsin Laurentide Ice Sheet and the subsequent recolonization of the province by species from
ice-free regions to the north and south. Five probable origins have been proposed for the extant
fauna. The Cypress Hills fauna has its affinities with the Rocky Mountains and was derived by a
connective bridge of flora and fauna which formed either after the Wisconsin or following earlier
glaciations. The fauna to the extreme north of the province has tundra elements and is likely
derived from a northwestern refugium. The boreal fauna is derived from southern North American
refugia in the northeast and northwest and the prairie fauna is derived chiefly from the
southcentral North American refugium.
ACKNOWLEDGEMENTS
Thanks are given to Mr. E. Driver and Mr. D.H. Smith for reviewing certain sections of this
manuscript and for providing helpful suggestions.
Special thanks are extended to Mr. Doug Smith who assisted in numerous aspects of this study
but especially in collecting and in sharing his knowledge of various techniques in rearing and in
illustrating.
Grateful appreciation is extended to John Waddington, Elaine Dribnenky and Alex Campbell
who assisted in various aspects of preparing photographs and drawings.
Several persons made their collections freely available for study, and to them my sincere
appreciations are extended: Mr. Ron Demaray for some Cypress Hills specimens.
Dr. W.W. Sawchyn for Churchill River and Cluff Creek specimens. Dr. D.J. Larson for
McDougal Creek and Scarth River specimens, and Mr. L. Kratt for Stoney Rapids material.
Special thanks are given to Margaret Height for assistance in the laboratory. We also thank
Mr. Peter Mason for collecting assistance.
For financial assistance received during the course of this study appreciation is extended to the
Institute for Northern Studies for a research scholarship. The study was also financed in part by a
Quaest. Ent., 1979, 15(1)
72
Dosdall& Lehmkuhl
National Research Council grant held by Dr. D.M. Lehmkuhl.
Sincere gratitude is extended to Dr. W.E. Ricker of the Fisheries Research Board of Canada,
Nanaimo, B.C., for verifying most of the species identifications and for the loan of some specimens
which were used for the drawings. For verification of Nemouridae material, we thank
Dr. R.W. Baumann of Brigham Young University, Provo, Utah.
Finally, special thanks are extended to Teresa Height-Dosdall for freely devoting much of her
time to several aspects of the study, but especially for assistance in collecting, preparing the plates,
typing, and for making innumerable sacrifices so that this project could be completed.
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Stoneflies of Saskatchewan
79
Figures 1-5. Nymphs of Perlidae. Fig. 1, Acroneuria abnormis; Fig. 2, A. lycorias; Fig. 3, Hesperoperla pacifica; Fig. 4,
Claassenia sabulosa; Fig. 5, Paragnetina media.
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Fig. 6-17. Fig. 6, Nymph of Pteronarcys dorsata (ventral) showing gills on the first two abdominal segments; Fig. 7, Nymph of
Pteronarcella hadia (ventral) showing gills on the first three abdominal segments; Fig. 8, Terminal abdominal segments of
Claassenia sabulosa (male, dorsal) showing hooks on the tenth tergite; Fig. 9, Terminal abdominal segments of A croneuria
abnormis (male, ventral) showing hammer (h); Fig. 10, Adult of Zapada cinctipes (lateral) showing cervical gill remnants;
Fig. 1 1, Nymph of Z. cinctipes (ventral) showing cervical gills; Fig. 12, Nymph of Amphinemura Undo (ventral) showing
cervical gills; Fig. 13, Nymph of Taeniopteryx nivalis (ventral) showing coxal gill (eg); Fig. 14, Nymph of Z. cinctipes (dorsal);
Fig. 15, Nymph of Capnia gracilaria (dorsal); Fig. 16, Rearing apparatus; Fig. 17, Rearing container.
Stoneflies of Saskatchewan
81
Figures 18-30. Fig. 18, Pro thoracic and meso thoracic sterna of Isocapnia crinita, showing the basisternum (Bs) and presternum
(prs); Fig. 19, Prothoracic and meso thoracic sterna of Paracapnia angulata, showing basisternum (Bs) and presternum (prs);
Fig. 20, Nymphal labium of Isogenoides colubrinus, showing paraglossa (pgl), glossa (gl), submentum (sm) and submental
gill (smg); Fig. 21, Nymphal labium of Leuctra ferruginea, showing paraglossa (pgl), glossa (gl) and labial palpus (Ip); Fig. 22,
Nymphal hindleg of Oernopteryx fosketti; Fig. 23, Nymphal hindleg of Capnia gracilaria; Fig. 24, Nymphal meso thoracic ridge
pattern of Isogenoides colubrinus, showing the furcal pit (fp); Fig. 25, Nymphal mesothoracic ridge pattern of Skwala parallela,
showing the furcal pit (fp); Fig. 26, Distal cereal segments of Isocapnia crinita nymph; Fig. 27, Nymphal abdomen of Leuctra
ferruginea; Fig 28, Nymphal mandible of Skwala parallela; Fig. 29, Nymphal labium of Zapada cinctipes; Fig. 30, Nymphal
maxilla of Diura bicaudata.
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32 H. brevis
Figures 31 — 39. Wings of Plecoptera. Fig. 31, Isoperla bilineata; Fig. 32, Hastaperla brevis; Fig. 33, Paracapnia angulata; Fig. 34,
Paraleuctra vershina; Fig. 35, Skwala parallela; Fig. 36, Leuctra ferruginea; Fig. 37, Capnia confusa; Fig. 38^ Isogenoides colubrinus;
Fig. 39, Isocapnia crinita.
Stoneflies of Saskatchewan
83
Figures 40 - 49. Genitalia of Taeniopterygidae and Pteronarcidae. Fig. 40, Taeniopteryx nivalis (male, lateral); Fig. 41, T. nivalis
(male, ventral with 9th stemite removed to show aedeagus and subanal lobes); Fig. 42, Oemopteryx fosketti (male, dorsal);
Fig. 43, T. nivalis (male, dorsal); Fig. 44, T. nivalis (female, vehtral); Fig. 45, O. fosketti (female, ventral); Fig. 46, Pteronarcys
dorsata (male, dorsal); Fig. 47, 48, P. dorsata (female, ventral showing variation in subgenital plates); Fig. 49, Pteronarcella badia
(female, ventral).
Quaest.Ent., 1979, 15(1)
Dosdall & Lehmkuhl
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50 U. trava
51 U. trava
53 c gracilaria
54 C. confusa
55 C. confusa
58 C. coloradensis
59 C. coloradensis
60 P- angulata
61 P angulata
Figures 50-61. Male genitalia of Capniidae. Fig. 50, Utacapnia trava (dorsal); Fig. 51, U. trava (lateral); Fig. 52, Capnia gracilaria
(dorsal); Fig. 53, C graciliaria (lateral); Fig. 54, C. confusa (dorsal); Fig. 55, C. confusa (lateral); Fig. 56, C vernalis (dorsal);
Fig. 57, C. vernalis (lateral); Fig. 58, C. coloradensis (dorsal); Fig. 59, C. coloradensis (lateral); Fig. 60, Paracapnia angulata (dorsal);
Fig. 61, P. angulata (lateral).
Stoneflies of Saskatchewan
85
63 /- crinita
64 /. crinita
65 I. crinita
66 /. missourii
67 C. vernaiis
68 P- angulata
69 U. trava
70 C. confusa
71 C. coioradensis
72 C. graciiaria
Figures 62 - 72. Genitalia of Capniidae. Fig. 62, Isocapnia crinita (male, ventral); Fig. 63, /. crinita (male, dorsal); Fig. 64, /. crinita
(male, lateral); Fig. 65, 1, crinita (female, ventral); Fig. 66, 1. missourii (female, ventral); Fig. 67, Capnia vernaiis (female, ventral);
Fig. 68, Paracapnia angulata (female, ventral); Fig. 69, Utacapnia trava (female, ventral); Fig. 70, Capnia confusa (female, ventral);
Fig. 71, C coioradensis (female, ventral). Fig. 72, C. graciiaria (female, ventral).
Quaest.Ent., 1979, 15(1)
86
Dosdall& Lehmkuhl
77 A- linda 78 A. linda
79 N. rickeri
80 N. rickeri
81 P. vershina
82 P. vershina 83 P. vershina 84 L ferruginea
Figures 73 - 84. Male genitalia of Nemouridae and Leuctridae. Fig. 73, Malenka californica (dorsal); Fig. 74, Zapada cinctipes
(dorsal); Fig. 75, Shipsa rotunda (dorsal); Fig. 76, Podmosta delicatula (dorsal); Fig. 77, Amphinemura linda (dorsal); Fig. 78,
A. linda (ventral); Fig. 79, Nemoura rickeri (dorsal); Fig. 80,7V. rickeri (ventral); Fig. 81, Paraleuctra vershina (dorsal); Fig. 82,
P. vershina (ventral); Fig. 83, P. vershina cercus (lateral); Fig. 84, Leuctra ferruginea (dorsal).
Stoneflies of Saskatchewan
87
85 M. californica
86 Z. cinctipes
87 S. rotunda
88 A. linda
S9 N. rickeri
90 P. delicatula
92 L. ferruginea
Figures 85 - 92. Female genitalia of Nemouridae and Leuctridae. Fig. 85, Malenka californica (ventral); Fig. 86, Zapada cinctipes
(ventral); Fig. 87, Sliipsa rotunda (ventral); Fig. 88, Amphinemura linda (ventral); Fig. 89, Nemoura rickeri (ventral); Fig, 90,
Podmosta delicatula (ventral); Fig. 91, Paraleuctra vershina (ventral); Fig. 92, Leuctra ferruginea (ventral).
Quaest.Ent., 1979, 15(1)
Dosdall& Lehmkuhl
88
96 H. brevis
97 T. signata
98 S. lineosa
Figures 93 - 98. Genitalia of Chloroperlidae. Fig. 93, Hastaperla brevis (male, dorsal); Fig. 94, Triznaka signata (male, dorsal)
Fig. 95, Suwallia lineosa (male, dorsal); Fig, 96, H. brevis (female, ventral); Fig. 97, T. signata (female, ventral); Fig. 98, S. lineosa
(female, ventral).
Stoneflies of Saskatchewan
89
104 H. pacifica
105 P- media
Figures 99 - 106. Genitalia of Perlidae. Fig. 99, Acroneuria abnormis (female, ventral); Fig. 100, A lycorias (female, ventral);
Fig. lOl, Hesperoperla pacifica (male, dorsal); Fig. 102,^1. lycorias (male, dorsal); Fig. 103, /4. abnormis (male, dorsal); Fig. 104,
H. pacifica (female, ventral); Fig. \Q5 , Paragnetina media (female, ventral); Fig. 106, Claassenia sabulosa (female, ventral).
Quaest.Ent., 1979, 15(1)
114 S. parallela
115 A compacta
116 /. colubrinus
Figures 107 - 116. Genitalia of Perlodinae and Isogeninae. Fig. 101, Diura bicaudata (male, dorsal); Fig. 108, 109, D. bicaudata
(female, ventral, showing variation in subgenital plates); Fig. 110, Isogenoides colubrinus (male, dorsal); Fig. 111,7. frontalis
(male, dorsal); Fig. 112, Skwala parallela (male, dorsal); Fig. \ \2>, Arcynopteryx compacta (male, dorsal); Fig. 114, S. parallela
(female, ventral); Fig. 115,^4. compacta (female, ventral); Fig. 116, 7 colubrinus (female, ventral).
Stoneflies of Saskatchewan
91
119 /. transmarina
Figures 117 — 124, Male genitalia of Isoperlinae. Fig. Ill , Isoperla bilineata (dorsal); Fig. 118,7. bilineata (ventral); Fig. 119,
I. transmarina (dorsal); Fig. 120, /. longiseta (dorsal); Fig. 121, 7. decolorata (lateral, cerci removed); Fig. 122, 7. patricia (dorsal);
Fig. 123, 7. petersoni (dorsal); Fig. 124, Aedeagal process of 7. petersoni (lateral).
Quaest . Ent 1979, 15(1)
92
Dosdall& Lehmkuhl
Figures 125 — 1 37. Female genitaUa of Isoperlinae. Fig. 125 - 128, Isoperla bilineata (ventral, showing variation in subgenital plates);
Fig. 129 - 132,/. longiseta (ventral, showing variation in subgenital plates); Fig. 133,/. decolorata (ventral); Fig. 134,/. petersoni
(ventral); Fig. 135,/. transmarina (ventral); Fig. 136,/. patricia (ventral); Fig. 137,/. marlynia (ventral).
Stoneflies of Saskatchewan
93
140 C. coloradensis
Figures 138 - 144. Nymphal setation and maxillae of Capniidae. Only the setae which can be seen in profile on the top of the
head, on the pronotum and on the abdomen have been indicated; the eye is outlined as a point of reference. Fig. 138, Capnia
gracilaria (male); Fig. 139, C. confusa (male); Fig. 140, C. coloradensis (female); Fig. 141, Maxilla of C vernalis; Fig. 142,
Maxilla of C. confusa; Fig. 143, Maxilla of C. gracilaria; Fig. 144, Maxilla of C. coloradensis.
Quaest.Ent., 1979, 15(1)
94
Dosdall & Lehmkuhl
Figures 145 - 153. Setation of Nemouridae nymphs. Fig. 145, Hindleg of Nemoura rickeri; Fig. 146, Right half of pronotum of
N. rickeri; Fig. 147, Left half of pronotum of Malenka californica; Fig. 148, Terminal abdominal tergites of TV. rickeri (male);
Fig. 149, Terminal abdominal tergites of M. californica (female); Fig. 150, Terminal abdominal tergites of Podmosta delicatula
(male); Fig. 151, Cercus of TV. rickeri (lateral); Fig. 152, Cercus of P. delicatula (lateral) ; Fig. 153, Cercus ofM. californica
(lateral).
Stoneflies of Saskatchewan
95
Figures 154 — 158. Nymphal mouthparts, basal antennal segments and abdominal setation of Oemopteryx fosketti. Fig. 154,
Maxilla; Fig. 155, Color pattern on basal antennal segments; Fig. 156, Mandible; Fig. 157, Labium; Fig. 158, Terminal abdominal
segments indicating bristles seen in profile (male).
Quaest.Ent., 1979, 15(1)
96
Dosdall& Lehmkuhl
Figures 159 - 162. Nymphal color pattern and mouthparts of Triznaka signata. Fig. 159, Mature nymph showing color pattern;
Fig. 160, Mandible; Fig. 161, Labium; Fig. 162, Maxilla and maxillary palpus (mp).
Stoneflies of Saskatchewan
97
Figures 163 - 167. Adult head patterns and nymphal mouthparts of Chloroperlidae. Fig. 163, Head pattern oi Suwallia lineosa;
Fig. 164, Head pattern of Triznaka signata; Fig. 165, Maxilla of S. lineosa; Fig. 166, Mandible of S. lineosa; Fig. 167; Labium of
S. lineosa.
Quaest.Ent., 1979, 15(1)
98
Dosdall & Lehmkuhl
Figures 168 - 170. Nymphal color pattern and mouthparts of Isoperla decolorata. Fig. 168, Mature nymph showing color pattern;
Fig. 169, Mandible; Fig. 170, Maxilla with maxillary palpus (mp).
Stoneilies of Saskatchewan
99
Figures 171 - 176. Adults, nymph and habitat oi Oemopteryx fosketti under field conditions. Fig. 171, Two males actively
searching out females on the snow near the North Saskatchewan River; Fig. 172, Female; Fig. 173, Two males competing for a
female; the male nearest the female had begun mating, and upon the arrival of the second mal? , a struggle ensued until one was
driven away; Fig. 174, Mating pair; Fig. 175, Mature male nymph; Fig. 176, North Saskatchewan River partially ice-covered near
Borden Bridge (Hwy. 5) at emergence time of O. fosketti.
Quaest . Ent 1979, 15(1)
100
Dosdall & Lehmkuhl
177
Fig. 177, Saskatchewan records for Capnia coloradensis and C. gracilaria.
Stoneflies of Saskatchewan
101
Fig. 178, Saskatchewan records for Capnia vernalis, Isocapnia missourii and Utacapnia trava.
Quaest.Ent., 1979, 15(1)
102
Dosdall & Lehmkuhl
179
Fig. 179, Saskatchewan records for Capnia confusa, Isocapnia crinita and Paracapnia angulata.
Stoneflies of Saskatchewan
103
Fig. 180, Saskatchewan records iot Pteronarcys dorsata and Pteronarcella badia.
Quaest.Ent., 1979, 15(1)
104
Dosdall & Lehmkuhl
Stoneflies of Saskatchewan
105
Fig. 182, Saskatchewan records for Amphinemura Undo, Podmosta delicatula and Malenka callfornica.
Quaest.Ent., 1979, 15(1)
106
Dosdall& Lehmkuhl
Fig. 183, Saskatchewan records for Nemoura rickeri, Zapada cinctipes and Shipsa rotunda.
Stoneflies of Saskatchewan
107
Fig. 184, Saskatchewan records iox Paraleuctra vershina and Leuctra ferruginea.
Quaest.Ent., 1979, 15(1)
108
Dosdall & Lehmkuhl
Fig. 185, Saskatchewan records ior Hastaperla brevis, Suwallia lineosa and Triznaka signata.
Stoneflies of Saskatchewan
109
Fig. 186, Saskatchewan records for Acroneuria lycorias, A. abnormis and Hesperoperta pacifica.
Quaest.Ent., 1979, 15(1)
no
Dosdall & Lehmkuhl
Fig. 187, Saskatchewan records for Paragnetina media, Claassenia sabulosa and Perlesta placida.
Stoneflies of Saskatchewan
111
Fig. 188, Saskatchewan records ioi Arcynopteryx compacta and Skwala parallela.
Quaest.Ent., 1979, 15(1)
112
Dosdall& Lehmkuhl
Fig. 189, Saskatchewan records for Isogenoides colubrinus, /. frontalis and Diura bicaudata.
Stoneflies of Saskatchewan
113
Fig. 190, Saskatchewan records for Isoperla bilineata and/, patricia.
Quaest.Ent., 1979, 15(1)
114
Dosdall & Lehmkuhl
Fig. 191, Saskatchewan records foi Isoperla marlynia and/, longiseta.
Stoneflies of Saskatchewan
115
Fig. 192, Saskatchewan records fox Isoperla transmarina, I. decolorata and/. petersonL
Quaest.Ent., 1979, 15(1)
116
Dosdall& Lehmkuhl
C. con f us a
Fig. 193, Seasonal succession of adults of some species of Plecoptera in Saskatchewan. Extreme dates of capture of adults in
1974 - 1976.
UPPER OLIGOCENE FOSSIL PUPAE AND LARVAE OF CHAOBORUS TERTIARIUS
(VON HEYDEN) (CH AOBORIDAE, DIPTERA) FROM WEST GERMANY
ART BORKENt
Department of Entomology
University of Alberta
Edmonton, Alberta, Canada
T6G2E3
Corrigenda. A. Borkent. Quaest. Ent. 14(4): 491-496.
page/ line
491 / 44 Change Culcites tertiarius von Heyden to Culicites tertiarius von Heyden
* Present Address: Biosystematics Research Institute, Research Branch, Agriculture Canada,
Ottawa, Ontario, Canada K1 A 0C6.
119
Editorial - To Compute or Not Compute
There have been many attempts to cast Shakespeare’s plays in 20th century format, some
successful, others not. Recently, I have read various attempts at rewriting, in modern English,
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modified) was: “To compute or not compute, that is the go/no-go decision’’.
The quotation is appropriate to Quaestiones Entomologicae, as the text for this, and I hope for
other issues, was and will be produced using a computer textformatting program. Why change
from our time-tested methods of production? Proofs and satisfactory copy for Quaestiones
Entomologicae were previously produced on a special, highly sophisticated typewriter. This
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As textformatting programs were available through an existing computer facility at the University
of Alberta, it seemed logical to make use of them, if we were to continue publication of
Quaestiones Entomologicae.
The Michigan Terminal System is now used for typing manuscripts. Text is typed using the
editor mode and proofs are run off on a computer line printer. When all corrections are made, the
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The decision to switch to textformatting for production of Quaestiones Entomologicae was
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Acting Editor much easier. Also, a special tribute and thank you to D. Reinhart of Printing
Services, University of Alberta, for designing the textformatting layout that we are using for
Quaestiones Entomologicae.
The quality of an editor’s life is further enhanced by those willing to review manuscripts and I
thank the following for reviewing papers in Volume 14 and this issue of Volume 15: G.E. Ball,
W.G. Evans, L.H. Herman, J. Kukalova-Peck, A.P. Nimmo and D.R. Whitehead. Thanks also to
H. Goulet for translating the abstracts into French.
Unfortunately, I have to announce an increase in page charges. We have always held these
charges as low as possible and they have been $1 1.00 per page for the past five years. However,
increases in production and printing costs now require a charge of $15.00 per page. Costs of
reprints will be $3.00 per page per 100 copies.
Until George Ball resumes editorship, I intend to maintain the standards set by him. Some
people no doubt are natural editors, others like me have the responsibility thrust upon them. I have
much to learn. I certainly have a greater appreciation now of the steps involved in getting a
manuscript published. In fact all who publish should serve time as an editor as it would improve
the quality of their subsequently submitted manuscripts.
Douglas A. Craig
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1
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
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Edmonton^ Alberta, Canada T6G 2E3
Quaestiones
Entomologicae
A periodical record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.
VOLUME 15
NUMBER 2
APRIL 1979
QVAESTIONES ENTOMOLOGICAE
ISSN 0033-5037
A periodical record of entomological investigation published at the Department of Entomology,
University of Alberta, Edmonton, Alberta.
Volume 15 Number 2 April 1979
CONTENTS
Borkent - Systematics and bionomics of the species of the subgenus Schadonophasma Dyar and
Shannon {Chaoborus, Chaoboridae, Diptera) 122
Jones and Heming - Effects of temperature and relative humidity on embryogcnesis in eggs of
Mamestra configurata (Walker) (Lepidoptera: Noctuidae) 257
FRONTISPIECE
i
!
J
J
(
■i
j
Chaoborus trivittatus adult male.
SYSTEMATICS AND BIONOMICS OF THE SPECIES OF THE SUBGENUS
SCHADOMOPHASMA DYAR AND SHANNON {CHAOBORUS, CHAOBORIDAE, DIPTERA)i
ART BORKENr-
Department of Entomology
University of Alberta
Edmonton, Alberta, Canada Quaestiones Entomologicae
T6G2E3 15: 122-255 1979
Study of structure, intraspecific variation, life cycles and bionomic features suggests the presence of
three species of the subgenus Schadonophasma. Chaoborus trivittatus (Loew) and Chaoborus cooki
Saether are restricted to North America while Chaoborus nyblaei (Zetterstedt) is known only from
Eennoscandia. Chaoborus brunskilli Saether and Chaoborus knabi (Dyar) are conspecific with C.
trivittatus
All stages of each species were studied except first, second and third larval instars of C. nyblaei
which are unknown. Eggs of C. cooki and C. nyblaei, unlike those of C. trivittatus, exhibit a thickened
exochorion. C. cooki eggs are laid in a spherical mass with little gelatinous matrix while those of C.
trivittatus are laid in a spiral arrangement in a disc of gelatinous matrix. Eirst instar larvae of C. cooki
possess a more pronounced egg burster than do those of C. trivittatus. All other stages can be
distinguished only by using a combination of characters. A compound character index is provided for
separation of fourth instar larvae of C. trivittatus and C. cooki and was used to test possibilities of
conspecificity of these two species. Male adults can be identified by the shape of the penis valves and a
ratio of the length of two wing veins.
The thickened exochorion of eggs of C. cooki, and probably C. nyblaei, is an adaptation to
overwintering as eggs in temporary ponds. C. cooki is univoltine. C. trivittatus immatures occur mostly
in permanent lentic habitats where this species overwinters as a fourth instar larva. This species may be
uni- or multivoltine or have a two year life cycle.
Behavioural differences are evident between C. trivittatus and C. cooki. Only C. cooki larvae are
capable of ingesting ostracods. Male adults of C. trivittatus form large swarms. Adult females are the
main dispersing agent of C. trivittatus while limited evidence for C. cooki suggests that both sexes of
this species disperse. Under laboratory conditions C. trivittatus male adults live up to seven to eight
days and female adults up to twelve days.
The pattern of intraspecific variation indicates that some of the variation of fourth instar larvae is
due to sex-associated, age-related and geographical variation. Geographical variation of characters of
fourth instar larvae and geographical distribution of C. trivittatus indicates that this species was
present in the Beringian refugium and those refugia south of the continental ice sheet during the
Wisconsin glaciation.
A reconstructed phylogeny of some Chaoborus species is provided. The subgenera Schadonophasma
and Chaoborus s. str. are both monophyletic and are sister groups. An upper Oligocene Chaoborus
fossil indicates the minimum age of the speciation events which gave rise to the ancestor of these two
groups, to be 25 million years.
^Based on a thesis submitted in partial fulfillment of the requirements for the degree of Master of
Science at the University of Alberta, Edmonton, Canada.
^Present address: c/o D.M. Wood, Biosystematics Research Institute, Research Branch, Agriculture
Canada, Ottawa, Ontario, Canada KIA 0C6
124
Borkent
A reconstructed phylogeny of Schadonophasma species indicates that C. cooki and C. nyblaei are
more closely related to each other than either is to C. trivittatus. Zoogeographic considerations suggest
two possible hypotheses for the origin of these species. The first invokes allopatric speciation between
the Nearctic and Palaearctic regions producing C. trivittatus and the ancestor of C. cooki and C.
nyblaei. Subsequent geographical isolation of populations of the latter lineage produced C. cooki and
C. nyblaei. The second hypothesis differs in proposing that the C. cooki-nyblaei lineage arose in
sympatry with C. trivittatus from individuals of a C. trivittatus-///:^ ancestor that involved an
overwintering egg stage which allowed exploitation of temporary habitats. A model for the
development of reproductive isolation in sympatry suggests the occurrence of a mutation resulting in a
diapausing egg in the ancestral population. Through temporal disjunction of mating periods,
homogamic mating, complete selection against overwintering larvae in temporary ponds and
overwintering eggs in permanent ponds, the two lineages which gave rise to C. trivittatus and the
ancestor of C. cooki-nyblaei became genetically isolated.
L’etude de la morphologie, des variations intraspecifiques, des cycles vitaux et de I’histoire naturelle suggere que le sous-genre
Schadonophasma consiste en trois esp'eces. Chaoborus trivittatus (Loew) et Chaoborus cooki Saether ne se recontrent qu’en Amerique
du Nord, alors que Chaoborus nyblaei (Zetterstedtj n’est connu que de la Scandinavie. Je considere Chaoborus brunskilli Saether et
Chaoborus knabi (Dyar) comme appartenant a C. trivittatus.
J’ai etudie tous les stades de chaque espece, sauf les premier, deuxieme et troisi'eme ages larvaires de C. nyblaei, qui sont
inconnus. Les oeufs de C. cooki et de C. nyblaei, contrairement a ceux de C. trivittatus, ont un exochorion epais. Les oeufs de C. cooki
sont deposes en une masse spherique entouree d’un peu de matiere gelatineuse, alors que ceux de C. trivittatus sont deposes en
spirale dans un disque de matiere gelatineuse. Les larves du premier age de C. cooki. ont un ovorupteur plus prononce que celles de
C. trivittatus. Tous les autres stades ne peuvent etre distingues que par des combinaisons de caracteres. Un index, defini par de telles
combinaisons, permet de separer les larves du quatrieme age de C. trivittatus et de C. cooki, et a ete utilise pour verifier que ces deux
esp'eces etaient bien distinctes Tune de Tautre. Les adultes males peuvent etre identifies par la forme des valves de Tedeage et par un
rapport entre les longueurs de deux des nervures alaires.
L’exochorion epais des oeufs de C. cooki, et probablement de C. nyblaei, est une adaptation pour Thibernation au stade d’oeuf
dans les etangs temporaires. C. cooki n'a qu’une seule generation par annee. Les immatures de C. trivittatus se rencontrent surtout
dans les etangs permanents oil les larves du quatrieme age hibernent. Cette espece peut avoir une ou plusieurs generations par annee,
ou avoir un cycle vital de deux ans.
II y a plusieurs differences entre le comportement de C. trivittatus et celui de C. cooki. Seules les larves de C. cooki peuvent se
nourrir d’ostracodes. Les adultes males de C. trivittatus s’assemblent en grands essaims. Les adultes femelles constituent le principal
agent de dispersion de C. trivittatus, alors que chez C. cooki, il semblerait que les deux sexes participent a la dispersion. En
laboratoire, les males de C. trivittatus vivent jusqu’d sept ou huit jours, alors que les femelles vivent Jusqu’d douze Jours.
La variabilite intraspecifique chez les larves du quatrieme age serait reliee au sexe et d Edge, ainsi qu'd des facteurs
geographiques. La variation geographique des caracteres du quatrieme age larvaire et la repartition geographique de C. trivittatus
indiquent que cette espece etait presente dans le refuge de Bering, ainsi que dans le refuge situe au sud de la calotte glaciaire
continentale durant la glaciation du Wisconsin.
J’ai reconstruit la phylogenie de quelques especes de Chaoborus. Les sous-genres Schadonophasma et Chaoborus s. str. sont
monophyletiques et apparentes. Un fossile de Chaoborus datant de TOligocene superieur suggere que le processus de speciation
ayant produit Tancetre de ces deux sous-genre a eu lieu il y a au moins 25 millions d'annees.
Une reconstruction de la phylogenie des especes de Schadonophasma montre que C. cooki et C. nyblaei sont plus etroitement
apparentes Tun d Tautre que chacun d'eux ne Test d C. trivittatus. Une etude biogeographique suggere deux hypotheses quant d
Torigine de ces esp'eces. La premiere, basee sur le principe d’evolution allopatrique, suggere que C. trivittatus et Tancetre de C.
cooki-nyblaei resulterent d’une separation de la population de leur ancetre commun en deux populations, une nearctique et une
palearctique. Une separation geographique subsequente de la population palearctique aurait produit C. cooki et C. nyblaei. Suivant
la seconde hypoth'ese, la lignee C. cooki-nyblaei aurait evolue en sympatrie avec C. trivittatus, et serait derivee d’individus
semblables d Tancetre de C. trivittatus, mais hibernant au stade d’oeuf, cette derniere adaptation ayant permis Texploitation des
habitats temporaires. J’explique Tevolution de Tisolement reproducteur entre ces deux lignees sympatriques comme suit: une
mutation aurait introduit la diapause au stade d’oeuf dans la population ancestrale; grace d une separation temporelle des periodes
reproductrices, permet t ant ainsi des accouplements homogamiques, et grace d une selection tot ale contre Thibernation d Tetat
larvaire dans les etangs temporaires d’une part, et contre Thibernation d Tetat d’oeuf dans les etangs permanents d’autre part, les
deux lignees qui produisirent C. trivittatus et Tancetre de C. cooki-nyblaei devinrent genetiquement isolees.
Systematics and Bionomics of Schadonophasma
125
TABLE OF CONTENTS
Introduction. 125
Materials and Methods 127
Classification 135
Analysis of Morphological Variation 1 55
Artificial variation 155
Characters varying between larval instars 1 56
Variation of characters of fourth instar larvae 161
Variation of characters of pupae 176
Variation of characters of male adults 1 77
Variation of characters of female adults 178
Descriptions of populations Incertae sedis 180
Bionomics 183
Lifecycle 183
Development and behavior 187
Mating Experiments 193
Phylogeny and Zoogeography 194
Evidence and recognition of three species of Schadonophasma 195
Phylogeny of Chaoborus 196
Fossil evidence 199
Phylogeny of Schadonophasma 200
Zoogeography and speciation events 201
Concluding Remarks 203
Acknowledgements 204
References 205
INTRODUCTION
For systematists interested in understanding the intraspecific and interspecific variation of species, a
frequent concern is the investigation of species which do not exhibit discrete morphological
characteristics allowing identification of individuals. This study is partially an investigation of such a
problem within the genus Chaoborus Lichtenstein. I first recognized that it was difficult to separate
species of the subgenus Schadonophasma Dyar and Shannon, while attempting to identify fourth instar
larvae, pupae and adults using Saether’s (1972) key. Some specimens exhibited combinations of
characters from different species, as delimited by Saether (1970, 1972). I therefore undertook a
reinterpretation of variation exhibited by members of this subgenus through examination of the
structural and bionomic features of eggs, all stages of larvae, pupae and adults.
Although Chaoborus is assigned species that have marked variation and show slight morphological
differences between species, few studies have attempted to precisely interpret this variation. Saether
(1967) showed that the characters previously thought to separate Chaoborus flavicans (Meigen) and
Chaoborus alpinus Peus feature overlapping variation and consequently he considered these forms to be
eonspecific. However, overlap of variation alone is not evidence of reproductive unity. Detailed analysis
of character variation, distributional data and bionomic information yields superior data from which to
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Borkent
decide species status. This study employs all three sources of data.
However recognition of members of different species is only the initial step permitting systematists to
investigate species. I therefore further undertook study of intraspecific variation of the species, including
analysis of sex-associated, age-related and geographical variation, and differences between larval instars.
Sources of variation of individual characters of Chaoborus species are not well understood. Sikorowa
(1973) showed that some variation of Chaoborus flavicans can be considered ecotypic and Cook (1956:
32) suggested the same for Chaoborus punctipennis (Say) larvae. Variation in number of mandibular
fan bristles, ratio of length to width of prelabral appendages, total length (Smith, 1960a: 398) and air
sac length (von Frankenberg, 1915: 514) of fourth instar Chaoborus crystallinus (De Geer) larvae is
sex-associated. Total length and color of Chaoborus americanus (Johannsen) fourth instar larvae is also
sex-associated (Bradshaw, 1973: 1249) as well as reflecting a polymorphism. Roth (1967: 66) noted
worn prelabral appendages of overwintered fourth instar larvae of Chaoborus albatus Johnson. Seasonal
differences of total length of mature fourth instar larvae of Chaoborus punctipennis was reported by
Muttkowski (1918: 407). Chaoborus punctipennis adult size (Cook, 1956: 32) and number of
mandibular fan bristles of Chaoborus anomalus Edwards fourth instar larvae (McGowan, 1972: 365)
may vary geographically. All these studies suggest possible sources of variation and I have attempted to
provide interpretations of variation of characters of species of Schadonophasma in light of these previous
studies.
Systematic studies should be based on evolutionary theory and the aforementioned analyses provide a
sufficient background from which to hypothesize the evolutionary history of the subgenus
Schadonophasma and to also propose a possible example of sympatric speciation. An interpretation of
the phylogeny of some Chaoborus species is provided.
Because of their importance as fish food (Goldspink and Scott, 1971; Green et al., 1973; Kruger,
1973; Lindquist et al. 1943; MacDonald, 1956; Pope et ai, 1973), as mosquito predators (Montchadsky,
1964; Sailer and Lienk, 1954; Skierska, 1969, 1974), in the prominent role they play in fresh water
communities (e.g. Bonomi, 1962; Fedorenko and Swift, 1972; Juday, 1921; Miyadi, 1932; Miyadi and
Hazema, 1932; Petr, 1972; Swiiste et al., 1973; Woodmansee and Grantham, 1961) and occasionally as
pests (Brydon, 1956; Herms, 1937; Hitchcock, 1965; Lindquist and Deonier, 1942a, 1942b; Lindquist et
al., 1951; MacDonald, 1953, 1956; Shemanchuk, 1959) many Chaoborus populations are better known
bionomically than systematically. For example, although Chaoborus flavicans has been studied
ecologically in detail (e.g. Balvay, 1977d; Parma, 1971b), no thorough distributional study has been
done on this species or any other species of the Holarctic region. This situation allows systematists
working on Chaoborus to take into account a wide array of ecological work and to draw systematic
inferences from such studies.
Chaoborus is composed of medium sized flies 2. 0-9. 3 mm in length. Adults of this genus are not
known to bite although the most primitive genus of the family, Corethrella Coquillett, includes species
with haematophagous adults (McKeever, 1977; Miyagi, 1974, 1975; Williams and Edman, 1968). In the
Holarctic region larvae may be commonly found in permanent and temporary lentic waters and
occasionally in rivers.
Chaoborus includes 37 described species in six subgenera {Neochaoborus Edwards, Sayomyia
Coquillett, Edwardsops Lane, Peusomyia Saether, Schadonophasma, and Chaoborus sensu stricto).
The genus has been recorded from most major land masses with the exception of New Zealand,
Antarctica, Greenland, and most oceanic islands. Edwards (1932) gave a world list of species and their
distribution. The Palaearctic species have been described most recently by Hirvenoja (1961), Martini
(1931), Parma (1969), Peus (1934), Prokesova (1959), and Sikorowa (1967a, 1973) and Nearctic
species by Dyar and Shannon (1924) and Cook (1956) who also provided a detailed morphological
Systematics and Bionomics of Schadonophasma
127
study. Saether (1970) gave the most recent revision of Palaearctic and Nearctic species and was first to
compare material of most species from both these regions. Palaearctic and Nearctic samples of
Chaoborus flavicans were studied by Saether (1967). Saether (1972) presented keys to fourth instar
larvae, pupae and adults of species of the Holarctic region and gave a synopsis of their way of life and
structure.
Lane (1953) and Saether (1976) have provided descriptions for most known species of Chaoborus in
South America. African species have been described by Edwards (1930), Green and Young (1976),
McGowan (1972, 1976), and Verbeke (1957, 1958) and Oriental and Australian species by Giles (1902,
1904), and Theobald (1901b, 1903). Fossil species have been described by Borkent (1978b), Edwards
(1923), von Heyden (1862), Meunier (1904), and Scudder (1890; questionable identification).
The objects of this study were species of the subgenus Schadonophasma, whose adults and larvae are
the largest of Chaoborus species. Species are restricted to northern boreal regions. Larvae of all species
sometimes occur in large numbers in permanent and temporary lentic waters. Adults and immatures are
poorly represented in museum collections.
Most descriptions are brief and bionomic information anecdotal (Allan, 1973; Anderson and
Raasveldt, 1974; Dickenson, 1944; Dodson, 1970; Dyar and Shannon, 1924; Edwards, 1932; Felt, 1904,
1905; Giles, 1902; Hamilton, 1971; James et al., 1969; James and Smith, 1958; Jenkins and Knight,
1950; Johannsen, 1903, 1934; Loew, 1862; Matheson, 1944, 1945; Pope et al., 1973; Smith, 1960b;
Theobald, 1901b; Tullock, 1934; Twinn et al., 1948; Zetterstedt, 1838, 1851). The feeding
characteristics and vertical migration of the larvae and the life cycle of Chaoborus trivittatus (Loew)
from Eunice Lake, British Columbia have been described by Fedorenko (1975a, 1975b, 1975c),
Federenko and Swift (1972), Swift (1975, 1976), and Swift and Fedorenko (1975).
Schadonophasma was erected as a subgenus by Dyar and Shannon (1924) to contain the North
American species Chaoborus trivittatus. Edwards (1932), Cook (1956), Matheson (1944), and Peus
(1967) considered Chaoborus nyblaei (Zetterstedt) to be conspecific with Chaoborus trivittatus.
Hirvenoja (1961), however, considered these to be two distinct species. Saether (1970) recognized, on
morphological grounds, Chaoborus nyblaei in Fennoscandia and Baffin Island, Canada, and Chaoborus
trivittatus, Chaoborus cooki Saether and Chaoborus brunskilli Saether in North America.
This study clarifies the above confusion concerning species of Schadonophasma.
MATERIALS AND METHODS
Materials
This study was based on examination of 293 eggs of Chaoborus cooki, more than 50 egg masses of
Chaoborus trivittatus, and of all species, 49 first instar, 58 second instar, 139 third instar, and 832
fourth instar larvae, 93 male and 80 female pupae or pupal exuviae and 31 1 male and 248 female adults.
Eggs and most adults were preserved in fluid and members of other stages and some adults were
mounted on microscope slides. Many more specimens of all stages, preserved in fluid, were studied. All
Schadonophasma material examined by Saether (1970), except the female pupa of Chaoborus
trivittatus, was reexamined. I have also examined material collected by Osten Sacken (MCZ) but,
because I could not interpret the locality labels, did not include the material in this study.
For comparative purposes (diagnostic and phylogenetic) I also examined various stages of other
species of Chaoborus, Mochlonyx Loew, and Eucorethra Underwood. (Table 1).
Because I wantpd to sample a wide geographical area and sampling lakes would have taken too much
time, I collected immatures only from smaller water bodies and depended on loans for material from
lakes. Place names in Alberta referred to in the text are indicated in Figure 1.
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128
Borkent
Table 1. Material other than Schadonophasma examined. ® - undescribed material.
Voucher specimens from this study and some of the material I collected are in the Stickland
Museum, Department of Entomology, University of Alberta, Canada. Other material collected is
deposited in museums as indicated in the list of localities at the end of species descriptions.
Nearly all immatures and about half of all adults were collected by myself, mostly in Alberta. The
remainder were borrowed from the following museums or persons designated in the text by the following
abbreviations. Names of curators are also given to facilitate inquiries about their collections.
ABOr A. Borkent, Department of Entomology, University of Alberta, Edmonton, Alberta,
Canada, T6G 2E3.
BMNH British Museum (Natural History), London, England, SW7 5BD; P.S. Cranston.
CAS California Academy of Sciences, San Francisco, California, USA, 94118; P.H. Arnaud.
CNC Biosystematics Research Institute, Research Branch, Agriculture Canada, Ottawa,
Ontario, KIA 0C6; D.M. Wood.
CUNY Department of Entomology, Cornell University, Ithaca, New York, USA, 14850;
L.L. Pechuman.
EFCo E.F. Cook, Department of Entomology, Fisheries and Wildlife, University of Minnesota,
St. Paul, Minnesota, USA, 55101.
FWI Freshwater Institute, 501 University Crescent, Winnipeg, Manitoba, Canada, R3T 2N6;
D.M. Rosenberg.
JDAl J.D. Allan, Department of Zoology, University of Maryland, College Park, Maryland,
USA, 20742.
MCZ Museum of Comparative Zoology, Cambridge, Massachusetts, USA, 02138;
J.F. Lawrence.
OASa O.A. Saether, Museum of Zoology, University of Bergen, N-5014 Bergen, Norway.
UASM Strickland Museum, Department of Entomology, University of Alberta, Edmonton,
Alberta, Canada, T6G 2E3; G.E. Ball.
USNM Systematic Entomology Laboratory, United States Department of Agriculture,
c/o United States National Museum, NHB 168, Washington, D.C., USA, 20560;
C.F. Thompson.
WSUP Department of Entomology, Washington State University, Pullman, Washington, USA,
99163; W.J. Turner.
ZMHF Zoological Museum, Division of Entomology, SF-00100, Helsinki 10, Finland;
B. Lindeberg.
Systematics and Bionomics of Schadonophasma
129
ZMLS Zoological Institute, Department of Systematics, Helgonavagen 3, S-223 62 Lund,
Sweden; H. Andersson.
Methods
Collection, preservation and dissection of material. - Egg masses of Chaoborus trivittatus, like those
of most other Chaoborus species, float at the surface of the water body in which they are laid. I collected
these by hand picking the floating masses from the water surface with soft-tipped tweezers and dropping
them into 70% ethanol. Although egg masses can be scooped from the water surface with a dip net, they
stick to the netting and attempts then made to pick them up result in their disintegration. This method
should be avoided if determinations of the number of eggs per egg mass are important.
Because Schadonophasma larvae concentrate in the shaded portions of ponds such as that provided
by Typha, surrounding trees, or willows, it is imperative that these areas be sampled. A common method
of collecting larvae is to briskly sweep an aquatic net through a water body. Many third and fourth
instar larvae are benthic during daylight hours, so stirring up the bottom is important to ensure adequate
samples of these stages.
First and second instar larvae, because of their small size, are best collected with a plankton net. In
this study, only ponds were sampled so that the plankton net, with a mesh size of 1 16 iim, was thrown
from the shore and hauled in. Contents of either aquatic or plankton net were washed out in a white
enamel pan with three to five centimeters of water to insure that all captured specimens were retrieved.
The immatures were then picked up with soft-tipped tweezers and dropped into 70% ethanol.
Pupae were collected in the same manner as larvae except that an overly brisk motion of either
plankton or aquatic net will result in specimens in which the pupal horns have been lost. Pupal exuviae
were commonly found on the leeward side of ponds and were collected by hand or aquatic net.
Adults were sampled by means of an aerial net. Females of Chaoborus trivittatus are sedentary and
although they can occasionally be collected with an aerial net, are best retrieved with an aspirator from
tree trunks and herbaceous plants at the periphery of ponds.
I attempted to collect adult Schadonophasma with light traps but with little success. Twenty-two
New Jersey light traps in and around Edmonton, Alberta from June 24 to September 4, 1974 yielded
only two female Chaoborus cooki but did yield 1065 adults of Chaoborus sensu stricto. A New Jersey
and CDC (Communicable Disease Centre) light trap beside a pond from which Chaoborus cooki adults
were emerging caught none. This experience corresponds to that found for mosquitoes, in which
woodland species are not attracted to lights but other species are (Bidlingmayer, 1971).
Quantitative samples of immatures for life cycle studies were obtained with the plankton net which
was thrown out at least six times in the same direction from the shore and dragged at least twice across
the bottom, resulting in the bottom being disturbed and thereby ensuring a representative sample. The
numbers of immatures in each sample were not meant to be temporally comparable, but were
interpreted to reflect only percentages of an instar present at a given time.
Material, with the exception of some pinned adults, was preserved in 70% ethanol. However,
retention in alcohol for long periods of time will fade specimens. Pinned specimens are required for
determination of adult coloration.
All immatures measured were prepared using the method described by Saether (1972) with the
following modifications. Larvae and pupal exuviae were not cleared in KOH as this is not necessary for
accurate measuring or examination and clearing has resulted in some faulty observations concerning
shape of the dorsal process of larvae. One antenna and both mandibles were removed from each larva
while in Canada Balsam to ensure retention of these parts, especially of younger instars whose
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130
Borkent
mouthparts are easily lost. All larvae were mounted laterally and all pupae and pupal exuviae were
mounted with their ventral surfaces up. Pupal exuviae should be mounted with glass chips supporting the
coverslip to prevent compression of the specimen. An alternative method is to mount the specimen in
Canada Balsam, which is allowed to dry for a few hours before application of the coverslip.
Schlee (1966) indicated the advantages of fluid preserved material for the examination of chironomid
adults. Mainly, dried material results in distortion of some key characters. As can be expected, this is
also true of chaoborid adults. Therefore, adult material which was to be examined in detail was mounted
in Canada Balsam. The two antennae, head, thorax, three left legs, wings and abdomen were
disarticulated. Heads, thoraces, and abdomens were cleared in hot 8% KOH, transferred to 15% acetic
acid for 15 minutes, and all parts treated consecutively for at least ten minutes each with 98% ethanol,
98% ethanol layered over cedarwood oil, cedarwood oil and Canada Balsam. Chaoborus adults need not
be so treated to be identified. Most adult material examined was pinned.
The abdomen of each pinned male, after clearing in hot 8% KOH, was placed in one of eight
arbitrarily numbered wells filled with glycerine on each of a number of 7.6 x 7.6 cm microscope slides.
One gonocoxite and gonostylus with the adjoining penis valve were dissected from the rest of the
abdomen and examined under both stereoscopic and compound microscopes. This ensured that no a
priori decisions were made concerning species differences. Maximum lengths of the penis valves were
estimated with the aid of a compound microscope at 200X magnification. After examination, abdomens
were stored in glycerine in a genitalia vial on the pin. Examination of the genitalia from different
orientations is imperative for the determination of interspecific differences of the penis valve.
Abdomens of some pinned females were treated with a weak solution (three to six crystals in three
drams of distilled water) of tri-sodium orthophosphate. This treatment restores dried, more or less
shrivelled parts to approximately natural size and also restores eggs, if present in the abdomen, to their
natural dimensions.
Number of eggs per female was determined using reared specimens killed in 70% ethanol. Eggs were
easily removed and counted while in this medium.
Rearing methods and mating experiments. - All live material was reared at 20 C and a long day
(18:6, light:dark) photocycle. Field collected cladocerans, copepods and chironomid larvae and
laboratory reared larvae of Culiseta inornata provided a continual food source for the larvae. Bearings
of larvae to pupae were completed with each larva in a separate Dominion Brand 473 ml wide mouthed
Mason jar filled with pond water to ensure that no larvae were lost due to cannibalism.
To determine length of pupal peiod, larvae were reared to adulthood in 22.7 liter aquarium filled with
pond water and an abundance of food. Pupae were removed on the day they appeared and kept in
separate Mason jars.
In general, rearings were completed from field collected fourth instar larvae, although earlier instars
were occasionally captured for rearing. Pupae were not used for rearings as capture often resulted in at
least one pupal horn being broken off and consequent reduced adult emergence (Parma 1971b; 46; pers.
obs.).
Adults could not be made to mate successfully in the laboratory except when decapitated males were
brought into contact with CO2 anaesthetized females, the method used for mating experiments. Adults
were reared from fourth instar larvae of Chaoborus trivittatus collected 2.4 km west of Edmonton,
Alberta, on 1 3-V- 1976, and 16 km west of Jasper, Alberta, on \ -1916, Chaoborus collected 1.4
km west of George Lake, Alberta, on 1 3-V- 1976. After mating, each female was placed in a three dram
vial with wet paper towelling and gauze was placed over the mouth of the vial. Few adults were used in
these experiments because of difficulty in synchronizing adult emergence of Chaoborus cooki and
Chaoborus trivittatus in the laboratory. For the purposes of this study only eggs which embryonated
were considered fertilized.
Systematics and Bionomics of Schadonophasma
131
Adult feeding experiment. - This experiment had two purposes; first, to see if adult Chaoborus
trivittatus drink water and second, to determine if water uptake affects longevity. Goff (1972) tested for
imbibition of water by adults of Chironomus riparius by coloring sucrose water with food coloring and
checking excreta for color changes. This method was used with Chaoborus trivittatus adults to
determine if imbibition of water took place.
Adult Chaoborus trivittatus were collected at 1500-1600 hours on May 6, 1976, 2.4 km west of
Edmonton, Alberta, with an aerial net and aspirator. They were transported to the laboratory in a
screened cage placed in a styrofoam container with wet paper towels. The adults were then
anaesthesized with CO2 and sorted into treatments by 2300 hours on the day of capture. Ten males were
placed in each 15 mason jars and four females in each of three jars. All were kept at 20 C with a long
day (18:6, L:D) photocycle. Each jar contained a 35x10 mm plastic petri dish containing 5 ml of
deionized water which was either: (1) clear and available, (2) 1% blue food coloring solution (Scott
Bathgate Ltd., Winnipeg, Toronto, Vancouver) and available, or (3) screened with netting to prevent the
flies from drinking. I assumed uniform humidity in all jars. Fifty males and four females were used in
each treatment. The low numbers of females used was due to their scarcity in the field. Some specimens
were lost during the experiments and are therefore not recorded in Figure 35.
Mortality, and the presence or absence of colored excreta, were recorded every 12 hours.
Measurements and statistics. - All measured specimens are listed at the end of species descriptions.
Most measurements were made with an ocular micrometer (120 divisions) through a Wild M20
compound microscope equipped with phase constrast. Total length of larvae and the Y/X wing ratio
were measured with an ocular micrometer (120 divisions) through a Wild M5 stereoscopic microscope.
Measurements were taken at as high a magnification as was practical to comply as far as possible with
Sokal and Rohlf(1969: 15).
Terms for structures follow those used by Cook (1956), Knight (1971a, 1971b) or Laffoon and
Knight (1971). It is likely that many structures of Chaoborus larvae are homologous to those of culcid
larvae. For example, what are termed here “antennal seta” and “antennal blades” are probably
homologous to the “antennal setae” of culicid larvae (Knight and Laffoon, 1971). However, until
detailed comparative studies are completed, I use traditional terms.
Measurements of larval structures were those recognized as taxonomically useful by Cook (1956),
Parma (1969), Saether (1970) and Sikorowa (1973). Although numbers of bristles in both mandibular
fans were counted, to reduce the consequences of possibly lost bristles, only the highest number was used
for analysis. Total length was measured as length of head capsule plus distance from the anterior edge of
the prothorax to the tip of the abdomen exclusive of the anal tubules. Anal fan setae which are broken
off at the bases can still be counted by noting the internal portion of the setae. Some anal fan setae of
some larvae were reduced as compared to the other setae and these were only counted if they were at
least 0.5 times as long as the longest anal fan seta. I attempted to study the chaetotaxy of larvae but
found most setae too difficult to see to obtain reliable, comparative results. I did not study the shape of
anal tubules. Studies by Wichard (1975) and Wigglesworth (1938) would suggest that these would
exhibit too large a degree of variation to be useful in distinguishing the morphologically similar species
of Schadonophasma.
Measurement of pupal features follows the treatment by Saether (1970). Pupal abdominal length
was measured from the anterior margin of abdominal tergite I to the posterior margin of abdominal
segment VIIL I found total length to be too difficult to measure accurately from slide mounted material.
Saether (1970) recorded 74 measurements and 20 ratios from males and 67 measurements and 15
ratios from females of adult Chaoborus cooki {6, n = 7;9, n = 4) and Chaoborus trivittatus n = 7;9,
n = 5). According to the results of this study these were all correctly identified, with the possible
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132
Borkent
exception of the adult male Chaoborus trivittatus from Lac Phillipe, Quebec. Those measurements or
ratios judged, on the basis of Saether’s study, to be of possible taxonomic use were studied from slide
material. Two of the most important characters, penis valve shape and Y /X wing ratio were also studied
from pinned material. Terms for structures of the penis valves are shown in Figure 3B. Total length of
the penis valve was measured from the base of the penis valve to the tip of the claw. The characterization
of penis valves was based on a study of these structures from many orientations. Because it was
impossible to standardize these, future comparisons should preferrably be based on orientations in which
the penis valves look similar to those illustrated in this work.
All thoracic setae, except scutellar setae, of the adults were counted on one side of the thorax. Setae
on tergite IX were counted for only one side.
Searches for additional characters of all stages to separate the species of Schadonophasma, except
eggs, previously undescribed, were generally fruitless. Features which yielded no criteria for separation
of the species were:
Adults. Structure of digestive tract; setal pattern on thorax; extent of coloration; sperm shape.
Male and female pupae. Reticulation of pupal horns; length/ width of respiratory horn; chaetotaxy of
abdominal segments II-VIII; coloration, including that of the ribs of the anal paddle; anal paddle shape;
degree of serration of outer rib of paddles; length of median rib; distance of plumose seta from base of
median rib; distance of plumose seta from base of median rib/fength of median rib; length, width and
length/width of genital lobes.
Four instar larvae. Microsculpture of head capsule; dorsal outline of head capsule; shape of antennae;
curvature of long antennal blades; length of postantennal filaments; length of labrum; width of labrum;
distance of long seta from anterior base of labrum; ratio of the distance of long seta from anterior base of
labrum to the length of labrum; shape of maxillae; length of seta on maxillary palpus; dorsal outline of
the dorsal process.
I assumed that characters, with the exception of the length of the postantennal filaments, not useful
for identification of fourth instar larvae would not be useful for younger instars.
Ratios were used to describe shapes or proportions statistically and some are complemented by
illustrations. Measurements used to construct ratios or ratios themselves are abbreviated as follows:
Adults
X = length of vein Rs between R-M and fork of Rsa and Rs (Fig. 3C)
Y = length of vein Rs between R-M and fork of R^ and R j (Fig. 3C)
Z = length of vein M between R-M and fork of M, and M^ (Fig. 3C)
R;j = length of vein R;j (Fig. 3C)
M, = length of vein M, (Fig. 3C)
Fe = femur length
Ti = tibia length
Ta,-Ta5= length of tarsomeres 1-5
JL = length of first tarsomere
R length of tibia
_P = length of penultimate antennal article
U length of ultimate antennal article
FI W = head width
PL
prementum length
Systematics and Bionomics of Schadonophasma
133
HV = length of male
gonostylus length X 10
HR = gonocoxite length
gonostylus length
Male and Female Pupae
WS = width of seventh abdominal segment(Fig. 3A)
LS length of seventh abdominal segment
Larvae
AS = distance of antennal seta from base of antenna (Fig. 2A)
AL antennal length
LB = length of long antennal blade (Fig. 2A)
AL antennal length
LB = length of long antennal blade (Fig. 2A)
SB length of short antennal blade
PAL = prelabral appendage length (Fig. 2A)
PAW prelabral appendage width
Legs are referred to by the following abbreviations.
P, = foreleg
P9 = midleg
P;j = hindleg
Statistical tests were used to compare species, larval instars and sexes, to understand geographieal
variation and to eomplement the species descriptions. Descriptive statistics of samples of eight or more
individuals are ranges, means, 1.5 standard deviations (SD), and two standard errors (SE) on either side
of the mean.These statisties are those graphieally presented in the Hubbs diagrams. For samples of less
than eight, only range and mean are given. Ranges only are given for samples of two. Total lengths of
larvae are expressed only as a range, as length is growth related and for the purposes of this study,
further statistics would not give additional information.
Beeause Schadonophasma fourth instar larvae could not be identified on the basis of discrete
eharacters, I employed a compound character index to reeognize fourth instar larvae of Chaoborus
trivittatus and Chaoborus cooki. Although this index has been used primarily to reeognize hybrids and
analyze zones of hybridization (Freitag, 1965, and papers cited therein; Hubbs and Peden, 1969; Rising,
1970) it is also useful to distinguish morphologieally similar speeies. Compound charaeter indiees
eompile an overall measure of differenee to test for morphological discontinuity (i.e. reproductive
isolation). Kim et al. (1966) have given a step by step analysis of how characters are best used in
combination to distinguish similar species and I have followed most of their suggestions. However,
because I was using the index to test for lack or presence of eonspeeifieity of samples and provide a
means of identifieation, I did not statistieally test homogeneity of samples and eould not determine,
beforehand, the amount of overlap of eharaeter variation between speeies.
Differenees between male and female larvae and pupae were estimated using the unpaired t-test.
Illustrations. - Most illustrations were made with the aid of a drawing tube on a Wild M20
eompound mieroseope from slide mounted material. They eomplement verbal or statistical descriptions
of the speeies and describe the eomplex shape and depict the various forms of adult penis valves. Shape
of the lateral outline of the dorsal process is also eharacterized with illustrations. Penis valves were
drawn with the head in a horizontal plane.
Quaest. Ent., 1979, 15 (2)
134
Borkent
Scanning electron photomicrographs illustrate some characters.
Distribution maps are provided. Dots represent collecting sites of all stages of a species. These
localities are a composite of all material examined and therefore may not accurately reflect present
distributions. Details of collecting sites and dates, and stage collected of measured specimens are
available at the end of each species description.
Criteria for species recognition. - I have accepted for the purposes of this study Mayr’s (1969)
species definition as ‘groups of actually (or potentially) interbreeding populations which are
reproductively isolated from such groups’. Reproductive isolation, reflected in discontinuities in variation
of characters and especially in circumstances of sympatry may be used to recognize species. However,
some discontinuities can be better explained by other interpretations. Circles of races, ecophenotypic or
polymorphic variation may be interpreted incorrectly as reflecting barriers to gene flow and the presence
of more than one species (Brown, 1959; Mayr, 1969; Ross, 1974).
When I initiated this study I suspected that Chaoborus trivittatus and Chaoborus cooki might be
conspecific. I therefore undertook a study of the structure, bionomics, behavior, and development of both
species and used these data as criteria for species recognition. In addition, I carried out mating
experiments with adults of Chaoborus trivittatus and Chaoborus cooki. In light of the resulting
concordant evidence I was better able to interpret discontinuities of morphological variation.
Lack of bionomic data and associated adults for several samples of immatures made me uncertain of
their specific status. I describe and discuss these under the heading ‘populations incertae sedis' to better
define and encourage further work on the problems posed by these samples.
Although I recognized the subspecies catagory as useful for the recognition of some distinctive
populations, the varied uses of this category has led to numerous misunderstandings of variaton within
species and I therefore prefer to describe variation of populations without formally classifying them.
Taxonomic Methods. - The structure of fourth instar larvae, pupae and adults of several populations
of Schadonophasma in Alberta indicate that these were referable to previously described Chaoborus
trivittatus and Chaoborus cooki. Mating experiments and study of behavior and bionomics were used to
test interpretation of morphological differences as a result of reproductive discontinuities. Study of such
differences was then applied to all available material to determine if these would support indications of
lack of conspecificity. A compound character index was used to test for morphological discontinuities of
fourth instar larvae of Schadonophasma and provided a means of identification of these larvae.
Identification of eggs and first, second and third instar larvae was based on either laboratory rearings or
association with identified later stages.
The limited material of Chaoborus nyblaei available from Europe presented some problems of
interpretation and, although the criterion for recognizing this as a sample of a distinct species was mostly
morphological, interpretation of functional significance of some morphological features was also used.
Geographical variation was studied for two purposes. One was to test my concepts of biological
species and the other, to better understand the extreme variation and evolutionary history of the species.
Paucity of material limited the stages studied for geographical variation of characters, to fourth
instar larvae and adults.
For the study of geographical variation of characters the most informative statistic is the mean. To
investigate patterns of variation I plotted mean values of a character for all samples which contained
eight or more individuals, on a map of North America or Alberta and visually judged if any patterns
were present. Samples of less than eight were lumped with the geographically closest sample.
The geographical variation of all characters of the fourth instar larvae used in the compound
character index, of the penis valve length, number of setae on tergite IX of the male adults, and for Y /X
wing ratio of both male and female adults, was studied in detail. Male and female adults were analyzed
separately.
Systematics and Bionomics of Schadonophasma
135
The correct naming of taxa was confirmed by examination of types.
I discuss my approach to the evolutionary history of species of Schadonophasma in the section on
phylogeny and zoogeography.
Literature. - Using the very complete bibliography dealing with Chaoborus compiled by Roth and
Parma (1970) as a basis, I attempted to trace every published account of species of Schadonophasma up
to at least September, 1977.
Because much ecological work is dependent on correct recognition of the species studied, 1 have
attempted to reidentify every published record either by examination of the original material or on the
basis of published distributional and ecological information.
CLASSIFICATION
Genus Chaoborus
Edwards (1932) gave a complete list of generic synonyms (with the exception of Edwardsops Lane) and
Cook (1956) and Saether (1972) described the general structure of Chaoborus species. Only additions
and corrections to those descriptions are given here. The first, second and third instar larvae are fully
described as they differ from fourth instar larvae. Within a species, earlier instars differ from later
instars in numerous characteristics, such as head capsule length, antennal length, postantennal filament
length, number of mandibular fan bristles and number of anal fan setae and these are described for
Schadonophasma larvae in a later section. Only qualitative or non-varying differences are described
here. Descriptions of earlier instars of Chaoborus species are given by Deonier (1943), McGowan
(1972), Parma (1969, 1971a), Prokesova (1959), Saether (1967, 1970), Sikorowa (1973) and Smith
(1960a). Eggs or egg masses have been described by Aitken (1954), Berg (1937), Herms (1937),
Lindquist and Deonier (1942b), MacDonald (1956), McGowan (1974, 1975, 1976), Parma (1971b),
Sikorowa (1973), Smith (1960a) and von Frankenberg (1937).
Saether’s comments on the subgenus Sayomyia appear to be based on only the North American
species Chaoborus punctipennis, Chaoborus astictopus Dyar and Shannon, Chaoborus albatus,
Chaoborus annulatus Cook, and Chaoborus maculipes Stone, and do not apply to at least some tropical
species. Saether’s (1970) calculation of FIR was incorrect and actually is the inverse of the data he
presented.
Some erroneous interpretations of characters differing between larval instars are discussed under the
heading ‘Characters Varying between Larval Instars’.
Keys to subgenera of Chaoborus of the Holarctic region for adults, pupae, and fourth instar larvae
have been provided by Saether (1972). More comprehensive keys must await analysis of species of the
Neotropical, Oriental, and Australian regions. Some corrections to Saether’s (1972) keys are presented
below in the key to species.
Diagnosis. — Adults. Second tarsal article shorter than first; male tarsal claws equal; tarsal claws not toothed; females with
three seminal capsules; vein R, terminated distal to vein Cu, apex.
Pupae. Respiratory horns with spiracular openings small, slit-like; paddles free, movable, each supported by one medial, two
marginal ribs (Fig. 2C,D).
Larvae (all instars). Plead capsule laterally compressed; antennae proximate; pair of air sacs in both thorax and abdominal
segment VII; no respiratory siphon; anal segment with ventral setal fan.
Description of Chaoborus. - Adults. Females with row of comb-like setae on anterior face of third tarsus of midlegs (Fig. 4A).
Pupae. Respiratory horns spindle-shaped (Saether, 1972; 272) to globular (Verbeke, 1957: 190); terminalia almost parallel
Quaest. Ent., 1979, 15 (2)
136
Borkent
(Fig. 5B; Blanchard, 1905: 39; Deckart, 1958: 269; Prashad, 1918: 19; Senior-White, 1927: 65; Wesenberg-Lund, 1943: 418), to
perpendicular (Fig. 5A) to longitudinal axis of body.
Fourth Instar Larvae. Length 4.4-20.3 mm; predaceous; developing adult eye visible (present or absent in Chaoborus
anomalus)', apex of antenna with six terminal setae (Fig. 2A); four long, one noticeably shorter, one much shorter; pair of setae
dorso-posterior of antennae dendritie; ten postantennal filaments; prelabral appendages one in front of other, setaceous or laterally
flattened; mandibular fan present; maxillae as shown by Parma (1971a: 175), Cook (1956: 76); anal fan setae plumose, not paired;
abdominal segment IX dorsal process flat, conical or with short article.
Third Instar Larvae. Most specimens of most species with developing adult eye visible, absent from Chaoborus anomalus\
antenna with six terminal setae; pair of setae dorso-posterior of antennae simple; ten postantennal filaments; prelabral appendages,
one anterior of other, setaceous or flattened; mandibular fan present; maxillae as shown by Parma (1971a: 175); anal fan setae
plumose, not paired.
Second Instar Larvae. Some specimens of some species with developing adult eye visible; antenna with six terminal setae;
eight postantennal filaments; prelabral appendages lateral to one another, setaceous; mandibular fan present; maxillae as shown by
Parma (1971a: 175); anal fan setae plumose, not paired.
First Instar Larvae. Developing adult eye not visible; egg burster on dorsum of head capsule; antenna with four terminal setae;
base of antenna without posterior eurve (except Chaoborus edulis Edwards; McGowan, 1976: 300); two postantennal filaments;
prelabral appendages lateral to one another, setaceous; mandibular fan absent; maxillae as shown by Parma (1971a: 175); anal fan
setae simple, in pairs.
Eggs. Outline oval; some species with eggs in jelly-like matrix; white when laid, later light to dark brown; chorion transparent;
at least for species of Chaoborus sensu stricto egg shell canoe-like in shape (Parma, 1971b: 32; pers. obs.) with dorsal longitudinal
slit (see also descriptions of Schadonophasma species).
Subgenus Schadonophasma
Schadonophasma Dyar and Shannon 1924: 209. TYPE SPECIES (by monotypy) Corethra trivittata
Loew 1862: 186. Cook 1956: 28. Edwards 1932: 26.
Schadanophasma Saether 1970: 12 (misspelling).
Saether (1970: 14) incorrectly recognized Chaoborus nyblaei as type species.
Especially considering the morphological similarity of known stages of Chaoborus species. Lane’s
(1951: 336) suggestion that Chaoborus subgenera be raised to generic rank does not seem to be justified
and, at least until the world fauna is better understood, subgeneric status should be retained.
Diagnosis. — Adult. Total length 5. 2-9. 3 mm; general coloration as given by Cook (1956); wings with several spots (Fig. 3C);
more darkly pigmented than rest of thorax-anterior pronotum, dorsal portion of posterior pronotum (some specimens not so darkly),
ventral half of preepisternum, anterior half of anterior anepisternum 2, pleural apophysis, anterior-dorsal edge of anepimeron, meron,
center of anepisternum 3, scutellum, postnotum, two pairs of vittae on scutum; Rsa present, Cu., , faint or absent; wing veins scaled;
legs each with apex of femur, apex and base of tibia, apex of first, second and third tarsomeres and, all of fourth and fifth tarsomeres
darkly pigmented; pulvilli well developed, at least half as long as claws; ommatidia free, dorsal part of eye at least as long as three
ommatidia (Saether, 1970: Fig. 9); no parascutellar setae. Male. Without lobe or stout setae on inner face of gonocoxite; penis valve
with apical claw. Female. Inner face of tarsomere three of midlegs and hindlegs with row of comb-like setae (Fig. 4A); segment X not
covered by tergite IX in dorsal aspect, neither apically bilobed (Cook, 1956: fig 15A); seminal capsule ovoid with short, slightly curved
neck (Cook, 1956: fig. 14J); antenna with 13 flagellar articles.
Pupa. Overall coloration from light to dark brown, females slightly darker than males; mature specimens more darkly pigmented
along posterior and anterior margin of each abdominal segment, also bases of some setae, bases of paddles; longitudinal stripes parallel
to outer margin of abdominal tergite and sternite; median rib of paddle darker than lateral ribs; in some, outer and middle ribs equally
pigmented but darker than inner rib; when pharate adult developed (abdominal setae of pharate adult visible) pigmented wings and
legs visible, penis valves of males with apical claw (Fig. 3B); WS/LS of both male and female 1 .20-1 .89; median rib with plumose seta
0.47-0.79 of its length from base and simple seta near apex; shagreening at base of paddles; outer rib of paddle with only sparse
shagreening; median rib almost extended to margin of paddle and curved apically (Fig. 2C,D); respiratory horns spindle-shaped;
length of respiratory horn/width of respiratory horn of male 3.30-4.71, of females 3.23-4.80; chaetotaxy as shown on Figure 6, setae
present as shown by Belkin et al. (1970) except seta 4-11 present; male genital sac length 932-1298 ^m, width 189-307 ;um,
length/width 3.24-5.05.
Fourth Instar Larva. Total length 11.4-20.3 mm; head capsule length 1404-2620 /um; seta on anterior face of antenna
inconspicuous; AS/AL 0.75-0.93; antenna with curve at posterior base (Fig. 7D); tentorium pigmented, thick, conspicuous (Fig. 2A);
dorsal process on abdominal segment IX flat (Fig. 2B, 8C, D); head capsule colorless to dark brown; more pigmented specimens with
posterior margin, dorsal surface on head capsule, area around anterior tentorial pit, and line of attachment of mandible conspicuously
Systematics and Bionomics of Schadonophasma
137
pigmented; lateral outline of dorsal surface of head capsule as shown by Saether (1970: fig. 11A,B,C) and Figure 2A and 71;
longest mandibular tooth elongate, with curve about 0.75 from base (Fig. 8B; Saether, 1970: fig. 1 1 K; Felt, 1904: fig. 99); head
capsule with microsculpture polygonal on posterior edge (Fig. 4B); additional features in key.
Third Instar Larva. As fourth instar larva except following: total length 7.0-13.0 mm; head capsule length 944-1440 jum;
lateral outline of dorsal surface of head capsule as in Figure 7H; microsculpture on posterior margin of head capsule not obviously
polygonal; antenna with curve at posterior base (Fig. 7c); AS/AL 0.53-0.90; longest mandibular tooth elongate, with or without
slight curve about 0.75 from base; additional features in key.
Second Instar Larva. As fourth instar larva except following: total length 4. 7-7. 3 mm; head capsule length 522-760 yum;
lateral outline of dorsal surface of head capsule as in Figure 7G: microsculpture on posterior margin of head capsule not polygonal;
antenna with only slight curve at posterior base (Fig. 7B); AS/AL 0.38-0.72; longest mandibular tooth elongate, without curve;
additional features in key.
First Instar Larva. As fourth instar larva except following: total length 2.0-4. 5 mm; head capsule length 294-494 fxm-, lateral
outline of dorsal surface of head capsule as shown in Figure 7E,F; microsculpture on posterior margin of head capsule not
polygonal; antenna without curve at posterior base (Fig. 7A); AS/AL 0.24-0.40; mandible as in Figure 8A; additional features in
key.
Description. - Description of the subgenus is provided in the description of Chaoborus, the diagnosis
of Schadonophasma, and the following keys and description of species.
Key to the species of Schadonophasma
Species of Schadonophasma are morphologically very similar and are therefore difficult to identify.
No single character can or should be relied upon to identify larva, pupae or adults to species; thus, to be
confident of a correct determination, all characters described should be examined. Because of this
overlap of range of variation it is best to identify several individuals from a sample and opposites of the
couplet should be compared.
Male adults
The shape of the penis valves is particularly important but difficult to use in identifying males. It is
most important that the penis valves be examined from a number of orientations until their shape
approximates those illustrated.
1 Penis valve head elongate, with claw mostly parallel to longitudinal axis (Fig. lOA-O);
penis valve length 145-232 /um; Y /X wing ratio 1.17-3.22; length of tarsomere five of
foreleg 186-273 )um; length of claw of foreleg 53-83 pm; coloration of vittae generally
black or very dark brown Chaoborus trivittatus (Canada, eastern and western U.S.A.)
Penis valve shape not as above (Fig. 1 1 A-N) or if similar (Fig. fl, 0,P), specimen from
Fennoscandia; penis valve length 191-232 pm; Y/X wing ratio 1.64-3.62; length of
tarsomere five of foreleg 220-284 ^m; length of claw of foreleg 65-88 pm; coloration of
vittae generally brown 2
2 Penis valve head bulbous, with claw mostly perpendicular to longitudinal axis (Fig. 1 1
A-N); Y/X wing ratio 1.64-3.62; length of claw of foreleg 65-88 pm
Chaoborus cooki (Canada)
Penis valve head elongate with claw mostly perpendicular to longitudinal axis (Fig. 1 1
0,P); Y/X wing ratio 1 .88-2. 13; length of claw of foreleg about 85 pm
Chaoborus nyblaei (Fennoscandia)
Female adults
Egg number and type, when present in the abdomen and in good condition, are consistent and reliable
characters for species determination.
1 Seminal capsules 71-102 pm in diameter; Y /X wing ratio 1.19-3.22; length of tarsomere
Quaest. Ent., 1979, 15 (2)
138
Borkent
five of foreleg 232-290 iim; length of claw of foreleg 64-81 jum; prementum length
325-510 ium; coloration of vittae generally black or very dark brown; distance from
anterior tip of longest ovary to apex of abdomen 0.71-1.1 1 times total length of abdomen
of teneral females; total number of eggs 150-329 in nulliparous individuals; eggs, if
present, of parous individuals with no obviously thick exochorion (Fig. 9A)
Chaoborus trivittatus (Canada, eastern and western U.S.A.)
Seminal capsules 65-79 /urn in diameter; Y /X wing ratio 2.03-5.83; length of tarsomere
five of foreleg 232-325 iim', length of claw of foreleg 64-104 )um; prementum length
458-574 jum; coloration of vittae generally brown; distance from anterior tip of longest
ovary to apex of abdomen 0.32-0.62 times total length of abdomen of teneral females
(unknown for Chaoborus nyblaei)\ total number of eggs 58-144 in nulliparous
individuals; eggs, if present, of parous individuals with obvious thick exochorion
(Fig. 9B,C) Chaoborus cooki (Canada) Chaoborus nyblaei (Fennoscandia)
Male and female pupae
Male and female pupae are distinguished from one another by the shape of their genital lobes (Fig.
2C,D), (Deonier, 1943: fig. 1,2).
Saether’s (1972) key to pupae should read ‘abdominal segment VIF instead of ‘abdominal segment
Vlir and couplet 7(4) (pg. 271) should read ‘Median rib of anal paddle with a minute seta at apex and a
single plumose seta medially (Fig. 12 O-Q); greatest width of thoracic organ at or below the middle’.
Male pupae
1 Width of adominal segment VII 1074-1794 )um; respiratory horn length 1.18-1.48 mm;
non-teneral live individuals with abdomen nearly vertical (Fig. 5B)
Chaoborus trivittatus (Canada, eastern and western U.S.A.)
Width of abdominal segment VII 1605-1947 /im; respiratory horn length 1.31-1.64 mm;
non-teneral live individuals with tip of abdomen nearly horizontal (unknown for
Chaoborus nyblaei (Fig. 5 A) 2
2 WS/LS 1.40-1.69 Chaoborus cooki (Canada)
WS/LS 1.63-1.74 Chaoborus nyblaei (Fennoscandia)
Female pupae
1 Width of abominal segment VII 1239-2266 jum; WS/LS 1.25-1.62; mature individuals
(pharate adult setae evident) with abdomen full of eggs; non-teneral live individuals with
abdomen nearly vertical (Fig. 5B)
Chaoborus trivittatus (Canada, eastern and western U.S.A.)
Width of abdominal segment VII 1605-2382 )um; WS/LS 1 .43-1 .89; mature individuals
(pharate adult setae evident) with ovaries extended to, at most, abdominal segment V
(unknown for Chaoborus nyblaei); non-teneral live individuals with tip of abdomen
nearly horizontal (unknown for Chaoborus nyblaei) (Fig. 5A) 2
2 Width of abdominal segment VII 1605-2230 /am; WS/LS 1.43-1.87
Chaoborus cooki (Canada)
Width of abdominal segment VII 2030-2384 ^m; WS/LS 1.71-1.89
Chaoborus nyblaei (Fennoscandia)
Systematics and Bionomics of Schadonophasma
139
Larvae
Roman numerals refer to larval instars. Chaoborus nyblaei I, II, and III are unknown.
1 Prelabral appendages one in front of other 5
Prelabral appendages lateral to one another 2
2 ( 1 ) Egg burster on dorsum of head capsule(Fig. 7E,F); antennae each with four terminal
setae; two postantennal filaments; mandibular fan absent; most anal fan setae arranged
in pairs.... 3
No egg burster on dorsum of head capsule (Fig. G-I); antennae each with six terminal
setae; eight postantennal filaments; mandibular fan present; no anal fan setae arranged in
pairs 4
3 (2) Egg burster on dorsum of head capsule not pigmented, shaped as in Figure IK; head
capsule length 294-368 jum; antennal length 79-1 15 jum; antennal blade length 176-232
/urn Chaoborus trivittatus I (Canada, eastern and western U.S.A.)
Egg burster on dorsum of head capsule pigmented, shaped as in Figure 7J; head capsule
length 347-493 iim; antennal length 103-132 pm; antennal blade length 208-288 pm
Chaoborus cooki 1 (Canada)
4 (2) Head capsule length 522-650 pm; LB/AL 1.02-1.16; 8-14 mandibular fan bristles;
16-21 anal fan setae Chaoborus trivittatus II (Canada, eastern and western U.S.A.)
Head capsule length 638-760 ^um; LB/AL 0.91-1.07; 7-9 mandibular fan bristles; 19-26
anal fan setae Chaoborus cooki 11 (Canada)
5 (1) Head capsule length 944-1440 pm; antennal length 423-673 pm; postantennal filament
length 354-578 /am 6
Head capsule length 1404-2620 pm; antennal length 623-1235 pm; postantennal
filament length 555-1015 /am 7
6 (5) AS/AL 0.58-0.90; PAL/PAW 3.36-15.00; 9-24 mandibular fan bristles; 19-28 anal fan
setae
Chaoborus trivittatus III (Canada, eastern and western U.S.A.)
AS/AL 0.53-0.81; PAL/PAW 3.56-6.60; 8-12 mandibular fan bristles; 22-23 anal fan
setae .Chaoborus cooki 111 (Canada)
7 (5) Head capsule length 1404-2384 /am; antennal length 632-1 195 /am; LB/SB 1.47-2.92;
PAL/PAW 2.63-9.88; 1 1-36 mandibular fan bristles; 21-33 anal fan setae
Chaoborus trivittatus IV (Canada, eastern and western U.S.A.)
Head capsule length 1707-2620 pm; antennal length 789-1235 pm; LB/SB 1.14-2.13;
PAL/PAW 2.00-5.30; 8-17 mandibular fan bristles; 28-40 anal fan setae 8
8 (7) Postantennal filament length 631-944 /am; prelabral appendage length 186-306 /am;
PAL/PAW less than 4.16; 28-38 anal fan setae Chaoborus cooki IV (Canada)
Postantennal filament length 684-1015 pm; prelabral appendage length 300-348 pm for
specimens with undamaged tip (tip not coming to flat end (Fig. 15E-H)); PAL/PAW for
such specimens 3.58-5.13; 34-40 anal fan setae Chaoborus nyblaei IV (Fennoscandia)
Eggs
1 Laid in spiral arrangement in flat jelly-like matrix; exochorion thin, without obvious
sculpturing (Fig. 9A; 4C); length/width 2.85-3.16
Chaoborus trivittatus (Canada, eastern and western U.S.A.)
Quaest. Ent., 1979, 15 (2)
140
Borkent
Laid in more or less spherical mass (unknown for Chaoborus nyblaei)-, exochorion thick,
with hexagonal sculpturing (Fig. 9B,C; 4D); length/width without exochorion 2.36-2.77
(unknown for Chaoborus nyblaei)
Chaoborus cooki (Canada), Chaoborus nyblaei (Fennoscandia)
Chaoborus trivittatus (Loew)
Corethra trivittata Loew 1862: 186. Holotype adult male, labelled: ‘Me’, ‘Loew Coll.’, ‘Type 4050’,
‘trivittata‘; (MCZ). Johannsen 1903: 398. Not Dyar 1902: 201.
Sayomyia trivittata; Felt 1904: 361. Felt 1905: 497.
Sayomyia knabi Dyar 1905: 16. Holotype third instar larva, labelled: ‘Sayomyia trivittata (trivittata
crossed out), Springfield Mass, F. Knab’, ‘96’. A figure of the prelabral appendage on the label is
identical to that drawn by Dyar 1905: 14; (USNM).
Chaoborus trivittatus; Dyar and Shannon 1924: 212. Saether 1970: 23. Johannsen 1934: 44. Dickinson
1944: 351.
Chaoborus trivattus Saether 1970: 23 (misspelling).
Chaoborus nyblaei; Cook 1956: 28 in part (not Zetterstedt). Saether 1970: 14 in part. Matheson
1944: 94.
Chaoborus brunskilli Saether 1970: 21. NEW SYNONOMY. Details of holotype and paratype given
by Saether (1970). Label of holotype reads ‘L. 244’ but this is probably incorrect and should read ‘L.
241’; (CNC).
Characters tested to distinguish adults of Schadonophasma species are described here to supplement
those descriptions by Saether (1970).
Descriptions. — Males. General coloration of most specimens dark grayish brown; specific pigmentation as other members of
subgenus; vittae generally black or very dark brown; penis valves as shown in Fig lOA-0. Measurements and proportions: see Table 2.
Females. Coloration as for males. Measurements and proportions: see Table 3.
Male pupae. Non-teneral live individuals with tip of abdomen nearly vertical when at rest (Fig. 5B); coloration as other members
of subgenus. Measurements and proportions: see Table 4.
Female pupae. Non-teneral live individuals with tip of abdomen nearly vertical when at rest. (Fig. 5B); mature individuals
(pharate adult setae evident) with ovaries extended to at least abdominal segment III; coloration as other members of subgenus.
Measurements and proportions: see Table 5.
Cook (1956: 3 1 ) reported respiratory horn length of specimens from Green Valley, California to be 1 .02-1 . 1 3 mm.
Fourth instar larvae. Coloration as other members of subgenus. Measurements and proportions: see Table 6.
Third instar larvae. Coloration as other members of subgenus. Measurements and proportions: see Table 7.
Second instar larvae. Coloration as other members of subgenus. Measurements and proportions: see Table 8.
First instar larvae. Egg burster on dorsum of head capsule not as prominent (Fig. 7F) as that of Chaoborus cooki first instar
larvae; pigmentation of egg tooth and posterior margin of head capsule not nearly as dark as that of Chaoborus cooki-, coloration of
head capsule very light. Measurements and proportions: see Table 9.
Eggs. Laid in spiral arrangement in disc of jelly-like matrix; egg with thin exochorion (Fig. 9A) with very fine sculpturing (Fig.
4E) restricted to dorsal surface. Measurements and proportions: see Table 10. Egg shell as in Figure 9D.
Bionomics. - Individuals overwinter as fourth instar larvae and are mostly restricted to permanent
lentic habitats ranging from small shaded ponds to large deep lakes in woodland. Although some larvae
live in temporary waters, they cannot successfully overwinter in these habitats. Adults emerge under
spring conditions in Alberta. Records from elsewhere indicate some multivoltine populations (p. 184).
Systematics and Bionomics of Schadonophasma
141
Table 2. Descriptive statistics for male adult Chaoborus trivittatus. Measurements in ^^m unless stated
otherwise.
Quaest. Ent., 1979, 15 (2)
142
Borkent
Table 4. Descriptive statistics for male pupae of Chaoborus trivittatus. Measurements in )um unless
stated otherwise.
Table 6. Descriptive statistics for fourth instar larvae of Chaoborus trivittatus. Measurements in jum.
Systematics and Bionomics of Schadonophasma
143
Table 7. Descriptive statistics for third instar larvae of Chaoborus trivittatus. Measurements in
Quaest. Ent., 1979, 15 (2)
144
Borkent
Derivation of specific epithet. - The name 'trivittatus' is derived from Latin (tri — three; vitta —
stripe) and refers to the three vittae (actually four but two are very proximate) on the scutum of the
adult.
Distribution. - Specimens of this species have been collected in woodland areas throughout Canada
but are restricted in the United States to California, Oregon, Washington and Montana and to the
northeastern States (Fig. 16,17). Lack of records from north central Canada and Alaska probably
reflect lack of collecting. Chaoborus trivittatus is clearly restricted to woodland. Records from southern
Alberta (Fig. 17), for example, are clearly the eastern limit of this species in this area. I attempted to
collect east of these localities, but with no success. In the Rocky Mountains, the species has not been
found above treeline. The single record from Baffin Island is suspect. No other records of
Schadonophasma are north of treeline. However, for the sake of completeness, I have included the
record on the distribution map.
Records of Chaoborus trivittatus from Wisconsin reported by Dickenson (1944) are undoubtedly
correct. Cook’s (1956) objection to the above report, in that he found one specimen to be culicine,
erroneously assumed only one specimen collected. However, Dickenson (1944) also recorded larvae
which could only be those of Chaoborus trivittatus (length of two centimeters). In addition, I have
examined specimens from Vilas County, Wisconsin, which are Chaoborus trivittatus.
Even though I have not examined specimens of Chaoborus trivittatus from Churchill, Manitoba, the
reports by James and Smith (1958) of overwintering larvae (see p. 184), and Twinn et al. (1948) of
swarming adults (see p. 189) would suggest this species to be present at that locality.
Cleugh and Hauser (1971) give the exact localities for the numbered lakes near Kenora, Ontario,
given below.
LOCALITIES
Male Adults
CANADA
ALBERTA: Banff, 24-VII-1918 (1 BMNH, 4 USNM); 10 mi. (16 km) W. Jasper, 4-VI-1976 (5 CNC, 5 USNM, 8 UASM, 20
ABOr); Bigoray River, oxbow lake, 25-V-1971 (1 OASa); 1.5 mi. (2.4 km) W. Edmonton, 3-V-1976 (5 CNC, 5 USNM, 25 ABOr, 7
UASM); Edmonton, 17-V-1974 (1 ABOr); BRITISH COLUMBIA: Kaslo, 13-21-VI-? (7 USNM); Terrace, 18-VI-1960 (5 CNC, 1
ABOr); Prince Rupert, 13-V-1919 (1 CNC); Atlin, 5-VIM955 (6 CNC, 1 ABOr); Lower Post, 20-VI-1948 (2, CNC)
NORTHWEST TERRITORIES: Aklavik, 28-V111-1932 (1 USNM); Lake Harbour, Baffin Island, 7-V111-1935 (1 CNC); NOVA
SCOTIA: 4-III-I935 (reared) (1 CNC); ONTARIO: North Burgess Twp., 1 5-20-V-1970, (6 CNC); Black Lk., 44°46'N 76°18'W,
26-IV-1973 (8 CNC, 1 ABOr); Kenora, Lk 241, 13-V1-1968 (reared) (1 CNC); QUEBEC: Lac de Jean Venne (Duncan Lk.),
Masham Twp., 20-IV-1973 (3 CNC), 5-14-V-1973 (12 CNC, 2 ABOR); YUKON TERRITORY: Whitehorse, 14-21-V-1949 (5
CNC, 1 ABOr).
UNITED STATES
CALIFORNIA: Alameda Co.,: Oakland, 4-V-1937 (2 CAS); Humboldt Co.: Mad River Beach, 13-V111-1948 (7 USNM); Monterey
Co.: Pacific Grove, 18-VII-1940 (2 USNM); San Luis Obispo Co.: Oceano Beach 20-VIII-1948 (1 USNM); Santa Clara Co.:
Stanford University, 7-11I-1903 (1 CUNY, 2 USNM), 28-V-1903 (1 USNM); MAINE: (1 MCZ); MASSACHUSETTS: Worcester
Co.: Worcester, 23-1V-? (2 MCZ); Andorra (specific locality unknown), (1 USNM); NEW HAMPSHIRE: Belknap Co.: Center
Harbor, (7 USNM); Rockingham Co.: Hampton, 21-1V-1906 (1 MCZ); NEW YORK: Essex Co.: Elizabethtown 10-13-V1-1904 (1
CUNY, 1 USNM); OREGON: Multnomah, Clackamas and Washington Co.: Portland, (1 USNM): WASHINGTON: Kitsap Co.:
Bremerton, 2-V-1924 (1 USNM); Port Madison, 3-11-1934 (reared) (2 USNM); Hoodsport, 3-9-V-1924 (2 BMNH, 6 USNM).
Female Adults
CANADA
ALBERTA: 10 mi. (16 km) W. Jasper, 4-V1-1975 (8 ABOr, 3 CNC); 1.5 mi. (2.4 km) W. Edmonton, 13-V-1975 (15 ABOr, 8 CNC,
10 UASM); BRITISH COLUMBIA: Kaslo, 13-20-VI-? (7 USNM); Victoria, 4-X-1922 (1 CNC); Canim Lk., 23-VI-1938 (2 CNC);
Terrace, 7-18-V1-1960 (8 CNC, 1 ABOr); Atlin, 5-VI1-1955 (5 CNC); Lower Post, 20-V1-1948 (1 CNC); NORTHWEST
Systematics and Bionomics of Schadonophasma
145
TERRITORIES: Aklavik, 28-VIII-1932 (1 USNM); ONTARIO; Cordova Mines, 7-IX-1962 (2 CNC); North Burgess Twp.,
15-20-V-1970 (1 CNC); Black Lk„ 44°46'N 76°18'W, 26-IV-1973 (4 CNC); QUEBEC; Old Chelsea, 20-IX-1961 (12 CNC, 1
ABOr); YUKON TERRITORY: Whitehorse, 15-V-1944 (1 CNC), 16-V-1949 (I CNC), 15-19-V-1950 (1 CNC).
UNITED STATES
CALIFORNIA: Alameda Co.: Berkeley, 5-V1-1948 (1 USNM); Oakland, 4-V-1937 (1 CAS); Humboldt Co.; Mad River Beach,
13-V1I1-1948 (7 USNM); Kings Co.: Hanford, 8-V1I-1947 (1 USNM); Madero Co.: Matadero Creek, 24-V-1937 (1 CAS);
Mariposa Co.: Yosemite, 15-V-1916 (1 USNM); Mendocino Co.: Garcia River, 30-VII-1948 (1 USNM); Placer Co.; NW of
Cisco, Nevada Co., 16-V-1948 (1 USNM) Placer and El Dorado Co.: Lake Tahoe, 17-1X-1915 (1 USNM); San Luis Obispo Co.:
Oceano Beach, 20-VIII-1948 (2 USNM); Santa Clara Co.: Stanford University, 111-1903 (1 CUNY, 6 USNM); MONTANA:
Glacier Co.: North Fork Rangers Station, Glacier National Park, 13-V-1926 (1 USNM), 5-V1-1926 (1 USNM); NEW
HAMPSHIRE: Belknap Co.: Center Harbor, (2 USNM); Rockingham Co.: Hampton, 19-VI-1906 (1 MCZ); NEW YORK: Essex
Co.: Elizabethtown, 10-1 l-Vl-1904 (2 USNM), 25-V1I1-1904 (1 CUNY); Rockland Co.: Bear Mnt., 25-IX-1927 (1 USNM);
OREGON; Multnomah, Clackamas and Washington Co.: Portland, 4-V1I-1934 (2 USNM); WASHINGTON; Kitsap Co.:
Manitou Beach, 13-111-1934 (reared) (1 USNM); Port Madison, 3-11-1934 (1 USNM); Mason Co.: Hoodsport, 5-9-V-1924 (4
USNM).
Male Pupae
CANADA
ALBERTA: 1.5 mi. (2.4 Km.) W. Edmonton, 8-V-1974 (2 ABOr, 1 UASM); BRITISH COLUMBIA: Kaslo (4 USNM); Eunice
Lk., 24-V-1971 (3 OASa); Lower Post, 1 9-20-Vl- 1 948 (20 CNC); NEWFOUNDLAND; 3 mi. (4.8 km) N. St. John’s, 4-V1-1975
(1 ABOr); ONTARIO: Kenora, Lk. 241, 13-V1-1968 (1 CNC); QUEBEC: Lac de Jean Venne (Duncan Lk.), Masham Twp.,
20-21-IV-1973 (12 CNC, 2 ABOr).
UNITED STATES
CALIFORNIA; Contra Costa Co.: Jewel Lk., ll-V-1948 (5 USNM); Humboldt Co.: Mad River Beach, 13-VIII-1948 (9 USNM);
MICHIGAN: Gogebic Co.: Hummingbird Lk., 15-VII1-1971 (I OASa); WASHINGTON: Snohomish Co.; Everett, 7-IV-1949 (3
WSUP); Hall Lk., 29-VI1I-1952 (11 WSUP).
Female Pupae
CANADA
ALBERTA: 53 mi. (85 km) N. Coleman, 3-VIII-1975 (2 ABOr, 2 UASM); 1.5 mi. (2.4 km) W. Edmonton, 8-V-1974 (4 UASM);
BRITISH COLUMBIA; Kaslo (4 USNM); Lower Post, 19-20-VI-1948 (8 CNC); NEWFOUNDLAND: 3 mi. (4.8 km) N. St.
John’s, 4-VI-1975 (1 ABOr); QUEBEC: Lac de Jean Venne (Duncan Lk.), Masham Twp., 20-IV-1973 (1 CNC), 5-6-V-1973 (3
CNC).
UNITED STATES
CALIFORNIA: Contra Costa Co.: Jewel Lk., ll-V-1948 (2 USNM); Humboldt Co.: Mad River Beach, 13-VI1I-1948 (8 USNM);
Snohomish Co.; Hall Lk., 29-VIII-1952 (4 WSUP).
Fourth Instar Larvae
CANADA
ALBERTA: 2.5 mi. (4.0 km).NW Edmonton, 13-1X-1970 (4 ABOr); 1.5 mi. (2.4 km) W. Edmonton, 26-IV-1975 (25 ABOr, 13
CNC, 5 UASM); 2 mi. (3.2 km) N. Devon, 23-IX-1972 (1 ABOr); 4.7 mi. (7.6 km) NW Mountain Park, 21-IX-1974 (20 ABOr,
7 CNC, 5 UASM); East Henry Pond, Jasper National Park, 12-IX-1968 (1 FWl, 4 CNC); 1.5 mi. (2.4 km) S. Robb, 23-VII-1975
(50 ABOr, 6 UASM); 53 mi. (85 km) N. Coleman, 3-V11I-1975 (10 ABOr, 2 UASM); 43 mi. (69 km) N. Coleman, 3-V1II-1975
(1 ABOr); 33 mi. (53 km) N. Swan Hills, l-IX-1975 (12 ABOr); 62 mi. (100 km) N. Coleman, 3-VI1I-1975 (15 ABOr, 7
UASM); 38 mi. (61 km) N. Swan Hills, l-IX-1975 (11 ABOr); 14 mi. (22 km) E. Fox Creek, 30-V1I1-1975 (11 UASM); 1 mi.
(1.6 km) N. Swan Hills, l-lX-1975 (11 ABOr); 23 mi. (37 km) E. Fox Creek, 30-VI1I-1975 (11 ABOr); 43 mi. (69 km) E. High
Prairie, l-IX-1975 (11 ABOr); 3 mi. (4.8 km) S. Hinton, 23-VI1-1975 (11 ABOr); 32.5 mi. (52 km) N. Banff, 8-XI-1976 (11
ABOr); Pond nr. Kinky Lk., ll-lX-1976 (10 ABOr); 26 mi. (42 km) N. Sundre, ll-X-1976 (11 ABOr); 47 mi. (76 km) N.
Nordegg, 7-IX-1976 (11 ABOr); 12 mi. (19 km) S. Seebe, 3-VIII-1975 (11 ABOr); 10 mi. (16 m) W. Jaspeer, 27-IV-1976 (10
ABOr); 58 mi. (93 km) S. Seebe, 3-VII1-1975 (11 ABOr); 28 mi. (45 km) E. Jasper, 27-IV-1976 (11 UASM); 4 mi. (6.4 km) W.
Jasper, 20-V-1975 (9 ABOr); 5 mi. (8.0 km) S. Jasper, 18-V1-1975 (12 ABOr); *Pond nr. Cameron Lk., 18-1X-1977 (17 ABOr);
*Pond nr. Wabamum Lk., 15-X-1977 (1 ABOr); *Opal, lO-X-1977 (6 ABOr); BRITISH COLUMBIA: 3 mi. (4.8 km) E.
Sicamous, lO-VI-1976 (11 ABOr); Prince Rupert, 13-V-1919 (15 USNM); Kaslo, (5 USNM); 4.8 mi. (7.7 km) N. Clearwater,
12-VI-1976 (11 ABOr); Eunice Lk., fall of 1971 (4 OASa, 10 ABOr); Lower Post, 19-VI-1948 (11 CNC); Gwendoline Lk.,
7-VI1I-1973 (10 ABOr); NEWFOUNDLAND: 3 mi. (4.8 km) N. St. John’s, 4-VI-1975 (11 ABOr); 0.4 mi. (0.6 km) W. Logy
Quaest. Ent., 1979, 15 (2)
146
Borkent
Bay, 8-V11-1967 (5 USNM); ONTARIO; Kenora, Lk. 241, 14-V-1969 (3 FWI, 8 CNC); Kenora, Lk. 81, 7-V-1969 (1 FWI, 1
CNC); Kenora, Lk. 230, I4-V-1969 (2 FWI, 2 CNC); Kenora, Lk., 240, 14-V-1969 (1 FWI); *10 mi (16 km) E. Thessalon,
19-1X-1978 (18 ABOr); *Kendel, 15-V-1973 (3 ABOr), 17-V1I-1973 (2 ABOr); QUEBEC; Lac de Jean Venne (Duncan Lk.),
Masham Twp., 21-IV-1973 (17 CNC); Lk. a la Croix, 26-VI-1971(2 OASa, 10 ABOr); Randin Lk., 19-VI-1974 (10 ABOr);
YUKON TERRITORY; Klutlan Glacier moraine, VII-1971 (6 EFCo, 3 ABOr).
UNITED STATES
CALIFORNIA; Contra Costa Co.; Jewel Lk., ll-V-1948 (4 USNM); Humboldt Co.; Mad River Beach, 13-VI11-1948 (12 USNM);
Santa Clara Co.; Standford University, 7-I1-1945 (3 EFCo, 11 USNM); MAINE; Piscataquis Co.; Milo, 6-1-1906 (1 USNM);
MASSACHUSETTS; Hampden Co.; Springfield, 7-V11-1903 (1 USNM); MONTANA; Meagher Co.; *27 mi. (43 km) N. White
Sulpher Springs, 27-X-1978; MICHIGAN; Gogebic Co.; Hummingbird Lk., I5-VIII-1971 (3 OASa, 8 JDAl); Ziesnis Bog,
24-VIII-1971 (1 OASa); WASHINGTON; Pacific Co.; 10 mi. (16 km) S. South Bend, I9-VI-1977 (1 ABOr); Snohomis Co.; Hall
Lk., 8-V-1953 (10 WSUP); WISCONSIN; Vilas Co.; Forest Service Bog, 24-VII1-1971 (1 OASa).
Third Instar Larvae
CANADA
ALBERTA; 62 mi. (100 km) N. Coleman, 3-VIII-1975 (7 ABOr, 2 UASM); 53 mi. (85 km) N. Coleman, 3-VIII-1975 (1 ABOr);
1.5 mi. (2.4 km) S. Robb, 23-VII-1975 (25 ABOr, 5 CNC); 4.7 mi. (7.6 km) NW Mountain Park, 21-IX-1974 (2 ABOr);
BRITISH COLUMBIA; 4.8 mi. (7.7 km) N. Clearwater, 12-VI-1976 (7 ABOr); ONTARIO; Kenora, Lk. 230, 14-V-1969 (2
FWI); Kenora, Lk. 241, 13-VI-I968 (I FWI, 8 CNC); YUKON TERRITORY; Klutlan Glacier moraine, Vll-1971 (22 EFCo).
UNITED STATES
MASSACHUSETTS; Hampden Co.; Springfield, (1 USNM), 7-V111-1903 (5 USNM).
Second Instar Larvae
ALBERTA; 62 mi (100 km) N. Coleman, 3-VIII-1975 (6 ABOr, 2 CNC, 2 UASM); East Henry Pond, Jasper National Park,
12-IX-I968 (4 FWI); NEWFOUNDLAND; 3 mi. (4.8 km) N. St. John’s, 4-VI-1975 (I ABOr); ONTARIO; Kenora, Lk. 230,
14-V-1969 (5 FWI); Kenora, Lk. 240, 14-V-1969 (1 FWI); QUEBEC; Randin Lk., 19-VI-1974 (2 ABOr).
First Instar Larvae
ALBERTA; 1.5 mi. (2.4 km) mi. W. Edmonton, 26-VI-1975 (10 ABOr, 4 CNC); 10 mi. (16 km) W. Jasper, 18-V1-1975 (8
ABOr, 4 UASM); 20 mi. (32 km) W. Edson 9-IX-1976 (1 ABOr); East Henry Pond, Jasper National Park, 12-IX-I968 (1 CNC);
NEWFOUNDLAND; 3 mi. (4.8 km) N. St. John’s, 4-VI-1975 (1 ABOr).
Eggs
ALBERTA; 10 mi. (16 km) W. Jasper, 31-V-1976 (12 egg masses ABOr, 2 egg masses CNC, 2 egg masses UASM).
*Material identified but not measured.
Taxonomic notes. - Historically, recognition of specimens of this taxon has presented difficulties.
Edwards (1932: 26) incorrectly synonymized Chaoborus trivittatus with Chaoborus nyblaei. Chaoborus
trivittatus and Chaoborus punctipennis were erroneously considered conspecific by Brunetti (1911:
229), Dyar (1902: 201), Giles (1902: 502) and Theobald (1901b: 296; 1905: 43). Distributional
information and/or type of habitat in which larvae were found suggest that Allan (1973; original
material examined), Dodson (1970), Main (1953; original material examined), Maleug and Hasler
(1967), Myklebust (1966), Stahl (1966) and Teraguchi and Northcote (1966) incorrectly identifed
specimens of Chaoborus trivittatus as Chaoborus nyblaei. Some specimens examined by Smith (1960b)
were probably larvae of Chaoborus trivittatus as indicated by the number of anal fan setae and
PAL/PAW. Similarly, data about overwintering larvae studied by James and Smith (1958) at
Churchill, Manitoba indicate that these were individuals of Chaoborus trivittatus. I was unable to
confirm the identificaion of some specimens from British Columbia collected by Hearle (1928) as
Chaoborus trivittatus.
Shape of the penis valve of the type specimen (Fig. 10 O) confirmed the naming of this species.
Examination of the type of Sayomyia knabi showed that the diagnostically (for that geographical
Systematics and Bionomics of Schadonophasma
147
region) thick tentorium of the larvae of Chaoborus trivittatus was present. All measurements were
within the range of the third instar larvae of Chaoborus trivittatus. In addition, the pharate fourth instar
larva was evident.
All diagnostic features of Chaoborus brunskilli (Saether, 1970: 21) are within range of variation of
Chaoborus trivittatus. Penis valves of the holotype, when reoriented (Fig. lOM), were inseparable from
those of Chaoborus trivittatus. Specimens of Chaoborus brunskilli mentioned by Anderson and
Raasveldt (1974) and Hamilton (1971) are therefore Chaoborus trivittatus.
The identification of the specimen from Lac Phillipe, Quebec, as Chaoborus trivittatus by Saether
(1970) could not be confirmed as the penis valves were distorted. I have not included the specimen in this
description.
Previous descriptions of pupae are given by Saether (1970) (as Chaoborus trivittatus and Chaoborus
brunskilli), by Cook (1956) (as Chaoborus nyblaei) and by Felt (1904).
The only useful previous descriptions of fourth instar larvae are those by Saether (1970) (as
Chaoborus trivittatus and Chaoborus brunskilli) and Felt (1904). Cook’s (1956) description of larvae
of Chaoborus nyblaei undoubtedly included specimens of both Chaoborus trivittatus And Chaoborus
cooki (see especially description of head capsule length and PAL/PAW).
Saether (1970) gave the only previous descriptions of first, second and third instar larvae of
Chaoborus trivittatus (as Chaoborus brunskilli).
Chaoborus cooki Saether
Chaoborus cooki Saether 1970:18. Details of holotype, allotype and paratypes given by Saether (1970);
(CNC).
Chaoborus nyblaei'. Cook 1956: 28, in part (not Zetterstedt).
Characters tested to distinguish between adults of Schadonophasma species are described here to
supplement the descriptions by Saether (1970).
Description. — Males. General coloration of most specimens light grayish brown; specific pigmentation as other members of
subgenus; vittae generally brown; penis valves as shown in Fig. I I A-N. Measurements and proportions; see Table I I .
Females. Coloration as for males. Measurements and proportions: see Table I 2.
Male pupae. Non-teneral individuals with tip of abdomen nearly horizontal when at rest (Fig. 5A); coloration as other members of
subgenus. Measurements and proportions: see Table I 3.
Female pupae. Non-teneral individuals with tip of abdomen nearly horizontal when at rest (Fig. 5A); mature individuals (pharate
adult setae evident) with ovaries extended at most to abdominal segment V. Measurements and proportions: see Table 14.
Fourth instar larvae. Coloration as other members of subgenus. Measurements and proportions; see Table I 5.
Third instar larvae. Coloratin as other members of subgenus. Measurements and proportions; see Table 16.
Second instar larvae. Coloration as other members of subgenus. Measurements and proportions: see Table I 7.
First instar larvae. Egg burster on dorsum of head capsule prominent (Fig. 7E), pigmented; posterior margin of head capsule
noticeably pigmented. Measurements and proportions: see Table 18.
Eggs. Laid in spherical mass with slight amount of jelly-lke matrix; egg with thick exochorion (Fig. 9c), with polygonal
sculpturing (Fig. 4D). Measurements and proportions: see Table 19. Egg shell as in Eig. 9E.
Bionomics. - Overwinters as egg, immatures restricted to temporary occasionally permanent ponds in
woodland. Larvae may be collected only during summer months (Fig. 25B). Adults generally emerge
later in season than do those of Chaoborus trivittatus.
Derivation of specific epithet. - Named after Dr. E.F. Cook.
Distribution. - Specimens of Chaoborus cooki have been collected only from Canada and Alaska
(Fig. 18,19). Collecting in Alberta has shown that the species is restricted to woodland. Paucity of
records, as compared to Chaoborus trivittatus is explained by lack of collecting in the restricted habitat
in which the species is found, and the presence of larvae only during the summer months.
Quaest. Ent., 1979, 15 (2)
148
Borkent
Systematics and Bionomics of Schadonophasma
149
Table 13. Descriptive statistics for male pupae of Chaoborus cooki. Measurements in pm unless stated
otherwise.
Table 14. Descriptive statistics for female pupae of Chaoborus cooki. Measurements in pm unless stated
otherwise.
Table 15. Descriptive statistics for fourth instar larvae of Chaoborus cooki. Measurements in pm.
Quaest. Ent., 1979, 15 (2)
150
Borkent
Table 16. Descriptive statistics for third instar larvae of Chaoborus cooki. Measurements in ^um.
Systematics and Bionomics of Schadonophasma
151
LOCALITIES
Male Adults
ALBERTA: nr. George Lk„ 30-VII-1974 (1 ABOr); 0.9 mi. (1.4 km) W. George Lk., 13-V-1976 (reared) (20 ABOr, 15 CNC, 10
UASM); Edmonton, l-VII-1974 (1 ABOr); MANITOBA: Gillam, 19-V1-1950 (1 OASa), 29-30-V1-1950 (6 CNC); Churehill,
24-VII-1947 (1 CNC), 19-VII-1951 (1 CNC), Vll-VlIl-1950 (4 CNC); NORTHWEST TERRITORIES: Yellowknife, 6-VII-1949
(2 CNC), 17-V1I-1949 (1 CNC); Reindeer Depot, 13-VII-1949 (2 CNC); ONTARIO: Chisholm, 5-VI-1959 (4 CNC), 19-VI-1959
(I CNC); QUEBEC: Great Whale River, 28-3 1 -VI I- 1 949 (6 CNC); 1 3-29-VIII-1949 (2 CNC, 1 OASa); YUKON TERRITORY:
Whitehorse, 1 1 -24-VI I- 1 950 (7 CNC); Mi. 87 (Km 140), Dempster Hwy., 27-30-VI- 1 973 (8 CNC), 8-1 3-VII-I973 (11 CNC),
16-17-VII-1973 (6 CNC),4-8-VIII-1973 (4 CNC).
Female Adults
ALBERTA: 0.9 mi. (1.4 km) W. George Lk., 13-V-1976 (reared) (10 ABOr, 5 CNC, 3 UASM); MANITOBA: Churchill,
22-30-VII-1948 (14 CNC), Vll-Vlll-1950 (10 CNC), 17-V1I1-1951 (1 CNC); ONTARIO: Chisholm, 19-VI-I957 (6 CNC),
3-VI-1959 (1 CNC); QUEBEC: Great Whale River, 29-VIII-1949 (1 CNC); Indian House Lake, 17-VI1-1954 (1 CNC); YUKON
TERRITORY: Whitehorse, 3-V1I-1919 (1 CNC), 7-22-VI I- 1 950 (5 CNC), l-Vlll-1950 (1 CNC); Mi. 87 (Km 140), Dempster
Hwy. 8-13-VI1-1973 (15 CNC), 1 6- 1 7-VI 1- 1 973 ( 1 7 CNC); 4-8-VI 1 1-1 973 (17 CNC).
Male Pupae
ALBERTA: 33 mi. (53 km) S. Jasper, 4-VI-1975 (3 ABOr, 3 UASM); 1.5 mi. (2.4 km) S. Robb, 23-V1I-1975 (3 ABOr);
NORTHWEST TERRITORIES: Yellowknife, 6-VI1-1949 (3 OASa), 14-VI1-1949 (1 CNC); YUKON TERRITORY: Mi. 87 (Km
140), Dempster Hwy., 1 3-VII-1973. ( I CNC).
Female Pupae
ALBERTA: 43 mi. (69 km) N. Coleman, 3-V1II-1975 (5 ABOr, 5 UASM); 20 mi. (32 km) W. Edson, 17-VI-1975 (4 ABOr, 2
CNC, 2 UASM); BRITISH COLUMBIA: 4.8 mi. (7.7 km) N. Clearwater, 12-VI-1976 (4 ABOr); YUKON TERRITORY: Mi. 87
(Km 140), Dempster Hwy., 13-VII-1973 (8 CNC).
Fourth Instar Larvae
ALBERTA: 43 mi. (69 km) N. Coleman, 3-VII1-I975 (13 ABOr); 33 mi. (53 km) S. Jasper, 4-VI-1975, (5 ABOr, 6 UASM); 1
mi. (1.6 km) S. Jasper, 4-V1-1975 (30 ABOr, 15 UASM, 17 CNC); 14.5 mi. (23.3 km) W. Jasper 21-VI-1975 (8 ABOr); 4 mi. (6.4
km) W. Jasper, 20-V-1975 (9 ABOr); 20 mi. (32 km) W. Edson, 5-V1-1975 (15 ABOr, 5 CNC, 4 UASM); 29-V-1976 (3 ABOr); 1.5
mi. (2.4 km) S. Robb, 23-V1I-1975 (2 ABOr); 1.5 mi. (2.4 km) W. Edmonton, 1 2-VI-1975 (5 ABOr, 3 UASM); 0.9 mi. (1.4 km) W.
George Lk., 12-V-1975 (5 ABOr, 5 CNC, 3 UASM); BRITISH COLUMBIA: 4.8 mi. (7.7 km) N. Clearwater, 12-VI-1976 (1
ABOr); MANITOBA: Gillam, IO-VI-1950 (1 CNC); Churchill, 1-1 l-VII-1943 (I USNM), 29-VI-1950 (1 CNC) *nr. Childs Lk.,
27-V-1978 (39 ABOr); NORTHWEST TERRITORIES: Pond nr. Harris River, 2-VI-1973 (1 CNC); ONTARIO: Aberfoyle,
14-V-1974 (1 ABOr); YUKON TERRITORY: Dempster Hwy., 23-30-VI- 1 972 (13 ABOr); Mi. 87 (Km 140), Dempster Hwy.,
13-VII-1973 (1 1 CNC); Klutlan Glacier moraine, Vll-1971 (20 EFCo, 6 ABOr); *01d Crow, 1 9-Vl I- 1 977 ( 1 ABOr).
UNITED STATES
ALASKA: Mi. 6 (Km 10), McKinley, 14-VI-1954 (1 USNM); Mi. 13.5 (Km 21.7) Cantwell Rd., I7-VI-1954 (1 USNM).
Third Instar Larvae
ALBERTA: 1.5 mi. (2.4 km) W. Edmonton, 26-V-I975 (8 ABOr, 4 UASM); 1 mi. (1.6 km) S. Jasper, 4-VI-1975 (5 ABOr, 5
CNC, 4 UASM); 20 mi. (32 km) W. Edson, 29-V-1976 (3 ABOr, 2 CNC, 2 UASM); 0.9 mi. (1.4 km) W. George Lk., 12-V-1975 (8
ABOr, 3 UASM); NORTHWEST TERRITORIES: Pond nr. Harris River, 2-VI-1973 (I CNC); YUKON TERRITORY; Klutlan
Glacier moraine, VII-1971 (6 EFCo).
Second Instar Larvae
ALBERTA: 1 mi. (1.6 km) S. Jasper 25-VI-1976 (3 ABOr); 7 mi. (11 km) E. Obed, 27-1V-1976 (8 ABOr, 4 UASM); 20 mi. (32
km) W. Edson, 27-IV-1976 (4 ABOr, 4 UASM); 0.9 mi. (1.4 km) W. George Lk., 24-1V-1976 (8 ABOr, 4 UASM).
First Instar Larvae
ALBERTA: 1 mi. (1.6 km) S. Jasper 25-1V-1976 (8 ABOr, 4 UASM); 20 mi. (32 km) W. Edson, 27-IV-1976 (4 ABOr, 4 CNC).
Eggs
From male and female adults collected as fourth instar larvae 0.9 mi. (1.4 km) W. George Lk., Alberta and reared and mated in
laboratory (ABor).
* Material identified but not measured.
Quaest. Ent., 1979, 15 (2)
152
Borkent
Taxonomic notes. - Because many specimens of Chaoborus cooki key out to Chaoborus nyblaei
using Saether’s (1972) keys, records of Chaoborus nyblaei by Anderson and Raasveldt (1974) probably
refer to specimens of Chaoborus cooki. The numbers of anal fan setae and PAL/ PAW recorded by
Smith (1960b) suggests that Chaoborus cooki larvae were examined. The suggestion by James and
Smith (1958) that some of the Chaoborus nyblaei population overwinters in the egg stage at Churchill,
Manitoba, indicates the presence of Chaoborus cooki.
The only previous description of Chaoborus cooki is that of the adults, pupae and fourth instar larvae
by Saether (1970). Cook (1956) undoubtedly included specimens of Chaoborus cooki in his description
of Chaoborus nyblaei. The larval head capsule length recorded by Cook (1956) as 2.78 mm, larger than
any recorded here, probably refers to a specimen of Chaoborus cooki. Contrary to Saether’s (1970: 26)
statement. Cook’s (1956) description of the abdomen probably included Chaoborus cooki.
Saether (1970: 20) incorrectly states that Chaoborus cooki larvae do not have a dorsal process on
abdominal segment IX (see page 157).
Chaoborus nyblaei (Zetterstedt)
Erioptera nyblaei Zetterstedt 1838: 830.Three syntypes, all females; LECTOTYPE HERE
DESIGNATED, labelled: ‘Corethr. nyblaei, Z. 9, Erioptera, (illegible word), Lapp Dovre’; one
syntype with no label and the other labelled ‘Corethr. nyblaei, Z.6, Erioptera, (illegible word) Lapp
Dovre.’; (ZMLS).
Corethra nyblaei; Zetterstedt 1851: 3794. Giles 1902: 502. Theobald 1901b: 291.
Chaoborus nyblaei; Edwards 1930: 533. Edwards 1932: 26. Hirvenoja 1961. Saether 1970: 14, in part.
?Corethra pilipes Gimmerthal 1845: 297. Location of male type unknown. Collected at Riga,
Latvijskaja S.S.R.
Description. — Males. (n = two, unless otherwise given in parentheses). Total length 7. 9-8. 5 mm; general body coloration light
grayish brown; vittae brown; specific pigmentation as other members of subgenus.
Antennae: pedicel width 302-319 jum; length of flagellar articles 1,2-1 1,12,13: 244-267 /xm, 148-151 fxm, 354-371 /um, 261 yum;
P/U 1.42-1.56 Mm.
Head: width 1096-1 125 Mm; width between eyes 313-389 Mm; clypeus length 447-464 Mm; prementum length 470-493 Mm; head
width/width between eyes 2.90-3.50; head width/clypeus length 2.42-2.45; head width/prementum length 2.22-2.40; number of
setae on vertex 106 ( 1 ); lengths of palpal articles two to five: 121-139 Mm, 249-261 Mm, 244-313 Mm, 528 Mm ( 1 ).
Thorax: number of setae: pronotal 24; postpronotal three or four; proepisternal seven or eight; preepisternal four to seven;
anepisternals 1 2 or 13; upper mesepimerals 1 4 or 15; scutellar 72-88; supraalar 37 or 38.
Wing: length 4.53-4.63 mm; width l.t)3 mm; length/width 4.41-4.51; wing length/head width 4.03-4.15; wing length/length of
femur of foreleg 1.68 (1); Y/X 1.88-2.13; Y/Z 1.35-1.59; Y/R;j 0.75-0.77; Z/M, 0.44-0.56; R;i/M', 0.94-0.99; number of setae on
squama 38 ( 1 ); length of Rsa 27 1 Mm ( 1 ).
Haltere: capitulum nearly spherical; two to three anterior setae; two to three posterior setae.
Legs: Foreleg: Fe 2690 Mm ( 1 ); Ti 1 25 1 Mm ( 1 ); Ta, 590 Mm ( 1 ); Ta,^ 249 Mm ( 1 ); Hindleg: Fe 2997 Mm ( 1 ). Length of foreleg claw
85 Mm (I); pulvilli length 53 Mm (1).
Genitalia, number of setae on tergite IX 10 (1); penis valve length 194-226 Mm; penis valve as shown in Fig. 11,0 and P;
gonocoxite length 702 Mm ( 1 ); gonostylus length 609-632 Mm; HR 1.15(1); HV 1 .25-1 .39.
Females. (n = three, unless otherwise given in parentheses). Total length 8.0 mm (1); general body coloration light grayish brown;
vittae brown; specific pigmentation as other members of subgenus.
Antennae: (n = 2) pedicel width 162 Mm; length of flagellar articles 1,2-11, 12,13: 192-203 Mm, 124-138 Mm, 180-191 Mm,
197-215 Mm; P/U 0.89-0.91 .
Head. (n = 2) head width 1102 Mm; width between eyes 360-400 Mm;; clypeus length 464-487 Mm; prementum length 493-528
Mm; head width/width between eyes 2.75-3.06; head width/clypeus length 2.26-2.38; head width/prementum length 2.09-2.24;
number of setae on vertex 134-146; length of palpal articles two to five: 151-157 Mm, 290-313 Mm, 302-331 Mm, 586-621 Mm.
Thorax. Coloration as in male.
Number of setae: pronotals 24-43; 70.0; postpronotals 3-1 1, 6.7; proepisternal 8-1 1, 9.3; preepisternal 8-10, 9.0; anepisternals
1 3-34, 24.0; upper mesepimerals 1 7-24, 1 9.3; scutellar 87-1 1 9, 98.3; supraalar 53-66, 59.7.
Systematics and Bionomics of Schadonophasma
153
Wing. (n = 6, unless otherwise given) coloration as other members of subgenus, length 5. 5-6. 3, 5.9 mm; width 1.4-1. 8, 1.6 mm;
length/width 3.55-4.09, 3.84; wing length/head width 5.17-5.25 (2); wing length/length of femur of foreleg 2.22-2.67 (2); Y/X
2.29-3.43, 3.10; Y/Z 1.33-1.67, 1.53; Y/R.j 0.70-0.93, 0.79; ZM, 0.42-0.56, 0.50; R;j/M, 0.91-1.01, 0.96; Rsa 126-302, 238 Mm.
Halteres: capitulum nearly spherical; two anterior setae (2); two to eight posterior setae (2).
Legs. (2), coloration as other members of the subgenus.
Foreleg: Fe 2608-2631 Mm; Ti 2761-2796 Mm; T, 1345-1369 Mm; Ta., 802 Mm; Ta,.j 625-627 Mm; Ta4 389-401 Mm; Ta,/Ti
0.49.
Midleg: Fe 2395-2537 Mm; Ti 2230-2384 Mm; Ta, 1121-1145 Mm; Ta^ 637 Mm; Ta,.; 496 Mm; Ta4 318-342Mm; Ta,^ 273-296
Mm; Ta,/Ti 0.47-0.51; 7 setae in Ta;j comb (1).
Hindleg: Fe 2974-3221 Mm; Ti 2620-2950 Mm; Ta, 1758-1782 Mm; Ta. 909-944 Mm; Ta;, 625-649 Mm; Ta4 366-413 Mm; Ta,^
319-325 Mm; Ta,/Ti 0.60-0.67; 7 setae in Ta-, comb (1).
Length of foreleg claw; 94 Mm; pulvilli length 56-61 Mm.
Genitalia: seminal capsule diameter 65-76; 71.4 Mm; 1.4 (4); cerci length 267-284, 274 Mm (4).
Male pupae. Measurements and proportions: see Table 20.
Female pupae. Measurements and proportions: see Table 21.
Fourth instar larvae. Measurements and proportions: see Table 22.
Eggs: only eggs dissected from abdomens of females were examined. Measurements are probably not, therefore, very accurate.
Eggs from lectotype with thick exochorionic layer (Fig. 9B); eggs from female from Abisko with exochorionic layer as relatively
thick as that shown in Figure 9C. Sculpturing of exochorion not discernable. Measurements and proportions: see Table 23.
Bionomics. - Probably overwinters in egg stage. Larvae have been collected from both permanent
and, possibly, temporary lentic habitats (Hirvenoja, 1961). This species may be able to invade temporary
ponds. Adults emerge about the end of July.
Derivation of specific epithet. - Named after Olavus Nyblaeus.
Distribution. - The distribution of Chaoborus nyblaei is shown in Fig. 20. Although this species is
known from certain records only from Fennoscandia, it may, especially if Chaoborus pilipes from Riga,
Latvijskaja S.S.R. is conspecific with Chaoborus nyblaei, be more widely distributed in the boreal
region of the Palaearctic region. If this species has requirements similar to those of Chaoborus
trivittatus and Chaoborus cooki, Chaoborus nyblaei is restricted to areas of boreal woodland.
LOCALITIES
Male Adults
FINLAND: 2-3 km SW Nuorgam, 24-V11-1960 ( 1 ZMHF); SWEDEN: Abisko, 1931 (1 BMNH).
Female Adults
FINLAND: 2-3 km SW Nuorgam, 24-VII-1960 (2 ZMHF); SWEDEN: Abisko, 1931 (I BMNH); NORWAY; Dovre, 30-VII-?
(2 ZMLS); unlabelled specimen from type series (1 ZMLS).
Male Pupae
FINLAND: 2-3 km SW Nuogram, 24-25-VI1-1960 (2 OASa; 2 ZMHF).
Female Pupae
FINLAND: 2-3 km SW Nuorgam, 24-25-VI1-1960 (4 OASa; 3 CNC; 4 ZMHF).
Fourth Instar Larvae
FINLAND: 2-3 km SW Nuorgam, 24-25-VII-1960 (2 OASa, 2 CNC; 20 ZMHF; 2 ABOr).
Eggs
From females collected as follows: SWEDEN: Abisko, 1931 (I BMNH); NORWAY: Dovre, 30-VII-? ( 1 ZMLS).
Taxonomic notes. - Considerable confusion has surrounded the name of this species. Edwards
(1920: 265), Martini (1931: 58), and Seguy (1942: 169) incorrectly considered Chaoborus nyblaei to be
eonspecific with Chaoborus pallidus (Fabricius). Many authors treated Schadonophasma as monotypic
Quaest. Ent., 1979, 15 (2)
154
Borkent
Table 20. Descriptive statistics for male pupae of Chaoborus nyblaei. Measurements in jum unless stated
otherwise.
Table 22. Descriptive statistics for fourth instar larvae of Chaoborus nyblaei. Measurements in fim.
Table 23. Descriptive statistics for eggs of Chaoborus nyblaei. Measurements in /im.
Systematics and Bionomics of Schadonophasma
155
or have followed Saether’s (1970) conclusion that Chaoborus nyblaei occurs in North America and,
consequently, have incorrectly applied this name to North American specimens of Schadonophasma
(Allan, 1973; Anderson and Raasveldt, 1974; Cook, 1956; Dodson, 1970; James and Smith, 1958; James
et al. 1969; Jenkins and Knight, 1950; Main, 1953; Maleug and Hasler, 1967; Matheson, 1944; McCloy,
1950; Myklebust, 1966; Pens, 1967; Smith, 1960b; Stahl, 1966; Teraguchi and Northcote, 1966).
The only previous detailed description of male adult Chaoborus nyblaei, by Saether (1970), was
based on a specimen of Chaoborus trivittatus from Baffin Island, Canada, and the hypopygium of a
specimen of Chaoborus nyblaei from 2-3 km SW Nuorgam, Finland. The main diagnostic character
was shape of the penis valve. Reorientation of the genitalia of these specimens, however, showed that the
characterization by Saether (1970) was incorrect. Penis valves of the specimen from Baffin Island are
typical of Chaoborus trivittatus. Those from the specimen from Finland are similar to, but different
from, penis valves of Chaoborus trivittatus. I was able to associate the hypopygium from Finland with
the rest of the specimen, and the description includes this male.
Adult characters described by Saether (1970), but not given here, were those which could not be
measured.
Pupae and fourth instar larvae have been previously described by Hirvenoja (1961) and Saether
(1970).
Of the three syntypes of Chaoborus nyblaei only measurements of the female genitalia of the
lectotype are included in this description. The lectotype and paralectotypes were used only to describe
color and wing characters. The features of the pharate female adult prepared from a pupa and described
by Saether (1970), are not included in this description.
Saether (1970: 16) incorrectly states that fourth instar larvae of Chaoborus nyblaei do not have a
noticeable dorsal process on abdominal segment IX (see page 156).
The first, second, and third instar larvae of Chaoborus nyblaei are unknown.
ANALYSIS OF MORPHOLOGICAL VARIATION
The following is a discussion of morphological variation of the stages of Schadonophasma species.
These results describe differences between species of eggs, all larval stages, pupae and adults.
Interpretation of some intraspecific morphological variation as the result of age-related, sex-associated
and geographical variation, and, as described for Chaoborus trivittatus, the result of past distributions,
is also provided. Detailed study of structure, especially when combined with bionomic and behavioral
observations, can allow interpretation of the functional significance of characters. These data thereby
provide an enhanced means of understanding intraspecific variation and phyletic relationships.
Comparison of results with studies of other Chaoborus species is provided to facilitate further
comparison between species of Chaoborus, to reinterpret some past results, and to suggest, in some
instances, more likely interpretation of data presented here.
Artificial variation
An important source of error in determining intraspecific and interspecific variation of species can
result from methods employed in the preservation or preparation of specimens. This factor accounts for
some variation previously described.
Cook (1956: 29) noted that thoracic coloration of adult males of Schadonophasma was related to age
of pinned specimens. Ground color had changed from light grey to yellowish-brown in specimens which
had been pinned for more than twenty years. I have not been able to confirm this observaion which may
be due, in part, to Cook’s (1956) inclusion in his analysis of adults of both Chaoborus trivittatus and
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156
Borkent
Chaoborus cooki, which do differ in thoracic coloration. In addition, glue had seeped unto the thorax of
some of the older pinned specimens producing a darker hue.
The color of all material, of any stage, appeared to bleach when preserved in ethanol for more than
about four or five years. In addition, clearing with KOH also bleaches the color of specimens. It is
uncertain, therefore, whether Saether (1967) adequately deseribed the color variation of KOH-treated
larvae of Chaoborus flavicans.
Saether (1970: 21) recognized the distinct conical dorsal process of Chaoborus brunskilli
{ = Chaoborus trivittatus) fourth instar larvae as diagnostic. Of all material examined, however, only
fourth instar larvae preserved in formalin possessed dorsal processes similar to those of Chaoborus
brunskilli drawn by Saether (1970: fig. 11Q,R) and ethanol-preserved material exhibited flatter dorsal
processes (Fig. 2B; 8C,D). Paratype larvae of Chaoborus brunskilli came from two localities. Hamilton
(1971) states that the specimens from near Kenora, Ontario were preserved in formalin. Anderson and
Raasveldt (1974) do not mention the preservative used to kill the specimens from East Henry Pond,
Jasper National Park, Alberta. Main (1953: 21) reported reduction in total length of 3.3-11.3% of 23
larvae of Chaoborus trivittatus or Chaoborus flavicans preserved for 24 hours in formalin. The
pronounced dorsal process diagnostic of Chaoborus brunskilli is therefore probably a result of shrinkage
in formalin.
Saether (1970) erroneously suggested that Chaoborus cook'i and Chaoborus nyblaei fourth instar
larvae do not exhibit pronounced dorsal processes. These observations were caused by coverslips
compressing cleared specimens and distorting the abdomens. All uncleared fourth instar larvae of
Chaoborus cooki subsequently examined had a developed dorsal process (Fig. 8D). Additional
specimens of Chaoborus nyblaei, which had been cleared but preserved in fluid, were available and I
mounted these under coverslips supported by glass chips. The abdomens retained their natural
dimensions and this allowed observation of the developed dorsal process (Fig. 8C) similar to those of
other fourth instar larvae of Schadonophasma.
McGowan (1976) reported six or eight postantennal filaments for second instar, and eight or ten
postantennal filaments for third instar larvae of Chaoborus edulis. However, the larvae of Chaoborus
pallidipes (Theobald), which are presently indistinguishable from those of Chaoborus edulis (compare
Green and Young, 1976; McGowan, 1976), have the two posterior postantennal filaments distinctly
separated (much more so than described for Chaoborus flavicans by Balvay (1977c)) from the rest of
the filaments and, if Chaoborus edulis also possesses this trait, McGowan’s (1976) results reflect the
breaking off of this separate pair and the variation is probably an artifact of preparation.
Saether’s (1970) misinterpretation of interspecific variation of penis valve shape of adult males
because of lack of standard orientation of these is described elsewhere (p. 177).
Characters varying between larval instars
Hennig (1966a) has discussed the importance of making comparisons only between comparable
semaphoronts in systematic work. The only difficulty in recognizing semaphorants of Schadonophasma
concerns the four larval instars; this section describes differences which permit recognition of each of
these.
Data on instar differences also allowed comparison of developmental patterns from instar to instar
between species and, for some characters, provided important clues for determining the polarity of
morphoclines for phyletic studies.
Differences between instars of Chaoborus have been deseribed by Balvay (1977a, 1977b, 1977c),
Deonier (1943), Fedorenko and Swift (1972), Green (1972), MacDonald (1956), Maleug (1966),
McGowan (1972, 1976), Parma (1969, 1971a), Prokesova (1959), Saether (1967, 1970), Sikorowa
Systematics and Bionomics of Schadonophasma
157
(1967b, 1970, 1973), Smith (1960a), Teraguchi and Northcote (1966), von Frankenberg (1915), and
Weismann (1866). Few workers have considered geographical variation in their analysis of instar
differences (Balvay, 1977a, 1977b, 1977c; Green 1972; Saether, 1967, 1970: 22; Sikorowa, 1973).
Sikorowa (1973) showed that Chaoborus flavicans larvae collected from ponds or lakes differed
significantly in a number of characteristics, suggesting that ecological factors produce at least some
variation. One character, the length of the longest antennal blade, differed significantly between first
instar larvae collected from these two habitat types, implying genetic differences.
It was not previously known, therefore, what characters could be used with certainty to distinguish
instars of material collected over an extensive geographical area, and an extensive range of habitats. This
paper provides that information for larvae of Chaoborus trivittatus and Chaoborus cooki. Although
some mistaken interpretations of characters are discussed here, qualitative differences between instars
are given in the description of Chaoborus and are not repeated here.
Mensural characters previously found to differ between instars, but not necessarily without some
overlap of variation, were total length, distance between anterior and posterior air sacs, head capsule
length, antennal length, distance of antennal seta from base of antenna, long antennal blade, short
antennal blade length, AS/AL, LB/SB, postantennal filament length, PAL/PAW, number of
mandibular fan bristles, and number of anal fan setae. Total length and distance between anterior and
posterior air sacs were not measured because they are clearly related to growth and exhibit continuous
variation (Balvay, 1977b; Eggleton, 1932; Fedorenko and Swift, 1972; Hongve, 1975; LaRow and
Marzolf, 1970; Lewis, 1975; MacDonald, 1956; Main, 1953; Malueg, 1966; McGowan, 1972; Parma,
1971a; Saether, 1967; Sikorowa, 1973; Smith, 1960a; Teraguchi and Northcote, 1966).
Smith’s (1960a) report that Chaoborus crystallinus first instar larvae have six terminal antennal
setae is probably erroneous. Sikorowa (1973: fig. 13) shows only four setae. Further differences between
first instar Chaoborus flavicans as described by Parma (1971a) and first instar Chaoborus crystallinus
as described by Smith (1960a) suggest that some second instar larvae contaminated Smith’s (1960a)
sample of first instar larvae.
Homologous structures in first and later larval instars of Schadonophasma. - To accurately depict
differences between larval instars, homologous structures should be compared. Failure to do so has led to
nomenclatural misinterpretation of certain structures in the past.
There has been confusion concerning the setae of the mandible of first instar larvae of Chaoborus.
Parma (1971a: 178) suggests that Chaoborus flavicans first instar larvae exhibit a mandibular fan
composed of three setae, but Sikorowa (1967b: 88) mentions the presence of four setae. Chaoborus
astictopus first instars do not have a mandibular fan (Deonier, 1943:385). Saether (1970: 22) states that
Chaoborus brunskilli { = Chaoborus trivittatus) first instar larvae have a mandibular fan of nine
bristles. Reexamination of specimen showed a mandible typical of first instars (Fig. 8A) and suggests
that the mandibular fan bristles counted were those of the pharate second instar which was present.
All first instar material examined exhibited mandibles similar to that in Figure 8A (also Balvay,
1977c: fig. 4; Weismann, 1866: fig. 22A).
The homologies of mandibular structures of Chaoborus larvae, as considered by all authors, are
summarized in Table 24.
Balvay (1977c) discusses at length the nomenclature of the mandible of different instars of
Chaoborus flavicans, and gives new names for all structures except the mandibular fan. He showed that
the mandibles of second, third, and fourth instar larvae were structurally similar to each other but were
all different from first instar larval mandibles. Balvay (1977c) considered the “mandibular fan’’ of first
instar larvae to be composed of three setae, and to be homologous to the mandibular fan of later instars,
but does not give any justification for doing so. A fourth articulated seta between the teeth of the
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158
Borkent
Table 24. Comparison of studies of homologies of mandibular structure of larvae of Chaoborus.
mandible and the “mandibular fan” was considered homologous to one of the mandibular setae
( = mandibular spur of Knight, 1971b) of later instars. Mandibular setae of the second instar developed
at the base of the mandibular seta of the first instar larval mandible. In addition, a nerve extends from
the mandibular seta base in the first instar to the bases of the two mandibular setae of the second instar.
The mandibular seta of the first instar larval mandible identified by Balvay (1977c) is therefore clearly
homologous to one of the mandibular setae of later instars.
Although Balvay (1977c) mentions that the mandibular setae of the pharate second instar larva
cannot develop inside the first instar mandibular seta because of the basal articulation, the tips of the
mandibular fan bristles of pharate second instar larvae of both Chaoborus trivittatus and Chaoborus
cooki do develop inside this mandibular seta.
Mandibles of first instar larvae of Chaoborus are similar to those of Mochlonyx (James, 1957: fig. 9)
which possess four (only three shown by James) setae, three of which are toothed and are very similar
and probably homologous to those of later instars of Mochlonyx (Cook, 1956: fig. 22E; Johannsen, 1934:
fig. 155; Meinert, 1886: 65). However, mandibles of second, third and fourth instar larvae of Mochlonyx
also bear mandibular fans undoubtedly homologous to the mandibular fans of Chaoborus. The three
setae on the mandible of first instar Chaoborus larvae are therefore not homologous with the mandibular
fan of later instars as suggested by Balvay (1977c), Parma (1971a), Sikorowa (1967b, 1973), and Smith
(1960a); they are mandibular spurs (Knight, 1971b). Only two of the mandibular spurs, each with a
similar pointed mandibular projection at their base, as all four setae of first instars, are retained by later
instars. I am not certain that the naming of the mandibular fan as the mandibular brush and comb by
Knight (1971b) is justified. I therefore retain the traditional name.
One of two additional setae on the mandible of first instar Chaoborus larvae (Fig. 8 A), called spines
Ej and E2 by Balvay (1977c), is probably homologous with seta O-MP (Knight, 1971b) of some culicid
larvae. Both setae are retained in later instars of Chaoborus (Fig. 8B; Balvay, 1977c: fig. 5,6,7). The
setae are undoubtedly homologous to the two setae on the mandible of first instar Mochlonyx (James,
1957: fig. 9) and Eucorethra underwoodi Underwood (pers. obs.) larvae. Also present in later instars of
Mochlonyx, these two setae are retained in later instars of Eucorethra underwoodi where they are
labelled as the pectinate brush by Knight (1971b: fig. Id). However, their placement, and similarity to
Systematics and Bionomics of Schadonophasma
159
those of the mandible of Chagasia bathana (Culicidae) (Knight, 1971b: fig. 2f), suggests one of these is
seta O-MP. I therefore label the large seta on the mandible of later instars of Chaoborus as such,
although it is not certain which of the two is actually seta O-MP. Knight (1971b) does not label the
second seta. Knight (1971b: fig. le,f) incorrectly labelled seta O-MP as one of the mandibular spurs on
the mandible of Chaoborus americanus (incorrectly identified as Chaoborus flavicans).
Sikorowa (1967b: fig. la,b) erroneously shows both second and fourth instar larvae of Chaoborus
flavicans with three mandibular spurs.
Previous authors have misinterpreted changes in the anal fan between first and second instar larvae.
Balvay (1977b) considered, without justification, each pair of anal fan setae of first instar larvae to
correspond to a single anal fan seta of the second instar. Sikorowa (1970) mentions that these pairs of
setae arise from a common base. This is not so in any of the material I have examined, but the sockets
for the two setae are placed laterally very close to one another. This paired arrangement of setae is
reflected in all later instars in which, although the setae are in an anterior-posterior plane (Fig. 2B), all
anal fan setae are placed alternately slightly one side or the other of the sagittal plane. This, and the fact
that the number of anal setae for first and second instar larvae is nearly identical for both Chaoborus
trivittatus and Chaoborus cooki, supports the view that each seta in the first instar corresponds to a
single seta in the second instar.
Head capsule length. - Variation in this measurement is useful to distinguish Chaoborus larval
instars (Fedorenko and Swift, 1972; Goldspink and Scott, 1971; LaRow and Marzolf, 1970; Malueg,
1966; MacDonald, 1956; Parma, 1969, 1971a: Prokesova, 1959; Saether, 1967, 1970; Sikorowa, 1973;
Teraguchi and Northcote, 1966; von Frankenberg, 1915). However, Balvay (1977c) found that the
range of head capsule length overlapped between third and fourth instar larvae of Chaoborus flavicans.
Variation in lengths of larval head capsules of Chaoborus trivittatus and Chaoborus cooki (Fig. 21 A)
shows, consistent with most previous work, no overlap of range between any of the instars within each
species, although ranges of third and fourth instar Chaoborus trivittatus are proximate. Head capsule
length is sex-associated in fourth instar larvae and therefore probably in third instars and any overlap
between these instars, as shown by Balvay (1977b), is probably due to a mixture of male fourth instar
and female third instar larvae.
Antennal length. - Antennal length is useful to discriminate all instars (or at least 99% of all
specimens when authors do not give ranges) collected at a single locality or in a limited geographical
region (Green, 1972; McGowan, 1972, 1974, 1976; Parma, 1971a; Saether, 1970; Sikorowa, 1973).
Saether (1967) reported a slight overlap between third (330-420 ^um) and fourth (420-640 pm) instar
antennal lengths of Chaoborus flavicans samples from throughout this species’ range. Balvay (1977b)
reported overlap of antennal lengths only between second and third instars of this species collected in
France. Range of variation for Chaoborus trivittatus and Chaoborus cooki (Fig. 21 B) shows that this
character can be used to discriminate all instars. Like head capsule length, to which this character is
correlated at least in male larvae, antennal length of third and fourth instar Chaoborus trivittatus larvae
overlap slightly. As this character also exhibits sex-associated variation in fourth instar larvae, the
proximity between antennal lengths of third and fourth instar larvae is probably due in part to this
source of variation. This, however, could not fully explain Balvay’s (1977b) finding of overlap between
second and third instars only.
Distance of antennal seta from base of antenna/ antennal length (AS/AL). - Parma (1971a: 177) for
Chaoborus flavicans, and Saether (1970: 22) for Chaoborus brunskilli { = Chaoborus trivittatus),
showed that differences in AS/AL distinguish all instars, although Balvay (1977b) showed overlap for
each instar of Chaoborus flavicans. Chaoborus trivittatus and Chaoborus cooki overlap between second
and third, and between third and fourth instars (Fig. 22A). Later in-stars of Chaoborus trivittatus
Quaest. Ent., 1979, 15 (2)
160
Borkent
exhibit, particularly between second and third instars, a substantial degree of overlap of range of
AS/AL. Therefore this character is useful to distinguish all first instar larvae and only some second,
third and fourth instar larvae of Chaoborus trivittatus and Chaoborus cooki. Change in the ratio from
instar to instar is more constant in Chaoborus cooki than in Chaoborus trivittatus, where there is a more
pronounced increase from first to second, and a lesser increase from second to third and third to fourth
instars.
Length of long antennal blade/ antennal length {LB/AL). - This ratio distinguishes all first instar
larvae of Chaoborus cooki and all first and second instars of Chaoborus trivittatus from all later instars
(Fig, 22B). The character can also be used to distinguish some fourth from third instar larvae of
Chaoborus trivittatus and some second, third and fourth instar larvae of Chaoborus cooki.
Length of long antennal blade/length of short antennal blade {LB/SB}. - First instar larvae do not
possess the short antennal blades of later instars. Inequality of length of antennal blades noted by Balvay
(1977b) and Parma (1971a; 176) for first instar larvae of Chaoborus flavicans is also present in later
instars where one of the long antennal blades is slightly shorter than the other three. Both Balvay
(1977b) and Parma (1971a: 177) have shown that, although there were statistical differences between
LB/SB of second, third and fourth instars of Chaoborus flavicans, there was a good deal of overlap of
range. Saether (1967: Table 1) showed overlap of variation of this ratio between third (0.43-0.64) and
fourth (0.50-0.60) instar larvae of Chaoborus flavicans.
Considerable overlap is exhibited between second, third and fourth instars of Chaoborus brunskilli
{ = Chaoborus trivittatus) (Saether, 1970: 22). LB/SB overlaps in range between the last three instars
of both Chaoborus trivittatus and Chaoborus cooki (Fig. 23A). There were differences in the pattern of
change of LB/SB from instar to instar between Chaoborus trivittatus and Chaoborus cooki. At least
some individuals of Chaoborus cooki can be identified to instar using this character where the ratio
decreases with each successive instar. The range of ratio for the three later instars of Chaoborus
trivittatus, however, are nearly identical to each other. There is only a slight increase in the means of the
ratio between second and third instars and a slight decrease between third and fourth instars.
As antennal blades are undoubtedly important in capture and ingestion of prey, differences in
progression of prey type from instar to instar between Chaoborus trivittatus and Chaoborus cooki may
be suggested. However, detailed interpretation of the adaptive significance of this difference must await
further study. Chaoborus trivittatus larvae (Fedorenko, 1975a, 1975b; Swift, 1976; Swift and
Fedorenko, 1975) probably show closer affinities in prey selection to Chaoborus flavicans larvae (Berg,
1937; Dodson, 1970; Parma, 1971b: 43; Sikorowa, 1973: Table 18; Swiiste et al. 1973) which exhibit
allometric change in LB/SB similar to that of Chaoborus cooki. However Chaoborus cooki, because of
the temporary ponds it inhabits, takes quite different types of prey (commonly ostracods, chironomid
larvae, culicid larvae). Therefore, an explanation of differing food types does not adequately explain the
differences in the pattern of change of LB/SB in Chaoborus trivittatus and Chaoborus cooki.
Postantennal filaments. - Parma (1971a), Saether (1967, 1970) and Sikorowa (1973) showed no
overlap of range between instars in postantennal filament length.
Results for Chaoborus trivittatus and Chaoborus cooki (Fig. 23B) show that this character is useful
to distinguish all individuals of each instar of Chaoborus cooki and all, except a few (due to one
specimen each) third and fourth Chaoborus trivittatus. The smallest antennal filament length of fourth
instar larvae is from specimens collected 85 km north of Coleman, Alberta from a high altitude (1975
m) pond which was not shaded. The extreme environmental conditions of this habitat possibly results in
smaller individuals.
Prelabral appendages. - Parma (1971a: 176), Saether (1967: Table 1; 1970: 22) and Sikorowa
(1973: fig. 9) have shown, in Chaoborus species with laterally flattened prelabral appendages in third
Systematics and Bionomics of Schadonophasma
161
and fourth instar larvae, that generally prelabral appendages of third instar larvae are more slender than
are those of fourth instar larvae. This is also true for these instars of Chaoborus trivittatus and
Chaoborus cooki (Fig. 24A). Amount of overlap of PAL/PAW between third and fourth instar larvae is
especially pronounced for Chaoborus trivittatus.
Number of mandibular fan bristles. - This character is useful to identify all specimens to instar in
only a few species (Balvay, 1977b; Deonier, 1943; Green, 1972; McGowan, 1972, 1976; Parma, 1971a;
Saether, 1967; Sikorowa, 1967b; 1973).
There is some overlap of range in number of mandibular fan bristles between second, third, and
fourth instar larvae of both Chaoborus trivittatus and Chaoborus cooki (Fig. 24B) and this character
alone can be used only to recognize some second and fourth instars of either Chaoborus trivittatus or
Chaoborus cooki. Range of number of mandibular fan bristles increases in successive instars at a greater
rate in Chaoborus trivittatus than in Chaoborus cooki.
Anal fan setae. - Differences in number of anal fan setae between larval instars have been described
for some Chaoborus species by Balvay (1977b). Deonier (1943), McGowan (1972, 1976), Parma
(1971a), Saether (1967, 1970), Smith (1960a) and Sikorowa (1973).
Variation of this character for Chaoborus trivittatus and Chaoborus cooki (Fig. 25 A) shows that
only some fourth instar larvae of either Chaoborus trivittatus or Chaoborus cooki can be distinguished
using the number of anal fan setae. First instar larvae may be distinguished from second instar larvae by
their paired arrangement of setae (Balvay, 1977b; Parma, 1971a; Sikorowa, 1970, 1973).
Variation of characters of fourth instar larvae
Age-related variation. - Age-related variation of fourth instar larvae was studied for Chaoborus
cooki only. These were collected 1.6 km south of Jasper on June 4 (n = 29) and June 17, 1975 (n = 33)
with third instar larvae and pupae, respectively. This ensured a sample of fourth instar larvae which
exhibited the total age range. Total length was interpreted as representative of age, and statistical
correlation of the following characters with total length was studied to determine age related variation:
antennal length, long antennal blade length, short antennal blade length, LB/SB, LB/AL, distance of
antennal seta from antennal base, AS/AL, postantennal filament length, prelabral appendage length
and width, PAL/PAW, number of mandibular fan bristles and number of anal fan setae. Shape of
dorsal process was also studied.
Male and female larvae were distinguished using the index described below in the section about
sex-associated variation. Probably because of small sample size, female larvae exhibited no characters
significantly correlated with body length. Three characters of male larvae were significantly negatively
correlated with growth: long antennal blade length, LB/AL, and prelabral appendage length. Because
long antennal blade length and antennal length are not correlated to each other in male larvae, the
correlation of LB/AL to total length probably reflects the correlation of the long antennal blades to total
length. Therefore only two remaining characters are meaningfully related to age. Relationship to total
length of both long antennal blade length (Fig. 26A) and prelabral appendage length (Fig. 26B)
indicates wear with age of these two structures. Wear of long antennal blade is probably associated with
prey capture. Roth (1967: 66) previously noted wear of prelabral appendages of overwintered
Chaoborus albatus larvae. Evidence of damage, particularly to the tip of prelabral appendages,
producing a blunt tip and therefore a shorter prelabral appendage, is common in fourth instar larvae of
Chaoborus trivittatus (Fig. 13G), Chaoborus cooki and Chaoborus nyblaei (Fig. 15A-DJ. Examination
of freshly captured, fluid-preserved material shows that wear of the prelabral appendages is not an
artifact of preparation.
Although the drawing of the prelabral appendage of Chaoborus nyblaei by Saether (1970: fig. 1 IG)
Quaest. Ent., 1979, 15 (2)
162
Borkent
does not show it, the tip had been broken off and the prelabral appendages of Chaoborus nyblaei fourth
instar larvae are in fact more elongate (Fig. 15E-H; Hirvenoja, 1961: fig. 2A). The breaking off of the
tip of the prelabral appendages of Chaoborus trivittatus and Chaoborus nyblaei makes them appear
similar to those of Chaoborus cooki (compare Fig. 15C, 14A-L). However, an undamaged prelabral
appendage of Chaoborus cooki can be separated from damaged ones of Chaoborus trivittatus or
Chaoborus nyblaei through examination of the form of the tip.
Sex-associated variation. - Inclusion of two different groups of semaphoronts in an analysis of
variation of characters can result in an overestimation of the amount of intraspecific variation.
Characters which display a large degree of sex-associated variation may therefore be mistakenly
interpreted as exhibiting a greater degree of intraspecific variation than is actually present, as shown by
the following analysis of some characters of fourth instar larvae of Chaoborus trivittatus and Chaoborus
cooki.
Little has been previously reported about larval sex-associated variation of Chaoborus species
(Bradshaw, 1973: 1249; Smith, 1960a: 198, von Frankenberg, 1915; 514). Teraguchi and Northcote
(1966: fig. 2) give the distribution of head capsule lengths of fourth instar larvae of Chaoborus flavicans
collected in September which exhibits a bimodality. If this represents male and female larvae, as might
be suggested by the results presented below, a comparison of the larvae collected in August and
September would suggest that male larvae molt to fourth instar earlier than do females.
Chaoborus trivittatus. Statistical comparison of 14 characters (Table 25) of 23 male and 21 female
larvae exuviae from reared fourth instars collected 2.4 km west of Edmonton, Alberta, April 26, 1975,
indicated that eight characters differed significantly between male and female fourth instar larvae but
none without some overlap.
To sex larvae without rearing, I constructed a compound character index (Table 26) in which range
of variation of head capsule length, antennal length, AS/AL, long antennal blade length, LB/SB, and
postantennal filament length were divided into numbered units, with typical male larvae given the lowest
unit values, and female larvae the highest. Summation of values of each character for each individual
showed that male and female larvae can be distinguished if the range of variation of each character in
the index is divided into ten units (Fig. 27A). The method, however, depends on a large enough sample
to express the bimodality.
Chaoborus cooki. Fourth instar larvae of Chaoborus cooki were collected 32 km west of Edson,
Alberta on June 5, 1975 and reared to adulthood. Of these, 15 male and nine female larval exuviae were
measured for 14 characters and statistically compared (Table 27). Five characters differed significantly
between male and female larvae with only postantennal filament length showing no overlap of range.
Because of proximity of ranges of postantennal filament length of male and female larvae, this character
cannot be used alone to sex larvae which have not been reared. A compound character index, similar to
that described above, was used to sex larvae (Table 28). Distance of antennal seta from antennal base
was not used in the index because it is so markedly correlated to antennal length (Fig. 29B). Resultant
index values for individual larvae (Fig. 27B) show that larvae can be sexed using this method. Fourth
instar larvae used to study growth related variation were also sexed using a compound character index
(Table 29). Results (Fig. 27C) were tested by examining distribution of characters used to sex larvae.
For example, length of postantennal filaments over-lapped only by one specimen between male and
female larvae.
It may have been due to smaller sample size used in the analysis of Chaoborus cooki that some
characters were sex-associated for Chaoborus trivittatus larvae, but not for those of Chaoborus cooki.
Chaoborus nyblaei. I did not have reared material of Chaoborus nyblaei and was, therefore, unable
to calculate sex-associated variation from fourth instar larvae of known sex. I attempted to sex the larvae
Table 25. Descriptive statistics for male and female fourth instar larval exuviae of Chaoborus trivittatus collected 2.4 km west of Edmonton,
Alberta.1 Measurements in /um.
Systematics and Bionomics of Schadonophasma
163
Quaest. Ent., 1‘979, 15 (2)
164
Borkent
by assuming that those larval characters exhibiting sex-associated variation for both Chaoborus
trivittatus and Chaoborus cooki were also sex-associated for Chaoborus nyblaei larvae and constructing
a compound character index similar to that for the larvae of Chaoborus trivittatus and Chaoborus
cooki, on the basis of head capsule length, antennal length, long antennal blade length, and postantennal
filament length. Presumably because of small sample size (n = 24) I could not confidently interpret the
slight bimodality present in resultant index values.
Table 26. Compound character index used to sex fourth instar larvae of Chaoborus trivittatus collected
2.4 km west of Edmonton, Alberta. Measurements in ixm. (- = no data)
Correlation of characters. - Analysis of correlation of characters gives important clues on how best to
interpret sources of variation, results of compound character indices, and characters used in phylogenetic
analysis. Ratios which are significantly correlated to another ratio with which they share one character,
or to one of the characters from which the ratio is composed are presented, but most of these correlations
are probably artifacts of analysis (Atchley et al, 1976).
Characters of male and female larvae were analyzed separately; this precluded use of specimens of
Chaoborus nyblaei. However, characters which were significantly correlated for male and female larvae
of both Chaoborus trivittatus and Chaoborus cooki were studied for Chaoborus nyblaei fourth instar
larvae. Characters were considered significantly correlated at the 95% confidence limit. Because of low
numbers of female Chaoborus cooki larvae studied, significance or lack of significance may be
fortuitous in some comparisons.
The Chaoborus cooki larvae that were studied for age-related variation and Chaoborus trivittatus
larvae that were used to determine sex-associated variation, were studied in this analysis. In addition, the
Chaoborus cooki larvae studied for sex-associated variation were used to confirm the results from the
aforementioned Chaoborus cooki larvae. However, presumably because of their greater numbers, some
characters were significantly correlated for the Chaoborus cooki larvae used to determine age-related
variation but were not so for the larvae used to study sex-associated variation.
Results for male and female fourth instar larvae of Chaoborus trivittatus and Chaoborus cooki are
given in Figure 28A-D. As for Chaoborus trivittatus and Chaoborus cooki (Fig. 29A,B), distance of
Table 27. Descriptive statistics of male and female fourth instar larval exuviae of Chaoborus cooki collected 32 km west of Edson,
Systematics and Bionomics of Schadonophasma
165
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Quaest. Ent., 1979, 15 (2)
166
Borkent
antennal seta from antennal base is strongly correlated to antennal length of fourth instar larvae of
Chaoborus nyblaei (Fig. 29C).
Correlation between two characters does not necessarily imply a genetic or functional relationship
between those characters. However, lack of correlation suggests little relationship, and I have used only
that information in subsequent analyses.
Geographical variation. - Study of geographical variation of characters of fourth instar larvae of
Chaoborus trivittatus and Chaoborus cooki explained some of the large intraspecific variation of some
characters, and provided evidence for inferring past distributions. In addition, geographical variation can
give supportive evidence of reproductive isolation. Discovery of differences in patterns of geographical
variation of head capsule length, AS/AL, and number of mandibular fan bristles supports the conclusion
from other analyses, that Chaoborus trivittatus and Chaoborus cooki are different species.
It is difficult to determine to what extent clines are products of past and present gene flow or
environmental gradients. However, analyses of intrapopulational correlation show which characters are
associated, and study of patterns of geographical variation in different areas can indicate which
characters are selected independently. Characters such as these, which in some locations show common
patterns, are most likely to reflect gene flow and, possibly, historic events.
The geographical variation of seven characters was studied. However, because head capsule length is
correlated with antennal length, at least in male larvae, and analysis of antennal length variation showed
almost exactly the same pattern as head capsule length for both Chaoborus trivittatus and Chaoborus
cooki, only results of variation of head capsule length are discussed here. Other characters are those used
in the compound character index (Fig. 43).
Chaoborus trivittatus. I studied 21 samples from Alberta and 14 samples, or grouped samples, from
the rest of North America. Within Alberta, only two areas showed geographical patterns of variation for
some of the characters. The samples, numbered 1-5 and 6-9 (Fig. 30), were collected from an area near
to or east of Jasper and north of Swan Hills, respectively.
The characters which show a common pattern are head capsule length (Table 30), AS/AL
(Table 31), PAL/PAW (Table 32) and number of mandibular fan bristles (Table 33). Generally there
is a linear increase or decrease in mean values of samples 1-5 or 6-9, corresponding to the generally
linearly arranged set of localities from which samples were collected. The PAL/ PAW of samples 1 and 6
do not fit the pattern suggested by other samples or other characters. In addition, direction of clines
differs between areas for different characters. Mean values for head capsule length and AS/AL
decrease in a west-east direction in the Jasper area and in a north-south direction in the Swan Hills area,
while mean number of mandibular fan bristles decreases and PAL/PAW increases in a west-east
direction in the Jasper area and south-north direction in the Swan Hills area. While reflecting gene flow,
this pattern suggests the non-correlation of local selection of these two character pairs. Lack of patterns
of variation of these four characters, particularly in southwestern Alberta (6 samples), is probably due to
either a complicating factor or lack of resolution.
Study of six samples, or clumped samples, from Ontario, Quebec, Newfoundland, Wisconsin and
Michigan indicated no apparent patterns of geographical variation, probably because of the few
numbers of samples studied from such a large area.
r could distinguish no patterns of geographical variation of head capsule length for samples from
outside of Alberta but, because this character can vary markedly in a relatively small area, as shown
above, this was probably due to a small number of samples. Therefore it may be significant that, of all
samples studied, specimens from Klutlan Glacier moraine, Yukon exhibited the highest mean head
capsule length (2157 yum).
Two characters, LB/SB and number of anal fan setae, did not exhibit geographical patterns within
Systematics and Bionomics of Schadonophasma
167
Alberta but did vary geographically on the west coast of North America. At least for number of anal fan
setae this discrepancy is explained by the overall homogeneity of samples from Alberta which exhibit,
for example, a total range of means in the Jasper region of 27.5-28.1.
Patterns of geographical variation are apparent along the west coast of North America for AS/AL
(Table 34), LB/SB (Table 35), PAL/PAW (Table 36), number of mandibular fan bristles (Table 37),
and number of anal fan setae (Table 38). Samples are numbered 1-7 (Fig. 31).
Variation in number of mandibular fan bristles and number of anal fan setae, exhibit a linear
north-south dine with mean values increasing in a northerly direction. The number of anal fan setae of
sample 6 (Table 38) is unexpectedly high, however. The other three characters, AS/AL, LB/SB, and
PAL/PAW, show a different pattern of variation, with the highest mean values near the middle of the
dine in the Washington or southwestern British Columbia region. None of these three characters show
any intrapopulational correlation. In addition, study of AS/AL and PAL/PAW in Alberta shows that at
least these two characters are independently selected for. Thus, concurrent patterns of geographical
variation of these three characters probably reflect the historic events discussed below.
It seems likely, considering the present distribution of Chaoborus trivittatus, that this species was
able to survive in both the Alaskan- Yukon refugium, and refugia south of the continental ice sheet
during the Wisconsin glaciation. Because Chaoborus trivittatus is now restricted to woodland conditions
(p. 140), discovery of woodland in parts of the Beringian refugium (Hopkins, 1972) would also suggest
that Chaoborus trivittatus could have survived in this region.
This inferred distribution during the Wisconsin glaciation adequately explains some geographic
variation. Even if an ice free corridor was present in Alberta (Reeves, 1973) and the Alaskan-Yukon
refugial population was not reproductively isolated from some populations south of the continental ice
sheet, this would not affect the argument presented here. Geographical patterns of variation along the
west coast of North America would be the result of resumed reproduction between northern and
southern populations along the west coast as the continental ice retreated (Prest, 1969).
The Yukon sample, from a pond on Klutlan Glacier moraine, exhibits the extreme of clinal variation
of five characters (AS/AL, LB/SB, PAL/PAW, number of mandibular fan bristles, number of anal fan
setae). These patterns are consistent with an hypothesis suggesting the past isolation of this population
(see below).
Johnson (1977) has recently suggested that coastal California had a climate during the last
continental glaciation similar to that now prevailing. There is little doubt that Chaoborus trivittatus
populations were present in the region at that time. It would be reasonable to suggest that clinal
variation of AS/AL, LB/SB and PAL/PAW along the west coast of the United States (i.e., south of the
area formerly occupied by the continental ice sheet) is similar to that existing during the Wisconsin
glaciation. Probably the peak of mean values of AS/AL, LB/SB, and PAL/PAW in Washington or
southwestern British Columbia represents the northern end of clinal variation of past populations.
Subsequent retreat of the continental ice opened a coastal corridor and allowed contact between
populations from the Alaskan-Yukon refugium and those immediately south of the ice sheet, thereby
resulting in intermediate clinal values (Prince Rupert sample).
Chaoborus cooki. I studied seven samples, number 1-7 (Fig. 32), of fourth instar larvae of
Chaoborus cooki in Alberta. Two additional samples from the Yukon, from along the Dempster
Highway and from a pond on Klutlan Glacier moraine, were so similar to each other and distant from
the Alberta samples, that I did not include them in this description.
A common geographical pattern is evident for four characters: head capsule length (Table 39),
AS/AL (Table 40), number of mandibular fan bristles (Table 41), and number of anal fan setae (Table
42). Of the three samples from near Jasper, sample 3 is closest geographically to sample 5. Samples 2, 3,
Quaest. Ent., 1979, 15 (2)
168
Borkent
and 4 are, although not in a straight line geographically, probably linearly arranged in termV of dispersal
of this species along a continuous valley bordered, except to the eastward side of sample 3, by high
mountains.
Table 28. Compound character index used to sex fourth instar larvae of Chaoborus cooki collected 32
km west of Edson, Alberta. Measurements in ixm.
Table 29. Compound character index used to sex fourth instar larvae of Chaoborus cooki collected 1.6
km south of Jasper, Alberta. Measurements in fim.
Systematics and Bionomics of Schadonophasma
169
Quaest. Ent., 1979, 15 (2)
170
Borkent
Table 33. Variation in numbers of mandibular fan bristles of fourth instar larvae of Chaoborus
trivittatus in Alberta. See Figure 30 for location of sample numbers.
Table 34. Variation in AS/AL of fourth instar larvae of Chaoborus trivittatus from the west coast of
North America. See Figure 3 1 for location of sample numbers.
Table 35. Variation in LB/SB of fourth instar larvae of Chaoborus trivittatus from the west coast of
North America. See Figure 31 for location of sample numbers.
Systematics and Bionomics of Schadonophasma
171
Table 36. Variation in PAL/PAW of fourth instar larvae of Chaoborus trivittatus from the west coast
of North America. See Figure 31 for location of sample numbers.
Table 37. Variation in numbers of mandibular fan bristles of fourth instar larvae of Chaoborus
trivittatus from the west coast of North America. See Figure 31 for location of sample numbers.
Table 38. Variation in numbers of anal fan setae of fourth instar larvae of Chaoborus trivittatus from
the west coast of North America. See Figure 31 for location of sample numbers.
Quaest. Ent., 1979, 15 (2)
172
Borkent
Table 39. Variation in head capsule length (in jum) of fourth instar larvae of Chaoborus cooki. See
Figure 32 for location of sample numbers.
Table 40. Variation in AS/AL of fourth instar larvae of Chaoborus cooki. See Figure 32 for location of
sample numbers.
Table 41. Variation in numbers of mandibular fan bristles of fourth instar larvae of Chaoborus cooki.
See Figure 32 for location of sample numbers
Systematics and Bionomics of Schadonophasma
173
Table 42. Variation in numbers of anal fan setae of fourth instar larvae of Chaoborus cooki. See Figure
32 for location of sample numbers.
The pattern of geographical variation for the characters is similar. Sample 1 exhibits the highest
average for all four characters with the exception of AS/AL. The three Jasper samples 2, 3 and 4, show
a successive increase and decrease in mean value of number of mandibular fan bristles and number of
anal fan setae, respectively. There is, generally, a successive increase of mean values of head capsule
length, AS/AL, number of mandibular fan bristles and number of anal fan setae for samples 3, 5, 6 and
7. However, the following are exceptions to this pattern. Sample 5 mean number of mandibular fan
bristles is less than that of sample 3. For AS/AL, the mean value of sample 6 is nearly identical to that
of sample 7. For number of anal fan setae, the mean values of samples 5, 6 and 7 show no significant
change.
Nevertheless, these overall patterns show that some of the greater differences among means may be
attributable to this source. In addition, variation of number of anal fan setae of samples 2, 3 and 4 shows
that characters vary significantly in a small geographic area and therefore, suggests either strong local
selection or restricted gene flow between populations.
Compound character index and characters differing between fourth instar larvae of species of
Schadonophasma. - Initially I had difficulty distinguishing between fourth instar larvae of Chaoborus
trivittatus and Chaoborus cooki. Although I could identify some populations using characters described
by Saether (1970), I found variation of all individual characters to present a confusing continuum (see
Fig. 2 1-25 A) and was uncertain which characters could, in combination, be used to identify members of
these two species. I therefore compared reared material of both sexes of Chaoborus trivittatus and
Chaoborus cooki, used in analysis of sex-associated variation, to discover which characters might differ
between the two species. Characters examined in the study of sex-associated variation differed
significantly, except length of postantennal filaments (see Tables 25, 27) between male larvae and
prelabral appendage width between female larvae. Identifications were confirmed through examination
of associated adults.
For the purpose of this study I treated these two samples as if they were sympatric. Testing for
species differences through initial comparison of samples collected from the same locality reduces
differences due to geographic and ecotypic variation. However, such samples were not available to me
when I initiated this study.
Six characters which differed between both males and females of Chaoborus trivittatus and
Quaest. Ent., 1979, 15 (2)
174
Borkent
Chaoborus cooki and not correlated to each other, were chosen in order to construct a compound
character index to give single values for individual specimens summarizing their features (Table 43).
The correlation between LB/SB and AS/AL for Chaoborus cooki females and between PAL/PAW and
number of mandibular fan bristles of male Chaoborus cooki were considered to be unimportant,
especially as the correlation did not appear in the opposite sex of Chaoborus cooki or in either sex of
Chaoborus trivittatus.
Range of each character was divided into 21 equal units numbered 0-20. The number 0 represented
the extreme of Chaoborus cooki variation of a character and number 20 the opposite extreme of
Chaoborus trivittatus variation. Chaoborus cooki larvae would therefore receive overall lower final
index values than those of Chaoborus trivittatus. To compare all Schadonophasma species, the index,
although based on differences between Chaoborus trivittatus and Chaoborus cooki, included the
variation of Chaoborus nyblaei. Consequently the lowest index values of the number of mandibular fan
bristles and two of the lowest values of number of anal fan setae, represent extremes of Chaoborus
nyblaei variation.
The values of two specimens of Chaoborus trivittatus went beyond the recorded range of LB/SB and
one each beyond the range of PAL/PAW and number of mandibular fan bristles. These specimens were
given the highest index value for that character. It seems unlikely that these changes in procedure
significantly affected the final results.
The distribution of index values for both Chaoborus trivittatus and Chaoborus cooki fourth instar
larvae (Fig. 33) shows that almost all specimens can be identified with confidence. In addition, if more
than seven specimens are examined, range of means of compound character values of Chaoborus cooki
(34.0 - 43.3) and Chaoborus trivittatus (63.8 - 81.0) allows identification of all samples. Although one
character was sex-associated for Chaoborus cooki and three for Chaoborus trivittatus, there was no
significant difference between index values of male and female Chaoborus trivittatus. There was a
significant difference between index values of male and female larvae of Chaoborus cooki but with a
large degree of overlap. I therefore did not consider it important to separate male and female larvae. In
addition, most samples were too small to confidently distinguish male and female larvae using a
compound character index.
Chaoborus nyblaei fourth instar larvae (n = 20) index values had a range of 16-47, mean of 36.4,
1.5SD of 11.6 and 2SE of 3.6. Therefore Chaoborus nyblaei fourth instar larvae can be distinguished
from those of Chaoborus trivittatus but not from those of Chaoborus cooki using this index.
Specimens from Russell Lake, Alberta (n=10, range = 43-57, mean = 52.3, 1.5SD = 7.1, 2SE =
3.0), from Mechant Lake, Quebec (n=ll, range = 50-62, mean = 57.1, 1.5SD = 6.8, 2SE = 2.7)
and from Gallienne Lake, Quebec (n=19, range = 45-66, mean = 55.3, 1.5SD = 8.4, 2SE = 2.6)
exhibited intermediate index values and could therefore not be identified. These samples are described
and discussed below under ‘Populations incertae sedis\
Mean index values did not exhibit geographical patterns of variation. This was, probably, partially
the result of independent patterns of geographical variation of individual characters evidenced, for
example, by some characters of Chaoborus trivittatus larvae in Alberta.
On the basis of the above results I could determine that variation of each character for all samples
studied, although with overlap, differed significantly between fourth instar larvae of Chaoborus
trivittatus and Chaoborus cooki (Fig. 2 1-25 A; Table 6, 15). Fourth instar larvae of Chaoborus nyblaei
were significantly different in all characters from those of Chaoborus trivittatus but were only
significantly different from those of Chaoborus cooki in postantennal filament length, number of
mandibular fan bristles and number of anal fan setae. Both second and third instar larvae of Chaoborus
trivittatus and Chaoborus cooki differed significantly for each character studied. First instars differed
Table 43. Compound character index used to recognize fourth instar larvae of Chaoborus trivittatus and Chaoborus cooki. Measurement in jum.
(• = no data)
Systematics and Bionomics of Schadonophasma
175
Quaest. Ent., 1979, 15 (2)
176
Borkent
in all characters except LB/AL and postantennal filament length.
Saether (1970: 21) recognized number of mandibular fan bristles (23-32), LB/SB (1.8 - 2.3) and
LB/AL (0.70 - 0.79) as diagnostic for Chaoborus brunskilli fourth instar larvae. However, it is clear
from the distribution of these characters of larvae of Chaoborus trivittatus (Fig. 34A-C) that these
characters do not exhibit any bimodality suggestive of another species, and cannot be used to distinguish
the larvae placed in Chaoborus brunskilli as members of a separate taxon.
Because some species of Chaoborus exhibit a limited range of number of mandibular fan bristles
Saether (1970: 18) argued that ‘At least it seems very unlikely that a variation of 1 1-32 setae, the total
variation within Schadanophasma (sic), may be accounted for by the presence of only one or two
species’. I can see no justification for such an argument. Determination of a large range of variation of a
character, such as the number of mandibular fan bristles of Chaoborus trivittatus reported here, is not a
valid criterion for suspecting the presence of more than one species.
Variation of characters of pupae
Pupal material was identified on the basis of reared material, of association with known larvae, of
geographical distribution of the species or, observations of pharate adult characters.
The most detailed, recent description of pupae of all species of Schadonophasma has been provided
by Saether (1970). All material was correctly identified with the exception of pupae of Chaoborus
brunskilli { = Chaoborus trivittatus). The pupa from Stanford, California was not reexamined but the
locality from which it was collected and its association with larvae of Chaoborus trivittatus shows
clearly that this specimen was correctly identified.
Saether (1970), however, did not take into account the marked sex-association variation. His
description of Chaoborus cooki pupae included four males, of Chaoborus brunskilli {^Chaoborus
trivittatus) one male, and of Chaoborus nyblaei two males and ten females. The sex of the pupa of
Chaoborus trivittatus is not known, but as Saether (1970) recorded genital sac length only for male
pupae, and did not do so for this specimen, suggests that it was a female.
Descriptive statistics for both sexes of each species for most characters described below are given in
the species descriptions and comparison of that data shows only those characters presented in the keys to
be useful for species determination.
Because of the small size of individual samples, to estimate sex associated variation of Chaoborus
trivittatus and Chaoborus cooki pupae I compared all males with all females within each species.
Compared material of Chaoborus nyblaei was from a single locality. Seven characters were studied:
abdomen length; length, width and length/ width of respiratory horns; .and length, width and
width/length of abdominal segment seven. The shape and size of the genital lobes differs markedly
between male and female Chaoborus pupae (Fig. 2C,D; Deonier, 1943: fig. 1,2). In addition, female
pupae are slightly darker in overall coloration than males collected from a single locality.
Five characters were sex associated in pupae of Chaoborus trivittatus: length and width of
respiratory horns; length and width of abdominal segment seven; and total abdominal length. Length,
width and length/ width of the respiratory horns; and length and width of abdominal segment seven were
sex associated for pupae of Chaoborus cooki; for pupae of Chaoborus nyblaei length and width of
respiratory horns; length, width and width/length of abdominal segment seven; and total abdominal
length were sex-associated.
It is reasonable to assume that length and width measurements are sex-associated because of a
difference in overall size of male and female pupae. Total abdominal length is sex-associated in pupae of
Chaoborus trivittatus but not in those of Chaoborus cooki. This probably reflects stage of development
of eggs, which may distend the abdomen and are more developed in older female pupae of Chaoborus
Systematics and Bionomics of Schadonophasma
177
trivittatus than in Chaoborus cooki. Because this character is sex-associated for pupae of Chaoborus
nyblaei this suggests that ovaries of this species develop in a similar fashion to Chaoborus trivittatus.
Saether (1970: 18) noted that width/length of abdominal segment VII (erroneously stated as VIII) is
1.4 for Chaoborus trivittatus, 1.5 for Chaoborus brunskilli {== Chaoborus trivittatus), 1.6 for
Chaoborus cooki, and 1.7-1. 9 for Chaoborus nyblaei pupae. Examination of Tables 4, 5, 13, 14, 20 and
21 shows extensive overlap of range of this character between males or females of each species.
Coloration differences noted by Saether (1970) were unreliable when more material was examined.
There is little doubt that ethanol preserved specimens lose at least some of their color with time. In
addition, Saether (1967) has shown that coloration of the anal paddle ribs varies extensively within
pupae of Chaoborus flavicans. However, his results may be biased through use of specimens treated with
KOH.
Differences of the angle of the abdomen in relation to the longitudinal axis of the body between pupae
of Chaoborus trivittatus and Chaoborus cooki (Fig. 5A,B) were observed from most of the samples as
they were collected and/or reared in the laboratory, from localities in Alberta during the course of this
study. The character was consistent for identification of pupae examined.
Differences in pupal ovarian development of Chaoborus trivittatus and Chaoborus cooki were
apparent (see key to female pupae) and this trait is also expressed in the stage of ovarian development in
teneral female adults (see below). In live female pupae of Chaoborus trivittatus at least 48 hours old (at
20 C) ovaries are macroscopically apparent as white elongate organs in the abdomen (see Parma, 1971b:
40). These are not apparent in female pupae of Chaoborus cooki.
Variation of characters of male adults
Saether (1970) noted two characters of diagnostic value for determination of adult males to species:
shape of penis valves and X/Y wing ratio. These two characters were studied in detail. Other differences
between males of the three species of Schadonophasma are available by comparison of the descriptions
(Table 2, 11). Characters differing significantly between male adults of Chaoborus trivittatus and
Chaoborus cooki are Rsa length, all leg measurements, gonocoxite and gonostylus length, Y /X, number
of tergite IX setae and penis valve length.
Differences in coloration of adults of Chaoborus trivittatus and Chaoborus cooki (see descriptions)
may in part be ecophenotypic. Bradshaw (1973: 1256) mentions that male adult Chaoborus americanus
are a darker color when reared at temperatures below 15 C, and this may also have an effect on color
variation of adult Schadonophasma (also see Rapoport, 1969; Vernberg, 1962).
Range of variation of Y /X of male adults of Chaoborus trivittatus (Table 2) and Chaoborus cooki
(Table 11), although significantly different, exhibits extensive overlap. Range of variation of Y/X of
Chaoborus nyblaei male adults (see description) is within the range of Y /X of both Chaoborus
trivittatus and Chaoborus cooki.
I studied geographical variation of this character for Chaoborus trivittatus and Chaoborus cooki but,
possibly because of the small number of samples, recognized no patterns. However, it is clear that the
ratio, at least for Chaoborus trivittatus, varies markedly in a relatively small geographical area. A
sample from 16 km west of Jasper, Alberta, (n = 38, range = 1.35 -2.18, mean = 1.705, 1.5SD =
0.310, 2SE = 0.067) differed significantly from a sample from 2.4 km west of Edmonton, Alberta,
(n = 40, range = 1.54-2.42, mean = 1.915, 1.5SD = 0.309, 2SE = 0.065).
Saether (1970) distinguished four types of penis valves characteristic of each of the, four species he
described. Apparently because of the similarity of penis valves of the male from Baffin Island with those
of the specimen from Finland, he identified the former specimen as Chaoborus nyblaei. In addition,
differences in the penis valves of the male from Lk. 241, Kenora, Ontario supported his recognition of
Quaest. Ent., 1979, 15 (2)
178
Borkent
the larvae from that area as indicative of the new species of Chaoborus brunskilli. However, these penis
valves were not examined in a standard orientation. Reorientation of the penis valves of the specimens
from Baffin Island and Lk. 241, Kenora resulted in forms indistinguishable from those typical of
Chaoborus trivittatus. The dark, very short tips of the penis valves of Chaoborus nyblaei, as drawn by
Saether (1970: fig. 7A,B), are actually the claws pointing downward. Outlines of a single penis valve of a
male Chaoborus trivittatus (Fig. 12A-E), drawn from different aspects, shows that one orientation (Fig.
12D) is very similar to that thought by Saether (1970: fig. 7A,B) to be typical of Chaoborus nyblaei,
and another (Fig. 12B) typical of Chaoborus brunskilli (Saether, 1970: fig. 7C). The effect of changed
orientations of a single penis valve of Chaoborus cooki (Fig. 12F-J) shows the importance of examining
the penis valves from different aspects. For example, one orientation of the valve of a male of Chaoborus
cooki (Fig. 12F), is extremely similar to one orientation of the penis valve of Chaoborus trivittatus (Fig.
12A). For the purposes of this study only Figures 12E and 12H show useful orientations. Of the material
I examined, no penis valves of Chaoborus trivittatus looked like those of Chaoborus cooki when in this
orientation.
The two types of penis valves recognized by Saether (1970: 27) to be typical of two forms of
Chaoborus americanus are also a result of examination of material from different aspects (pers. obs.).
Because of the seemingly impossible prospects of completely standard orientation of penis valves,
even though all specimens were drawn with the penis valve head in a horizontal plane, it is not known to
what degree this affected the drawings of intraspecific and interspecific variation of penis valves of
Chaoborus trivittatus (Eig. lOA-O) and Chaoborus cooki (Fig. IIA-N). Although a comparison
between amount of intrapopulational and interpopulational variation of penis valves of either Chaoborus
trivittatus or Chaoborus cooki do not show very marked differences in these figures, there is actually
more interpopulational than intrapopulational variation.
The penis valves of male Chaoborus nyblaei are illustrated in Fig. 1 1 0,P.
Length of penis valves differed significantly between Chaoborus trivittatus and Chaoborus cooki,
even though the entire range of variation of that of Chaoborus cooki was contained in the range of
variation of length of penis valves of Chaoborus trivittatus. Length of penis valves of Chaoborus nyblaei
was within the range of both Chaoborus cooki and Chaoborus trivittatus.
The only apparent pattern of geographical variation of the length of penis valves was found in
specimens of Chaoborus trivittatus from along the west coast of North America (Table 44). Although
this linear increase in mean values cannot be confidently interpreted on its own, geographical variation
in five larval characters in this region suggests that the pattern is significant (see section on geographical
variation of fourth instar larvae).
Number of setae on one side of tergite IX was studied only because it was convenient to do so when
examining penis valves. Saether (1970) has already shown a large amount of overlap of variation of this
character between Chaoborus trivittatus and Chaoborus cooki, although analysis here does indicate
them to be significantly different. I could detect no patterns of geographical variation of this character
for either species.
Variation of characters of female adults
The only diagnostic feature of females given by Saether (1970) was Y/X wing ratio of Chaoborus
trivittatus and Chaoborus cooki. Eemale adults were not available for Chaoborus nyblaei.
I identified females on the basis of their temporal and geographical association with males,
geographical distribution or type of ovarian development and egg type. I studied only variation of Y/X
wing ratio in detail. Other differences between females of these species are in the descriptions.
Systematics and Bionomics of Schadonophasma
179
Table 44. Variation of penis valve length (in pm) of male adult Chaoborus trivittatus.
Sample 1 from Pacific Grove, Stanford, Oakland and Mad River Beach, California. Sample 2
from Hoodsport, Port Madison and Bremerton, Washington, and Kaslo, British Columbia.
Sample 3 from Prince Rupert and Terrace, British Columbia.
Characters which differed significantly between Chaoborus trivittatus and Chaoborus cooki were
length of penultimate antennal article, prementum length, HW/Pl, number of anepisternal setae, all leg
measurements, and seminal capsule diameter. Number of setae on the comb of the third tarsomere of
either midleg or hindleg has not been previously recorded but was found to differ significantly between
the two species.
Although means of the ratio Y/X of female adult Chaoborus trivittatus and Chaoborus cooki differ
significantly, ranges overlap extensive (Table 3, 12). Mean values did not exhibit geographical patterns
of variation. As with males, this character of females can vary markedly in a relatively short distance. A
sample from 16 km west of Jasper, Alberta, (n=10, range = 2.03-2.86, mean = 2.414, 1.5SD =
0.425, 2SE == 0.179) differed significantly from a sample 2.4 km west of Edmonton, Alberta, (n = 28,
range = 1.57-3.22, mean = 2.267, 1.5SD = 0.538, 2SE = 0.136). However, a sample from eight
localities in California (n=19, range = 1.19-2.95, mean - 1.919, 1.5SD = 0.653, 2SE = 0.200) did
not differ significantly from a sample from Terrace and Atlin, British Columbia and Whitehorse,
Yukon, (n= 15, range = 1.46-2.56, mean = 2.011, 1.5SD = 0.444, 2SE = 0.153).
Females of Chaoborus trivittatus and Chaoborus cooki differ in ovarial development. Female
Chaoborus trivittatus less than one hour old, reared from fourth instar larvae collected 2.4 km west of
Edmonton, Alberta, had abdomens full of eggs . Ovaries of some females extended into the thorax. A
ratio of distance of the anterior tip of the longest ovary from the end of the abdomen/abdominal length
was 0.71-1.11 (mean = 0.967). The ovaries together contained 251-329 (mean = 279) eggs which, at
this stage, were each at least 348 pm long. Fedorenko (1975c: 3102) recorded 161 ± 7.4 SE eggs per
female pupae from Eunice Lake, British Columbia.
Females of Chaoborus cooki less than 12 hours old, collected as early instars 1.6 km south of Jasper,
or as fourth instar larvae, 1.4 km west of George Lake, Alberta, had much smaller ovaries with a ratio of
the distance of the anterior tip of the longest ovary from the tip of the abdomen/abdominal length of
0.32-0.62 (mean == 0.444). Total number of eggs present was 74-95 (mean = 86). Each egg in a
teneral female is no more than 120 pm long. Teneral females of these two species can therefore be
distinguished from one another by size of ovaries. A nulliparous female can be identified by number of
eggs present. If an individual with at least some eggs present is suspected of being parous, the eggs are
developed enough to distinguish the type. A thick exochorion is typical of Chaoborus cooki (Fig. 9C)
and a thin exochorion of Chaoborus trivittatus eggs (Fig. 9A).
I had available only two females of Chaoborus nyblaei which still had eggs in their abdomens. One
was the lectotype and its abdomen contained eggs with a thick exochorion (Fig. 9B). I could not count
the number of eggs present. The other female, from Abisko, Sweden, had the same type of egg and its
abdomen contained 58 eggs. Implications of differences in egg type are discussed below in the section on
development of eggs.
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180
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Descriptions of populations incertae sedis
Three samples of larvae and one of pupae could not be identified with certainty. The three larval
samples exhibited compound character index values (see p. 174) intermediate between those of
Chaoborus trivittatus and Chaoborus cooki. Although the two samples from Mechant and Gallienne
Lakes, Quebec are here described separately, they possibly belong to the same reproductive population,
considering the close proximity of the lakes. Pope et al. (1973) recognized samples from these two lakes
as members of a new species or subspecies of Schadonophasma but did not describe them.
Several interpretations of these samples seem equally reasonable. Individuals may belong to one or
more new species or they may be hybrids between Chaoborus trivittatus and Chaoborus cooki, although
number of specimens would seem too large to be the product of hybridization (see section on mating
experiments). Also, specimens may be members of Chaoborus cooki. Data from the sample from Russell
Lake, Alberta, shape of prelabral appendages of all larvae of all samples, and shape of penis valves of
pupae from Gallienne Lake, Quebec, suggests the likelihood of this (see below for details). None of the
specimens in the analysis of Chaoborus cooki came from lakes. Chaoborus cooki may possibly invade
this habitat and resultant individuals may therefore be phenotypically different from individuals of
Chaoborus cooki described in this study. Sikorowa (1973) has shown that habitat influences variation of
characters of larvae of Chaoborus flavicans. Examination of adults, study of geographical variation once
more material is available, especially from eastern Canada, study of the life cycle, and developmental
studies could be used to test this possibility.
Sample from Russell Lake, Alberta. - All specimens collected (1 -VIII- 1969) were fourth instar
larvae. The population from which this sample was taken is within the geographical range of both
Chaoborus trivittatus and Chaoborus cooki. Coloration is as other members of the subgenus.
Measurements and proportions are given in Table 45.
Table 45. Descriptive statistics for fourth instar larvae of Schadonophasma from Russell Lake, Alberta.
Measurements in ixm.
Variation of most characters gave no clues as to whether this sample might be composed of larvae of
either Chaoborus trivittatus or Chaoborus cooki, or of a third undescribed species, because the range of
these characters was within the range of variation of both Chaoborus trivittatus and Chaoborus cooki.
Low PAL/PAW values would suggest the sample is conspecific with Chaoborus cooki. However, some
specimens exhibit LB/AL values, number of mandibular fan bristles, and number of anal fan setae
wh.ch are beyond the range of Chaoborus cooki and into the range of variation of Chaoborus trivittatus.
Systematics and Bionomics of Schadonophasma
181
All characters were examined in relation to clinal variation of these characters of both Chaoborus
trivittatus and Chaoborus cooki for clues of conspeeificity. Only mean number of mandibular fan
bristles may possibly be explained by clinal variation of this character of Chaoborus cooki fourth instar
larvae.
All material deposited with ABOr.
Sample from Mechant Lake, Quebec. - All specimens were collected (14-VII-1975) as fourth instar
larvae. Although Pope et al. (1973) report that a population represented by this sample coexisted with
larvae of Chaoborus trivittatus, I did not examine any material of Chaoborus trivittatus whieh would
have given better clues concerning possibilities of the two being conspecific. However the recognition of
two forms in a single lake does not negate possibilities of them being conspecifie. Fedorenko and Swift
(1972) described two forms of Chaoborus trivittatus which differed in size and color but merely
represented two overlapping generations.
Larvae are darker than any other Chaoborus larvae I have examined. Head capsules were notably
darker. In particular, dorsum of head capsule, area around the anterior tentorial pits and dorsal from
each pit to dorsum of head capsule, posterior edge of head capsule including ocular-antennal segment,
ventral edge of head capsule between antennae and labrum, antennae, maxillary palpus and seta, and
anal fan setae were all more darkly pigmented. All thoracic and abdominal segments were infuscated
dorsally. Measurements and proportions are given in Table 46.
Some individuals have AS/AL and PAL/PAW values within the range of Chaoborus cooki and
beyond the range of Chaoborus trivittatus. Conversely, some individuals exhibited head capsule length,
antennal length, LB/SB, LB/AL, and number of mandibular fan bristles within the range of variation
of those characters of Chaoborus trivittatus and beyond the range of those of Chaoborus cooki.
All material deposited with ABOr.
Sample from Gallienne Lake, Quebec. - All specimens were collected (13-VIII-1971) as fourth
instar larvae. Color of larvae was similar to that of larvae from Mechant Lake, Quebec. For
measurements and porportions see Table 47.
Some individuals exhibited LB/SB, LB/AL ratios and number of anal fan setae beyond the range of
variation of these characters for Chaoborus cooki but within the range of those of Chaoborus trivittatus.
Prelabral appendages were typical of those of Chaoborus cooki larvae. Head capsule, antennal and •
postantennal filament length, and number of mandibular fan bristles of this sample differed significantly
from those of larvae from Mechant Lake. However, this might be explained by geographical variation or
differences in habitat. No ratios significantly differed between the two samples.
Pupae were darker than any other Chaoborus material examined. Measurements and proportions of
male pupae are given in Table 48, and of female pupae in Table 49. Nearly all measurements of both
male and female pupae were within, or less than, the range of variation of Chaoborus trivittatus. Only
the WS/LS of male pupae was within the range of WS/LS of both Chaoborus trivittatus and
Chaoborus cooki.
Penis valves were developed in three male pupae but, because these could not be removed to study
from different orientations without destroying the rest of the specimen, they were difficult to interpret.
Penis valves of one of these individuals were most similar to those of Chaoborus cooki adults. However,
length of penis valves ranged from 156.6-162.4 iim. Because material had been previously cleared I
could not determine the degree of ovarian development of two pharate female adults. One specimen had
well developed seminal capsules with a diameter of 50.0 pm.
All material deposited with OASa.
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182
Borkent
Table 46. Descriptive statistics for fourth instar larvae of Schadonophasma from Mechant Lake,
Quebec. Measurements in im\.
Table 49. Descriptive statistics for female pupae of Schadonophasma from Gailienne Lake, Quebec.
Measurements in pm unless stated otherwise.
Systematics and Bionomics of Schadonophasma
183
BIONOMICS
Studies of life histories, developmental characteristics, and behavior give important information on
questions of conspecificity, and suggest how species closely related retain their reproductive unity, such
studies are also important in interpretation of characters used to infer the evolutionary history of such
species and the events pertaining thereto. In this section I present observations made on Chaoborus
trivittatus and Chaoborus cooki in Alberta. These are used to infer some probable bionomic features of
Chaoborus nyblaei in Fennoscandia. For the following analysis observations were made of events which
I had the opportunity and time to deal with and which I believed to be most informative to elucidate
species differences and their evolution. A more complete, general account of such bionomic information
has been presented by Parma (1971b).
Life cycle
All temperate Chaoborus species previously studied exhibit a life cycle in which fourth instar larvae
overwinter, pupate in spring, emerge as adults which lay eggs and develop to fourth instar larvae by
winter (Balvay, 1977d; Bradshaw, 1973; Fedorenko and Swift, 1972; Parma, 1971b; Sikorowa, 1973). In
multivoltine populations these fourth instar larvae result in another adult emergence and subsequent
larvae but, regardless of number of generations per year, every temperate species studied overwinters as
fourth instar larvae. For this reason, immatures of these species are usually present only in permanent
waters. Studies of tropical species (Corbet, 1958; Lewis, 1975; MacDonald, 1956; McGowan, 1974,
1975; Tjonneland, 1958; Verbeke, 1957) suggest that these are also restricted to permanent waters.
Records of Chaoborus larvae in temporary waters in the tropics (Belkin and Heinemann, 1975, 1976;
Heinemann and Belkin, 1977a, 1977b; Panday, 1975) can be explained as incidental as McGowan
(1974: 498) observed adult females of Chaoborus anomalus and Chaoborus ceratopogones (Theobald),
species restricted to permanent waters in Africa, laying eggs in dishes of water in which the larvae
undoubtedly would not survive.
Records of larvae of temperate Chaoborus species from temporary waters (e.g. Anderson and
Raasveldt, 1974; 16) are likely from either a single generation of a multivoltine population which
overwintered elsewhere in permanent waters, a population which eventually became extinct (see p. 186),
or are larvae which were transferred from over-flowing ponds upstream on a river to downstream
inundation pools. Merely the presence of Chaoborus larvae in an aquatic habitat is not sufficient
evidence to suppose survival of the population in this habitat. Records of Chaoborus americanus and
Chaoborus flavicans fourth instar larvae from the Red Deer river, near Innisfail, Alberta, when the river
still had ice on its edge, showed all specimens with empty guts, suggestive of their inability to
successfully survive to adulthood in this habitat. These larvae probably were washed into this permanent
river from ponds further upstream. Records of Chaoborus larvae from the Hudson River, New York
(Beck et al., 1975) are undoubtedly larvae from impoundments further upstream. Peus’s (1934) often
cited record of Chaoborus flavicans from inundation pools along rivers in Germany and Reisen’s (1973)
observation of a single specimen of Chaoborus punctipennis from an inundation pool near a river in
Oklahoma are therefore probably from larvae washed downstream from flooded ponds. Retreat of an
over-flowing river in inundation areas can leave intact a population of Chaoborus larvae (Heinemann
and Belkin, 1977a: 272, Roback, 1966; see p. 185).
Chaoborus trivittatus. Fedorenko and Swift (1972) found a population of Chaoborus trivittatus in an
oligotrophic lake in southwestern British Columbia to be composed of two overlapping generations.
Fourth instar larvae overwintered for two consecutive winters before pupating. Although adults emerged
every spring, larvae which had overwintered only one winter did not pupate. Continual presence of
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184
Borkent
fourth instar larvae throughout and after the pupal emergence period is therefore indicative of this two
year life cycle.
I studied the life cycle of Chaoborus trivittatus in a pond 2.4 km west of Edmonton which measured
approximately 18 by 85 meters, had a maximum depth of about two to three meters, and was densely
shaded by mature stands of Populus balsamifera and Picea mariana. Sampling during 1975 (Table 50)
produced a few fourth instar larvae on May 13, all of which pupated by May 26. These data indicate a
univoltine population of Chaoborus trivittatus. Changes in pupal and pupal exuvial sex ratios is
explained by earlier pupation of male larvae (Table 51) and less time for development of male pupae
(p. 193). Emergence of adults was restricted to later in May. Eggs hatched in three to four days under
laboratory conditions.
Table 50. Number of Chaoborus trivittatus immatures collected 2.4 km west of Edmonton, Alberta.
Table 51. Numbers of pupae resulting from 45 individually reared fourth instar larvae of Chaoborus
trivittatus collected 2.4 km west of Edmonton, Alberta, 25-IV-1975.
Overwintering of fourth instar larvae as a feature of Chaoborus trivittatus is reflected in the range of
collection dates of material examined (Fig. 25B). Gaps in the record undoubtedly reflect lack of
collected material (Fedorenko and Swift, 1972; Main, 1953). James and Smith (1958) also observed
overwintering of some Chaoborus trivittatus larvae (as Chaoborus nyblaei).
Parma (1971b) has discussed how univoltine or multivoltine Chaoborus populations are a result of
environmental factors and vary in response to type of habitat and geographical area. These observations
explain the differences between the bivoltine life cycle of Chaoborus trivittatus described by Fedorenko
and Swift (1972) and the univoltine life cycle described here. In addition, temporal bimodality of records
of field collected adults in southern Ontario and Quebec and northeastern United States (Table 52)
suggest that some populations of the species are multivoltine in this region (see Balvay, 1977c;
Hirvenoja, 1960, 1965; Parma, 1971b).
Systematics and Bionomics of Schadonophasma
185
Table 52. Records of field collected adult Chaoborus trivittatus from southern Ontario and Quebec and
northeastern United States. Roman numerals represent months and Arabic numerals are:
l=Days 1-10,2-Days 1 1-20; 3 = Days 21-30(31).
Chaoborus cooki. The only previous allusion to the life cycle of Chaoborus cooki has been by James
and Smith (1958) who mention that some of the populations of Chaoborus nyblaei (probably =
Chaoborus trivittatus and Chaoborus cooki) at Churchill, Manitoba, overwintered as eggs.
The life cycle of Chaoborus cooki was studied at three locations. A pond 1.6 km south of Jasper
measured 10 by 41 meters with a maximum depth of 0.9 m during most of its existence. Shading on the
north, south and west sides of the pond was provided by Salix sp. and Picea mariana. The pond was
located about 40 m from the south bank of the Miette River and, although it did not initially receive its
water from the river, I observed overflow of its banks and flooding of the pond to about six times its
normal surface area on two occasions, on 19-VI-1976 and 30-VI-1976. Retreat of the river from this
inundation zone seemed to leave most of the population of Chaoborus cooki intact. However, I did
collect several larvae stranded in isolated pools which drained within one day after retreat of the river.
This pond was dry by October 8, 1976.
A pond 32 km west of Edson measured 1 1 by 29 meters, with a maxinium depth of 0.7 m during most
of its existence and was well shaded by dense brush and trees of Picea mariana and Populus
tremuloides. The pond was dry by October 8, 1976.
A third pond, 1.4 km west of George Lake, measured 15 by 19 meters and had a maximum depth of
0.9 m during most of its existence. Although surrounded by dense brush and trees of Picea mariana and
Populus tremuloides on its north, east and west sides the pond was often in direct sunlight. However,
larvae were almost always located in the shade of numerous Typha sp. The pond dried up on two
occasions, 23-VII-1976 and 30-X-1976.
Tables 53, 54 and 55 give numbers of animals collected from these three sites. A common type of life
cycle is apparent. There is a progression within one growing season, of first through fourth larval instars,
pupation, and, observed at 1.4 km west of George Lake and 1.6 km south of Jasper, emergence of adults.
Lack of records of larvae after the pupal period until the next spring suggests that Chaoborus cooki
overwinters in these temporary aquatic habitats as eggs. This conclusion is supported by observed
diapause of Chaoborus cooki eggs in the laboratory and their resistance to desiccation (p. 191).
However, in spite of this adaptation to temporary waters, Chaoborus cooki larvae may occasionally live
in permanent ponds (e.g. 2.4 km west of Edmonton).
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Borkent
This univoltine life cycle, in which the eggs overwinter, is reflected in the restricted temporal
distribution of records of fourth instar larvae of Chaoborus cooki (Fig. 25 B).
It is clear that this species circumvents the problems presented by temporary habitats to other
Chaoborus species. Larvae of Chaoborus americanus were collected late in the season from each of the
three sites studied (Tables 53, 54, 55) and those remaining undoubtedly died when the ponds became
dry. Both live and dead Chaoborus trivittatus larvae and a live Chaoborus americanus fourth instar
larva were taken off wet mud at the location 1.4 km west of George Lake on 18-IX-1976.
The unique life cycle of Chaoborus cooki in temporary ponds allows this species to avoid competition
with other sympatric Chaoborus species, except when some of these occasionally inhabit the same
temporary ponds. Even so, these would be as younger instars than those of Chaoborus cooki.
Chaoborus nyblaei. There is no direct evidence of the type of life cycle of Chaoborus nyblaei.
However, a thick exochorion, similar to that of Chaoborus cooki would suggest that the egg is similarly
adapted to resist desiccation and that this species also overwinters as eggs. Chaoborus nyblaei larvae
have been collected from both a large permanent pond, and a small pool a few meters in diameter, in
northern Fennoscandia (Hirvenoja, 1961).
Low number of eggs per female would support the conclusion that Chaoborus nyblaei immatures
inhabit temporary ponds. Collection of adults only in the later part of July agrees with this
interpretation of the type of life cycle.
Table 53. Numbers of Chaoborus cooki immatures collected 1.6 km south of Jasper, Alberta.
symbolizes dry conditions.
*Chaoborus americanus larvae also present.
Systematics and Bionomics of Schadonophasma
187
Table 54. Numbers of Chaoborus cooki immatures collected 32 km west of Edson, Alberta.
symbolizes dry conditions.
*Seven Chaoborus trivittatus and one Chaoborus americanus fourth instai larvae collected.
Development and behavior
Adults. — Parma (1971b: 45) described the emergence of adult Chaoborus flavicans and Chaoborus
crystallinus. Most observations of Chaoborus trivittatus and Chaoborus cooki agree with that
description, but the following are additions or differences. Wings are released from the exuviae after the
antennae but before the legs become free. Each of the forelegs and midlegs are released one after the
other, but the hindlegs are pulled out together. While the hindlegs are being freed, the wings are folded
over the abdomen. Emergence, from the time of the dorsal split to complete release from the exuviae is,
under laboratory conditions, 44-95 seconds (n = 28) for male and 47-92 seconds (n = 25) for female
Chaoborus trivittatus and 39-70 seconds (n = 30) for male and 38-51 seconds (n=13) for female
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Borkent
Chaoborus cooki. All adult Chaoborus trivittatus emergences in the laboratory occurred during
daylight conditions, and those of Chaoborus cooki only took place during night. Myconial fluid was
released in 5-21 drops. Males and females flew readily after release of at least some of the myconial
fluid but, while females flew only a short distance before landing again, males would fly much further.
At 20 C both Chaoborus trivittatus and Chaoborus cooki males completed rotation of their genitalia
within four hours.
Males of both species possess at least some mature sperm upon emergence. Differences in ovarial
development between teneral females of Chaoborus trivittatus and Chaoborus cooki are described
elsewhere (p. 179).
Chaoboridae are commonly designated as non-biting mosquitoes. However, Cook (1956:6) states that
“Chaoborinae ( = Chaoboridae) have all structures necessary for piercing and for feeding on fluids”. Felt
(1904), Pucat (1965) and Theobald (1901a: 62) suggested that adult Chaoboridae feed on plants but
these claims were unsupported. Martini (1931) also recognized adults to be capable of feeding but gave
no actual observations.
Both mammalian and avian blood have been found in the guts of female Corethrella brakeleyi
(Coquillett) and Corethrella wirthi Stone (Williams and Edman, 1968). Miyagi (1974) reported the
presence of a blood meal of unknown origin in the gut of a female Corethrella japonica Miyagi. Its
mandibles and hypopharynx are serrated, as are those of fem‘ale Corethrella brakeleyi, Corethrella
wirthi and Corethrella appendiculata Grabham (Miyagi, 1975). McKeever (1977) has recorded
Corethrella females feeding on Hyla tree frogs and was able to attract adult Corethrella brakeleyi to a
tape recording of frog calls.
Jenkins and Knight (1950) reported an adult male Eucorethra underwoodi probing or feeding at
pistillate flowers of willow.
Adult Mochlonyx cinctipes (Coquillett) did not feed on proffered prunes, raisins, honey-water, or
wild flowers, although imbibition of water was observed in the field (O’Conner, 1959). Mochlonyx
velutinus (Ruthe) took diluted honey (James, 1957).
Some workers have reported on adult life span of chaoborids (summarized by Borkent 1978a: 162)
but most, unfortunately, do not report their experimental conditions.
While collecting adult Chaoborus trivittatus 2.4 km west of Edmonton, Alberta, on May 13, 1975, I
observed that male and female adults often alighted on the pond surface and applied their mouthparts to
the water for about 30 seconds, rested for a short period, and then flew away. Air temperature was 17 C.
In addition, two males alighted on the ground and walked with their front legs moving alternately in
sweeping motions similar to that described by Downes (1974) for adult chironomids searching for
honeydew. At this time of year no aphids were present to provide this source of nutrition. Adults
commonly exhibit this searching behavior under laboratory conditions if kept in sealed mason jars for at
least 24 hours without water.
These observations led me to suspect that adults drink water and that this may be an important factor
in prolonging adult life span, consequently influencing adult dispersal capabilities.
I carried out a feeding experiment to see if adults drink water and to determine if water uptake
affects longevity.
Blue excreta from adults in jars supplied with dyed water showed that both males and females imbibe
water. Longevity results are shown in Figures 35A,B. These results were compared statistically using a
Duncan’s Multiple Range test (p<0.05) and it was found that: 1) there is no significant difference in
longevity between males or females given either clear or dyed water; 2) males which could not drink
lived for a significantly shorter period than males which could drink; 3) females which could not drink
lived for a significantly shorter period than females with dyed water; and 4) although there was no
Systematics and Bionomics of Schadonophasma
189
significant difference between females without access to water and those with clear water, this is
explained by the low numbers in each treatment. Comparison of females without access to water to
combined results of females with dyed or undyed water available, using a T-test, revealed them to be
significantly different at the 98% level.
These results show that Chaoborus trivittatus adults can imbibe water and that this significantly
increases their life span. Longevity of adults of most other species may have been seriously
underestimated.
During the springs of 1974-1977 I made daytime observations of behavior of Chaoborus trivittatus
adults 2.4 km west of Edmonton, and 16 km west of Jasper, Alberta. Males were more often seen
throughout the emergence periods. They formed columnar swarms (n = 8) of 1-132 individuals at the
periphery of ponds at heights of about 0.3-1. 8 m. Females were sedentary on surrounding vegetation,
particularly on tree trunks. When disturbed, they flew and were easily seen. However, on May 13, 1975,
after 1.5 hours of collecting with an aspirator and aerial net from tree trunks and low vegetation at 2.4
km west of Edmonton, only 32 females were procured. Concurrently, males in swarms were extremely
numerous. Similarly, 25 females and 218 males were collected with an aspirator from tree trunks at this
locality on May 7, 1977. Female pupae and exuviae, collected May 13, 1975, composed 59.6% (n = 460)
and 40.7% (n = 329) of a sample respectively. The possibility that the over-abundance of males was due
to their earlier emergence, as might be suggested from laboratory rearings, is therefore unlikely.
When females made one of their occasional flights through a male swarm, they were invariably
seized front to front by a male. The pair interlocked genitalia and fell immediately to the ground or
water surface, where they assumed an end to end position. Matings lasted 51-64 seconds (x = 58.7,
n = 5). Nine other matings observed required a similar period of time but were not measured exactly.
These data suggest the following pattern. Most adult males emerge earlier than females and form
large swarms at the periphery of the ponds from which they emerged. Females, after emergence and an
unknown period of time, fly through this ‘barrier’ of males and are mated. Because the sex ratio of
emerging adults is nearly 1:1 (Table 51), the far greater proportion of male adults at pond peripheries
suggests that females are the main dispersing agent of this species. Well developed ovaries at emergence
are probably correlated to the stable habitat which this species inhabits and suggests that Chaoborus
trivittatus female adults cannot disperse as far with their fully developed eggs as can Chaoborus cooki
female adults (see below).
If a sex ratio of nearly 1:1, and observations of concentrated male swarms at the periphery of water
bodies are alone indicative of this type of adult behavior (i.e., females mated at the pond or lake
periphery and subsequently dispersing), observations of adults of the species Chaoborus sensu stricto
suggest that these exhibit similar behavior. Bradshaw (1973) reported males to compose 57% of
populations of Chaoborus americanus. I have observed large male swarms of this species at the
periphery of numerous ponds near Edmonton, Alberta. Males make up 43-45% of Chaoborus flavicans
populations (Parma, 1971b) and swarms have been observed at a number of localities (Berg, 1937).
Parma (1971b) noted the concentration of male Chaoborus flavicans at the periphery of the water
bodies he studied. Parma (1971b) reported males to comprise 39-60% of Chaoborus crystallinus
populations and both he and Nielsen and Greve (1950) have observed male swarms of this species.
Neither sex ratio or swarming behavior have been reported for Chaoborus obscuripes (van der Wulp).
These data suggest that at least three of the four species of Chaoborus sensu stricto share a common
type of adult reproductive behavior and dispersal with Chaoborus trivittatus. The significance of this is
discussed in the section on zoogeography and speciation.
I have little data on adult behavior of Chaoborus cooki in the field. In J 975 and 1976, at the pond 1.4
km west of George Lake, and in 1976, at the pond 1.6 km south of Jasper, Alberta, emergence of adult
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Borkent
Chaoborus cooki was observed only during the night. Although both New Jersey and CDC light traps
were present at the edges of the ponds, adults that emerged were not trapped. Upon emergence, adults
flew upward out of the field of vision. On the day after each of the nights I observed emergence, I swept,
with an aerial net, all surrounding vegetation for a least 100 m in all directions of both ponds, but with
no captures. I also visually examined the immediate area and climbed the trees surrounding the ponds
but neither collected nor saw any adults.
Timing of laboratory matings of Chaoborus trivittatus adults (45-65 seconds, n = 6) approximated
those observed in the field (see above). This would suggest that timings of matings of Chaoborus cooki
adults in the laboratory (91-122 seconds, n = 8) also approximates those under field conditions.
These data, a sex ratio of males to females of about 3-5 to 1 (see p. 193) and retarded ovarial
development, indicate that Chaoborus cooki adult behavior differs substantially from that of Chaoborus
trivittatus. This suggests that Chaoborus cooki adults do not remain near the pond from which they
emerge and both males and females disperse. This behavior would ensure the survival of at least part of
the total population. Temporary ponds which became dry before adult emergence would undoubtedly
kill the immatures present.
Female adults probably exhibit underdeveloped ovaries on emergence so that fat body can be used as
energy for flight should a suitable habitat not be located and would guarantee at least the production of
a few eggs once a suitable habitat is found (Roff, 1977).
Lack of records of adults in the area from which they just emerged suggests that the adults mate
after a period of dispersal. Greater numbers of males than females would be more conducive to
successful mating after dispersal and might explain the inequality in numbers of males and females.
Differences in dispersal capabilities and behavior of Chaoborus trivittatus and Chaoborus cooki
concurs with the suggestion by Southwood (1962) that, generally, species found in unstable habitats
show more intensive dispersal behavior than do those in more stable habitats. For species of Chaoborus,
degree of ovarial development in teneral adults (Fedorenko, 1975c; Parma, 1971b) and number of eggs
per female (Fedorenko, 1975c; McGowan, 1974; Parma, 1971b; Sikorowa, 1973) is also correlated to
stability of habitat.
Egg rafts of Chaoborus trivittatus are laid in a similar way to those of Chaoborus flavicans and
Chaoborus crystallinus (Parma, 1971b: 28). I have observed oviposition in the field during daylight
hours, but did not measure the rate of egg laying.
I have no observations of oviposition by females of Chaoborus cooki.
Eggs. - Egg rafts of Chaoborus trivittatus, like those of species of Chaoborus sensu stricto (McCloy,
1950 (as Chaoborus crystallinus = Chaoborus americanus)\ Parma, 1971b; Sikorowa, 1973) are in the
form of floating discs with the eggs arranged in a spiral. My observations of egg rafts and egg
development are similar to those reported by Parma (1971b: 30). I found many egg rafts stuck to
floating debris at the water surface. Eggs in the laboratory turn dark brown after one or two days. The
pharate first instar larvae were positioned as shown by Parma (1971b: fig. 12) for Chaoborus flavicans.
As Parma (1971b: 29) has pointed out, oviposition can be disrupted; thus I did not consider number of
eggs per raft to be a significant measurement. I have found an egg mass, for example, of Chaoborus
trivittatus composed of nine eggs. This clearly does not reflect the number of eggs per female (p. 179)
and therefore cannot be used to interpret interspecific or intraspecific variation.
Eggs of Chaoborus cooki were obtained only from laboratory mated females. These were laid in a
semispherical mass on walls of vials, which were used to retain adults. The eggs were held together by
minute amounts of gelatinous matrix. The exochorion of each individual was also sticky. Endochorions of
fertilized eggs turned dark brown within two to three days. The endochorion of non-fertilized eggs
turned slightly brown in places.
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191
1 attempted to collect eggs of Chaoborus cooki from the site 1.4 km west of George Lake, after the
pond had become dry on October 30, 1976, when the surface soil was frozen to a depth of about one to
three centimeters. Soil samples were taken from each of 50 by 50 cm grids on the entire area covered by
the high water mark of the temporary pond. When Typha sp. stems were present in a grid a sample was
taken of these, cut off below the soil surface. In addition, samples of brush that immediately surrounded
the pond were taken. In the laboratory soil samples were sieved and vegetation scanned under the
binocular microscope. Nine soil samples were treated with floatation techniques using NaCl. I was
unable to retreive any eggs or egg shells.
I have observed hatching of eggs only for Chaoborus trivittatus from 2.4 km west of Edmonton, and
16 km west of Jasper, Alberta. The events of hatching were similar to those described by Parma (1971b:
32). Eggs of Chaoborus trivittatus hatch within three days, usually all of one egg mass hatching within a
few hours. A few eggs were sieved from a bottom sample taken from a pond 16 km west of Jasper,
Alberta, on June 18, 1976, after the adult emergence period was completed. All eggs were white,
suggesting that they were not fertilized, and by June 24, 1976, all had deteriorated in the laboratory.
Outer eggs on the egg raft hatch first. Egg shells of Chaoborus trivittatus split longitudinally and the
posterior part of each larva is freed. A few were caught with their head capsule stuck in the egg shell but
most, with violent struggling, freed themselves. Unlike the egg shells of Chaoborus crystallinus and
Chaoborus flavicans (Parma, 1971b: fig. 13), and Chaoborus americanus (pers. obs.), in which the egg
shell on each side of the longitudinal slit folds inwards, the sides of the egg shell of Chaoborus trivittatus
fold outward (Fig. 9D). This unique character may have evolved to compensate for the large size of
Chaoborus trivittatus first instar larvae and therefore allow for fewer numbers of first instar larvae
being stuck in the egg shell.
All eggs of Chaoborus trivittatus collapsed under atmospheric conditions and therefore are not
resistant to desiccation.
Eggs of Chaoborus cooki do not hatch within a few days, as do those of other Chaoborus species
(Deonier, 1943; Herms, 1937; McGowan, 1974: 498; Sikorowa, 1973: 76). Endochorions of these eggs,
white when laid, become extremely dark brown within 2-3 days. Larvae develop to at least a stage where
head capsule and air sacs are visible, but do not hatch at 20 C. After three months at this temperature,
and an L:D photoperiod of 18:6, some eggs were treated with either three successive floodings, three
days apart, of deoxygenated water, or one treatment with xylene vapor, but they failed to hatch. Other
eggs were treated to 20 C temperatures and an L:D photoperiod of 18:6 for two months, 2 C and total
darkness for four months and then returned to former conditions but with no evidence of hatching. A few
eggs did hatch after being kept at 20 C and an L:D photoperiod of 18:6 for two months and then total
darkness at 2 C for ten months. Study of the egg shell showed that the exochorion was gone (as it was in
some unhatched eggs), and the chorion had split to release the larva in a way different (Fig. 9E) to that
described for other species.
Eggs of Chaoborus cooki were treated to one month of atmospheric conditions without collapse. The
thick exochorion is apparently a protective layer against desiccation. The highly sculptured exochorion
probably functions as a plastron under alternately dry and wet conditions (Hinton, 1969; Hinton and
Service, 1969).
Larvae. - I observed horizontal clumping of second, third, and fourth instar larvae in ponds which
were not entirely shaded from direct sunlight. Clumping was related to presence of shade provided either
by surrounding vegetation, topographical features, or emergent aquatic vegetation. This is clearly a
factor which influences sampling results of Chaoborus trivittatus and Chaoborus cooki larvae from
shallow water bodies. For example, a pond measuring about 5 by 14 m, 7.6 km northwest of Mountain
Park, Alberta, was sampled successively with a dip net along the north side and a portion of the south
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192
Borkent
side which were exposed to direct sunlight, without collecting a single larva. A prominent boulder on the
south shore provided shade in part of the pond and in this region I collected 109 fourth instar larvae of
Chaoborus trivittatus in a single sweep of the net.
Similar response to shade is probably exhibited by Chaoborus pallidus (Akehurst, 1922: 352;
Edwards, 1920; Sikorowa, 1964)
This factor is probably one which influences the restriction of Schadonophasma species to woodland
conditions. In addition, it is possible that this resonse to shading is a modification of the response to light
which is important in vertical migration of some Chaoborus species (Chaston, 1969; LaRow, 1968,
1969).
I observed grooming behavior of fourth instar larvae of Chaoborus trivittatus and Chaoborus cooki.
Larvae often lashed at the posterior portions of their bodies with their antennae and mandibles.
Occasionally larvae place the posterior portion of the abdomen in their oral cavity, the anal fan
completely in the mouth. Even the slightest disturbance resulted in return to a normal position. However,
on one occasion I observed a larva of Chaoborus trivittatus keep its tail end in its mouth for over 2.5
hours. This behavior is undoubtedly an important adaptation for removal of epibionts.
I did not observe larvae of Chaoborus trivittatus or Chaoborus cooki bury themselves in the bottom
substrate in aquaria, although, when disturbed, they would dart toward the bottom and often cling with
mouthparts to bottom debris.
Although differences in prey type were evident between Chaoborus trivittatus and Chaoborus cooki,
I did not quantify this information because of biases produced by preservation techniques (Swift and
Fedorenko, 1973). Although Fedorenko (1975a, 1975b) and Swift and Fedorenko (1975) have recorded
the diet of Chaoborus trivittatus fourth instar larvae, from an oligotrophic lake in southwestern British
Columbia, as composed of copepods and cladocerans, it is not known to what extend another habitat,
such as a pond, affects prey type. For example, I observed Chaoborus trivittatus fourth instar larvae
capturing chironomid larvae. I did not find ostracod remains in the gut of Chaoborus trivittatus fourth
instar larvae and, in the laboratory they lashed out at, but rejected ostracods as food. Contrary to this I
found numerous Chaoborus cooki larvae with ostracods in their crops. Chironomid and, in Chaoborus
cooki larvae from some ponds, culicid larvae were also common. To a certain degree, differences of prey
type must reflect habitat differences, but these observations of ostracod ingestion suggest species
differences in ability to ingest at least this prey type.
Male larvae of both species pupate earlier than do female larvae (Table 51, 56). Chaoborus cooki
larvae have a more extended pupation period than do those of Chaoborus trivittatus under laboratory
conditions.
Pupae. - Pupation of individuals of Chaoborus species has not previously been observed. The
following is derived from observations of a single pupation for each of Chaoborus trivittatus and
Chaoborus cooki. Under laboratory conditions pupation required about two or three seconds. The fourth
instar larva suddenly contracted along its entire body length in a quivering motion. The larva turned into
a vertical position, the pupal horns popped through the thoracic suture, followed by the thorax and head
of the pupa. The larval exuviae slipped down the length of the pupal abdomen and was thrown off the tip
by a forward flick of the abdomen.
Parma’s (1971b: 38) description of macroscopic color changes during pupal development also apply
to pupae of Chaoborus trivittatus and Chaoborus cooki.
Under laboratory conditions, the pupal period of Chaoborus trivittatus is three (n=l) to four
(n = 85) days for males and four (n = 22) to five (n = 4) days for females. The pupal period of male
Chaoborus cooki lasted two to six days (n= 1,2,6,28,21 respectively), and of females three to seven days
(n= 1,13,13,1 1,4 respectively).
Systematics and Bionomics of Schadonophasma
193
Table 56. Numbers of pupae resulting from 42 individually reared fourth instar larvae of Chaoborus
cooki collected 32 km west of Edson, Alberta. Pupation recorded only on dates presented.
Differences in development of ovaries between pupae of Chaoborus trivittatus, Chaoborus cooki and
Chaoborus nyblaei are described elsewhere (p. 179).
To compare the sex ratio of Chaoborus trivittatus and Chaoborus cooki I collected fourth instar
larvae before any indication of pupation in the field, from 2.4 km west of Edmonton and 32 km west of
Edson, Alberta (not the same locality from which the life cycle of Chaoborus cooki was studied),
respectively. Individual rearings of 44 larvae of Chaoborus trivittatus produced 23 male and 21 female
pupae and 42 Chaoborus cooki larvae produced 33 male and nine female pupae. The use of a compound
character index to sex fourth instar larvae of Chaoborus cooki (p. 161), from 1.6 km south of Jasper,
indicated a sex ratio of 52 males to 10 females.
MATING EXPERIMENTS
Observations of forced matings in the laboratory between members of two populations give
indications of their reproductive computability and, therefore, their conspecificity. I tested my
interpretation of morphological, bionomic and temporal discontinuities as indicative of the presence of
two species of Schadonophasma in North America by attempting to cross individuals of Chaoborus
trivittatus and Chaoborus cooki.
Results of crossing (Table 57) show several important features. Crossings of male Chaoborus
trivittatus and female Chaoborus cooki, as compared to intraspecific crossings, showed reduced number
of eggs, number of fertilized eggs, and reduced fertility. Four of the six crosses resulted in no eggs being
laid. Reduced fertility and low number of eggs from some of the intraspecific crosses must be the result
of experimental conditions.
Crosses between male Chaoborus cooki and female Chaoborus trivittatus resulted in no eggs. It was
apparent during the crossings that male Chaoborus cooki had difficulty in coupling with the female
genitalia. In an additional six attempted couplings the penis valves of male Chaoborus cooki could not
insert into the female genitalia of Chaoborus trivittatus and copulation did not take place. This may be
due to the more bulbous nature of the head of penis valves of Chaoborus cooki which may not fit into the
proper receptacle of the female genitalia.
These observations indicate that reproductive isolation between Chaoborus trivittatus and
Chaoborus cooki is additionally maintained by mechanical and genetic incompatibilities.
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Borkent
Table 57. Results of intraspecific and interspecific crossings of adult Chaoborus trivittatus and
Chaoborus cooki.
PHYTOGENY AND ZOOGEOGRAPHY
Phylogenetic and zoogeographic considerations are’ logical conclusions to systematic studies. In this
section I give my working principles for such considerations, evidence for the recognition of three species
of Schadonophasma, and hypothesize their evolutionary history. Because the evolution of species of
Schadonophasma can be best understood in the context of ex-group comparisons I also consider the
evolution of the genus Chaoborus.
In recent years there has been much debate concerning the principles and methods of cladistic
analysis as used to infer phyletic relationships (e.g. Ashlock, 1974; Brundin, 1972; Darlington, 1970;
Griffiths, 1972, 1974; Hull, 1970; Mayr, 1974; Schlee, 1975; Sneath and Sokal, 1973; Sokal, 1975; and
papers cited in these). Although adherance to methods themselves can blind investigators to more
reasonable alternative interpretations (Darlington, 1970), Hennigian principles (see Kavanaugh, 1972)
have broad applicability in the determination of evolutionary relationships. I have, therefore, used them
to infer the evolutionary history of species of the genus Chaoborus.
One potential source of error in the application of cladistic analysis concerns the weighting of
characters on a superficial basis. Hecht and Edwards (1977) and Szalay (1977) have discussed the
importance of detailed investigation of individual characters used in phylogenetic analysis. Although this
may not be possible in many studies, this investigation provided a basis from which to better interpret
the evolutionary significance of most characters used and therefore improved the use of cladistic
techniques. These data allowed a more probable interpretation of the evolution of some Chaoborus
species and, in particular, of the species of Schadonophasma.
Systematics and Bionomics of Schadonophasma
195
Most systematic studies, some because of a lack of bionomic information, only consider allopatric
speciation as the means of evolution of lineages. In a recent study Tauber and Tauber (1977) have
proposed a genetic model for sympatric speciation of more general applicability than that proposed by
Bush (1975a, 1975b). However, most studies suggesting interpretations of sympatric speciation lack
study of the phyletic relationships between the organisms concerned. My interpretation of possible
speciation events in the history of Schadonophasma species provides an example of how phyletic
relationships and zoogeographical considerations may give supportive evidence for inferring an allopatric
or sympatric speciation model.
Darlington (1970: 3) has criticized assumptions of dichotomous speciation and described several
situations in which branching was possibly polychotomous. I have not found evidence of trichotomous
speciation of the three species of Schadonophasma. The presence of synapomorphies for two of the
species supports only dichotomous speciation in the history of Schadonophasma species.
Evidence and Recognition of three species of Schadonophasma
The basic taxonomic unit used for inference of phylogenies is the species or species group. A
phylogeny therefore depends on correct interpretation of the diversity it attempts to explain. The
following is a discussion of evidence for the recognition of three species of Schadonophasma.
Morphological, bionomic, developmental and behavioral incongruities, as well as differences in
geographic variation and distribution among groups of populations, are logical criteria for recognition of
species if the biological species concept is accepted as a testable hypothesis. Only experimental tests of
reproductive incompatibilities, however, might provide direct evidence of reproductive isolation. It is the
concordance of such indirect and direct evidence of reproductive isolation that allows systematists to
interpret the former as the result of such reproductive isolation. The results of this study, as do many
others, confirm the validity of this assumption.
From the results of this study I consider discontinuities in the following features to support my
conclusion that there are two species of Schadonophasma in North America recognizable as Chaoborus
trivittatus and Chaoborus cooki:
1 . Structure of eggs, all stages of larvae, pupae and adults
2. Pattern of development of LB/AL from first through fourth instar larvae.
3. Characters of fourth instar larvae which are sex-associated in one species but not the
other.
4. Correlations of characters of fourth instar larvae.
5. Patterns of geographical variation of characters of fourth instar larvae in Alberta.
6. Rates of development of eggs, larvae and ovaries of adults.
7. Life cycle and types of habitat commonly occupied.
8. Larval prey type.
9. Adult behavior.
10. Reproductive compatibilities, when adults from different populations were crossed in the
laboratory.
1 1 . Lack of sympatry in western and eastern coastal regions of the United States.
I do not have nearly the same amount of evidence to justify recognition of the third species,
Chaoborus nyblaei, in Fennoscandia. Because of similarity of compared semaphoronts, I assumed that
all specimens from Fennoscandia were conspecific. Adult material collected 2-3 km SW of Nuorgam,
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Borkent
Finland, on July 24, 1960, was probably conspecific with larval and pupal material collected there.
Most characters of the immatures and color of adults of Chaoborus nyblaei are similar to those of
Chaoborus cooki. However, shape of the adult male penis valve and prelabral appendage of fourth instar
larvae are most similar to those of Chaoborus trivittatus. This combination of features suggests the
presence of a third species, and this is further supported by extremes of variation of certain characters of
the fourth instar larvae of Chaoborus nyblaei (number of mandibular fan bristles and anal fan setae).
Because of difficulties in interpreting morphologically similar, allopatric populations separated by a
geographical area from which specimens are not known (Mayr, 1969), it is impossible to be certain that
Chaoborus nyblaei is a separate species. Therefore it might be conspecific with either Chaoborus
trivittatus or Chaoborus cooki. Overall morphological similarity would indicate, if this were so, that
Chaoborus nyblaei and Chaoborus cooki are conspecific. However, patterns of geographical variation of
Chaoborus cooki would not suggest that the higher mean values of number of anal fan setae, and lower
mean value of number of anal fan setae of the larvae of Schadonophasma in Fennoscandia are the result
of variation of those characters of Chaoborus cooki. Nevertheless, the possibilities of character
displacement producing such features as exhibited by the population in Fennoscandia cannot be
presently negated. Further study of Chaoborus nyblaei populations to complement the data given in this
study for Chaoborus trivittatus and Chaoborus cooki, further investigation of the range of Chaoborus
nyblaei and detailed analysis of geographical variation of Chaob'orus nyblaei and Chaoborus cooki once
further samples become available, would give further clues to questions of conspecificity. Not
withstanding, it is probable that the Bering Strait area is devoid of Schadonophasma populations and
presents at least a recent barrier between North American, and should they exist, eastern Asian
populations of Schadonophasma.
Phylogeny of Chaoborus
Saether (1970), using cladistic methods, proposed a phylogeny of Chaoborus species and chaoborid
genera but dealt only with those of the Holarctic region. He used 29 characters to determine
relationships among subgenera of Chaoborus. However, his interpretation of the polarity of many of
these characters is questionable.
A number of characters are used by Saether (1970), for which he gives no criteria for determining
the apomorphic and plesiomorphic ends of the morphoclines and for which there are none apparent
(Maslin, 1952). For example, the character HW/WBE is considered plesiomorphic if 2. 5-4. 2 and
apomorphic if 1.5-1. 8. However, the closest sister group to Chaoborus for which these data are available
is Mochlonyx which, if all species are considered, has a total range of 1.8-2. 4 (Cook, 1956). If this
feature represents the plesiomorphic condition of Chaoborus, both ends of the morphocline recognized
by Saether would be apomorphic. However, HB/WBE ratio of male adult Eucorethra underwoodi is
8. 3-8. 5 (Cook, 1956) and this may also reflect plesiomorphy. Until further ex-group comparison has
been made, phylogenetic interpretation of this character is conjectural.
It is also not clear how Saether interpreted the polarity of morphoclines of values of Y/R3 and Y/X
of adults (males?), width/length of abdominal segment VII of pupae and AS/AL of larvae, when these
data were available only for some species of Chaoborus.
It is well known that character states which involve loss of a feature are particularly suspect of
convergent or parallel evolution (e.g. it could be as likely for species to independently lose features as for
a single ancestor of species to do so). Interpretation of the following character states as synapomorphic
provides at best an uncertain basis for reconstructing a phylogeny: loss of parascutellar setae and
reduction of number of mesepimeral and pronotal seta of adults; loss of color of the pupal paddle
membrane; reduction of the relative length of the median rib of the pupal paddle; and reduction of the
Systematics and Bionomics of Schadonophasma
197
larval tentorium.
Shape of the apex of the male adult gonostylus and loss of a pair of larval labral setae are
autapomorphies for Chaoborus pallidus, and cannot be used to determine relationships between species
or groups of species.
Saether (1970: 47, Trend 46) considered the placement of the simple seta toward the apex of the
median rib of the pupal paddle to be apomorphic within Chaoborus. This condition, however, most
closely approximates the condition featured by Mochlonyx and Eucorethra pupae suggesting that it is
more likely to be the plesiomorphic character state.
Following Saether’s (1970) publication, larvae of some African species have been well described
(Green and Young, 1976; McGowan, 1972, 1976). In addition, Saether (1976) has redescribed adults,
pupae and fourth instar larvae of Chaoborus brasiliensis (Theobald) and Chaoborus magnificus Lane
from Venezuela. Using these descriptions and my own observations, I have constructed a phylogeny of
those species of Chaoborus for which sufficient information is available. The characters used and
interpretation of their plesiomorphic and apomorphic states are given in Table 58. All character states
were judged plesiomorphic or apomorphic on the basis of ex-group comparisons (especially Mochlonyx
and Eucorethra) with the exception of prelabral appendage shape. The interpretation of several
characters requires special comment.
Two characters used are interpreted as apomorphic in their reduced state: relative length of the
median rib of the pupal paddle and, relative degree of sclerotization and size of the larval tentorium.
These were used because of a shortage of characters in this analysis. As proposed here, reduction of the
larval tentorium has occurred twice.
The prelabral appendage of Chaoborus larvae is undoubtedly a seta (Balvay, 1977c). Although it is
not known which seta of other chaoborids is homologous to this prelabral appendage, it is reasonable to
assume that ontogenetic development of the prelabral appendage from a simple seta in the first two
instars of all Chaoborus species to a laterally flattened appendage in the last two instars of some species,
corresponds to the evolution of these setae (Akehurst, 1922: 352).
The only clues for determining the polarity of AS/AL of Chaoborus larvae are provided by the larvae
of Mochlonyx velutinus and Eucorethra underwoodi. Mochlonyx velutinus fourth instar larvae possess
an antennal seta near to the base of the antenna but none were measured exactly. The AS/AL of three
fourth instar larvae of Eucorethra underwoodi was 0.51-0.59 (mean = 0.558). In the construction of the
phylogeny I have used this feature only to interpret the extreme values of AS/AL of Schadonophasma
as apomorphic. Change of this ratio, from a low value in first instar larvae to a high ratio in fourth
instars (Fig. 22A), would support such an interpretation.
The polarity of two characters, coloration of adult legs and state of the larval dorsal process, has
apparently been misinterpreted by Saether (1970). He suggested that ringed legs of adults is a
plesiomorphic state. This is probably only partially true. Although bases and apices of leg segments and
articles are dark in a number of ex-groups, and is therefore probably plesiomorphic, spotted femur and
tibia of legs of some African spcies of Chaoborus (Verbeke, 1958), and some species of the subgenus
Sayomyia (e.g. Chaoborus punctipennis, Chaoborus astictopus, (per. obs.) Chaoborus brasiliensis), is
probably apomorphic.
Saether (1970: 50) also considered the two segmented dorsal process of the fourth instar larvae of
some Chaoborus species to be the plesiomorphic condition for the genus and implies (Trend 67) that the
dorsal process is actually the evolutionary vestige of the respiratory siphon. However, as pointed out by
Cook (1956: 14) ‘the annulation on which this “siphon vestige” occurs has a musculature which
indicates that it is a segment. This then is segment 9, and the siphon occurs on segment 8. Thus, the term
“siphon vestige” seems to be inaccurate’. In addition, Parma (1971a) followed the shape of the dorsal
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Borkent
Table 58. Characters and character states within the genus Chaoborus.
process from instar to instar of Chaoborus flavicans and found the pointed dorsal process to be present
only in the later instars. If this feature represents the siphon vestige, one would expect it also to be
present in earlier instars. I further tested this hypothesis by examining the position of the tracheal trunks
of fourth instar larvae of Chaoborus cooki. I found that thawing previously frozen larvae resulted in the
trachea filling with gas, allowing them to be clearly observed. The tracheal trunks showed a notable
dorsal bend in the eighth abdominal segment (Fig. 8D) similar to the dorsal bend of the tracheal trunks
of Mochlonyx larvae into the siphon (Montchadsky, 1953: fig. 1).
These observations indicate that the small dorsal article of the ninth abdominal segment of larvae of
some Chaoborus species is in fact a newly evolved feature and not the vestige of the respiratory siphon.
Absence of such an article is therefore the plesiomorphic condition. The pointed dorsal process exhibited
by Chaoborus flavicans and Chaoborus crystallinus fourth instar larvae is also probably an
independently derived characteristic.
The most probable phylogeny of species of Chaoborus, using the characters in Table 58, is presented
in Figure 36. 1 accept Saether’s (1970) conclusion concerning the phyletic relationships between the four
species of Chaoborus sensu stricto. This phylogeny differs from that hypothesized by Saether (1970,
Systematics and Bionomics of Schadonophasma
199
1976) primarily in considering Chaoborus albatus as possibly the sister species of Chaoborus
magnificus and these species plus Chaoborus brasiliensis, Chaoborus punctipennis, and Chaoborus
astictopus (hereafter designated as ‘ambpa’) the sister group of Schadonophasma plus Chaoborus sensu
stricto. The placement of Chaoborus pallidipes is uncertain. This species could equally well be the sister
group of Schadonophasma plus Chaoborus sensu stricto, or of the species group ‘ambpa’.
The grouping of Chaoborus anomalus, Chaoborus ceratopogones, and Chaoborus pallidus is
tentative. These are recognized as monophyletic on the basis of leg coloration which, as here
hypothesized, shows convergence with the species group Chaoborus brasiliensis, Chaoborus
punctipennis, and Chaoborus astictopus. Any of Chaoborus anomalus, Chaoborus ceratopogones or
Chaoborus pallidus could be considered a sister group of the species group ‘ambpa’ if the following
events are assumed to have occurred: 1) the basal curvature of the larval antenna was secondarily lost in
these species, or the curvature evolved independently in species groups ‘ambpa’ and Schadonophasma
plus Chaoborus sensu stricto; and 2) Chaoborus albatus and Chaoborus magnificus secondarily lost the
derived coloration of adult legs. However, these events provide a less parsimonious explanation than does
the convergence of leg coloration as presented here. Similarly the development of a lobe on the inner face
of the gonocoxite of both Chaoborus anomalus and the species group Chaoborus brasiliensis,
Chaoborus punctipennis, and Chaoborus astictopus, is considered a result of convergent evolution.
A character which may be synapomorphous for Chaoborus anomalus, Chaoborus ceratopogones and
Chaoborus pallidus, except for which observations were not available for all other species, is the basally
swollen and highly curved postantennal filaments (Saether, 1970: fig. 6C). Verbeke (1958: 43) also
considered Chaoborus ceratopogones and Chaoborus pallidus to be closely related on the basis of
similarity of adults, pupae and larvae.
Although the broadening of the prelabral appendage may have evolved independently four times, I
here consider it to have occurred only once. The setose appearance of the prelabral appendages of
Chaoborus punctipennis and Chaoborus astictopus is therefore a derived state and is only superficially
similar to the primitive condition. This explains why the setaceous prelabral appendages of these two
species have spines directly only anteriorly and posteriorly.
Fossil evidence
The study of fossils can give important evidence for the correct interpretation of the evolutionary
history of a group of extant organisms. However, few systematists studying arthropods are fortunate to
have such fossils available to them.
A number of authors have described or mentioned Chaoborus subfossils (Alhonen and Haavisto,
1969; Deevy, 1942; Frey, 1955, 1962, 1976); Goulden, 1966a, 1966b; Hofmann, 1971, 1978; Marland,
1967; Stahl, 1959, 1969) or fossils (Borkent, 1978b; Edwards, 1923; von Heyden, 1862; Hope, 1847;
Meunier, 1904; Scudder, 1890 (questionable identification); Serres, 18^9: 268). Most of the fossil
material is in need of redescription. I examined the fossil pupa and disarticulated pupal parts described
by von Heyden (1862) as Culicites (= Chaoborus) tertiarius and have published (Borkent, 1978b), a
redescription of those fossils and of some larval fragments (mandibles and anal fans) also present on the
brown, paper-coal. The pupal paddles exhibit median ribs which extend to the edge of the paddle and are
curved apically. In addition the subordinate tooth at the base of the two larger teeth of some of the larval
mandible is very similar to that of Chaoborus flavicans. It is not certain that the larval fragments are
from the same species as the pupa and pupal parts. Nevertheless, mandible structure suggests that they
are from a population conspecific with Chaoborus flavicans {Chaoborus sensu stricto), or from a species
which is the sister species of Chaoborus flavicans. The apical curvature of the median rib of the pupal
paddles is a synapomorphous character which serves to group Schadonophasma and Chaoborus sensu
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200
Borkent
stricto. Although I cannot confidently place the fossil pupa or pupal parts with one of these lineages, it is
clear that if convergence has not taken place this species shares a common ancestry with, or within,
either of these two subgenera.
According to Magdefrau (1968), Edwards (1923) correctly determined the age of the fossils as
Upper Oligocene. This would indicate that speciation giving rise to Chaoborus flavicans, and its sister
species Chaoborus crystallinus, took place at least 25 million years B.P., and that the speciation event
which gave rise to the lineages represented by extant species of Schadonophasma and Chaoborus sensu
stricto took place before that time.
The finding of these fossils at Rott, Siebengebirge, West Germany is consistent with a hypothesis that
the species group of Schadonophasma plus Chaoborus sensu stricto evolved under temperate conditions
(Magdefrau, 1968).
Phylogeny of Schadonophasma
Saether (1970) proposed a phylogeny for the species of Schadonophasma based on six characters. I
have already discussed the difficulties of interpretation of four of these (number of mesepimeral and
pronotal setae; ratio of Y/X; WS/LS; shape of dorsal process). Saether’s phylogenetic interpretation of
variation of the penis valve shape seems to be correct for Schadonophasma. The interpretation of body
size variation is probably incorrect but is discussed below.
Although the phylogeny I propose for the species of Schadonophasma is identical to that of Saether
(1970), except for his consideration of Chaoborus brunskilli { = Chaoborus trivittatus) as the sister
species of Chaoborus trivittatus, my approach to the problem is substantially different. For the analysis
I have chosen seven characters (Table 59), some of which are discussed below.
Table 59. Characters and character states within the subgenus Schadonophasma.
Saether (1970: 43) in Trend 2, recognized the largest chaoborid, Eucorethra underwoodi to represent
the plesiomorphic condition and the smallest, the apomorphic condition. He therefore considers the large
size of Schadonophasma spp. and within the subgenus Chaoborus cooki and Chaoborus nyblaei, to be
plesiomorphic. However, his determination of polarity of this morphocline is almost certainly incorrect.
Systematics and Bionomics of Schadonophasma
201
Schadonophasma species are the largest of all described Chaoborus, and are larger than all species of
Cryophila Edwards, Mochlonyx, Promochlonyx Edwards, and Australomochlonyx Freeman; genera to
which Chaoborus is more closely related than to Eucorethra. It is almost certain therefore that the large
size of Schadonophasma members as a group, and those of Chaoborus cooki and Chaoborus nyblaei in
particular, represents an apomorphic condition. In this presentation I have used larval head capsule
length to represent the overall size of the species.
I have already discussed above the interpretation of AS/AL and prelabral appendage shape.
The high number of mandibular fan bristles of some members of Chaoborus trivittatus probably
represents the apomorphic end of a morphocline. Mochlonyx fourth instar larvae have six to seven
bristles in the mandibular fan (Cook, 1956; O’Conner, 1959), and no other Chaoborus species have
members exhibiting such large numbers of mandibular fan bristles.
Some fourth instar larvae of Chaoborus trivittatus and Chaoborus cooki and all those of Chaoborus
nyblaei have a larger number of anal fan setae than other Chaoborus species. Mochlonyx fourth instar
larvae have 28-30 anal fan setae, Eucorethra underwoodi has 32 (Cook, 1956). These ex-group
comparisons would suggest that only the high number of anal fan setae of Chaoborus cooki and
Chaoborus nyblaei are apomorphic. However, some evidence suggests that the evolution of changes in
number of anal fan setae is more complex. From observations of the swimming behavior of Chaoborus
larvae it is clear that the anal fan functions as a fulcrum against which the larvae can produce a forward
motion. It would seem reasonable therefore to suggest that if there are developmental restrictions on the
size of individual anal fan setae, the number of anal fan setae may be related to the size of the species
and that these two characters coevolved. Distribution of number of anal fan setae, relative to range of
head capsule length for species of Chaoborus for which these data are available (Fig. 37), would support
this hypothesis. These two characters would generally, therefore, not give independent evidence of
phyletic relationships. However, it is also apparent from Figure 37 that the relationship is not a constant
one. Although this may be due to inadequate sampling for some species, it is probable that other factors
are important in the evolution of number of anal fan setae. For example, even though there is no
significant difference in head capsule length of Chaoborus cooki and Chaoborus nyblaei fourth instar
larvae, number of anal fan setae does significantly differ. I have therefore interpreted the large number
of anal fan setae as autapomorphic for Chaoborus nyblaei.
The egg structure and type of egg mass of Chaoborus trivittatus is extremely similar to those of
species of Chaoborus sensu stricto. Because Chaoborus sensu stricto is the sister group of
Schadonophasma it is probable that the condition found in Chaoborus trivittatus represents the
plesiomorphic state within Schadonophasma. However, it is possible that the character state of
Chaoborus trivittatus represents a convergence with species of Chaoborus sensu stricto. A ‘layer of
jelly-like substance’ or ‘thin gelatinous sheath’ around individual eggs (= thick exochorion?) of some
species has been described (MacDonald, 1956; McGowan, 1976; Sikorowa, 1973) and, if these features
are homologous with those of Chaoborus cooki and Chaoborus nyblaei, they might be considered
plesiomorphic within Schadonophasma. However, the eggs of these other species of Chaoborus need to
be better studied before such inferences can be made.
The proposed phylogeny of the species of Schadonophasma is given in Figure 38.
Zoogeography and speciation events
Hennig (1966b) has shown that zoogeographic inferences are dependent upon accurate interpretation
.of the evolutionary relationships of organisms. Because it seems likely that detailed study of Chaoborus
species, both extant and fossil, will alter the phylogeny of Chaoborus as proposed here, I will not discuss
the zoogeography of the genus. However, the inferred sister group relationship between
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202
Borkent
Schadonophasma and Chaoborus sensu stricto is probably correct.
Because all extant species of Schadonophasma and Chaoborus sensu stricto are found only in north
temperate regions it would be reasonable to assume that the ancestor which gave rise to extant species of
Schadonophasma was present in the north temperate region. Distributions of species of Chaoborus
sensu stricto give no clues as to where specifically this ancestor may have arisen.
In the following discussion of the speciation event giving rise to Chaoborus cooki and Chaoborus
nyblaei, I have assumed that Chaoborus nyblaei has had or does have a continuous Palaearctic
distribution. However, future work may indicate that Chaoborus nyblaei has been, and is presently,
restricted to Fennoscandia. A more likely explanation of the speciation event giving rise to Chaoborus
cooki and Chaoborus nyblaei would suggest that dispersal of adults (or possibly desiccation resistent
eggs on the feet of migrating shorebirds) took place over the North Atlantic during a period of climatic
amelioration.
There are two equally probable zoogeographic interpretations of the history of Schadonophasma
species (Figure 39).
The first suggests that the ancestor of extant Schadonophasma species was Holarctic in distribution.
A split of its populations into Nearctic and Palaearctic elements resulted in the lineage giving rise to
Chaoborus trivittatus and to the ancestor of Chaoborus cooki and Chaoborus nyblaei (hereafter
referred to as Chaoborus cooki-nyblaei). The Palaearctic species then reinvaded North America and
subsequently was split into Palaearctic and Nearctic populations which gave rise to Chaoborus nyblaei
and Chaoborus cooki respectively. The model only requires allopatric speciation and dispersal to have
taken place. A land bridge between Asia and North America is known to have appeared and disappeared
a number of times during the Tertiary and Quaternary (Hopkins, 1967, 1972) which could have
provided, during certain periods, a barrier to contact between Asian and North American populations.
The second zoogeographic hypothesis suggests that the ancestor of extant Schadonophasma species
arose in North America and speciated there to give rise to the lineages resulting in Chaoborus trivittatus
and Chaoborus cooki-nyblaei. Although an allopatric speciation model might explain the speciation
event in North America resulting in the lineages which gave rise to Chaoborus trivittatus and
Chaoborus cooki-nyblaei, it seems most likely that the invasion into temporary waters by Chaoborus
cooki-nyblaei was not the result of geographical isolation. Although the lineage might have invaded
temporary ponds because of competition with other Chaoborus species as a geographic isolate, it seems
more likely that, especially considering the degree to which Chaoborus trivittatus and other Chaoborus
species can coexist (Stahl, 1966), this would not result in a speciation event. It is also difficult to imagine
two isolated geographical areas which contained populations of the ancestral species, one of which
contained only temporary waters conducive to the evolution of a temporary pond species.
The available data for Schadonophasma species allow for a hypothesis of sympatric speciation. The
following are hypothesized events, as indicated by this study, for Schadonophasma species and features
of species of Chaoborus sensu stricto.
The Nearctic species ancestral to extant Schadonophasma species inhabited permanent waters with
fourth instar larvae as the over-wintering stage. Upon emergence of adults in spring, the males formed
swarms on the periphery of the permanent water body and mated with females after these emerged.
These are features of Chaoborus trivittatus and those species of Chaoborus sensu stricto for which
information is available, and are therefore, probably plesiomorphic characteristics of Schadonophasma.
As previously shown for the two species of Chaoborus trivittatus and Chaoborus americanus, which
overwinter as fourth instar larvae in permanent ponds (p. 186), the dispersing females occasionally laid
eggs in temporary ponds. Eggs which hatched resulted- in larvae which could not survive overwintering in
this habitat. This type of egg was strongly selected against, as all larvae which hatched from them in
Systematics and Bionomics of Schadonophasma
203
temporary ponds would die. Tauber et al. (1977) have shown that mutation of a single allele can result
in the appearance of diapause in individuals of a population. If this feature is controlled by a single gene
in Schadonophasma species, crosses of male Chaoborus trivittatus and female Chaoborus cooki
resulting in diapausing eggs would suggest that the allele for diapausing eggs is dominant. I have
assumed this to be the situation in the following discussion. The primitive non-diapausing egg is
represented by autosomal recessive alleles aa. Mutation resulted in the production of a dominant Aa,
producing diapausing eggs. A female carrying these genes laid her eggs in a temporary pond, which then
diapaused in this habitat. Resistance to complete desiccation might not have evolved at this stage
because, at least for the temporary ponds I investigated, the bottoms were quite moist; probably, in some
ponds, moist enough to prevent desiccation of eggs. Larvae which hatched out in spring from these
overwintered eggs would mature to adulthood at a later time than would those in permanent ponds,
which emerge as adults early in spring. This might give at least a certain degree of temporal isolation.
However, as discussed above (p. 184), Chaoborus trivittatus might be multivoltine in certain regions and
this may have provided some temporal overlap between two populations of the ancestral species.
The adults which emerged from the temporary pond would, as did the parent population in
permanent ponds or lakes, have swarmed and mated at the pond periphery. This homogamic behavior
led to further segregation of populations and ensured an increase in numbers of individuals carrying this
gene. The resultant adults would carry the alleles AA or Aa. Any aa eggs laid in temporary ponds would
be eliminated as larvae. AA or Aa eggs subsequently laid in permanent ponds would survive and hatch in
the spring as first instars, but any other Chaoborus larvae, if present, would be fourth instar and these
first instars would probably be cannibalized. To hypothesize the removal of heterozygotes from the
temporary pond lineage (and therefore the continued production of aa), I suggest that the allele A
became associated with co-dominant or recessive features conducive to temporary pond existence and
therefore, as the temporary pond populations adapted to the temporary pond environment, heterozygotes
were selected against. Adult behavior and comparatively reduced ovarian development of extant
Chaoborus cooki, may be examples of such adaptations, but these also may have evolved after the
speciation had occurred.
Through temporal disjunction of mating periods, homogamic mating, complete selection against
overwintering larvae in temporary ponds, and partial selection against heterozygotes in temporary ponds
and overwintering eggs in permanent ponds, the two lineages giving rise to Chaoborus trivittatus and
Chaoborus cooki-nyblaei became genetically isolated. These suggested events are summarized in Figure
40.
Chaoborus cooki-nyblaei invaded the Palaearctic region, and subsequent barriers to reproduction, as
described for the first zoogeographic hypothesis, resulted in isolated populations which gave rise to
Chaoborus cooki and Chaoborus nyblaei.
The sympatric speciation model is dependent at least on the assumption that Chaoborus nyblaei does
in fact overwinter as an egg. Although this was inferred from available data, I have no direct evidence
that this is so. Study of the life cycle of Chaoborus nyblaei would therefore provide a test of this
hypothesis. If study of the populations I have described as ‘populations incertae sedis" show these to be
one or more species, this may provide additional evidence for testing this proposal of sympatric
speciation.
CONCLUDING REMARKS
On the basis of their work, taxonomists should indicate particularly fruitful areas for future research:
those concerned primarily with the group studied in particular, and those concerned with systematic
problems in general.
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Borkent
Because of the paucity of museum material of specimens of Schadonophasma, future endeavors
relating to the systematics of the subgenus must await further collecting. Some details concerning the
bionomics of species may bear further investigation, as indicated in the text. In particular, investigation
of the life cycle of Chaoborus nyblaei and the specific affinities of the ‘populations incertae sedis’ may
provide additional insights into the systematics of species of Schadonophasma. The phyletic
relationships of species of Chaoborus are yet poorly known. Fossil species and extant species from the
tropics, are particularly in need of study and the entire genus is worthy of a complete revision.
In general, this study suggests that investigation of the bionomics of species of some groups may
provide additional tests of the sympatric speciation model. Because of domination of evolutionary
concepts by the allopatric speciation model, the full implications and requirements of the sympatric
speciation model are poorly understood. These aspects of the model can only become clear when
additionaT information on bionomics of organisms which may have speciated sympatrically become
available.
As is well illustrated by the work of Bush (1975b) on tephritid fruit flies, a sympatric speciation
model can only be justifiably invoked from a solid base of bionomic data. Although allopatric speciation
has in the past been the favored hypothesis to explain differentiation of organisms, it is also evident that
only morphological and distributional data are available for most organisms. Therefore, until adequate
bionomic data are available for a large array of taxonomic groXips, it will be difficult to estimate the
frequency of occurrence of sympatric speciation. As discussed by Bush (1975a), sympatric speciation
occurs under special genetic and environmental conditions. It seems clear that, in certain groups
(mammals, birds, and carabid beetles), such conditions are rarely, if ever met. In the genus Chaoborus,
considering the similarity of the life cycle of most species, it appears that sympatric speciation occurs
with less frequency than does allopatric speciation. Nevertheless, some groups, like some species groups
of tephritid flies, appear to be more prone to speciate sympatrically. This pattern suggests, therefore,
that systematists should not apply the most common mode of speciation to all taxonomic groups under
investigation. Rather, each speciation model should be based on data available for that group.
An aspect not explicitly clear in previous discussions of sympatric speciation concerns the phyletic
relationships and zoogeography of the organisms concerned — essential considerations in any speciation
model. This study of Schadonophasma provides an example of how zoogeographic considerations may
provide additional evidence for elucidation of sympatric speciation.
In conclusion, it is the intensive study of only a few closely related species, as here reported for the
subgenus Schadonophasma, which can most fruitfully provide the basic information necessary to test
hypotheses of sympatric speciation.
ACKNOWLEDGEMENTS
Many individuals have contributed to this study and I extend my thanks to them all. In particular, I
express my sincere appreciation to my supervisor G.E. Ball, not only for cultivating my interests in
entomology and, particularly, in systematics, since our first encounter when I was yet in high school, but
also for his cheerful enthusiasm and thoughtful direction throughout the course of this study. He allowed
me to choose and made financially possible this study (including support from National Research
Council grant A- 1399).
I thank D.A. Craig for many enthusiastic discussions of some of the results of this work.
I also express my appreciation to my wife Annette, who shared numerous field trips and has helped
with many of the tasks involved in this study.
Systematics and Bionomics of Schadonophasma
205
To my fellow students with whom I have had numerous discussions about the principles and
foundations of systematics and about this work, I express my gratitude: in particular S. Ashe,
D. Lafontaine, R. Longair, and R. Roughley.
I thank G. Braybrook for technical assistance with the scanning electron microscope and J. Scott for
his helpful suggestions concerning drafting techniques and for completing some of the figures. I
acknowledge D.E. Griffiths for her masterful drawing of the frontispiece.
I thank all the individuals and institutions noted in the text for loan of material.
I express my appreciation to Parks Canada for collecting permits for Jasper and Banff National
Parks.
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Quaest. Ent., 1979, 15 (2)
216
Borkent
0
h
Figure 1. Place names in Alberta cited in text.
Systematics and Bionomics of Schadonophasma
217
2
Figure 2. A. Characters and measurements of head capsule of fourth instar larva: A. Head capsule length; B. Antennal length; C
Long antennal blade length; D. Short antennal blade length; E. Antennal seta; F. Postantennal filaments; G. Prelabral appendages;
H. Mandible; total length of mandibular fan bristles and labral brush setae is not shown; B. Characters of terminal abdominal
segments of fourth instar larva. C. Terminalia of male pupa. D. Terminalia of female pupa. All drawings from specimens of
Chaoborus cooki.
Quaest. Ent., 1979, 15 (2)
218
Borkent
Figure 3. Terms and measurements A. Terminal abdominal segments of pupa: A. Length of abdominal segment VII; B. Width
of abdominal segment VII. B. Male adult penis valve. C. Wing of adult (male).
Systematics and Bionomics of Schadonophasma
219
Figure 4. Scale line = 20Atm for Figures A-D and l^m for Figure E. A. Third tarsomere of midlcg of female adult. B.
Microsculplure of dorsal margin of head capsule of fourth instar larva. C. Exochorion of egg of Chaoborus trivittatus.
D. Exochorion of egg of Chaoborus cooki. E. Sculpturing of dorsum of exochorion of Chaoborus trivittatus.
Quaest. Ent., 1979, 15 (2)
220
Borkent
Figure 5. Posture of live pupae. A. Chaoborus cooki. B. Chaoborus trivittatus. Drawings from photographs of live pupae.
Systematics and Bionomics of Schadonophasma
221
Figure 6. Abdominal chaetotaxy of pupa of Schadonophasma. Setae numbered after Belkin et al. (i970). S; sensilla; sp: spiracular
scar.
Quaest. Ent., 1979, 15 (2)
Figure 7. Scale line = 0.1 mm. Antenna and dorsal outline of head capsule of larval instars of Schadonophasnia. Antennae: A.
First instar; B. Second instar; C. Third instar; D. Fourth instar. Dorsal outline of head capsule: E. First instar of Chaoborus cooki;
F. First instar of Chaoborus trivittatus\ G. Second instar; FI. Third instar; 1. Fourth instar. J. Egg burster of first instar of
Chaoborus cooki. K. Egg burster of first instar of Chaoborus trivittatus.
Systematics and Bionomics of Schadonophasma
223
SEGMENT VIII
DORSAL PROCESS .*
c
D
8
Figure 8. Except where indicated, scale line = 0.1 mm. A. Mandible of first instar larva. B. Mandible of fourth instar larva
(mandibular fan not shown). C. Dorsal process of fourth instar larva of Chaoborus nyblaei. D. Terminal abdominal segment of
fourth instar larva of Chaoborus cooki showing position of tracheae.
Quaest. Ent., 1979, 15 (2)
224
Borkent
Figure 9. Scale line = 0.1 mm. Egg structure. A. Chaoborus trivittatus. B. Chaoborus nyblaei. C. Chaoborus cooki. Egg shell. D.
Chaoborus trivittatus. E. Chaoborus cooki (fracture line indicated).
10
Figure 10. Scale line = 0.1 mm. Penis valves of adult male Chaoborus trivittatus. A-G. Intrapopulational variation of specimens
from 2.4 km west of Edmonton, Alberta. H-O. Interpopulational variation of specimens from: H. Banff, Alberta; 1. Hoodsport,
Washington; J. North Burgess Twp., Ontario; K. Whitehorse, Yukon Territory; L. Old Chelsea, Quebec; M. Lake 241, Kenora,
Ontario (holotype of Chaoborus brunskilli)-, N. Stanford, California; O. Maine (holotype of Chaoborus trivittatus).
Quaest. Ent., 1979, 15 (2)
226
Borkent
Figure 1 1. Scale line = 0.1 mm. Penis valves of adult male Chaoborus cooki. A-H. Intrapopulational variation of specimens from
1.4 km west of George Lake, Alberta. 1-N. Interpopulational variation of specimens from: I. Km. 140, Dempster Highway, Yukon
Territory; J. Churchill, Manitoba; K. Gillam, Manitoba; L. Whitehorse, Yukon Territory; M. Yellowknife, Northwest Territories;
N. Chisholm, Ontario. Penis valves of adult male Chaoborus nyblaei. Specimens from: O. 2-3 km SW Nuorgam, Finland; P.
Abisko, Sweden.
Systematics and Bionomics of Schadonophasma
111
Figure 12. Scale line = 0.1 mm. Variation of penis valves of adult males due to orientation. A-E. Chaohorus irivitlatus from 2.4
km west of Edmonton, Alberta. F-J. Chaoborus cooki from 32 km west of Edson, Alberta.
Quaest. Ent., 1979, 15 (2)
228
Borkent
Figure 13. Scale line = 0.1 mm. Prelabral appendages of fourth instar larvae of Chaoborus trivittatus. A-G. Intrapopulational
variation of specimens from 2.4 km west of Edmonton, Alberta. H-O. Interpopulational variation of specimens from: H. 4.8 km
north of St. John’s, Newfoundland; I. 2.4 km west of Edmonton, Alberta; J. 2.4 km south of Robb, Alberta; K. Gwendoline Lake,
British Columbia; L. 4.8 km east of Sicamous, British Columbia; M. 45 km east of Jasper, Alberta; N. Lake 241, Kenora, Ontario;
O. East Henry Pond, Jasper National Park, Alberta.
Figure 14. Scale line = 0.1 mm. Prelabral appendages of fourth instar larvae of Chaohorus couki. A-F. Intrapopulational
variation of specimens from 1.6 km south of Jasper, Alberta. G-L. Interpopulational variation of specimens from: G. Churchill,
Manitoba; H. Pond nr. Harris River, Northwest Territories; I. Dempster Highway, Yukon Territory; J. 1.4 km west of George
Lake, Alberta; K. Klutlan Glacier moraine, Yukon Territory; L. 23.3 km west of Jasper, Alberta.
Quaest. Ent., 1979, 15 (2)
230
Borkent
15
Figure I 5. Scale line = O.l mm. Prelabral appendages of fourth instar larvae of Chaoborus nyblaei from 2-3 km southwest of
Nuorgam, Finland.
Systematics and Bionomics of Schadonophasma
231
Quaest. Ent., 1979, 15 (2)
232
Borkent
Figure 17. Known distribution of Chaoborus trivittatus in Alberta.
Systematics and Bionomics of Schadonophasma
233
Figure 18. Known distribution of Chaoborus cooki.
Quaest. Ent., 1979, 15 (2)
234
Borkent
Figure 19. Known distribution of Chaoborus cooki in Alberta.
Systematics and Bionomics of Schadonophasma
235
Figure 20. Known distribution of Chaoborus nyblaei.
Quaest. Ent., 1979, 15 (2)
236
Borkent
lil n = 26
ril n = 18
r#i " = 2o
I ^ I n = 35
Chaoborus trivittatus
n = 80
Chaoborus cooki
] n = 49
n =474
n=210
300 600 900 1200 1500 1800
Head Capsule Length (jiim)
A
2100 2400 2700
In stars
1
2
3
4
1
2
3
4
it " = 28
Chaoborus trivittatus
J±_ "=2>
^ I n = 83
it n = 20
rin n = 33
Chaoborus cooki
n= 50
n= 493
n=210
0 150 300 450 600 750 900 1050 1200
Antennal Length (^m)
B
21
Figure 21. Variation of characters of larval instars of Chaoborus trivittatus and Chaoborus cooki. A. Head capsule length. B.
Antennal length.
Systematics and Bionomics of Schadonophasma
237
n = 27
Chaoborus trivittatus
n=20
■ ' I nr 20
I ~l " = 83
r~#~i
Chaoborus cooki
nr 33
nr 50
rin "=210
H 1 1-
02 0 3 0 4 0 5 0 6 0 7
AS/AL
09 10
Instars
1
2
3
4
1
2
3
4
7^
0 4
Chaoborus trivittatus
nr 28
nr 21
nr 82
nr 513
nr 20
n r 34
n r 49
Chaoborus cooki
nr 201
055 0 7 0-85 10 115 13 1-85 2 3 275
LB/AL
B
22
Figure 22. Variation of characters of larval instars of Chaoborus trivittatus and Chaoborus cooki. A. AS/AL. B. LB/AL.
Quaest. Ent., 1979, 15 (2)
238
Borkent
Instars
2
3
4
2
3
4
I
±
Chaoborus trivittatus
^ 1 n=46 Chaoborus cooki
n=201
-H ^ 1 1 1 1 1 h-
1-25 V5 1-75 2 0 2 25 2-5 275 30
LB/SB
A
Instars
1 A n = 15
2 [tl n.2l
3
4
1 \k\ n = 20
2 n = 34
3
4
Chaoborus trivittatus
] n =88
"I n = 500
Chaoborus cooki
^ 1
n = 210
0 125 250 375 500 625 750 875 1000
Postantennal Filament Length (pm)
B
23
Figure 23. Variation of characters of larval instars of Chaoborus trivittatus and Chaoborus cooki. A. LB/SB. B. Postantennal
filament length.
Systematics and Bionomics of Schadonophasma
239
Instars
3
4
3
4
i=L
Chaoborus trivittatus
nr 88
n =46
nr 210
H ^ ^ ^ h-
20 4 0 60 80 100
PAL / PAW
nr 490
Chaoborus cooki
12-0
140
160
Instars
2
3
4
2
3
4
n r 19
rtl n= 34
rki n =50
r-i-1
Chaoborus trivittatus
n.82
n =517
Chaoborus cooki
n = 213
10 15 20 25 30
Number of Mandibular Fan Bristles
B
35
24
Figure 24. Variation of characters of larval instars of Chaoborus trivittatus and Chaoborus cooki. A. PAL/PAW. B. Number of
mandibular fan bristles.
Quaest. Ent., 1979, 15 (2)
240
Borkent
Insfars
1
2
3
4
1
2
3
4
r ^ ~i " = 26
L- L
Chaoborus triviftafus
] n = 87
i I nr513
^ Chaoborus cooki
n =35
15
-H- ^ 1 1 h
20 25 30 35 40
Number of Anal Fan Setae
A
______ __ C. TRIVITTATUS
C. COOKI
I II III IV V VI VII VIII IX X XI XII
B
25
Figure 25. Variation of characters of larval instars of Chaoborus trivittatus and Chaoborus cooki. A. Number of anal fan setae. B.
Temporal distribution of fourth instar larvae of Chaoborus trivittatus and Chaoborus cooki.
Systematics and Bionomics of Schadonophasma
241
720
680
640
600
560
•• •
• •
• •
• •
• •
• 4
•
• •• •
1:^ ^
0 11.0 12.0 13.0 14.0 15.0
TOTAL LENGTH (in mm.)
16.0
325
300
c
g 275
250
225
200
• •
• •
f •
» •
•• • •
o'U
• m • •
• •
• •
• • •
11.0 12.0 13.0 14.0 15.0 16.0
TOTAL LENGTH (in mm.)
B
26
Figure 26. Age-related variation of male fourth instar larvae of Chaoborus cooki. A. Relationship between total length and long
antennal blade length. B. Relationship between total length and prelabral appendage length.
Quaest. Ent., 1979, 15 (2)
242
Borkent
MALES
FEMALES
10
MALES
FEMALES (-1
n
15
20
25
30
35
40
COMPOUND CHARACTER INDEX YALUE
B
7
MALES
FEMALES
c
Figure 27. Results of compound character index used to sex fourth instar larvae. A. Of Chaoborus trivittatus from 2.4 km west of
Edmonton, Alberta. B. Of Chaoborus cooki from 32 km west of Edson, Alberta. C. Of Chaoborus cooki from 1.6 km south of
Jasper, Alberta.
Systematics and Bionomics of Schadonophasma
243
28
Figure 28. Correlation of characters of fourth instar larvae. A. Male Chaoborus trivittatus B. Female Chaoborus trivittatus. C.
Male Chaoborus cooki. D. Female Chaoborus cooki. Open spaces represent lack of significant correlation; + and - signify
positively and negatively significant correlations, respectively.
Quaest. Ent., 1979, 15 (2)
244
Borkent
1000 1100 1200
ANTENNAL LENGTH (in pm)
c
1300
29
Figure 29. Relationship between antennal length and distance of antennal seta from base of antenna of fourth instar larvae. A.
Chaoborus trivittatus. B. Chaoborus cooki. C. Chaoborus nyblaei.
Systematics and Bionomics of Schadonophasma
245
alberta
30
Figure 30. Localities of samples used to study geographical variation of characters of fourth instar larvae of Chaoborus trivittatus
in Alberta. Samples from; 1. 16 km west of Jasper; 2. 45 km east of Jasper; 3. Pond nr. Kinky Lake; 4. 4.8 km south of Hinton; 5.
2.4 km south of Robb; 6. 69 km east of High Prairie; 7. 61 km north of Swan Hills; 8. 53 km north of Swan Hills; 9. 1.6 km north
of Swan Hills.
Quaest. Ent., 1979, 15 (2)
246
Borkent
/
/
/
/
CANADA
states
500 KMS
— I
31
Figure 31. Localities of samples used to study geographical variation of characters of fourth instar larvae of Chaoborus trivittatus
from the west coast of North America. Samples from: 1. Klutlan Glacier moraine, Yukon Territory; 2. Prince Rupert, British
Columbia; 3. Gwendoline Lake, British Columbia; 4. Eunice Lake, British Columbia; 5. Hall Lake, Washington; 6. Mad River,
California; 7. Stanford and Jewel Lake, California.
Systematics and Bionomics of Schadonophasma
247
ALBERTA
Figure 32. Localities of samples used to study geographical variation of characters of fourth instar larvae of Chauhunis cooki in
Alberta. Samples from: 1. 69 km north of Coleman; 2. 53 km south of Jasper; 3. 1.6 km south of Jasper; 4. 6.4 and 23.3 km west
of Jasper; 5. 32 km west of Edson; 6. 1.4 km west of George Lake; 7. 2.4 km west of Edmonton.
Quaest. Ent., 1979, 15 (2)
TRIVITTATUS
248
Borkent
STvnaiAioNi do mmm
Systematics and Bionomics of Schadonophasma
249
Figure 34. Variation of characters of fourth instar larvae of Chaoborus trivitlatus. A. Number of mandibular fan bristles. B.
LB/SB. C. LB/AL.
Quaest. Ent., 1979, 15 (2)
250
Borkent
NUMBER OF DAYS
A
Figure 35. Mortality of adult Chaoborus trivittatus. A. Males. B. Females.
C ceratopogones
C.pallidus
Systematics and Bionomics of Schadonophasma
251
Quaest. Ent., 1979, 15 (2)
Figure 36. Phylogeny of species of Chaoborus. Black circles signify apomorphic states, open circles plesiomorphic states. M.Y.B.P. indicates million years before present.
252
Borkent
No. Anal Fan Setae
15 20 25 30 35 40
^ H ^ \ ^ ^
C. brasiliensis (0 79-0 83)
— C.anomalus (0-71-0 87)
C.ceratopogones (0 74-0 90)
C.magnificus (0 91-0 96)
— C.pallidipes (0 85 -103)
— C. punctipennis (0 85 -111)
C.astictopus (0 80-1 11)
C.albatus (0 89-112)
C.pallidus (100-150)
C.americanus (l-OO-l 30)
C. flavicans (0 90 - 1-45)
C.crystallinus (112-165)
C.obscuripes (0 98-182)
C.trivittatus (140-2 38)
C.cooki (1 71-2-62)
C.nyblaei (185-260)
37
Figure 37. Relationship between number of anal fan setae and head capsule length (given in parentheses as mm) of fourth instar
larvae of Chaoborus species.
Systematics and Bionomics of Schadonophasma
253
C trivittatus
C. cooki
C. nyblaei
Figure 38. Phylogeny of species oi Schadonophasma. Black circles signify apomorphic states, open circles plesiomorphic states.
Quaest. Ent., 1979, 15 (2)
TRIVITTATUS COOKI NYBLAEI TRIVITTATUS COOKI NYBLAEI
254
Borkent
Figure 39. Schematic representation of alternative zoogeographic hypotheses for Schadonophasnia species.
Systematics and Bionomics of Schadonophasma
255
O = PERMANENT POND
□ = TEMPORARY POND
Figure 40. Schematic representation of events of sympatric speciation of lineages giving rise to Chauhorus trivittatus and
Chaoburus cooki-nyblaei.
Quaest. Ent., 1979, 15 (2)
i.'„ M ■
^rnr]
mm r:'iMj9^m Q
T^a'.E:pjdrvtUi/!'F
-- r : 1-
L
-■^ iq I' I:
i sv. . - 4'
MvJi ;ij'Ul4 0
>,fJ r
'’ivi'i’f .Vp , \ i
./
. ' j") L.
EFFECTS OF TEMPERATURE AND RELATIVE HUMIDITY ON EMBRYOGENESIS IN EGGS
OF MAMESTRA CONFIGURATA (WALKER) (LEPIDOPTERA: NOCTUIDAE)
M.P. JONES^
Department of Entomology
University of Alberta
Edmonton, Alberta
T6G 2E3
B.S. HEMING
Department of Entomology
University of Alberta
Edmonton, Alberta
T6G 2E3
Effects of various combinations of temperature and relative humidity on embryogenesis in eggs of
Mamestra configurata (Walker) were investigated. The following temperature thresholds were
determined for some stages of embryogenesis: developmental-hatching, (8.5 C), hatching, (5.0 C),
developmental (between 0.0 and 2.0 C) and high temperature developmental-hatching threshold,
(30.0 C). Temperature and rate of development curves were derived using three different relative
humidities(0, 60 and 98% } and a range of temperatures (8.5-30.0 C). Length of exposure to 35.0 and
5.0 C required to produce mortality of 50% and 95%o, was determined for eggs of various ages. The
age of eggs exposed to 35.0 C did not appear to influence mortality, but was very important in eggs
exposed to 5.0 C. The older the eggs, the longer the exposure required to produce 50% and 95%o
mortality. The effects of daily exposure to 35.0 C and 5.0 C were studied for eggs of various ages.
Older eggs could tolerate longer daily exposure to 35.0 C without high mortality than could younger
eggs but, when total length of exposure was determined, there was no significant difference in tolerance
between older and younger eggs. Daily exposure to 5.0 C had little effect on mortality but lengthened
development.
The developmental rate and temperature curve for eggs of Mamestra configurata is J-shaped.
Practical application of this curve to field populations will increase the accuracy of larval surveys.
Nous avons etudie les effets de diverses combinaisons de temperature et d’humidite relative sur I’embryogenese des oeufs de
Mamestra configurata Walker. Nous avons determine les seuils de temperature pour quelques stades de I’embryogen'ese: la
temperature minimale permettant le developpement complete de I’oeuf et son eclosion est 8.5 C, et la temperature maximale est
30.0 C; la temperature minimale pour I’eclosion seulement est 5.0 C. la temperature minimale a laquelle I’embryon se developpe
(sans eclore) se situe entre 0.0 et 2.0 C. Nous avons etudie le taux de developpement en fonction de la temperature (entre 8.5 et
30.0 C) sous trois conditions d’humidite relative (0, 60 et 98%). Nous avons determine, pour des oeufs d'dges divers, les durees
d’exposition a 35.0 C et 5.0 C causant 50%> et 95% de mortalite. A 35.0 C, Cage des oeufs ne paralt pas affect er la mortal ite, mais a
5.0 C, son effet est important: plus les oeufs sont ages, plus longue est la periode d’exposition requise pour obtenir 50% et 95%> de
mortalite. Nous avons etudie les effets d’expositions journalieres a 35.0 C et a 5.0 C sur des oeufs d’dges divers. Les oeufs plus ages
peuvent tolerer une plus longue exposition journaliere a 35.0 C que les jeunes oeufs, sans qu’en resulte une mortalite elevee; mais
lorsqu’on considere la duree totale d’exposition, on n’observe aucune difference significative entre la tolerance des oeufs ages et celle
des jeunes oeufs. L’exposition journaliere a 5.0 C affecte peu la mortalite, mais prolonge le developpement embryonnaire.
Le taux de developpement des oeufs de M. configurata en fonction de la temperature suit une courbe en forme de "J”.
L’application d’une telle courbe aux populations naturelles permettra d’obtenir des estimes plus precis de I’etat des populations
larvaires.
Quaestiones Entomologicae
15:257-294 1979
^Present address: South Peace Senior Secondary School, Dawson Creek, British Columbia.
258
Jones & Heming
CONTENTS
Introduction 258
General Methods 259
Experiments and Results 260
Discussion and Conclusions 285
Acknowledgements 291
References 292
INTRODUCTION
Moths of the species Mamestra configurata (Walker) (Lepidoptera: Noctuidae), the Bertha
armyworm, occur from Mexico City, Mexico in the south (King, 1928) to Keg River, Alberta in the
north (Philip, pers. comm). Within this range, the species is of economic importance only in Western
Canada and in the State of Washington.
The species was first cited in the economic entomological literature in 1928 in a paper by King
(1928). That paper dealt with external structure of the various lifestages, with some aspects of its life
history including geographic range and host plants, (rape was not included), and included descriptions of
larval damage on various crop plants. His larval and pupal descriptions are still helpful in separating
specimens of Mamestra configurata from those of other noctuids causing damage to rape.
In Alberta, the species has one generation per year (Beirne, 1971). Eggs are deposited by the females
on leaves of host plants early in July. Larvae feed on these until about three-quarters grown and then
chew into the flowers, seed pods, bolls or fruit in August and early September. Larvae pupate in the soil
and overwinter, emerging as adults the following'June and July.
Embryogenesis in eggs of Mamestra configurata, from pre-fertilization to eclosion, has been
described by Rempel (1951) who mentioned ovipositional habits of this moth and enlarged upon King’s
(1928) observations of its eggs. One of the more important observations Rempel made, which had
important implications for our research was that fertilization at room temperature (20.0 C - 21.0 C)
occurred in the second half hour after oviposition.
The serious outbreaks in 1971 and 1972 of larvae of this species on rape {Brassica campestris L. and
Brassica napus L.) stimulated numerous studies, the results of which are now being published.
The male pheromones of Mamestra configurata have been studied by Clearwater (1975a,b) and
those of females by Struble et ai, (1975). Clearwater (1975c) also described the structure and
postembryogenesis of the male pheromone system. Bodnaryk (1978) investigated factors affecting
diapause development and survival of pupae and Bailey (1976), the effects of temperature on
non-diapause development.
In Western Canada, the importance of Mamestra configurata increased in 1971 and 1972, partly
because of a large increase in rape acreage. After the first serious outbreak of this insect in 1971, the
Alberta Department of Agriculture instituted a series of four annual surveys to determine, first, the
areas where densities of Mamestra configurata were high and where the potential for economic damage
great, and second, to follow the changing distribution of these populations in Alberta each year.
A fall pupal survey was conducted in areas where outbreaks occurred that year, and its results
enabled estimates to be made about the initial size of overwintering populations. A second pupal survey
was conducted in the spring to ascertain winter mortality of overwintering populations. A third survey.
Temperature, RH and embryogenesis in Mamestra configurata
259
conducted from mid-May until the end of September, involved the use of black light traps to monitor
adult emergence and abundance. Larval surveys were conducted in July in areas where light trap
captures were considered high, and first and second instar larval populations were monitored 14-21 days
after peak adult activity. Based on results of these surveys, a series of maps were prepared which
indicated areas of potential economic damage for that crop year. The most important survey was the
final one because this confirmed the presence of the damaging larval stage in the field.
The principal weakness of the final survey is its timing. For it to be effective, it should be conducted
at a time when the majority of eggs have hatched, but before the larvae have reached a size where they
are causing economic damage. If the survey is conducted too early before most of the eggs have hatched,
the population estimate resulting may be too low. If left until too late, it may be impossible to devise and
implement control measures before considerable damage has occurred.
There were three principal objectives in undertaking this study: (1) to remove some of the guess-work
involved in timing the egg and larval survey by developing temperature curves for embryogenesis which
could aid in predicting probable hatching time of eggs in the field, (2) to determine the effects on
development and viability of eggs exposed to various periods of unfavourable temperature, and (3) to
determine the effect of relative humidity on embryonic development in this insect.
GENERAL METHODS
Handling and collecting eggs
Collecting large numbers of eggs of known age was essential for much of the experimental work. The
following procedures were used to facilitate collection and aging of these eggs. All eggs used in the
following experiment were obtained from moths reared from field-collected pupae.
Eggs for stock use. - Moths were placed in cages containing four rape plants and subjected to a
photoperiod of 16L:8D. Eggs were collected once a day at the end of the dark period by removing leaves
containing egg clusters. Often it proved necessary to break the clusters into smaller units. The adhesive
which binds the eggs to the leaves was softened with distilled water and individual eggs were removed by
gently pushing them with a camel hair brush.
Collecting accurately-timed eggs. - A stock culture of male and female moths was caged in a growth
chamber at 20.0 C and approximately 60% RH (for some experiments a culture was maintained at
15.0 C). The moths were fed a 10% honey and water solution from wicked containers, the solution being
changed twice a week. Since adults of Mamestra configurata generally oviposit at night, we reversed the
normal photoperiod so that darkness occurred between 0800 and 1600 hours.
Females prefer a rough substrate for oviposition, but, given no choice, will oviposit on almost any
surface. Plastic sandwich wrap was avoided by females, probably because they were unable to find a
purchase on it. If the cage is lined with this material, the moths show strong ovipositional preference for
paper towellling or other rough textured materials. Paper toweling was chosen as the ovipositional
substrate because of its availability and ease of handling. The towelling was folded in such a manner that
it was able to stand on its own. It was introduced into the cage approximately one hour after dark and
replaced 1 3/4 hours later.
Once the paper towelling was removed, egg clusters were examined at XI 2 magnification under a
binocular microscope and were separated into groups of the desired number of eggs with a razor blade.
The strips containing eggs were quickly examined and any damaged eggs were discarded. Each strip was
then placed in a clean, 1 mm cap vial, ready to be used in the experiment. The top of each vial was
covered with plastic screening, held secure by a rubber band. Any eggs remaining from an egg cluster
were placed in a vial marked to indicate their origin and were used to check fertility for that group.
Quaest. Ent., 1979, 15 (2)
260
Jones & Heming
EXPERIMENTS AND RESULTS
Development of eggs
Eggs of Mamestra configurata are generally deposited on the underside of leaves of the host plant in
a tight, single-layered cluster and are oriented with their anterior, micropylar ends pointed away from
the substrate (Rempel, 1951). They are yellowish white when first deposited, but become off-white
several hours later. Approximately 24 hours later (at 20.0 C), a band of brown pigment appears around
the equator of fertile eggs with additional small patches developing in the micropylar area (Rempel,
1951). Approximately eight hours prior to hatching, the egg turns jet black as the larval head capsule
becomes visible through the transparent chorion — a stage referred to as the “black spot stage” by
Peterson (1964). A more detailed description of the egg was given by Jones (1977).
Viability and size of field-collected versus laboratory deposited eggs
The main purpose of this experiment was to determine if laboratory rearing and handling techniques
influenced egg viability or numbers laid when compared to eggs deposited under field conditions.
Methods. - A laboratory culture of adult moths was maintained in a growth chamber at 20.0 C ±
0.5 C and approximately 60% RH. The photoperiod maintained was 16L:8D, a close approximation of
field conditions at the time of the experiment. The moths were provided with potted rape plants for
oviposition, these being at approximately the same stage of development as were field sown plants. Ten
egg clusters of unknown age were removed at random from the caged culture by removing the leaf with
the cluster. Each cluster was then divided carefully into groups of 15 eggs as previously mentioned and
these placed in 1 mm cap vials. When less than 15 eggs remained, the remaining group was also placed
in a 1 mm cap vial. Eggs from each cluster were kept separate so that mortality and number of eggs
could be recorded for each cluster.
Field samples of eggs were studied in a field of rape near Lacombe, Alberta. Ten egg clusters, of
unknown age, were located and counted and the leaf with each was encased in a fibre-glass screen
having a mesh size of 16 threads per cm to prevent access of predators, parasites and larval escape.
These cages were inspected twice daily and the number of hatched larvae recorded for each cage.
Results. - The results of this experiment are recorded in Table 1. Although egg viability (97%) did
not differ significantly between the two groups, the average size of the laboratory clusters (146.5) was
significantly larger than that of field clusters (80.8). Bailey (1976) reported that females of Mamestra
configurata reared for ten generations on an artificial diet deposited 40-60 eggs per cluster with viability
of eggs ranging from 90-97% at temperatures from 8.0-28.0 C.
Temperature thresholds for embryogenesis
Insect embryogenesis is influenced by many external factors which, acting separately or together,
restrict developmental potential. Within the range of each of these factors are points designated as
thresholds. Continual exposure of an embryo to the factors beyond these points eventually results in
death.
One of the most important and easily studied factors influencing embryogenesis is temperature.
Insects, being poikilothermic, are greatly affected by ambient temperature. However, only the egg and
the quiescent pupal stage are unable to move to a different microclimate to avoid temperature extremes.
For this reason, eggs and non-mobile pupae are superior to other stages for study of temperature
thresholds.
Temperature, RH and embryogenesis in Mamestra configurata
261
Table 1. Comparison of size of egg clusters and viability between laboratory and field collected eggs of
Mamestra configurata.
*means significantly different based on T-tests (P <C 0.01).
The developmental threshold. - The developmental threshold is the temperature at which, on the
descending scale, development definitely ceases, and at which, on the ascending scale, development
begins (Peairs, 1927). Knowledge of this threshold is vital if one is attempting to predict development on
the basis of temperature. An experiment was conducted to determine this threshold for embryos of
Mamestra configurata. Histological studies were made simultaneously.
Methods. Eggs used in this experiment were collected from clusters of over 100 deposited by females
maintained at 15 ± 0.5 C. Eggs were placed at experimental temperatures of 0 ± 0.5 C, 2 ± 0.5 C and
4 ± 0.5 C when less than 30 minutes old, thus ensuring (based on Rempel’s 1951 paper) that no prior
development had occurred.
Two separate clusters of eggs were placed at each temperature. The eggs from each cluster were
divided into 20 groups of five eggs each, each group of five being placed in a separate 1 mm cap vial (the
eggs of the two clusters were kept separate). All but two vials were placed on a platform in a 160 mm
desiccator containing a saturated salt (KNO3) solution (Winston and Bates, 1960) to maintain the
humidity at about 96%. Eggs in one of the remaining vials were fixed immediately in hot, alcoholic
Bouins solution while the other vial was removed and placed at room temperature to determine if all its
contained eggs were fertile.
Hot alcoholic Bouins was poured into one vial from each group daily for the first five days. After that
time eggs were fixed at five day intervals. Standard histological techniques were used to prepare the eggs
for staining with Delafield’s Haematoxylin and Mallory’s Triple Stain (Humason, 1972).
The experiment was concluded when histological signs of development were found in two or more
eggs from both vials.
Results. The temperature developmental threshold for embryogenesis of Mamestra configurata is
between 0.0 and 2.0 C.
Eggs at 4.0 C showed recognizable development after ten days and at 2.0 C after 15 days. No sign of
development occurred at 0.0 C even after 40 days (data from eggs kept after end of experiment).
Quaest. Ent., 1979, 15 (2)
262
Jones & Heming
The hatching threshold. - The hatching threshold is the lowest temperature at which hatching of a
fully developed larva can occur (Johnson, 1940).
The egg of Mamestra configurata is an excellent subject for this type of experiment because
darkening of the head capsule always indicates that development is complete and that hatching is soon to
follow.
Methods. Three egg clusters, less than one day old, and each containing in excess of 150 individuals,
were collected over a period of two days from the stock culture. Each cluster was divided into groups
of 50. Surplus eggs from each cluster were placed in 1 mm cap vials and labelled according to their
cluster of origin. These eggs were used as spares to be substituted for any infertile eggs in the
experimental group. Each of these nine groups of 50 eggs was placed in a 4 mm glass cap vial and
labelled so that groups from individual clusters could be identified. All vials were then placed on a
platform, at 20.0 C in a 160 mm desiccator which had been partially filled with distilled water to raise
the humidity to greater than 90%.
The eggs were observed closely and were allowed to develop to the black head capsule stage. Then, to
prevent larval escape, the tops of the vials were covered with a piece of plastic screening, secured by a
rubber band. One vial from each egg cluster was then quickly transferred to identical, water-filled
desiccators located in incubators set at temperatures of 7.5 ± 0.5 C, 5.0 ± 0.05 C and 2.5 ± 0.5 C.
Eggs were observed twice daily and total number of hatched eggs recorded. The eggs were maintained at
the experimental temperature for a maximum of 30 days after which they were returned to 20.0 C to
determine if the remaining eggs would hatch.
Results. The results of this experiment are summarized in Table 2. The first group of eggs to hatch
were those at 7.5 C. The lowest experimental temperature at which eggs hatched was 5.0 C. No eggs
hatched at 2.5 C even after 30 days and these failed to hatch even after being returned to 20 C.
Examination of these eggs showed that the embryos had died and dried up. There was a greater
percentage hatch at 7.5 C (92.2%) than at 5.0 C (66.0%).
Table 2. The hatching threshold for eggs of Mamestra configurata.
The developmental-hatching threshold. - The developmental-hatching threshold is the lowest
temperature at which complete development from fertilization to eclosion can occur (Johnson, 1940).
This threshold is the most important of those so far discussed, because, for eggs of some species, it can be
used as an aid in determining distribution. Obviously, areas in which the daily high temperature is
always below this threshold for a particular insect, will not have that insect present.
This experiment was designed to determine the lowest temperature at which complete development of
eggs of Mamestra configurata could occur. Three different relative humidities were used to show if
Temperature, RH and embryogenesis in Mamestra configurata
263
humidity had any influence on development. Previous research had shown that complete embryogenesis
could occur at 10.0 C but not at 5.0 C.
Methods. Eggs were collected over a period of three days from a culture of adults maintained at
15.0 ± 0.5 C and 60% RH. The methods used for collecting and handling eggs were identical to those
described previously, with the exception that they were collected when less than 1 hour old to insure that
little, if any, development had occurred.
For each daily collection, the eggs were divided into ten groups of 30, each group being placed in a
1 mm cap vial. One of the groups was placed at 20.0 ± 0.5 C and left for the remainder of the
experiment as a test of egg viability. The remaining nine groups formed the first of three replicates. Each-
of the nine vials was placed under a different experimental condition. Vials were placed at temperatures
of 6.5 ± 0.25 C, 7.5 ± 0.25 C, and 8.5 ± 0.5 C and at relative humidities of about 0%, 60% and 98%
for each temperature. Temperatures were maintained in an incubator (6.5 C), in a refrigerated water
bath (7.5 C) and in a growth chamber (8.5 C). Relative humidity was controlled in 6 mm cap vials using
saturated salt solutions (P2O5 for 0%, Na2Cr207: H2O for 60% and K2SO4 for 98%) (Winston and Bates,
1960).
Eggs were observed daily for sign of development. When embryos reached the black head capsule
stage, observations were made every two hours.
Results. Percentage hatch for each control was 90% or greater. In the experimental groups, hatching
occurred only at 8.5 C and 98% RH. Development to the black head capsule stage occurred in some
individuals at 8.5 C and 60% RH and at 7.5 C and 98% RH. Dissection of a portion of these embryos
several days later revealed no outward sign of structural disorder.
The high temperature developmental-hatching threshold. - Mamestra configurata has a reported
geographic range that spans vastly different climatic regions extending from Keg River in northern
Alberta (Philip, pers. obs.) to Mexico City, Mexico (King, 1928). Populations occurring in central and
northern Alberta are exposed to cool springs, warm short summers, and long cold winters, their eggs
experiencing temperatures ranging from 5.0 C to just above 30.0 C. However, in the southern part of its
range, it is likely that eggs of Mamestra configurata are exposed to a much higher range of
temperatures.
The purpose of this experiment was to determine the high temperature developmental-hatching
threshold for eggs of Mamestra configurata and to find out whether eggs of the cold adapted
populations of Alberta retain some resistance to high temperature (30 C and above).
Methods. This experiment was performed essentially like the previous ones, (developmental-hatching
threshold) except that temperatures of 30.0 ± 0.5 C, 31.5 ± 0.5 C, 32.5 ± 0.5 C and 33.5 ± 0.5 C
were used. Temperatures of 30.0 C and 35.5 C were maintained in a heated water bath and of 31.5 C
and 32.5 C in incubators. Humidities were maintained using saturated salt solutions of P2O5 for 0% RH,
NH4NO3 for approximately 60% RH and K2Cr20y for 98% RH (Winston and Bates, 1960).
Results. The results of this experiment are summarized in Table 3. The listings are the totals of three
replicates of 30 eggs. Unlike eggs in the previous experiment, relative humidity did not have a significant
effect on hatching. Of the four temperatures tested, hatching occurred only at 30.0 C regardless of
humidity. Some development occurred in eggs at all other experimental temperatures. All eggs (with the
exception of some, probably infertile) developed the characteristic brown pigment of the equator and
micropylar areas.
Eggs at 32.5 C developed to the point where larvae were clearly visible through the chorion, however,
sclerotization and pigment deposition did not occur.
Quaest. Ent., 1979, 15 (2)
264
Jones & Heming
Table 3. High temperature developmental threshold of eggs of Mamestra configurata and effects on it
by relative humidity.
No. hatching (maximum 90)*
Temperature in ° C 0% RH 60% RH 98% RH
* Figures = totals of three replicates of 30 eggs.
Effects of constant temperature and relative humidity on embryogenesis
Effects of exposure to constant temperature and relative humidities of 0%, 60% and 98% on
developmental rate. - When dealing with an insect pest species, knowledge of the effects of climatic
factors, particularly those of temperature on various stages of its life cycle is imperative if reliable
predictions of outbreaks are to be made and if successful control procedures are to be implemented. The
most accurate way of determining individual effects of various field conditions oh a particular instar of
an insect is to duplicate every possible combination of those conditions in the laboratory. This, however,
is usually impossible to do or is too time consuming. The usual alternative (although it lacks the same
degree of accuracy) is to use a range of constant temperatures, encompassing the range found in nature
and to extrapolate from these results to development in the field.
Few authors have attempted to evaluate the effects of relative humidity acting in conjunction with
constant tC'mperatures on insect embryogenesis. This is rather surprising when one considers the effect
relative humidity can have on development (Buxton, 1932; Ludwick, 1945; and Bursell, 1974). Since
Mamestra configurata has a large geographic range (King, 1928), it is probably exposed to a
considerable range of relative humidities. This experiment was conducted to determine the effects that
relative humidity might have on embryogenesis under constant temperature conditions.
Methods. Ten different temperatures combined with three different relative humidities were used to
determine the effect of temperature and relative humidity on embryogenesis. The lowest temperature
used was 8.5 C (ie., the developmental-hatching threshold); the highest 30.0 C (ie., the high temperature
developmental threshold). The remaining temperatures began at 10.0 C and increased at 2.5 C
increments. Growth chambers were used to maintain the temperatures of 8.5 C, 12.5 C, 20.0 C, and
25.0 C, and incubators for the remainder. The temperature in each incubator was monitored and found
to vary no more than ± 0.5 C when the door was closed. For most experiments, the doors were opened
briefly, once a day, to check egg development. Temperatures returned to equilibrium in less than 30
minutes after the door was closed.
Dry P2O5 was used to provide 0% RH, and KNO3 to provide 98% RH at all temperatures used.
NaBr:2H20 was used to approximate 60% RH for temperatures of 10.0 C, 12.5 C, 15.0 C, 17.5 C and
20.0 C, Na2Cr20y:H20 for a temperature of 8.5 C and NH4NO3 for the remainder (Winston and Bates,
1960). The saturated salt solutions were prepared and placed at the desired temperature 30 days prior to
being used to allow them to equilibrate. Humidities in the chambers were checked prior to, during and
after the experiment and were always found to be within 6% of the desired humidity.
Eggs used in this experiment were collected as previously described from a culture of adults kept at
20.0 ± 0.5 C and 60% RH. Thirty eggs were placed in each vial. These vials were exposed to the
Temperature, RH and embryogenesis in Mamestra configurata
265
experimental conditions within one hour of collection. The experiment was replicated three times. Early
in the experiment, it became apparent that insufficient numbers of eggs could be collected at any one
time to complete a replicate for the temperature treatments. Instead, the eggs of each collection were
used in replicating the three relative humidity treatments used at each temperature.
The eggs were examined once a day until they had reached the black head capsule stage at which
time observations were recorded at two hour intervals until hatching was complete.
Results. Results of these experiments are summarized in Table 4 and Figures 1-3. Hatching occurred
at all temperatures tested. Rate of development (the reciprocal of the total development time measured
in hours) showed a strong positive correlation with temperature (correlation coefficients of 0.978, 0.989
and 0.982 were recorded for 0%, 60%, and 98% RH respectively). The fastest development occurred at
30.0 C- the slowest at 8.5 C.
Humidity had considerable effect on development. Development but no hatching occurred at 0% RH
for temperatures of 12.5 C and lower, and at 8.5 C for 60% RH. Hatching always occurred first in those
eggs at 98% RH followed by those at 60% RH and 0% RH.
Figures 1 to 3 demonstrate the effects of relative humidity and temperature on development. The
straight lines (B) are drawn on the basis of the linear regression equation Y = 7.43 + l,966.44x;
Y = 8.15 + l,761.23x and Y = 8.22 + l,708.85x for 0%, 60% and 98% RH respectively. These lines are
the velocity lines and represent the percentage of total development that occurs during one hour at that
temperature. The points on and around these lines are the calculated values based on the reciprocal of
the mean development time (listed in Table 4).
The hyperbolic curve (A) is based on the mean development time in hours at the experimental
temperatures and humidities indicated. The squares on the graph represent the mean development time
in hours for that temperature and humidity.
Effects of constant exposure to a temperature of 35.0 C on development of eggs of different ages. -
Results of preliminary experiments had suggested that Mamestra configurata is imperfectly adapted
to the climatic regime found in the prairie provinces (Putnam, 1972). To perfectly adapt to this climate,
the pupal population of Mamestra configurata should have an obligatory diapause (diapause
development in this insect has been studied by Bailey (1976) and Bodnaryk (1978)). This however, does
not always happen since flights of newly-emerged adults have been taken in blacklight traps in late fall,
and some field-collected pupae have been shown to develop without exposure to cold (Philip, pers.
comm.). These observations suggest that at least part of the population has either a facultative diapause
or lacks it completely. The existence of this partial late second generation in the prairie provinces
suggests that Mamestra configurata may be bi- or multivoltine in more southerly parts of its range.
The main purposes of this experiment were to determine if eggs of Mamestra configurata
demonstrate a tolerance to high temperature and to discover the length of exposure required to produce
50 and 95% mortality. A temperature of 35.0 C was chosen for two reasons; because previous
experiments had shown that this was above the upper developmental-hatching threshold and because
temperaures of 35.0 C or greater for periods over 16 hours per day have been recorded in Arizona cotton
fields by Fry and Surber (1971), well within Mamestra configurata's range. Their study of the effects of
high temperature on embryonic mortality included eggs of Estigmene acrea, a species often found in
conjunction with Mamestra configurata, and which has a similar life cycle in Alberta (Beirne, 1971).
Methods. Eggs used in this experiment were collected from a culture of adults reared at 20.0 ±
0.5 C, as previously described.
Eggs of four different ages (3 hours, 24 hours, 48 hours, and 96 hours) were exposed to 35.0 C
continuously for periods of 13, 20, 30, 45 and 67.5 hours. Six replicates of 20 eggs were used for each of
the four different ages.
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Table 4. Effect of temperature and three relative humidities on mean development time and
development rate of eggs of Mamestra configurata.
(x — did not complete development to eclosion)
(** means of 3 replicates of 30 eggs per replicate)
% DEVELOPMENT / HR.
Temperature, RH and embryogenesis in Mamestra configurata
267
Figure 1. The developmental curve -A; and the velocity line-B; at 0% RH for embryogenesis of eggs of Mamestra configurata.
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% DEVELOPMENT / HR.
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Figure 2. The developmental curve-A; and the velocity line-B; at 60% RH for embryogenesis of Mamestra configuraia.
HOURS
% DEVELOPMENT /HR.
Temperature, RH and embryogenesis in Mamestra configurata
269
Figure 3. The developmental curve-A; and the velocity line-B; at 98% RH for embryogenesis of eggs of Mamestra configurata.
Quaest. Ent., 1979, 15 (2)
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The eggs were collected daily when less than two hours old, divided into groups of six to form
replicates, and the remaining vials were used as a check to determine natural mortality. Eggs that were
to be used later were left at 20.0 ± 0.5 C and approximately 60% RH until they reached the desired age.
When this occurred the eggs were placed in a 160 mm desiccator located in an incubator set at 35.0 C.
The humidity in the desiccator was maintained at approximately 60% with a saturated salt solution of
NH4NO3 (Winston and Bates, 1960).
The temperature in the incubator never varied more than 0.5 C. The temperature in the desiccator
and within the vials was checked periodically and showed little variance (0.25 C) from that of the
incubator. The incubator was opened only when it was necessary to place or remove vials.
Once the treatment was completed, the eggs in the vials were returned to 20.0 ± 0.5 C and 60% RH,
and the tops of the vials were covered with plastic screening to prevent larval escape. The numbers
hatching were recorded twice daily. Abbot’s formula (Abbot, 1925) was used to determine the net
percent mortality. Natural mortality was considered to be the highest mortality that was not a function
of the treatment.
Variance analysis was used to determine the amount of variance caused by replication and error.
Results. The results of this experiment are summarized in Tables 5 to 8.
The mean time in hours required to produce 50% mortality were 34.45 ± 1.4, 33.19 ± 3.49,
32.8 ± 1.33 and 35.17 ± 1.11 for 3 hour, 24 hour, 48 hour and 96 hour old eggs respectively; and to
produce 95% mortality were 51.27 ± 3.55, 48.31 ± 3.34, 50.03 ± 3.4 and 54.95 ± 4.14 for 3 hour, 24
hour, 48 hour and 96 hour old eggs respectively. From this data it can be concluded that exposures of
greater than 37 hours to a temperature of 35.0 C will result in 50% or greater mortality and exposures of
60 hours will result in 95% or greater mortality, to eggs of Mamestra configurata regardless of their age.
Variance analysis and the resulting F-values showed that length of exposure was highly significant in
deterining mortality (Tables 5 to 8).
Effects of constant exposure to a temperature of 5.0 C on development of eggs of different ages. -
When we began this research, we found that Mamestra configurata was difficult to rear continuously in
the laboratory. Part of the problem was feeding the large number of larvae produced. The main purpose
of this experiment was to determine the length of time eggs of Mamestra configurata could be stored at
5.0 C without causing excessive mortality. The temperature of 5.0 C was chosen for two reasons: (1)
under natural conditions, eggs could be exposed to this temperature or lower (Average minimum
temperature for July in rape growing areas in Central Alberta are: Stettler 10.2 C, Vermilion 9.6 C,
Edmonton 9.1 C and Calgary 9.5 C (Philip, in note to Heming). Therefore, it is doubtful whether eggs
would be exposed very long to temperatures of 5.0 C), and (2) 5.0 C is below the
developmental-hatching threshold (8.5 C), but above the developmental threshold (between 0.0 and
2.0 C). A by-product of this research was to determine the length of exposure to 5.0 C required to
produce 50% and 95% mortality.
Methods. Procedures used in this experiment were identical to those of the experiment with a
constant temperature of 35.0 C , with the following exceptions: (1) the exposure was to 5 C; (2)
NaBr.2H20 (Winston and Bates, 1960) was used to maintain approximately 60% RH, and (3) exposure
times of 3, 45, 67, 101, 151, 227, 340, 510 and 765 hours were used. Additional times of 83, 189,
273 hours were used for eggs 48 hours old when initial mortality was high.
Probit anaysis was used to transform sigmoidal dosage mortality curves into straight lines (Bliss,
1935). Variance analysis was used to determine the sources of variation and the effects of the treatment.
Results. Results of this experiment are summarized in Tables 9 to 12 and Figures 4 to 7.
Generally, resistance to cold appeared to increase as the age of the egg increased. Initial mortality
occurred after 67 hours with 3 hour old eggs, after 151 hours with 24 hour old eggs, after 227 hours with
Temperature, RH and embryogenesis in Mamestra configurata
271
Table 5. Effect of continuous exposure to 35.0 C on three hour old eggs of Mamestra configurata.
Treatment F-value 433.06 (F at 1% 4.43) with 4 DF
Between Treatment F-value 1.72 (F at 5% 2.71) with 20 DF
Mean number of hours of exposure at 35 C required to produce 50%amd 95% mortality are:
34.45 ± 1.40 and 51.27 ± 3.55 respectively. Variance analysis showed that variation in per-
centage mortality was a function of treatment.
Table 6. Effect of continuous exposure to 35.0 C on 24 hour old eggs of Mamestra configurata.
Treatment F-value 404.1 (F at 1% level 4.43) with 4 DF
Between Treatment F-value 0.77 (F at 5% level 2.71) with 20 DF
Mean number of hours of exposure to 35 C required to produce 50% and 95% mortality are
33.19 ± 3.49 and 48.31 ± 3.34 respectively. Variance analysis showed that variation in percentage
mortality was a function of treatment.
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Table 7. Effect of continuous exposure to 35.0 C on 48 hour old eggs of Mamestra configurata.
Treatment F-value 805 (F at 1% level 4.43) with 4 DF
Between treatment F-value 0.82 (F at 5% level 2.71) with 20 DF
Mean number of hours of exposure to 35 C required to produce 50% and 95% mortality are
32.8 ± 1.33 and 50.03 ± 3.4 respectively. Variance analysis showed that variation in percentage
was a function of treatment.
Table 8. Effect of continuous exposure to 35.0 C on 96 hour old eggs of Mamestra configurata.
Treatment F-value 426.78 (F at 1% level 4.43) with 4 DF
Between tratment F-value 0.05 (F at 5% level 2.71) with 20 DF
Mean number of hours of exposure to 35 C required to produce 50% and 95% mortality are
35.17 ± 1.11 and 54.95 ± 4.14 hours respectively. Variance analysis showed that variation in
percentage mortality was a function of treatment.
Temperature, RH and embryogenesis in Mamestra configurata
273
Table 9. Effect of continuous exposure to 5.0 C on three hour old eggs of Mamestra configurata.
Treatment F-value 733.5 (F at 1% level 3.7) with 7 DF
Between treatments 5.13 (F at 5% level 2.42) with 35 DF
Mean number of hours of exposure to 5 C required to produce 50% and 95% mortality are
193.06 ± 5.09 and 252.93 ± 13.14 respectively. Variance analysis showed that variation in
percentage mortality was a function of treatment.
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Table 1 1. Effect of continuous exposure to 5.0 C on 48 hour old eggs of Mamestra configurata.
Treatment F-value 445.7 (F at 1% level 3.51) with 8 DF
Between treatments F-value 2.19 (F at 5% level 2.18) with 40 DF
Mean number of hours of exposure to 5 C required to produce 50% and 95% mortality are
271.64 ± 8.26 and 389.05 ± 17.82 respectively. Variance analysis showed that variation in
percentage mortality was a function of treatment.
Table 12. Effect of continuous exposure to 5.0 C on 96 hour old eggs of Mamestra configurata.
Treatment F-value 81 1.8 (F at 1% level 3.51) with 8 DF
Between treatments F-value 1.04 (F at 5% level 2.18) with 40 DF
Mean number of hours of exposure to 5 C required to produce .50% and 95% mortality are
337.29 ± 17.52 and 633.87 ± 55.77 respectively. Variance analysis showed that variation in
percentage mortality was a function of treatment.
Temperature, RH and embryogenesis in Mamestra configurata
275
Figure 4. Probit regression line showing the effects of various dosages of 5.0 C on 3 hour old eggs of Mamestra configurata.
Figure 5. Probit regression line showing the effects of various dosages of 5.0 C on 24 old eggs of Mamestra configurata.
Quaest. Ent., 1979, 15 (2)
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igression line, showing the effects of various dosages of 5.0 C on 48 hour old eggs of Mamestra configurata.
Figure 7. Probit regression line, showing the effects of various dosages of 5.0 C on 96 hour old eggs of Mamestra configurata.
Temperature, RH and embryogenesis in Mamestra configurata
111
48 hour old eggs and after 151 hours with 96 hour old eggs. However, after 227 hours the 48 hour old
eggs suffered 11.4% mortality while, after time period, the 96 hour old eggs had only 9.1% mortality.
The mortality difference is even greater after 340 hours exposure with 79.8% and 53.2% respectively for
the 48 hour and 96 hour old eggs. Probably, there is a developmental age where exposure to 5.0 C is
more critical than at other stages.
The increased resistance to cold is better illustrated when LD 50’s and LD 95’s are compared. Eggs 3
hours old had an LD 50 and an LD 95 of respectively 90.57 ± 2.32 hours and 118.85 ± 5.87 hours.
These values for 24 hour old eggs were respectively 193.06 ± 5.09 hours and 252.93 ± 13.14 hours.
Eggs aged 48 hours and 96 hours accrued 50% mortality after 271.64 ± 8.26 hours and 337.29 ± 17.52.
hours and 95% mortality after 389.05 ± 17.82 hours and 633.87 ± 55.77 hours.
The F-values between error and treatment were highly significant (Tables 9-12), but variance
between treatments was significant only for the 24 hour group.
Effects of alternating temperature on development
Alternating temperatures are used in biological temperature research to determine effects that
cannot be ascertained using constant temperature alone. For example, under alternating temperatures,
eggs of Oncopeltus fasciatus (Heteroptera) can complete embryogenesis at a mean temperature several
degrees lower than the lowest constant temperature at which complete embryogenesis can occur (Lin et
al. 1954).
During development, organisms are often subjected to extreme temperatures that would be fatal to
them if they were maintained over long periods. Knowledge of the effects of alternating temperature on
mortality and rate of development aids us in understanding naturally occurring temperatures effects on
development.
Two experiments were conducted on eggs of Mamestra configurata to determine the effects of
varying daily exposures to extreme temperatures on development rate. The first experiment was
designed to determine the effects of daily exposure to 35.0 C on rate of development; the second had
5.0 C as the experimental temperature. In both experiments, the alternate temperature used was 20.0 C.
Effects of daily alternation between temperatures of 20.0 C and 35.0 C on eggs of different ages. -
The main purpose of this experiment was to determine if eggs of the experimental population could
withstand daily exposure to 35.0 C. In Alberta, the ovipositional period of females of Mamestra
configurata extends from late June to mid-July so that its eggs would rarely be exposed to a temperature
of 35.0 C. However, since Mamestra configurata is partially bi-voltine (i.e., it lacks either an obligatory
or a facultative diapause in some members of its populations), its eggs may be expected to show partial
resistance to daily temperatures of 35.0 C.
The other purpose of this experiment was to determine if alternation of temperature influenced
development rate. Alternation of a high temperature with a moderate temperature often results in a
decrease in the development rate of an insect compared with its performance at a constant temperature
equal to the mean of the alternating temperatures (Johnson, 1940).
Methods. Eggs were collected from a culture reared at 20.0 ± 0.5 C and approximately 60% RH,
divided into groups of 10 and each group was placed in a 1 mm cap vial.
Four groups of ten individuals were used at each of 13 experimental exposures (from 1 to 22.5 hours
daily at 35.0 C; see Table 13 to 16 for details). Eggs of four different ages (3, 24, 48, and 96 hours) were
used to determine the effect of age on tolerance to 35.0 C.
Eggs from any one collection period were divided into as many groups of ten as their number allowed,
and the resulting groups were subdivided into units of four. Each of these units was then given daily
exposure of 35 C. One vial from each collection was left at 20.0 ± 0.5 C to determine fertility.
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During exposure, the vials were placed on a platform in a desiccator. A saturated salt solution of
NH4NO3 was used to produce an RH of appoximately 60% (Winston and Bates, 1960). First exposure
to 35.0 ± 0.5 C was made immediately after the eggs had reached the desired age and was repeated at
the same time each subsequent day until completion of the experiment. After exposure, the eggs were
returned to 20.0 ± 0.5 C for the remainder of the day.
Eggs were observed daily until they reached the black head capsule stage at which time observations
were made at two hour intervals. The number hatching and the total length of the development period
for each egg was then recorded.
The mean hatching time for each exposure and the mean temperature for each group was
determined. This allowed a comparison to be made between mean development time of eggs maintained
at constant temperatures and those maintained under alternating temperatures.
Fry and Surber’s (1971) experiment on the effects of exposure to 35.0 C and 40% RH on eggs of
Estigmene acrea was used as a model.
Results. Results of the various treatments are summarized in Tables 13 to 16. In general, daily
exposures of one hour did not result in significant change in development time (Daily exposures of 2.9
hours or greater resulted in a significant change in development time.) regardless of the number of
exposures.
Table 13. Effect of daily exposure to 35.0 C on development in three hour old eggs of
Mamestra configurata.
Treatment F-value 1 14.08 (F at 1% level 2.32) with 9 DF
*means followed by the same letter (A or B) are not significantly different (based on Duncan’s
New Multiple Range Test).
**Not significant at the 5% level using T-tests. Variance analysis showed that the difference
noticed in development times was a product of the various treatments.
Temperature, RH and embryogenesis in Mamestra configurata
219
Table 14. Effects of daily exposure to 35.0 C on development in 24 hour old eggs of
Mamestra configurata.
Treatment F-value 174.89 (F at 1% level 2.32) with 10 DF
*means followed by the same letter (A) are not significantly different (based on Duncan’s
New Multiple Range Test).
**not significant at the 5% level using T-tests. Variance analysis showed that the difference
noticed in development times was a product of the various treatments.
Duncan’s New Multiple Range Test was used to determine if individual treatments within each
experimental group varied significantly from each other. The difference between exposures of one hour
and 1.3 hour daily to 35.0 C was not significant in the 3 hour old groups. Non-significant differences
were shown also between daily exposures to 4.8 and 6.2 hours for both the 3 hour and 24 hour old
groups. Eggs 48 hours old showed no significant difference in development time for daily treatments of
2.9, 3.7, and 4.8 hours, nor between treatments of 4.8 and 6.2 hours and 1.7 and 2.2 hours. Eggs 96
hours old showed no significant difference in development time for daily treatments of 1 .0, 1 .3, 1 .7 or 2.2
hours.' No significant variation occurred between treatments of 3.7 and 4.8 or 8.0 and 10.4 hours.
Variance analysis was also used to determine if the variation noted in development times was due to
treatment. The resulting values showed that the treatments were highly significant in this.
A comparison between development times observed under constant temperature and under equivalent
alternating temperatures showed that development at constant temperatures of 22.5 and 25.0 C occurred
more rapidly than at an equivalent temperature produced by daily alternation between 35.0 C and
20.0 C but more slowly at a constant temperature of 20.0 C (Table 17).
Our results were difficult to reconcile with those of Fry and Surber (1971). A single exposure to
35.0 C and 40% RH for 20 hours, resulted in only a 1.6% hatch of salt marsh caterpillar , Estimene
acrea, eggs whereas all eggs of Mamestra configurata hatched that were given a similar exposure to
35.0 C and 60% RH for 22.8 hours. However, three exposures of 16 hours to 35.0 C and 40% RH
resulted in 65.7% hatch for salt marsh caterpillar eggs while three similar exposures of 17.5 hours to
35.0 C and 60% RH resulted in only 27.5% hatch for Mamestra configurata eggs.
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Table 15. Effects of daily exposure to 35.0 C on development of 48 hour old eggs of
Mamestra configurata.
Treatment F-value 105.36 (F at 1% level 2.32) with 11 DF
*means followed by the same letter (A,B,C) are not significantly different (based on Duncan’s
New Multiple Range Test).
**Not significant at the 5% level using T-test. Variance analysis showed that the difference
noticed in development times was a product of the various treatments.
Effects of daily alternation between temperatures of 5.0 C and 20.0 C on eggs of different ages. - In
the province of Alberta, Mamestra configurata has been recorded from almost all rapejrowing areas, a
territory extending from south of Fethbridge to north of Keg River, a distance of about 966 km (Philip,
pers comm.). Over much of this range, temperatures during the ovipositional period (June to July) could
fall to 5.0 C or lower (i.e. below the developmental-hatching threshold, 8.5 C) for short periods
(Canadian Department of Transport Meteorological Records, 1971-1974). The primary purpose of this
experiment was to determine the effects of a varying exposure to 5.0 C on development time of eggs. A
secondary purpose was to determine if alternation of temperatures would produce an acceleration in
development over those of eggs maintained at a constant equivalent temperature.
Methods. Methods and materials used in this experiment were identical to those of the previous one
except that (1) daily exposure was to 5 C, (2) the saturated salt solution used to maintain 60% RH was
Na2Cr207.H20, and (3) 25 rather than ten individuals were used in experiments involving three hour old
eggs.
Results. The effects of the various treatments are summarized in Tables 18 to 21.
T-tests were used to determine the significance of the effect oftreatment on development time. In all
cases, the treatments produced a significant delay in development as compared with that of controls.
Variance analysis was used to determine if the difference noted in development times was a function of
treatment. In all cases, the difference noted between development times was highly significant at the 1%
confidence level.
Duncan’s New Multiple Range Test was used to determine if the individual treatments produced a
significant difference in development times within individual test groups. In all cases, this test showed
that the treatments differed at the 5% confidence level.
Temperature, RH and embryogenesis in Mamestra configurata
281
When the rates of development of eggs under constant temperature and under its alternate
temperature equivalent were compared, those under the alternating temperatures developed more
rapidly (Table 22).
Table 16. Effects on daily exposure to 35.0 C on development of 96 hour old eggs of Mamestra
configurata.
Treatment F-value 56.39 (F at 1% level 2.32) with 12 DF
*means followed by the same letter (A,B,C) are not significantly different (based on Duncan’s New
Multiple Range Test).
**Not significant at the 5% level using T-tests.
Variance analysis showed that the difference noticed in development times was a product of
the various treatments.
Table 17. Comparison of development rate in hours and in hour-degrees at constant temperature and at
an equivalent alternating temperature (35.0 and 20.0 C) of eggs of Mamestra configurata.
Development time in hours
*Numbers in brackets are equivalent development rates in hour-degrees.
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(continued on page 283)
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283
Table 19. (Continued)
Treatment F-value 63,708 (F at 1% level 2.32) with 1 1 DF
**level of significance comparing mean development times of the combined replicates and the
control group using T-tests.
The difference in development time between treatments was analyzed using variance and
found to be highly significant and, almost completely, a function of treatment.
Table 20. Effects of daily exposure to 5.0 C on development in 48 hour old eggs of Mamestra
configurata.
Treatment F-value 39,499 (F at 1% level 2.32) with 1 1 DF
**level of significance comparing mean development times of thexombined replicates and the
control group using T-tests.
The difference in development time between treatments was analyzed using variance analysis
and was found to be highly significant and, almost completely, a function of treatment.
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Table 21. Effects of daily exposure to 5.0 C on development of 96 hour old eggs of Mamestra
configurata.
Treatment F-value 78,120 (F at 1% level 2.18) with 12 DF
*means followed by the same letter (A) are not significantly different based on Duncan’s New
Multiple Range Test.
**level of significance comparing mean development times of thejcombined replicates and
the control group using T-tests.
The difference in development times between treatments was analysed using variance
analysis and was found to be highly significant and, almost completely, a function of
treatment.
Table 22. Comparison of development rate in hours and in hour degrees at constant temperature and at
an equivalent alternating temperature (5 and 20 C) of eggs of Mamestra configurata.
*Numbers in brackets are equivalent development rates in hour-degrees.
Temperature, RH and embryogenesis in Mamestra configurata
285
DISCUSSION AND CONCLUSIONS
Temperature thresholds for embryogenesis
Study of temperature effects on organisms began when Reamur (1735, see Belehradek, 1930)
recognized that a relationship exists between temperature and the activity of an animal. Since the, these
relationships have been examined by numerous authors and reviewed by Crozier (1926), Belehradek
(1930), Uvarov (1931), Janisch (1932), Howe (1967), Bursell (1974) and several others. The results of-
these studies led to the development of the concept of various temperature thresholds for developmental
stages.
These are two principal types of temperature threshold; low temperature thresholds involving
temperatures too low for certain developmental processes to be completed, and high temperature
thresholds which consider temperatures too high for normal development. Determination of these
thresholds, in embryogenesis, has been made primarily of the first type, probably because of the larger
temperature coefficients existing between the various thresholds at lower temperatures.
Presently, four low temperature thresholds are recognized as affecting embryogenesis. Peairs (1927)
defined the developmental thresold as “the temperature at which, on the descending scale, development
ceases, and at which, on the ascending scale development is initiated”. Johnson (1940) introduced the
hatching threshold and the developmental-hatching threshold which are respectively, the lowest
temperature at which hatching of a fully developed larva can occur, and the lowest temperature at which
complete development from fertilization to eclosion can occur. The hatching-survival threshold was
suggested by Hodson and A1 Rawy (1956) and they chose Allee et al.'s (1949) definition of the “lowest
temperature at which a given stage in the life history can be carried through to completion” for a
definition.
The only high temperature threshold recognized for embryogenesis is the high temperature
equivalent of the developmental-hatching threshold. It has also been suggested that there exists a high
temperature threshold equivalent of the developmental threshold, but Uvarov (1931) questioned whether
this temperature is distinct from the upper lethal temperature. He further stated that if there was an
appreciable difference in temperature between the upper lethal point and the upper developmental
threshold, then there should be a quiescent stage similar to the quiescence observed at temperatures
below the developmental threshold but above the lower lethal point. This stage, referred to as heat
stupor, has been recognized in relation to activity but, as yet has not been described for development.
Theoretically, this stage could exist, although the range between it and the upper lethal temperature
may be so restricted that its discovery will be of limited practical value.
The concept of the developmental threshold of Peairs (1927), has been given numerous names by
various authors in attempts to convey with greater clarity the process that occurs at that particular
temperature. It has been called “the critical point”, “physiological zero” and “the minimum effective
temperature“ (Uvarov, 1931). More recent authors (e.g. Lin et al., 1954) have chosen to ignore the
original definition of this threshold and have used instead Johnson’s (1940) developrnental-hatching
threshold definition as if it were synonymous. The developmental threshold is seldom determined
analytically, but is generally arrived at by extrapolation from temperature and rate of development
curves. This method assumes that the effects of temperature on duration of embryogenesis can best be
represented, mathematically, by a hyperbolic function (Sanderson and Peairs, 1914). This function,
when transformed into its reciprocal, produces a straight line. This line is often called the “velocity line“,
and represents the effects of temperature on rate of development. When plotted on a graph, the point
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where the velocity line intercepts the temperature axis is the hypothetical zero of the hyperbola or the
developmental threshold. This method is used extensively to derive the developmental threshold; for
example it is 1 1.5 C for the corn earworm, Heliothis zea (Boddie), (Luckmann, 1963) and 1 1.0 C for
the European corn borer Ostrinia nubilalis (Hubner) (Matteson and Decker, 1965).
The difficulty in using a straight line to describe rate of development was first pointed out by Krogh
(1914) and later by Shelford (1927) and Peairs (1927). Peairs noted that the development rate deviated
from a straight line near the extremes of the temperature range. Thus, thresholds determined using this
method were higher than the actual threshold.
Johnson (1940) emphasized the need for determining thresholds empirically and the necessity of
recognizing different thresholds for various stages of embryogenesis. He suggested that because there
are distinct stages in embryogenesis, there should also be equally distinct temperature thresholds for
development of these stages. He suggested that both hatching and developmental hatching thresholds be
used. However, with the exception of a study on eggs of the milkweed bug Oncopeltus fasciatus (Dallas)
(which showed them to hatch at a temperature 2.0 C lower than their developmental-hatching threshold
if 90% of the previous embryonic development occurred at 20.0 C (Lin et al., 1954)) most researchers
have chosen to disregard both the hatching and the development-hatching thresholds.
The hatching survivial threshold of Hodson and Al Rawy (1956) has also been ignored by recent
workers, probably because of the extensive amount of time required for rearing immatures through to
adulthood.
The upper temperature developmental limit can be considered to be the upper equivalent of the
developmental-hatching threshold or the highest temperature at which complete development and
eclosion can occur. This upper threshold has been described in representative species of many orders of
insects, e.g. (Collembola) Onychiurus furci/erus (Borner) (Choudhuri, 1960); (Hemiptera) {Geocoris
atricolour Montd. (Dunbar and Bacon, 1927); (Diptera) Phormia regina Meig. (Melvin, 1934);
(Coleopera) Epilachna corrupta Mulsant (Pyenson and Sweetman, 1931); and (Lepidoptera) Telea
polyphemus Cramer (Ludwick and Anderson, 1942). Bursell (1974) listed the upper limit for various
insect species, the highest, 40.0 C , being recorded for Ptinus tectus Boield; the lowest, 28.0 C, for
Tribolium confusum (Herbst), both Coleoptera.
The various temperature thresholds demonstrated for embryogenesis in eggs of Mamestra
configurata show that these are well adapted to Alberta climatic conditions occurring at the time of
oviposition. Some development occurs at temperatures as low as 2.0 C and fully developed eggs can
hatch at temperatures as low as 5.0 C (3.5 C below the developmental-hatching threshold). Eggs can
develop completely and hatch at temperatures ranging from 8.5 C to 30.0 C, temperatures similar to
those recorded by Bailey (1976) for eggs of this insect. However, mortality at the
developmental-hatching threshold (8.5 C) is high (77.8%). Normal temperatures (mean, maximum and
minimum) for the ovipositional period rarely fall outside this developmental range (Department of
Transport Meteorological Records).
Similar high mortality at the development-hatching threshold has been demonstrated for eggs of
Oncopeltus fasciatus by Richards and Suannaksa (1962). It is probable that the larvae that hatched
would not have reached maturity. Their hatching behaviour showed the same anomalies as those
reported by Lin et al. (1954) for nymphs of Oncopeltus fasciatus. They found that even if these nymphs
were tranferred to ideal conditions, very few reached maturity. Larvae hatching under these conditions
appear debilitated and have difficulty moving effectively. It is possible that these larvae have internal
structural defects that result in early death.
The upper developmental-hatching threshold for eggs of Mamestra configurata recorded here
(between 30.0 C and 31.5 C) is comparable to the 32.0 C value given by Bailey (1976) for eggs of this
Temperature, RH and embryogenesis in Mamestra configurata
287
species and resembles that of eggs of the dirmyyNorm, Pseudaletia unipuncta (Howe) (Guppy, 1969). He
found that the rate of development for eggs of this species began to decrease when temperature was
increased above 29.0 C. Eggs of Mamestra configurata probably follow a smilar course of development.
The lack of data supporting this belief probably arises from experimental error. Sample sizes (50
individuals) may have been too small to reveal the small percentage of individuals that might have
hatched at temperatures above 30.0 C. Also, the interval between 30.0 C and the next temperature
(31.5 C) might have been too large. Mortality in insects increases rapidly as temperature increases
above their optimum temperatures (Stinner et al. 1974). Even if complete development is curtailed by
temperatures in excess of 30.0 C, the occurrence of these temperatures in Western Canada during the-
incubation period is rare and would have little effect on development.
Effects of constant temperature and relative humidity on embryogenesis
Research conducted on the effects of temperature on insect embryogenesis can be broadly classed
into two groups. The first treats the effects of constant temperature (often applied at different relative
humidities); the second, the effects of alternating or varying temperatures on embryonic development.
This subject has been reviewed by Sanderson (1910), Uvarov (1931), Janisch (1932), Howe (1967) and
Bursell (1974) . The effects of relative humidity have been reviewed by Buxton (1932) and Ludwick
(1945).
One of the principal reasons that the effects of constant temperature on insect development are
studied is the hope that the resulting developmental curves can be used to predict insect development in
the field. Numerous attempts have been made to derive a general mathematical function or equation
that describes the relationship between temperature and development in insects. These attempts have
been reviewed by Crozier (1926), Uvarov (1931), Janisch (1932), Davidson (1944), and Howe (1967).
The more widely used of these concepts are: day-degrees; thermal summation and summation of
development units.
Improper use of these concepts often leads to grossly inaccurate results. This occurs when fluctuating
temperatures are assessed in terms of a simple average rather than weighted according to the effects of
each individual temperature on development (Bursell, 1974). The inaccuracy of these concepts arises
from the assumption that the best way to represent mathematically the effect of temperature on rate of
development, is through use of a straight line. Near the extremes of the temperature range for a species,
the rate of development deviates from a simple linear relationship. At the lower extreme, the rate of
development declines more slowly; at the upper extreme, more rapidly than would be expected from a
strictly linear relationship. The resulting temperature and rate of development curve is thus closer to
being sigmoidal than linear. However, when the temperature range between the maximum rate of
development and the upper lethal limit is slight, the resulting curve is J-shaped (Howe, 1967).
Relative humidity can apparently have no effect on insect embryogenesis; or it can cause changes in
development time, and influence mortality rates and developmental thresholds. These effects and others
on invertebrates have been reviewed by Buxton (1932) and Ludwick (1945).
The rate of embryogenesis in eggs of Mamestra configurata showed a strong linear relationship with
temperature over the range of temperatures used in this experiment. Deviation from this relationship
occurred only at temperatures of 10.0 C and lower. Complete development and hatching occurred at
temperatures ranging from 8.5 C to 30.0 C.
Low humidity retarded development regardless of temperature. It is possible that eggs of Mamestra
configurata normally absorb from the atmosphere at least a portion of the water used during
development. Extremely low humidity would inhibit this absorption and force development to rely upon
already present reserves, thus slowing down rate of development.
Quaest. Ent., 1979, 15 (2)
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Jones & Heming
Lower temperatures prolonged exposure to low humidities and resulted in increased mortality.
Possible causes for this are (1) death due to desiccation as the eggs had insufficient water to complete
development, (2) weakening of larvae through water loss, making the act of hatching more difficult, and
(3) hardening of the chorion caused by desiccation, inhibiting hatching, or (4) by a combination of these
factors.
The fastest rate of development always occurred at 98% RH followed by 60% RH and 0% RH. This
suggests that eggs of Mamestra configurata absorb water during development and that water may
become the limiting factor during some part of embryogenesis. Water absorption would be faclitated
most in high humidity, and would decrease as humidity declines. It is unfortunate that we failed to
monitor micro-climatic conditions within blossoming rape fields since such data would have faclitated
speculation of this kind.
Effects of exposure to constant temperature and relative humidities of 0%, 60% and 90%) on
developmental rate. - In general, low relative humidity retards rate of development in insect eggs. We
have shown this to be true too for eggs of Mamestra configurata. Regardless of the temperature used,
hatching always occurred first at 98% RH followed by 60% and 0%. Relative humidity appeared to be
more important in the lower temperature range (15.0 to 8.5 C). At 0% RH, hatching was inhibited at
temperatures below 15.0 C. At the developmental-hatching threshold (8.5 C) hatching occurred only at
98% RH.
Bailey (1976) reared eggs of Mamestra configurata at 75% RH and at temperatures of from 6.0 to
36.0 C and found the time for completion of embryogenesis and hatching to decrease from 28 days at
8.0 C to 3 days at 28.0 and 32.0 C. These figures are similar to those recorded here: 36.6 days at 8.5 C
and 98% RH, 3.2 days at 30.0 C and 98% and 3.3 days at 30.0 C and 60% RH (Table 4). These results
agree with those reported for other Lepidoptera (Ludwick and Anderson, 1942).
Evidence from our experiments to suggest that both mechanical and physiological barriers retard and
reduce hatching is that the number of individuals first to eclose at each of the humidities used remained
relatively constant regardless of temperature. Differences of eight to ten hours occurred in eggs exposed
to RH’s of 0% and 98%. This evidence also suggests that the larvae were weakened and thus took longer
to hatch. There is also evidence suggesting a physiological delay. Eggs exposed to 0% RH always
required a longer time to develop brown pigmentation than did those exposed to higher humidities. This
suggests that early stages of development are impaired by lack of access to atmospheric moisture. In
many insects, development is enhanced by absorption of atmospheric moisture through a hydropyle
(Wigglesworth, 1972). Although examination of eggs of Mamestra configurata failed to locate a similar
structure, the numerous aeropyles present in the chorion could act in a similar capacity (Jones, 1979).
The experiments we conducted were not designed to determine the mechanism by which humidity
affects development. Experiments using a greater range of humidities and tests of egg shell tensile
strength would aid in determining which of physiological or mechanical barriers have the greater effect
on mortality and rate of development.
Effects of constant exposure to a temperature of 35.0 C on development of eggs of different ages. -
Constant exposure of eggs to 35.0 C produced approximately 50% mortality after 37 hours and 95%
mortality after 20 hours regardless of egg age. (Tables 5-8). Comparison of the 50% and 95% mortality
times of eggs of different ages demonstrated that older eggs had slightly more tolerance than their
younger counterparts. However, the difference was slight and only much larger sample sizes and a
greater number of exposures would demonstrate if the difference is significant.
Sensitivity to high temperature decreased in eggs of the silkworm Bombyx mori L., after meiosis but
before syngamy (Ostryakova-Varshaver, 1958). The exact cause of death in these eggs at high
temperatures was unknown. Denaturing of proteins, an upset in the balance of one or more metabolic
Temperature, RH and embryogenesis in Mamestra configurata
289
processes leading to accumulation of some toxic product of metabolism more quickly than it can be
removed, desiccation, or starvation have all been suggested as contributing to heat death (Bursell 1974).
It is possible that older eggs of Mamestra configurata, i.e. those 24, 48, and 96 hour old, might have
shown greater resistance to heat if they had been kept at 30.0 C prior to testing them at 35.0 C. For
example, eggs of Drosophila spp., show considerable variation in survival time depending upon the
temperature at which the flies were reared. Rearing at 25.0 C more than doubled their survival time at
33.5 C compared with that of counterparts reared at 15.0 C (Chapman, 1971).
Two types of acclimation have been demonstrated in insects: a longlasting “developmental”
acclimation and a transitory “physiological” one (Bursell, 1974). The first of these depends upon the
temperature at which the insect was raised prior to treatment. Insects raised at higher temperature
require a longer exposure to a particular high temperature to produce mortality than do their
counterparts raised at a lower temperature. This acclimation appears to be permanent and is not
affected by exposure to lower temperatures. Physiological acclimation, is readly reversible, its
effectiveness being a function of both the temperature and length of exposure experienced prior to
exposure to the experimental temperature (Bursell, 1974). For example, in one insect, the chalcid
parasite Dahlbominus fuscipennis (Zett.) Baldwin and Riodin (1956) {in Bursell, 1974) found that the
greatest amount of acclimation occurred after two hours exposure and declined to insignificant levels in
the next 12 hours.
Effects of constant exposure to a temperature of 5.0 C on development of eggs of different ages. -
Constant exposure of eggs of Mamestra configurata to 5.0 C had varying results (Tables 9-12).
Resistance to cold increased with increasing age, suggesting that early stages of embryogenesis are more
sensitive to low temperature. Thus, such stages may require daily exposure to favourable temperature
before they can complete development (Lin et al., 1954). Two day old eggs of the cabbage looper,
Trichoplusia ni exposed to 1 1.0 C showed increased mortality compared to that of one or three day old
eggs (Kishabo and Henneberry, 1966).
Death of embryos at low tempeature may result from structural abnomality. For example, some
embryos of Bombyx mori, stored at low temperatures, showed the following abnormalities; the amniotic
cavity was larger than normal and was broken in some individuals. Others showed incomplete dorsal
closure, resulting in parts of the alimentary canal being left outside the body (Totani, 1960). It thus
appears that constant exposure to low temperature can disrupt metabolism and subsequent development
of the embryo.
Another cause of death caused by continuous exposure to low temperature may be a lack of sufficient
food reserves to allow for complete development once the eggs are returned to a favourable temperature
(Richards, 1964). This does not appear to be the cause of death for three hour old eggs of Mamestra
configurata since none of these showed any deposition of pigment (a sign that they had not developed
significantly at the end of treatment). As in the previous experiment different results would have
occurred if the eggs had been pre-conditioned prior to exposure to the experimental temperature.
Effects of alternating temperature on development
Research on the effects of alternating temperature on insect embryogenesis are of the following types:
(1) a low temperature is alternated with a medial temperature (medial temperatures are a range of
temperatures for an insect species, where the rate of development most closely approaches a linear
relationship with temperature), (2) a high is alternated with a medial, (3) two medial temperatures are
alternated with each other, or (4) a temperature series based on field conditions is used to simulate
natural conditions.
These effects have been reviewed by Uvarov (1931), Howe (1967), Wigglesworth (1972) and others.
Quaest. Ent., 1979, 15 (2)
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Jones & Heming
In general, such treatment can (1) accelerate development rate, (2) decrease development rate, (3)
influence the developmental-hatching threshold, (4) influence mortality, or (5) have no effects.
Exposure to alternating temperatures appeared to either increase or decrease development rate in
eggs of Mamestra configurata depending on the situation. Alternation between 5.0 C and 20.0 C
apparently accelerated development compared with that of eggs experiencing an equivalent constant
temperature whereas alternation between 20.0 C and 35.0 C seemed to decrease development rate.
Effect of daily alternation between temperatures of 20.0 C and 35.0 C on eggs of different ages. -
Eggs of Mamestra configurata demonstrated an increasing tolerance to longer daily exposures to 35 0 C
as they matured (Tables 13-16). Eggs two days old at the beginning of the treatments withstood longer
daily exposures than did younger eggs. However, this does not mean that they showed a greater overall
resistance to temperatures of 35.0 C. When the total time spent at 35.0 C is compared, it shows that
regardless of age no eggs hatched from any group exposed to more than 55 hours. This data, combined
with that of the experiment on constant exposure to 35.0 C (Tables 5-8) suggest that the effects of
35.0 C may be cumulative. If this is true, it is unlikely that any one stage of embryogenesis is the most
susceptible. Rather, it suggests that the whole of metabolism is disrupted, perhaps leading to a build up
of toxic materials which, in turn, causes death (Bursell, 1974).
The time required for complete development was greater under alternating than under constant
temperature (Table 17). This agrees with similar results recorded for eggs of the Japanese beetle,
Popillia japonica by Ludwick (1938), for those of Drosophila melanogaster Meigen by Ludwick and
Cable (1933) and for those of other fruit flies by Messenger and Flitters (1958). The reason for this is
the sigmoidal nature of the temprature and rate of development curves. Above the optimum temperature
(30.0 C for eggs of Mamestra configurata), rate of development begins to decline, this decline increasing
rapidly as temperature increases. Thus, the actual rate of development at 35.0 C is probably similar to
that occurring at 30.0 C or lower. This means that the time spent at 35.0 C is equivalant to an identical
time spent at 30.0 C and that this should be calculated on this basis rather than on a hypothetical linear
relationship supposedly existing between temperature and development curves. If the actual sigmoidal
relationship is used, neither a retardation nor acceleration of development would appear to occur.
Eggs of Mamestra configurata react similarily to those of the saltmarsh caterpillar, Estigmene acrea,
when exposed to 35.0 C (Fry and Surber, 1971). This suggests that eggs of Mamestra configurata are
sufficiently tolerant to survive temperatures found in the south-western United States if the ovipositional
period of Mamestra configurata there is similar to that of Estigmene acrea.
Effects of daily alternation between temperatures of 5.0 C and 20.0 C on eggs of different ages. -
The effects of daily exposure to 5.0 C are more applicable to conditions found in Alberta. This
experiment showed that eggs of Mamestra configurata are well adjusted to Alberta climatic conditions.
Temperatures of 5.0 C, when alternated with a favourable temperature (20.0 C), do not have an
appreciable effect on mortality (Tables 18-21). The only instance of high mortality was when eggs were
exposed to 5.0 C for more than 22 hours daily. In these, hatching occurred only in the 96 hour group
(Table 21) even though development to the black head capsule stage occurred in all eggs tested. Failure
of these eggs to hatch was probably because the larvae within were debilitated by long exposure to 5.0 C.
These results compare favourably with those of Lin et al. (1954) who found that as little as one hour per
day spent at a favourable temperature allowed complete development of eggs of Oncopeltus fasciatus.
Time required for development was shorter under alternating than under constant temperature
(Table 22). Similar results have been recorded for eggs of the noctuids Agrotis orthogonia and
Chorizagrotis auxiliaris (Cook, 1927). There are two reasons for this apparent acceleration: (1) some
development is taking place at 5.0 C (see Ms p 13), and (2) due to the sigmoidal nature of the
temperature-rate of development curve, lower temperatures (5.0 C) contribute less to total development
Temperature, RH and embryogenesis in Mamestra configurata
291
than do those above the mean (Johnson, 1940).
Evolutionary and practical considerations
It is probable that the species Mamestra configurata originated further south and has gradually
expanded its range northward. This would explain the existence of the partial second generation which
still occurs in Alberta. Eggs of Mamestra configurata also show some resistance to temperatures that
they are unlikely to experience in Alberta.
Results of this study, hopefully, will be used to remove some of the guesswork in making larval
surveys of this insect. Even working with only mean, minimum and maximum temperatures, it will be
possible to estimate hatching with some accuracy. Proper timing of the survey will allow for
implementation of a more efficient control program. Up until now, control was not usually begun until
the larvae had reached at least the fourth instar and caused considerable damage. Control measures
directed against earlier instars will result in less damage occurring and in better control.
Further investigation should be conducted to determine if 30.0 C is actually the upper
development-hatehing threshold. Larvae which survived either constant high or low temperatures during
embryogenesis should be reared to adulthood to determine what, if any, effects these treatments have on
larvae, pupae, and adults.
The effects of humidity should be studied in greater detail to find out the mechanism or mechanisms
by which development is affected. A possible starting point would be to determine if eggs of Mamestra
configurata absorb moisture and if so to find out whether it is an active or a passive process. Additional
field research is necessary to determine the relationships existing between air temperature, relative
humidity, and micro-climate in rape fields and the effects of these on embryonic and larval development
in Mamestra configurata. Predictions based only on general air temperature and relative humidity data
may not be as reliable (Bursell, 1974).
ACKNOWLEDGEMENTS
This paper was extracted from an MSe thesis submitted to the Department of Entomology at the
University of Alberta (Jones, 1977). The work was begun in the facilities of the Crop Entomology
Section, Research Station, Canada Agriculture, Lethbridge, Alberta and finished in those of the Plant
Industry Laboratory, Alberta Agrieulture, Edmonton. The staffs of both of these institutions are
thanked for their patience, assistance and advice.
Dr. Barbara Chernick, Department of Zoology, University of Alberta, provided advice and criticism
on matters statistical and Drs. D.A. Craig, Department of Entomology, and G.E. Swailes of the
Lethbridge laboratory, trenchant criticism of an earlier draft of this manuscript. H.G. Philip of the Plant
Industry Laboratory reviewed the final draft and pointed out several minor errors as well as suggesting
ideas for additional research. Dr. B.S. Heming, provided guidance throughout the work and prepared the
manuscript for publication. The manuscript was typed by Ticia Brayton. Thanks to all of these
individuals for their help.
Financial support from AART Grant No. 55-28158 to the late Dr. Brian Hocking of the Department
of Entomology is gratefully acknowledged.
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Jones & Heming
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Shelford V.E. 1927. An experimental investigation of the relation of the codling moth to weather and
climate. Illinois Natural History Survey Bulletin 16: 307-440.
Stinner R.E., A.P. Gutierrez and G.D. Butler Jr. 1974. An algorithm for temperature dependant growth
rate simulation. Canadian Entomologist 106(5): 519-524.
Struble D.L., M. Jacobson, N. Green and J.D. Warthen. 1975. Bertha armyworm (Lepidoptera:
Noctuidae). Detection of a sex pheromone and the stimulatory effect of some synthetic chemicals.
Canadian Entomologist 107: 355-359.
Totani K. 1960. Histological observation on abnormal embryos in the silkworm. Research Report of the
Faculty of Textile and Sericulture, Shinshu University 10: 1 18-123.
Uvarov B.P. 1931. Insects and climate. Transactions of the Royal Entomological Society of London
79(1): 1-247.
Wigglesworth V.B. 1972. The Principles of Insect Physiology. Methuen and Co. Seventh Ed. revised
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Winston P.W. and D.H. Bates. 1960. Saturated solutions for the control of humidity in biological
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?
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QL
‘101
uaestiones^
Entomologicde
A periodical record of ent’omolo9icol inrestigafions,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.
VOLUME 15
NUMBER 3
JULY 1979
QUAESTIONES ENTOMOLOGICAE
ISSN 0033-5037
A periodical record of entomological investigation published at the Department of Entomology,
University of Alberta, Edmonton, Alberta.
Volume 15 Number 3 July 1979
CONTENTS
Guest Editorial 295
Kuster -Comparative structure of compound eyes of Cicindelidae and Carabide (Coleoptera):
Evolution of scotopy and photopy 297
Wong - Insect damage to old oak beams at Lower Fort Garry, Manitoba 335
Shorthouse - Observations on the snow scorpionfly Boreus brumalis Fitch
(Boreidae: Mecoptera) in Sudbury, Ontario 341
Straneo - Notes about classification of the South American Pterostichini with a key for
determination of subtribes, genera and subgenera (Coloptera: Carabidae) 345
Kurtak - Food of black fly larvae (Diptera: Simulidae): seasonal changes in gut contents and
suspended material at several sites in a single watershed 357
Guest Editorial - New Tools in Publishing
Although the acting editor of Quaestiones Entomologicae announced in his editorial “To compute or
not compute” {Quaestiones Entomologicae Vol. 15(1): 119) that Quaestiones Entomologicae is now
produced by the computer, you may not be aware that techniques used to produce this journal are among
the more innovative in the publishing industry. Since Quaestiones Entomologicae was the first
periodical at The University of Alberta to change to the new system, I would like to give an explanation
of what is involved.
The traditional approach to publishing is fairly costly, due to the number of times a document must
be typed. A manuscript, for example, may be typed several times before the author is satisfied.
Reviewers may suggest further changes; and then it must be typed again for submission to a journal.
Once the manuscript is received by the publisher, the cycle starts over again. Although Quaestiones
Entomologicae had previously been prepared by a typewriter which produced high quality copy, it was
not much ‘smarter’ than the average typewriter. The manuscript was typed, page numbers applied, and
then it was returned to the author to be proofread. At this point, further changes or corrections had to be
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Clearly, the possiblility for error increased each time a document was reworked. This meant that
proofreading was necessary at each stage, making the entire job quite tedious.
The decision to turn to the computer is often prompted by the rumor that ‘the computer does it
automatically’. Anyone who is apprehensive about this statement should be — the amount of time and
effort required to make a system ‘automatic’ can be considerable. However, the ultimate benefit lies in
the computer’s power to eliminate repetitive tasks. For a manuscript, entire pages need not be retyped to
accommodate extra words or editorial changes and repositioning of each page is under program control.
296
A text formatting program reads material that has been entered into the computer from terminals or
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italicize text, or store words for the index. The program decides how much text will fit on a line, and
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entries are sorted alphabetically and printed.
If text is subsequently altered or edited, lines or paragraphs may fall on a different page. The
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the computer, it never needs to be entirely retyped. Corrections or additions are made where necessary;
the program makes necessary adjustments to the format of pages.
In a journal such as Quaestiones Entomologicae, one of the greatest benefits is the automatic
indexing. At the end of the year, when the entire volume has been entered into the computer, the index
will be complete. The program will process all the manuscripts as one, and compile a comprehensive
index.
Quaestiones Entomologicae is prepared using the TEXTFORM® program, which was implemented
by a group in the Department of Computing Services at the University of Alberta. In 1975, when plans
for TEXTFORM began, several text formatting programs were already available. Most, however,
seemed inflexible, or were tailored specifically for typesetting. TEXTFORM was designed to format
documents on a variety of devices, without changes to the instructions used in the text. Documents
ranging from short reports to large scholarly texts are now produced on computer printers, plotting
devices, and phototypesetters. The computing and typesetting experience of the users varies from novice
to sophisticated.
As an author, you may feel that the new procedures will have little effect on you. However, you can
be of assistance if you are near a computing centre and are able to supply to Quaestiones Entomologicae
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manner. As long as the tape is 9-track, containing alphanumeric data coded in EBCDIC or ASCII, or a
7-track tape coded in BCD, it can be read at this computing centre. Be sure to indicate “DO NOT
X-RAY” on the outside of the package, if you send a magnetic tape through the mail.
Further developments may also affect the traditional publishing process. Technology is now available
to allow computers in Canada, the United States, and eventually other countries throughout the world,
to communicate via telephone lines. If you are near a computing centre which is taking part in these
developments, you may be able to access the computer at this university to check the final copy of your
manuscript, rather than waiting for it to be sent through the mail.
Procedures in the publishing industry have been relatively constant in the past several hundred years.
Changes are now taking place which will have a dramatic effect on information sharing. From seemingly
remote locations, we may be able to browse through publications housed in other centres.
When the decision was made, converting to the computer may have been a drastic step for
Quaestiones Entomologicae. However, if present publishing trends continue (as they undoubtly will) the
recent change at Quaestiones Entomologicae will not be the last.
Debra Reinhart
Publications Consultant
Department of Printing Services
University of Alberta
Edmonton, Alberta, Canada
COMPARATIVE STRUCTURE OF COMPOUND EYES OF CICINDELIDAE AND CARABIDAE
(COLEOPTERA): EVOLUTION OF SCOTOPY AND PHOTOPY
J.E. KUSTER^
Department of Entomology
University of Alberta
Edmonton, Alberta, Canada
T6G 2E3
Qiiaestiones Entomologicae
15 297-334 1979
Compound eyes of males of Amblycheila schwarzi Horn, Omus californicus californicus Horn,
Megacephala Carolina mexicana Gray, and Cicindela tranquebarica Herbst, North American
Cicindelidae, were examined by light and scanning electron microscopy. Intergeneric statistical
analyses were made using data from visual field areas and from measurements of eye structures.
Comparisons based on eye size showed two groups: small eyes, noctural A. schwarzi and noctural
O. californicus; and large eyes, crepuscular M. Carolina and diurnal C. tranquebarica adults. Three
categories for probable eye function were shown: scotopic A, A. schwarzi and M. Carolina; scotopic B.
O. californicus; and photopic, C. tranquebarica adults. Photopic eyes also occur in these other
cicindelids examined: Cicindela belfragei Salle, Cicindela limbata nympha Casey, Cicindela limbalis
Klug, Cicindela repanda repanda Dejean, and Cicindela longilabris Say. However, eyes of crepuscular
adults of Cicindela lepida Dejean are scotopic A, although these beetles are in the large eye group. The
plesiotypic character state of eye structure and function in cicindelid adults is scotopic A; the apotypic
state is photopic. C. lepida adults have secondarily evolved scotopic A eyes.
Cicindelid eye structure and probable function was compared with that of two representatives of
their sister family, the Carabidae. Adult nocturnal Pterostichus melanarius Illiger are small-eyed and
in the scotopic B functional category; diurnal Elaphrus americanus Dejean are large-eyed and photopic.
It is concluded that scotopy and photopy have evolved through parallelism in these sister taxa.
All beetle eyes examined are eucone and have a “subcorneal layer” between corneal lenses and
crystalline cones. They have a distal rhabdomere composed of microvilli from only retinula cell seven,
a more proximal, rectangular fused rhabdom formed from six retinula cells, and a basal eighth
retinula cell with a rhabdomere. Large bulbous eyes of diurnal and crepuscular beetles have
interfacetal mechanoreceptors.
Les yeux composees des males de quatre especes nord-americaines de Cicindelidae, Amblycheila schwarzi W. Horn. Omus
californicus californicus W. Horn, Megacephala Carolina mexicana Gray et Cicindela tranquebarica Herbst, sont etudies au
microscope optique et au microscope a balayage electronique. Les aires de champ visuel ainsi que differentes mesures structurales
des yeux sont comparees statistiquement entre les genres. L’analyse de la grosseur des yeux revele deux groupes; pet its yeux chez les
especes nocturnes A. schwarzi et O. californicus, et grands yeux chex I’espece crepusculaire M. Carolina et chez I’esp'ece diurne C.
tranquebarica. Trois categories sont mises en evidence quant a la fonction probable des yeux: yeux scotopiques A chez A. schwarzi et
M. Carolina, yeux scotopiques B chez O. californicus, et yeux photopiques chez C. tranquebarica. C. belfragei, Salle. C. limbalis
Klug, C. repanda repanda Dejean, C. longilabris Say et C. limbata nympha Casey ont aussi des yeux photopiques. Cependant les
adultes de I’espece crepusculaire C. lepida Dejean ont des yeux scotopiques A, bien qu’ils se classent parmi le groupe a grands yeux.
La structure et la fonction scotopique A consistuent la condition plesiotypique des yeux des Cicindelidae adultes, alors que le type
photopique est la condition apotypique. Le type scotopique A que possedent les adultes de C. lepida a evolue secondairement.
^Present address: Department of Biology, York University, Downsview, Ontario, M3J 1P3
298
Kuster
La structure et la fonction probable des yeux Cicindelidae out ete comparees a ceux de deux representants des Carabidae,
leur fatnille apparentee. Les adultes de I’espece nocturne Pterostichus melanarius llliger ont de petits yeux du type scotopique II;
I’espece diurne Elaphrus americanus Dejean a de grands yeux photopiques. II est conclu que la vision scotopique et la vision
photopique ont evolue parallelement chez ces taxons apparentes.
Tous les yeux de Coleopteres examines sont eucones et ont une “couche subcorneenne” entre les lentilles corneenes et les
cones crystallins. Ils ont un rhabdomere distal, compose de microvillosites a partir de la septieme cellue retinienne seulement, un
rhabdome plus proximal, rectangulaire et fusionne, forme de six cellules retiniennes, et un huitieme cellue retinienne basale,
possedant un rhabdomere. Les grands yeux globuleux des especes diurnes et crepusculaires ont des mechanorecepteurs entre leurs
facettes.
INTRODUCTION
On the basis of ecological correlations, Exner (1891) classified insect compound eyes into two
structural and functional categories. Apposition eyes are characteristic of diurnal insects active in bright
sunlight and superposition eyes are adapted for vision of crepuscular and nocturnal insects. Recently
functional categories for compound eyes have been borrowed from terms used for cone and rod visual
systems of vertebrate retina, (Goldsmith and Bernard, 1974), respectively, photopic and scotopic. In
scotopic eyes, a transparent zone or “crystalline tract” is formed either as an extension of Semper cells
(Horridge, 1968, 1969), or by the distal non-rhabdomeric portions of the retinula cells (Miller et ai,
1968; and Doving and Miller, 1969). Horridge (1971) showed that in clear zone scotopic eyes, light
entering many lenses is scattered upon several rhabdoms thus increasing light intensity, but decreasing
image resolution of individual ommatidia (Horridge et ai, 1972; and Horridge, 1975). In photopic eyes
such scattering of light does not occur (Varela and Wiitanen, 1970) and less than one percent of the
light captured by a rhabdom is received through neighbouring lenses (Shaw, 1969).
Eyes of males of one species of each of the four North American genera of Cicindela (Coleoptera)
have been examined: Amblycheila schwarzi Horn; Omus californicus californicus Horn; Megacephala
Carolina mexicana Gray; and Cicindela tranquebarica Herbst. Since adults of Cicindela lepida Dejean
and Cicindela belfragei Salle have apparently become secondarily crepuscular, their eye structures are
also described to determine if these eyes have evolved in response to this diel behavioural adaptation.
The question arises as to why tiger beetles were chosen for a detailed examination of eye structure
and function from an evolutionary approach? This bias is based on my hypothesis, that since there is a
behavioural transformation series from a plesiotypic (ancestral) nocturnal through crepuscular to the
apotypic (derived) diurnal diel activity within the four North American genera of cicindelids, that there
may also be a parallel transformation series in structure and function of their compound eyes. I therefore
believe this to be evolution of eye structure and function in relation to diel activity.
The only detailed research on larval stemmata and adult compound eye structure and function of
some species of Cicindela is that by Friedrichs (1931). On questioning the structural attributes of eyes of
individuals of other cicindelid genera, he wrote (translated from the German): “It would be particularly
interesting to establish in what manner the eyes of these nocturnal and crepuscular cicindelines have
been adapted to their way of life: It may well be assumed that superposition [scotopic] eyes with pigment
displacement have been formed, while the day-running or flying cicindelines possess apposition
[photopic] eyes (like Cicindela)C To answer some of Friedrichs’ questions, this paper provides
descriptions of eye structures based on histological examination; descriptions of the relationships of eye
size groups, eye function categories, and diel activities in terms of a reconstructed phylogeny of the
Cicindelidae.
Structure and function of cicindelid eyes are then compared to eyes of individuals of their sister
family, the Carabidae, to determine if carabids with similar diel activity have evolved similar eye
structures. To answer this question, eyes of adults of nocturnal Pterostichus melanarius llliger, and
Compound eyes of Cicindelidae and Carabidae
299
diurnal Elaphrus americanus Dejean are described. Eye structure is then related to eye size groups and
eye function categories of the cicindelids and the phylogeny of these sister taxa.
MATERIALS AND METHODS
For scanning electron microscopy (SEM), beetle heads were washed in Tide*^ laundry detergent,
rinsed in distilled water, then fixed in 5% formalin. After ethanolic dehydration, heads Were cleared in
xylol and air-dried overnight (Hollenberg and Erickson, 1973). The heads were carbon and gold coated
to a thickness of 15-20 nm using an Edwards 12E vacuum evaporator, then examined with a Cambridge
Stereoscan S4 Scanning Electron Microscope at accelerating voltages of 20-30 kV. Histological
material for light microscopy (LM) was fixed in hot 80% ethanolic Bouin’s Duboscq (Pantin, 1962).
Dark-adapted beetles were deprived of light for five days prior to fixation. Excised eyes were dehydrated
in tertiary butanol then double-embedded using Peterfi’s celloidin-parraffin technique (Pantin, 1962).
To facilitate sectioning of these hard beetle heads, the knife and wax blocks were chilled. Sections were
cut at 10-12 pm using a Leitz Wetzlar microtome. Longitudinal and transverse sections were treated in
a saturated mercuric chloride containing 5% acetic acid mordant solution (Pantin, 1962). Precipitations
of mercurous chloride and metallic mercury were removed using Gram’s variation of Lugol’s iodine
solution. A 5% sodium thiosulfate solution removed Lugol’s solution (Humason, 1962). Sections were
stained with Mallory’s triple stain (Pantin, 1962) and mounted with Canada balsam. Representative
photographs were taken using a Carl Zeitz Ultraphot II.
Measurements of structures for ratios were randomly chosen and calculated as x ± SE for a sample
size of five. The retinulae were assumed to be a cylinder consisting of three portions: the clear zone,
rhabdom zone, and basal zone. These volumes and the volumes of the rhabdom zone of the basal retinula
cell were calculated as cylinders. Volumes of the rhabdom of the retinula rhabdom zones were calculated
as a solid rectangle. Comparative measurement data were statistically analyzed using computer
programs for One-Way Analysis of Variance and Duncan’s New Multiple Range Test of Means (Sokal
and Rohlf, 1969). Using the statistical groupings resulting from Duncan’s test, measurements were
either tabulated in the eye size or eye functional category.
RESULTS
Structure of eyes of one species of each of the four North American genera of cicindelid adults
Eyes of the nocturnal genera, Amblycheila schwarzi (Fig. 1) and Omus californicus (Fig. 2) have
small, relatively flat eyes compared with the large bulbous eyes of crepuscular Megacephala Carolina
(Fig. 3) and diurnal Cicindela tranquebarica adults (Fig. 4). The verticies (v) of the latter two beetle
heads are concave, allowing the eyes to extend above the top of the heads. Figures 5-8 (Kuster, 1975)
show that representative compound eyes of all four genera are convex and outer surfaces consist of
convex, hexagonal corneal lenses (1). A ring of cuticle, the ocular sclerite (os), defines the border of the
eyes. Because of eye size and shape differences, each beetle has a variable anterior, posterior, and dorsal
stereoscopic area of the visual field (Kuster, 1978).
Table 1 shows that adults of nocturnal cicindelids have fewer ommatidia than have adults of
diurnal-crepuscular beetles. In representatives of nocturnal genera, eyes span less than one-third the
head width, but in crepuscular and diurnal genera, eyes occupy approximately one-half of the head
width. From values comparing eye height to head height, neither eyes of Amblycheila schwarzi or Omus
californicus adults extend above the vertex as do those of Megacephala Carolina and Cicindela
tranquebarica. It is possible therefore to assume that both Cicindela tranquebarica and Megacephala
Quaest. Ent., 1979, 15 (3)
300
Kuster
Carolina adults see above the head, but that vision above the vertex is less for eyes of Amblycheila
schwarzi and Omus californicus adults. Ratios of head width to pronotum width indicate that neither
adults of Amblycheila schwarzi nor Omus californicus can see behind their pronota. However, both
representatives of Megacephala Carolina and Cicindela tranquebarica have this ability. None of these
adult tiger beetles can see behind their elytral margins. However, the list of ratios (Table 1) does not
indicate the absolute limits of vision. Tiger beetles display an alert behavioural stance by rearing up on
the prothoracic legs so that the abdomen is pressed to the substrate (Swiecimski, 1957; Willis, 1967).
Such a stance may permit the beetles to see more of their environment in the anterior and posterior
directions.
Ommatidia of insect compound eyes can be divided into two distinct structural and functional
regions: the light receiving or dioptric apparatus, with its associated primary pigment cells and the
retinula with its associated secondary and basal pigment cells.
Figures 9-12 are representative longitudinal, and figures 13-16, representative transverse sections,
through compound eyes of one species from each of the four cicindelid genera. These figures show
lamellated corneal lenses (1) having distal acidophilic thin corneal layer (t), and crystalline cones (c)
with four quadrants. Normally, the dioptric apparatus of eucone eyes {sensu Grenacher, 1879) consists
solely of these two structures. However, in cicindelid beetle eyes, an acidophilic, lamellated third layer
has been discovered between the lens and the cone. This layer is termed the “subcorneal layer” (cl)
because of its position and structural similarity to the corneal lens. These beetle eyes therefore, have a
three layered dioptric apparatus. Figures 17-20 show that corneal lenses (1) are apparently convex
distally and hexagonal in shape. None of the lenses have corneal nipples (Bernhard et al., 1965).
Scattered between lenses of adult eyes of Megacephala Carolina (Fig. 19) and Cicindela tranquebarica
(Fig. 20) are conical interfacetal cuticular pegs (cp). There is approximately one peg per 20 ommatidia
with a total of approximately 210 per eye in adults of Megacephala Carolina and one peg per 15
ommatidia (total 260) on eyes of Cicindela tranquebarica adults. Pegs are slightly taller and wider in
eyes of Cicindela tranquebarica (Table 1).
Although not resolvable in figures 9-16, two primary pigment cells which are devoid of pigment
granules, surround the crystalline cones. Oblique light rays entering the eyes, which cannot be refracted
by the dioptric apparatus, are absorbed laterally by pigment granules in secondary pigment cells (2p).
Secondary pigment granules are more densely aggregated and appear black in eyes of nocturnal
Amblycheila schwarzi (Fig. 9,13) and Omus californicus adults (Fig. 10,14), compared to the less dense
brown pigment granules in eyes of Megacephala Carolina (Fig. 11,15) and Cicindela tranquebarica
(Fig. 12,16).
The dioptric apparatus is connected to the retinula by a crystalline thread which is shrouded by
secondary pigment cells. This thread is an extension from each of the four Semper cells which surround
the crystalline cone quadrants.
The retinula extends proximally from the proximal tip of the crystalline thread to the basement
membrane (bm). A cluster of seven neurons of retinula cells constitute an ommatidial retinula. In all
cicindelid beetle eyes examined, microvilli from retinula cell seven form a distal rhabdomere. Retinula
cells of Amblycheila schwarzi (Fig. 9,13) and of Megacephala Carolina adults (Fig. 11,15) consist of a
clear retinula zone (cr) and a proximal retinula rhabdom zone (rr). Retinula of eyes of Omus
californicus (Fig. 10,14) and of Cicindela tranquebarica (Fig. 12,16) have no clear retinula zone. All
have a basal retinula zone (br) of an eighth retinula cell with a rhabdomere.
The rhabdom zone consists of a rectangular, fused rhabdom (r) in the centre of six retinula cells (rt)
(Fig. 21-24). Two retinula cells contribute microvilli to form the rhabdom of the wide sides; one cell
contributes to each short side. The rhabdom occupies a greater percentage of retinula cell surface area
Compound eyes of Cicindelidae and Carabidae
301
and volume (Table 3) in eyes of Amblycheila schwarzi (Fig. 21) and Megacephala Carolina (Fig. 23)
than in Omus californicus (Fig. 22) or Cicindela tranquebarica adults (Fig. 24). The retinula cytoplasm
in eyes of Cicindela tranquebarica adults is distinctly visible. In all four beetle eyes, the vacuolated
seventh retinula cell is positioned lateral to the rhabdom and does not contribute a rhabdomere at this
level. Sixteen secondary pigment cells (2p) surround the retinula and four basal pigment cells (bp)
surround the basal retinula cell.
Each of the eight retinula cells extends an axon (a) to interneurons in the lamina ganglionaris (Ig)
(Fig. 9-12). Eight axons from each ommatidium penetrate a single circular fenestration in the
tracheole-rich basement membrane, and are aggregated with axons of five adjacent ommatidia in the
form of axonal bundles distal to the lamina ganglionaris. Evident from figures 25-28, axonal bundles
(ab) of eyes of Amblycheila schwarzi adults are much longer than those in other beetle eyes. Glial cells
(gl) surround axons. The probable neuronal pathway through the brain is suggested in these figures.
Following synapsis lamina interneurons, axons cross over at the first optic chiasmata (Ic), then extend to
the medulla (md), the second synaptic site of the optic lobe. Visual axons again cross over at the second
optic chiasmata (2c), followed by proximal synapsis in the third region of the optic lobe, the lobula (lo).
Optic lobes consist of a connective tissue sheath, the neurilemma (nl), an underlying cellular
perineurium (pn) with glial and neuronal cell bodies, and a central neuropile of axons and dendrites. A
large pigment accumulation (pa) is on the ventral aspect of the interface of the lamina and medulla of
the optic lobe (see also Fig. 12).
After dark adaptation for five days, structures of the eyes of Cicindela tranquebarica adults were
examined. Only minor changes occured when compared to light-adapted eyes. In eyes of these beetles,
pigment granules in secondary pigment cells migrated distaly around crystalline cones and proximally
around basal retinula cells leaving little pigmentation surrounding retinulae. This is assuming that the
same pattern of orientation of pigments is not altered by fixation and dehydration. Shortening of the
crystalline threads to approximately half their length in the light-adapted state was the most striking
change.
Structures of representative ommatidia are summarized diagrammatically in figures 29-32.
Table 1 and 3 provide measurements from five ommatidia chosen randomly.
Structure of eyes of Cicindela lepida and Cicindela belfragei adults
Figure 33, of the head of an Cicindela lepida adult shows large bulbous eyes, similar in shape
(Fig. 34) to those of Cicindela tranquebarica adults (Fig. 4). Corneal lenses (1) (Fig. 35) and
interfacetal pegs (cp) (Fig. 36) are typical of Cicindela adults. The corneal layer (t) is relatively thin.
From longitudinal (Fig. 37) and transverse sections (Fig. 38) of the eye, the cellular organization is
similar to that of eyes of Megacephala Carolina adults (Fig. 1 1,15). A clear retinula zone (cr) is present.
The surface area of the rhabdom (r) (Fig. 39) is moderately large.
Light-adapted eyes of Cicindela lepida adults show lengthening of crystalline threads, shortening,
but not disappearance of the clear retinula zone. A more even distribution of pigment granules in
secondary pigment cells also occurs along the length of retinulae compared to dark-adapted Cicindela
lepida eyes collected at twilight.
Eye shape (Fig. 40,41), corneal lenses (1) (Fig. 42) and cuticular pegs (cp) (Fig. 43) of eyes of
Cicindela belfragei adults are similar to those of other Cicindela adults. Cellular organization for vision
(Fig. 44,45) is similar to that of Cicindela tranquebarica eyes (Fig. 12,16). There is no clear retinula
zone. The surface area of the rhabdom (r) (Fig. 46) is small.
Quaest. Ent., 1979, 15 (3)
302
Kuster
Structure of eyes of Pterostichus melanarius and Elaphrus americanus carabid adults
The head of Pterostichus melanarius adult (Fig. 47) has a convex vertex like that of Amblycheila
schwarzi and of Omus californicus adults (Fig. 1,2). Eyes are small and spherical (Fig. 48) as are those
of Amblycheila schwarzi adults (Fig. 5). Hexagonal, convex corneal lenses (1) (Fig. 49) have a thin
corneal layer, but no interfacetal pegs. Material (x) secreted from dermal glands (Fig. 50) may be used
as grooming lubricant to clean the eye, or may contribute to the composition of the thin corneal layer.
These eyes have no clear retinula zone (Fig. 51,52) like eyes of Omus californicus adults (Fig. 10,14),
but the rhabdom (r) has a large cross-sectional surface area (Fig. 53) compared to that of the rhabdom
of Amblycheila schwarzi eyes (Fig. 24).
Vertices of Elaphrus americanus adults are convex, however, their eyes are bulbous and extend above
the vertex (Fig. 54). They are similar in shape (Fig. 55) to eyes of Megacephala Carolina adults (Fig. 7),
and those of other Cicindela adults (Fig. 8,34,41). Hexagonal corneal lenses (1) are well defined
(Fig. 56) due to their degree of convexity, and are similar to those of Cicindela adults (Fig. 20,35,42).
Interfacetal pegs (cp) (Fig. 57) are present. There is no clear retinula zone (Fig. 58,59) and these eyes
have a similar cellular organization to eyes of Cicindela tranquebarica (Fig. 12,16) and Cicindela
belfragei (Fig. 44,45). The rhabdom (r) (Fig. 60) has a small surface area.
Eye size groups and functional categories of cicindelid beetle eyes based on measurements of structures
From statistical inference using One-Way Analysis of Variance and Duncan’s Range Test of Means,
measurement data were grouped either into eye size or eye function categories.
Eye size groups. - When structural measurements are related to eye size, adults of the four North
American genera of Cicindelidae can be divided into two groups (Table 1): Small Eye Group: eyes of
adults of Amblycheila schwarzi; Omus californicus. Large Eye Group: adults of Megacephala Carolina ;
Cicindela tranquebarica; Cicindela lepida and Cicindela belfragei.
For clarification of small eye size relationships of cicindelid taxa, the similarity matrix (Table 2) is
included. The data for Table 2 are summations of statistically similar structures at P = 0.01% from
Table 1. Based on these totals, there are trends in similarities within eye size groups and differences
between these two groups among the cicindelids. Note that of 39 characters, small eyes of nocturnal
Amblycheila schwarzi and Omus californicus share 21 characters; large eyes of crepuscular
Megacephala Carolina and diurnal Cicindela tranquebarica adults share 16 characters. Also eyes of the
adults of Cicindela spp. share several attributes.
Unlike the diurnal and crepusclar beetles, nocturnal cicindelids possess small eyes with fewer
ommatida and no interfacetal pegs. Smaller visual fields are characteristic of eyes of these beetles, as
demonstrated by head, thorax, and elytral ratios, and from forward and dorsal Mollweide
homolographic projections. (Kuster, 1978). Corneal lenses are long in these eyes, while crystalline cones
of diurnal-crepuscular large eyes occupy a larger percentage of dioptric apparatus lengths. The dioptric
apparatus occupies over half the ommatidial length in small-eyed beetles; but only approximately
one-third the ommatidial length in the large eye group. Characteristic of large cicindelid eyes are
crystalline threads almost twice as long as in the small eye group. Retinulae extend only slightly over
one-third the ommatidial length of the small eye group but over half this length in the large eye group.
Retinulae extend only slightly over one-third the ommatidial length of the small eye group but over half
this length in the large eye group. Basal retinula zones are longer in the small eye group. There is also a
similarity in nocturnal beetles concerning rhabdom zone volume and retinula and rhabdomeric volumes
of ommatida and compound eyes, all of which are smaller than volumes of the long retinula and
rhabdoms of large eyes.
Compound eyes of Cicindelidae and Carabidae
303
Table 1. Arrangement of six eieindelid beetles into two groups based on eye size. The values are x for
n = 5 for each species. Solid underscore represents no statistically significant difference at
P = 0.05%. Dashed underscore represents no statistically significant difference at P = 0.01%.
Absence of an underscore indicates statistically difference. 0 indicates no such structure exists
for that species and — indicates no measurement made.
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Table 1. (continued)
Table 2. Similarity matrix for six cicindelid beetles based on eye size groups. For Amblycheila schwarzi,
Omus californicus, Megacephala Carolina, Cicindela tranquebarica there are 39 characters.
When all six beetles are compared there are 20 characters in common available.
Compound eyes of Cicindelidae and Carabidae
305
Eye functional categories. - When structures involved with function of cicindelid compound eyes are
statistically analyzed, three functional categories can be inferred (Table 3); Scotopic A; eyes of
representative adults of Amblycheila schwarzi\ Megacephala Carolina and Cicindela lepida.
Scotopic B: eyes of adults of Omus californicus. Photopic: eyes of adults of Cicindela tranquebarica, and
Cicindela belfragei.
For clarification of functional eye categories of cicindelid taxa, the similarity matrix (Table 4) is
included. The data for Table 4 are summations of statistically similar structures at P = 0.01% from
Table 3. Based on these totals, there are trends in similarities within categories of eye function and
differences among these three categories within the cicindelids.
Beetles included in the scotopic A functional category have relatively long antennae which may
permit increased touch and olfactory stimulation in addition to sight. The thin corneal layers of these
eyes are relatively thick, but the subcorneal layers are relatively thin. Eyes of adult Amblycheila
schwarzi, Megacephala Carolina and Cicindela limbalis have clear retinula zones and although less than
half these retinula lengths are rhabdomeric, these rhabdoms have very large surface areas. Volume of
rhabdom zones are greater in eyes of Amblycheila schwarzi than those of Omus californicus, its
small-eyed counterpart. It is larger in eyes of Megacephala Carolina than its large-eyed counterparts,
Cicindela tranquebarica, Cicindela limbalis and Cicindela belfragei. Percentage rhabdom zone volume
of retinulae are smaller in scotopic A eyes due to the presence of clear retinula zones. However, total
volume of the rhabdom per ommatidium is larger in scotopic A eyes as is percentage of rhabdom volume
to retinula volume since the rhabdom has such a large surface area.
Eyes of Omus californicus adults are scotopic B. Individuals of this species have short antennae and
although their eyes possess many small-eyed structural similarities with those of Amblycheila schwarzi
adults (Table 1), they can be grouped into a separate functional category. Like scotopic A eyes, these
eyes have thin subcorneal layers, but thinner, thin corneal layers. Unlike the scotopic A eyes, there is no
clear retinula zone. Although almost twice the retinula lengths are occupied by the rhabdom zone,
surface areas and volumes of the rhabdom are smaller, as is percentage volume of the rhabdom to
retinula volume in scotopic B and scotopic A eyes. Consequently, percentage volume of the retinula
around the rhabdom and percentage volume of the rhabdom are larger in scotopic B ommatidia, but
total retinula and rhabdom volumes are less in the whole scotopic B eye.
Cicindela tranquebarica and Cicindela belfragei adults have photopic eyes. Like adults of Omus
californicus, these beetles have short antennae. But based on eye size, these eyes share structural
similarities to eyes of Megacephala Carolina adults since they are in the large eye group (Table 1).
Photopic eyes of Cicindela tranquebarica and Cicindela belfragei adults have thick subcorneal layers,
but like scotopic B eyes, have thin corneal layers, no clear retinula zone, and rhabdoms occupying almost
the complete retinula length. Surface areas of rhabdoms are the smallest and rhabdom volumes are
small considering retinulae lengths. Percentage rhabdom volume to retinula volume is very small but
percentage of retinula volume surrounding the rhabdom is very large.
Eye size groups and functional categories of carabid beetle eyes based on measurements of structures
Eye size groups. - Cicindelid beetle eyes examined here can be placed into two groups based on eye
size and into three functional categories. To test convergence of eye structure and function based on eye
size, measurement of eyes of two carabid adults were statistically compared to those of four cicindelid
sister taxa (for basic data, see Kuster, 1978).
According to eye size, cicindelid and carabid adults have similar eye structures (Table 5). Eyes of
Pterostichus melanarius fit the small eye group; Elaphrus americanus, the large eye group. The data for
Table 5 are summations of statistically similar structures at P = 0.01%. Based on these totals, there are
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Table 3. Arrangement of six cicindelid beetles into three categories based on eye function. The values
are T for n = 5 for each, species. Solid underscore represents no statistically significance
difference at P = 0.05%. Dashed underscore represents no statistically significant difference at
P = 0.01%. Absence of an underscore indicates statistically difference. 0 indicates no such
structure exists of that species and — indicates no measurement made.
Compound eyes of Cieindelidae and Carabidae
307
trends in similarities within eye size groups and differences between these two groups between the
cicindelids and carabids.
Although eyes of the carabids have fewer ommatidia, eyes of diurnal Elaphrus americanus adults
have more ommatidia than eyes of nocturnal Pterostichus melanarius. Corneal lenses and crystalline
cones are longer and no interfacetal pegs are present in the small of this nocturnal, carabid eyes. Lengths
of crystalline threads of eyes of Omus californicus and Elaphrus americanus are similar, and crystalline
threads of Pterostichus melanarius, and Megacephala Carolina eyes are smaller in length. Basal retinula
zone lengths and diameters are similar in all these beetle eyes except in those of Elaphrus americanus
where they are half as large. Eyes of Pterostichus melanarius adults have longer retinulae similar to
those of Omus californicus adults. Volume of the rhabdom zone of the retinula and retinula volume per
eye are similar in Omus californicus and Pterostichus melanarius adults, and adults of Elaphrus
americanus, Amblycheila schwarzi and Omus californicus since all these eyes have relatively short
retinulae. Indicative of small eyes and nocturnal behaviour, both adults of Amblycheila schwarzi and
Pterostichus melanarius have similar rhabdom volume per eye but the carabid has a rhabdom volume
statistically similar to eyes of Cicindela tranquebarica adults because retinulae of this carabid are so
short. Rhabdom volume of Elaphrus americanus adults is exceedingly small.
Eye functional categories. - Comparisons of functional aspects of the cellular organization for vision
of cicindelid and carabid beetles show similarities (Table 6) (for basic data, see Kuster, 1978). Eyes of
Pterostichus melanarius adults are grouped with eyes of Omus californicus adults in the scotopic B
category; eyes of Elaphrus americanus adults in the photopic category with Cicindela tranquebarica
adults. The data for Table 6 are summations of statistically similar structures at P = 0.01%. Based on
these totals, there are trends in similarities within the eye functional categories and differences among
these three categories among the cicindelids and carabids.
Thickness of the thin corneal layer places eyes of Pterostichus melanarius close to those of
Amblycheila schwarzi while the subcorneal layer of eyes of Elaphrus americanus adults, though thicker
than that of Pterostichus melanarius is similar to that in eyes of adults of Amblycheila schwarzi, Omus
californicus and Megacephala Carolina. Diameters of retinulae of Pterostichus melanarius eyes are
similar to those of Omus californicus, while basal zone diameters of the two carabid eyes are similar.
Both lengths and widths of rhabdoms of Pterostichus melanarius eyes are similar to those of
Amblycheila schwarzi adults, but the rhabdom of Elaphrus americanus like that of Cicindela
tranquebarica adults, is exceedingly small with minimum surface area and volume.
DISCUSSION AND CONCLUSIONS
Dioptric apparatus
Adult eyes of representative species of North American genera of Cieindelidae and Carabidae have a
eucone, three-layered dioptric apparatus. Although Gissler (1879) observed the corneal lens of adult
Omus sp. and Cicindela sp. to be biconvex, and the cornea of adult Amblycheila sp. to be convex only
interiorly, I have shown that adult eyes of species of these genera to have biconvex lenses. Confusion
regarding corneal lens shape possibly resulted because the lenses of Amblycheila schwarzi and
Megacephala Carolina have relatively thick corneal layers which externally appear smooth. Thickness of
this layer may be important in understanding difference in eye function. A thick layer may scatter
incident light over many lenses so that light is shared by adjacent ommatidia. Because this corneal layer
is thinner in eyes of Omus californicus, Cicindela tranquebarica, Cicindela limbalis and Cicindela
belfragei adults, individual lenses are more distinctly separated and optical isolation is maintained
between adjacent ommatidia, possible resulting in enhanced visual acuity. Scratches on this layer may
result from burrowing or less likely from grooming activities.
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Kuster
Table 4. Similarity matrix for six cicindelid beetles based on eye function. For Amblycheila schwarzi,
Omus californicus, Megacephala Carolina, Cicindela tranquebarica there are 17 characters.
When all six beetles are compared there are only 16 characters in common available.
Table 5. Similarity matrix for four cicindelid and two carabid beetles based on eye size. There are 20
characters.
Table 6. Similarity matrix for four cicindelid and two carabid beetles based on eye function. There are
16 characters.
All cicindelid and carabid beetle eyes examined have a structurally distinct layer between the lens
and cone termed the subcorneal layer. Eyes of adult Notiophilus biguttatus F. and Loricera pilicornis F.
(Carabidae) also have this structure which Home (1976) terms the “proximal corneal layer”. From
observations sing Nomarski differential interference contrast microscopy, this layer is of intermediate
refractive index between the lens and cone and therefore may function to bend incident light rays
medially toward the crystalline cone (Kuster, 1978).
Compound eyes of Cicindelidae and Carabidae
309
Interfacetal pegs
Both crepuscular and diurnal adult cicindelids and the diurnal carabid have interfacetal pegs between
some corneal lenses. Nocturnal flightless cicindelids and the nocturnal carabid lack them. Since the pegs
appear to lie on a cuticular articulating membrane and since there is no visible hole at the apex, it is
assumed that these structures function as mechanoreceptors. Other adult beetles, capable of flight, such
as Creophilus erythrocephalus F. and Sartallus signatus Sharp (Staphylinidae also have interfacetal
pegs (Meyer-Rochow, 1972) similar in size and shape to those described here. According to Nesse
(1965, 1966) for Apis mellifica {=Apis mellifera Apidae) and Chi and Carlson (1976) for Musca
domestica (Muscidae), and Honegger (1977) for Gryllus campestris L. (Gryllidae), these interfacetal
mechanoreceptors sense the direction and relative velocity of passing over the eyes during flight and may
play a role in eye cleaning behavior (Honegger)
Retinula cells and rhabdoms
It is important to emphasize that the difference in retinula and rhabdom structure of the cicindelid
and carabid eyes investigated is not one of change in cell number, but is a difference in cellular
organization which results in varied functional abilities of these eyes.
Scotopic A eyes
Retinulae of eyes of adults of Amblycheila schwarzi, Megacephala Carolina and Cicindela lepida
have a clear retinula zone or crystalline tract {sensu Doving and Miller, 1969) consisting of seven
retinula cells which do not have a rhabdom at this level. Such a scotopic A retinula organization has also
been observed in adult carabid beetle eyes such as those of Carabus auratus L. (Kirchoffer, 1905, 1908;
Bernard, 1932; and Hasselmann, 1962), Steropus madidus Fab., and Eutrichomerus terricola Herbst
(Bernard, 1932); and Notonomus sp. (Horridge and Giddings, 1971); and in the following dytiscids;
Dytiscus sp. and Cybister sp. adults (Grenacher, 1879; Exner, 1891; Kirchoffer, 1908; Horridge, 1969;
Horridge et ai, 1970; and Meyer-Rochow, 1973, 1975).
Scarab adults, Melontha volgaris F. (Kirchoffer, 1908), Oryctes rhinoceros (Bugnion and Popoff,
1914), and others (Grenacher, 1879) also have scotopic A functional eyes. Based on research on
Repsimus manicatus Lea (Scarabaeidae) adults, Horridge and Giddings (1971) define the
“neuropteran” type of compound eye as having a crystalline thread in the light-adapted state only, with
retinula cell bodies extending to the tip of the cone only in the dark-adapted state. Eyes of
Anoplognathus pallidicollis Blanch (Scarabaeidae) are also scotopic A and have a basal retinula cell
(Meyer-Rochow and Horridge, 1975). Although dark and light adaptation experiments here were
preformed on scotopic A eyes only of Cicindela lepida, it is possible to assume that eyes of Amblycheila
schwarzi, Megacephala Carolina and Cicindela lepida adults are also of the neuropteran type, {sensu
Horridge and Giddings, 1971). Evolution of a clear retinula zone in these cicindelid eyes is probably an
adaptation to permit a further increase in sensitivity in the dark-adapted state by allowing an increase in
the acceptance angle of lenses and in the cross-sectional area of the rhabdoms without prejudice to
acuity of the light-adapted eye. Optical mechanisms of summation of scattered light in clear zone
compound eyes are reviewed by Horridge, 1971; Kunze, 1972; Horridge et al., 1972; Diesendorf and
Horridge, 1973; and Horridge, 1975.
Scotopic B eyes
Like eyes of adults of the closely-related genus Amblycheila sp. in the same subtribe Omina,
scotopic B eyes of Omus californicus adults are eucone and have a thick dioptric apparatus and a
crystalline thread, but importantly, they have no clear retinula zone. Instead, the rhabdom extends the
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Kuster
full length of the retinula from the distal seventh rhabdomere to the eighth basal retinula cell. Although
adephagans usually have a neuropteran type of scotopic eye as defined by Horridge and Giddings
(1971), these authors state that some adephagan eyes have long fused rhabdoms. In longitudinal section,
ommatidia of Pterostichus melanarius (this study), Procrustes coriaceus L., Carabus glabratus Payk.,
and Broscus cephalotes L. (Carabidae) (Kirchoffer, 1908) also have broad fused rhabdoms and no clear
retinula zones. Dorsal and ventral divided eyes of Gyrinus nator subtraitus Steph., (Bott, 1928), Gyrinus
subtriatus, (Wachmann and Schroer, 1975), Gryinus natator L., (Burghause, 1976), and dorsal eyes of
Dineutes assimilis adults, (Gryinidae) (Pappas and Larsen, 1973) are also of the scotopic B functional
category.
Photopic eyes
Other adult carabid eyes have rhabdoms extending the full retinula length (Bernard, 1932; Home,
1976) These eyes have three levels of rhabdom organization similar to those of Dytiscus marginalis
adult eyes (Horridge, 1969). However, like eyes of Omus calif ornicus adults, there is no clear retinula
zone but, importantly, the rhabdoms have less surface area.
A greater reduction of rhabdomeric surface area and volume occurs in photopic eyes of diurnal
cicindelid adults of the genus Cicindela. From histological examination of adult eyes of diurnal
Cicindela campestris L, Cicindela silvatica Latr., and Cicindela hybrida L., Kirchoffer (1905) made
descriptions and in 1908 figured ommatidia of the first two species. Further examination of eyes of
Cicindela campestris by Friedrichs (1931) and Home (1976) confirmed the slender fused rectangular
rhabdom structure. Swiecimski (1957) reported a similar retinula organization in eyes of Cicindela
hybrida adults, and I have also observed this cellular organization in eyes of adults of the following
diurnal cicindelids. Cicindela tranquebarica Herbst, Cicindela belfragei Salle, Cicindela limbalis Klug,
Cicindela longilabris Say, Cicindela limbata nympha Casey, and Cicindela repanda repanda Dejean.
Since these ommatidia do not have a clear retinula zone or a broad fused rhabdom, light is not scattered
over adjacent rhabdoms and the eyes are photopic. Eyes of Elaphrus americanus also are photopic and
although Elaphrus cupreus Duftschmid are active in the shade they have photopic eyes (Kirchoffer,
1908; Bauer, 1974; and Home, 1976) as do heliophilus adults of Elaphrus riparius Linnaeus (Bauer,
1974) . Possibly, photopic eyes have a greater spectral sensitivity than scotopic eyes (review: Menzel,
1975) and have the ability to detect polarized light (reviews: Snyder, 1973; Wehner, 1976).
Pigment cells
In dark-adapted scotopic A eyes of Amblycheila schwarzi adults, pigment is concentrated in distal
portions of the secondary pigment cells surrounding the crystalline cones and retinulae extend to the
cone tips. The clear retinula zone is devoid of pigment, allowing light to be scattered on adjacent
rhabdoms for increased light intensity. Such a cellular organization corresponds to the dark-adapted
scotopic eye of the neuropteran type {sensu Horridge and Giddings, 1971). Light-adapted scotopic A
eyes of Megacephala Carolina and Cicindela lepida have crystalline threads to direct light to individual
rhabdoms, but the long clear zones are not surrounded by secondary pigment granules which suggests
that light is scattered over adjacent rhabdoms. Dark-adapted photopic B eyes of Omus californicus and
Pterostichus melanarius and light-adapted photopic eyes of Cicindela tranquebarica, Cicindela
belfragei and Elaphrus americanus adults have distal aggregations of pigment granules surrounding
crystalline cones and threads. Like photopic eyes of Apis mellifica L. (Kolb and Autrum, 1972),
Cicindela tranquebarica and Cicindela belfragei eyes also have pigment granules along the retinula
length. As postulated for these apid eyes (Varela and Wiitanen, 1970), I suggest that parallel light
entering photopic cicindelid and carabid eyes is directed to the rhabdom for phototransduction and
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311
oblique rays are absorbed at the level of the dioptric apparatus by secondary pigment granules. Optical
isolation at the retinula level is maintained by an envelope of pigment along its length which prevents
stimulation of the rhabdom by light coming from adjacent ommatidia. This presumably results in finer
resolution of the image.
Large pigment aggregations on the ventral aspect of the lamina ganglionaris and medulla interface
are postulated to be reminants of six larval stemmata similar to that in other adult insect eyes
(Weber, 1933). To prove this, an analysis of tissue organization of the pharate pupa would be required.
Functionally, this pigment and glial cell pigment may prevent stimulation of the retinula by light
entering the eye antidromically through thin cuticular regions.
Retinula cell axons
I did not determine from light microscope studies if axons of similar colour sensitivity in an axonal
bundle synapse in the lamina cartridge as observed by Braitenberg (1967) in eyes of Musca domestica
Meig. (Muscidae). Why the axons are comparatively longer in eyes of Amblycheila schwarzi is not
understood, but a similar arrangement is also observed in nocturnal scotopic B eyes of Pterostichus
melanarius and Steropus madidus Fab. adults (Carabidae) (Bernard, 1932). Axons of the other
cicindelid and carabid beetle eyes are shorter, and these eyes have a lamina, medulla, and lobula broadly
similar to photopic eyes of the honey bee Apis mellifera L. (Ribi, 1975). To determine exact neural
connections, Golgi silver impregnation (Ribi, 1974) of axons and interneurones would be required.
Significance of evolution of character states of cicindelid and carabid beetle compound eyes
In this section, significance of differences in structure and function of compound eyes is approached
through a phylogenetic analysis of tiger beetles. This is followed by consideration of taxa representing
other families of adephagans. A general pattern is sought and its outlines are explained in terms of the
relationship between ecology and diversification.
Also evolution of character states of cicindelid and carabid beetle compound eyes are related to the
reconstructed phylogeny (Fig. 61). For readers interested in keys, descriptions and diagnoses of
character states of tiger beetle taxa, see Schaupp (1883); Leng (1902, 1920); Bradley (1930); and
Arnett (1968). For a discussion of character states determining cleavage points between tribes, see Horn
(1908-1915; 1926); Bradley (1930); and Arnett (1968); for subtribes, see Thompson (1857); Horn
(1908-1915); Leng (1920); and Wallis (1961); for genera within the subtribe Omina, see Lacordaire
(1843); Thompson (1857); Brous (1877); Schaupp (1883); Casey (1897); Leng (1902); Bradley (1930);
Arnett (1946, 1968); and Vaurie (1955); the genus Megacephala. See Thompson (1857); Schaupp
(1883); Horn (1908-1915); Arnett (1946); and Willis (1969); the genus Cicindela, see Leconte (1857);
Schaupp (1883); Leng (1902, 1920); Horn (1908-1915); Bradley (1930); Arnett (1946, 1968); Rivalier
(1954); Wallis (1961); and Willis (1968).
Ancestral stock of the Cicindelidae was probably related to Carabini of the family Carabidae. These
primitive cicindelines invaded an ecological zone probably involving hunting of relatively large, active,
heavily sclerotized prey, and larvae seizing prey from a fixed hiding place. Adults were probably
nocturnal hunters and basically ground beetle-like in behaviour. They did not fly actively. Early
divergence produced two lines, one retained the plesiotypic small, scotopic A eyes and nocturnal
behaviours (the Megacephalini); the other acquired large eyes (ancestors of the Cicindelini.
Within the Megacephalini, two major lineages developed; the Omina, whose adults retained small
eyes, and mainly nocturanl behaviour; and the Megacephalina, whose adults became crepuscular
acquired large eyes for stereopsis, but remained functionally scotopic A. Within the Omina, adults are
secondarily flightless. Adults of Amblycheila plesiotypically have small scotopic A eyes. However, eyes
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of Omus adults have evolved scotopic B eyes capable of finer image resolution for vision during more
frequent diurnal activity periods.
The Cicindelini became divergent and probably initially diversified in the shade of tropical forests,
where representatives of many cicindeline genera now live. Early lineages moved into more open areas
(initially, perhaps, along stream margins), developed quick flight, which could have been a correlate of
the superior binocular vision afforded by large eyes. A lineage with such properties could have been
ancestral to Cicindela, whose species became diurnal, and adapted for life in open areas. This taxon
underwent an evolutionary flowering that produced an abundance of species on all continents (except
Antarctica).
Among the species of Cicindela I examined, I found two functional eye types: plesiotypic scotopic A;
and apotypic photopic. Given only this information, one would be tempted to think that the taxa with
scotopic A eyes were ancestral to those with photopic eyes. However, 1 believe that the reverse is true,
based on the following consideration. Photopic eyes and diurnal activity are characteristic of groups
possessing more primitive male genitalia, and hence believed to represent more more primitive lineages
of the genus. These species and the groups to which they belong (indicated by numbers in Freitag, 1974),
based on Rivalier, (1954) are: group lA - Cicindela repanda and Cicindela limbata\ Group IB -
Cicindela longilabris; Group 1C - Cicindela limbalis\ Group III - Cicindela tranquebarica. On the
other hand, adults of some taxa characterized by highly derived genitalia are crepuscular as well as
diurnal, and have either photopic or scotopic A eyes. These are: Group X - Cicindela belfragei, eyes
photopic; Group XII - Cicindela lepida eyes scotopic A. Adults of Cicindelida pilatei (Group X) and
Cicindelida lemniscata (Group XI) are active both in full light and in dim light, but their eyes have not
been examined histologically.
Because of the nature of the correlations, I infer that diurnal activity and photopic eyes are
plesiotypic in Cicindela, and that crepuscular activity and scotopic A eyes are apotypic. Therefore,
presence of the latter type of eyes in Cicindela represents an evolutionary reversal.
Basically this phylogenetic framework provides a satisfactory continuity of evolution of eye function
through nocturnal to crepuscular, and diurnal to crepuscular diel activity transitions. However, one
abrupt change from nocturnal to diurnal is involved in the divergence of the Cicindelini from the
Megacephalini. It must be mentioned that within the Cicindelini there are four subtribes containing a
total of 16 genera which are more primitive than Cicindela (Horn, 1926). Eyes of adults of these genera
may provide a smooth transition from ancestral small scotopic A eyes through large scotopic A eyes to
still larger photopic eyes of Cicindela adults.
Based on earlier classification (Lacordaire, 1843, 1854; and Thompson, 1857), an alternative
reconstructed phytogeny can be provided. This places the crepuscular Megacephalina as the sister group
of the diurnal Cicindelini. One can then propose that the ancestors of these two taxa were crepuscular,
like the extant members of the Megacephalina. Thus a smooth transition is provided for evolution of
photopic eyes as suggested above. I believe that Horn’s hypothesis is more correct and suggest that in the
strict sense ancestors of Cicindelini had crepuscular eyes. This hypothesis should be tested by
examination of eyes of the more primitive taxa of Cicindelini.
Using Horn’s classification, several assumptions are required for the following events of evolution of
cicindelid compound eyes: divergence in eye size; divergence in eye function; divergence in eye size and
function; parallel acquisition of enlarged eyes; and reversion in function (but not in eye size) to an
ancestral condition. Divergence in eye size alone is exhibited by evolution of large eyes in the
Megacephalina; divergence in function alone, by acquisition of scotopic B eyes in adults of Omus\
divergence in eye size and function by evolving eyes of ancestral Cicindelini. Parallel evolution of eye
size is exhibited by independent acquisition of large eyes in both Megacephalina and Cicindelini.
Compound eyes of Cicindelidae and Carabidae
313
Reversal in function is exhibited by evolution of scotopic A eyes by a highly derived lineage of Cicindela
lepida. Also, in highly derived Cicindela belfragei, there is a reversal from diurnal to crepuscular diel
activity, without change in eye function.
Table 7 shows that, based on my three functional categories, cellular organization in adephagan
beetle eyes has undergone parallel evolution. Parallelism in function is identified in independent
acquisition of scotopic B eyes among Cicindelidae {Omus spp.), Carabidae (Pterostichus melanarius
Table 7. Functional eye categories of adephagan beetle adults.
Quaest. Ent., 1979, 15 (3)
314
Kuster
and other taxa), and Gyrinidae {Gyrinus spp.). Parallelism in eye size and function related to diurnal
activity is shown by Cicindelidae {Cicindela spp.) and Carabidae {Elaphrus spp.). All families but
Gyrinidae have living taxa with ancestral scotopic A eyes. The impression is that parallel acquisition of
the derived types of eyes occurred many times. Reversion to an ancestral functional condition might be
common, though probably less frequent than parallelism.
It is important to recapitulate that modifications are based on eye size and on an alteration of cellular
organization not on a change in cell number in ommatidia. Coadapted to nocturnal activity are small
scotopic A eyes, scotopic B eyes to nocturnal but more frequent diurnal activity; to crepuscular activity,
large scotopic A eyes (except large photopic eyes of Cicindela belfragei); and to diurnal activity, large
photopic eyes.
The mechanism used to evolve large eyes from small eyes is addition of number of ommatidia with an
accompanying shortening of the dioptric apparatus and increased retinula length. The transition from
scotopic A to scotopic B eyes involves elimination of the clear retinula zone by extension of the rhabdom
the complete retinula length. Such a structural modification only involves shortening of the retinula
cells. Changes involved with elimination of the clear retinula zone and reduction of rhabdom surface
area and volume, evolve photopic eyes from large scotopic A eyes, and converse relationships are
required for the opposite transition.
Because slight changes in internal structure have profound effects on function, it is fairly easy for
evolving groups to move from one adaptive zone to another, and back again. Such shifts are generally
correlated with speciation. This means an increase in diversity when such shifts occur, and ultimately
they involve change in eye function. Therefore, it seems likely that the ability of eyes to respond quickly
to selection is an integral component of evolution of diversity among the Adephaga in particular, and
perhaps among insects in general.
ACKNOWLEGEMENTS
For helpful advice and discussion, I wish to thank my supervisor, D.A. Craig, and G.E. Ball. Several
people were involved in collection of beetles used in this investigation. I thank G.E. Ball for collection of
Megacephala Carolina mexicana Gray in Mexico, and D.H. Kavanaugh, California Academy of
Science, for field assistance at Point Reyes National Seashore, California to collect Omus californicus
californicus. Also, I thank E.L. Sleeper, California State University, Long Beach, for his enthusiasm for
my project and his student. M.E. Mispagel for a memorable collection of Amblycheila schwarzi adults
in Joshua Tree National Monument, California (Kuster, 1976). For displaying his world cicindelid
collection, I thank W.D. Sumlin III of Riverside, California. I thank R.R. Murray, Texas A & M
University for specimens of Cicindela belfragei, my colleague, G.J. Hilchie for specimens of Cicindela
lepida from Empress, Alberta, and K.A. Shaw for local field assistance to collect adults of Cicindela.
This work was supported by a Gulf Oil Limited graduate student fellowship, and is based on part of my
Ph.D. thesis.
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Compound eyes of Cicindelidae and Carabidae
319
Figures I - 4. SEM of the frontal aspect of heads of cicindelid beetles, showing variation in eye size and shape. Note verticies (v).
Scale = 500 jam.
Quaest. Ent., 1979, 15 (3)
320
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Figures 5-8. Lateral view of compound eyes of cicindelid beetles, showing hexagonal corneal lenses (1) and ocular sclcrite (os).
Scale = 200 fim. Fig. 1,5; Amblycheila schwarzi\ Fig. 2,6: Omus californicus: Fig. 3,7; Megacephala Carolina-, and Fig. 4,6:
Cicindela tranquebarica.
Compound eyes of Cicindelidae and Carabidae
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Figures 13- 16. LM of transverse sections of compound eyes ofcicindelid beetles. Structural component abbreviations as in Fig. 9-12.
Scale = 100 ^m. Fig. 13: Amblycheila schwarzi; Fig. 14: Omus californicus\ Fig. 15: Megacephala Carolina., and Fig. 16: Cicindela
tranquebarica.
Compound eyes of Cicindelidae and Carabidae
323
Figs. 17-24. SEM of convex, hexagonal corneal lenses (1) of cicindelid beetles. Note cuticular pegs (cp) between some lenses. Scale
= 10 ^m. Figures 21-24. LM of transverse sections through the retinula cells (rt) and rhabdom (r).
Scale = 20 ^m. Fig. 17,21: Amblycheila schwarzi\ Fig. 18,22: Omus californicus; Fig. 19,23: Megacephala Carolina; and Fig. 20,24:
Cicindela tranquebarica.
Quaest. Ent., 1979, 15 (3)
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Figures 25 - 28. Frontal sections through optic lobes of cicindelid beetles, showing axonal bundles (ab); glial cells (gl); lamina
ganglionaris (Ig); first optic chiasmata (Ic); medulla (md); second optic chiasmata (2c); lobula (lo); neurilemma (nl); and perineurium
(pn). Note dense pigment accumulation (pa) on the ventral aspect of optic lobes. Scale = 100 )um. Fig. 25: Amblycheila schwarzi\
Fig. 26: Omus californicus\ Fig. 27: Megacephala Carolina ; Fig. 28 Cicindela tranquebarica.
325
Compound eyes of Cicindelidae and Carabidae
Figures 29 - 32. Diagrammatic longitudinal sections of representative ommatidia and transverse sections of proximal rhabdoms of
four cicindelid beetles, showing thin corneal layer (t); corneal lens (1); subcorneal layer (cl); crystalline cone (c); Semper cells (s);
crystalline thread (ct); distal rhabdom (dr) of retinula cell seven (7); clear retinula zone (cr); proximal rhabdom (pr) of six retinula
cells; basal retinula cell (b) with rhabdomere (br) secondary pigment cells (2p); basal pigment cells (bp); basement membrane (bm);
and eight axons (a). Longitudinal section scale = 50 )um. Transverse section scale = 20 jum. Fig. 29; Amblycheila schwarzi
(scotopic A); Fig. 30; Omus californicus (scotopic B); Fig. 31; Megacephala Carolina (scotopic A); and Fig. 32; Cicindela
tranquebarica (photopic).
Quaest. Ent., 1979, 15 (3)
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Figures 33 - 36. SEM of the frontal aspect of the head of a Cicindela lepida adult, showing large bulbous eyes.
Scale = 500 Fig. 34. Same, of a lateral view of the left compound eye, showing hexagonal corneal lenses (1) and ocular sclerite
(os). Vertex positioned at the left. Scale = 200 Fig. 35. Same, of convex distal surfaces of hexagonal corneal lenses (1). Note
cuticular pegs (cp) between some lenses. Scale = 10 Mm. Fig. 36. Same, of a cuticular peg (cp) of an interfacetal mechanoreceptor.
Note ecdysial scar (es). Scale = 1 ^m.
Compound eyes of Cicindelidae and Carabidae
327
Figures 37 - 39. LM of longitudinal section of the eye of a Cicindela lepida adult. Shown are: thin corneal layer (t); corneal lens (1);
subcorneal layer (cl); crystalline cone (c); clear retinula zone (cr); retinula rhabdom zone (rr); basal retinula zone (br); basement
membrane (bm); secondary pigment cells (2p); basal pigment cells (bp); axons (a); and lamina ganglionaris (Ig).
Scale = 100 ^m. Fig. 38. LM of transverse section of the eye. Structural component abbreviations as above. Scale = 200 /nm. Fig. 39.
Same, through the retinula rhabdom zone, showing retinula cells (rt) and rhabdom (r). Scale = 10 mhi.
Quaest. Ent., 1979, 15 (3)
328
Kuster
Figure 40 - 43. SEM of the frontal aspect of the head of a Cicindela belfragei adult, showing large bulbous eyes.
Scale = 500 /xm. Fig. 41. Same, of a lateral view of the left compound eye, showing hexagonal corneal lenses (1) and ocular sclerite
(os). Vertex positioned at the left. Scale = 200 ^m. Fig. 42. Same, of convex distal surfaces of hexagonal corneal lenses (1). Note
cuticular pegs (cp) between some lenses. Scale = 10 /um. Fig. 43. Same, of a cuticular peg (cp) of an interfacetal mechanoreceptor.
Scale = 1 Mm.
Compound eyes of Cicindelidae and Carabidae 329
Figure 44 - 46. LM of longitudinal section of the eye of a Cicindela belfragei adult. Shown are; thin corneal layer (t); corneal lens (1);
subcorneal layer (cl); crystalline cone (c): retinula rhabdom zone (rr); basal retinula zone (br); basement membrane (bm); secondary
pigment cells (2p); basal pigment cells (bp); axons (a); and lamina ganglionairis (Ig). Scale = 100 ^m. Fig. 45. LM of transverse
section of the eye. Structural component abbreviations as above. Scale = 200 /urn. Fig. 46. Same, through the retinula rhabdom zone,
showing retinula cells (rt) and rhabdom (r). Scale = 10 Mm.
Quaest. Ent., 1979, 15 (3)
330
Kuster
Figure 47 - 50. SEM of the frontal aspect of the head of a Pterostichus melanarius adult, showing relatively flat eyes.
Scale = 500 /iim. Fig. 48. Same, of a lateral view of the left compound eye, showing hexagonal corneal lenses (1) and ocular sclerite
(os). Vertex at the top. Scale = 200 ^m. Fig. 49. Same, of convex distal surfaces of hexagonal corneal lenses (1). No interfacetal pegs
are present. Scale = 10 /um. Fig. 50. Same, of dermal glands surrounding the eye. Glands secrete a material (x) which spreads over
the ocular sclerite (os) and some corneal lenses (1). Scale = \0 nm.
Compound eyes of Cicindelidae and Carabidae
331
Figures 51 - 53. LM of longitudinal section of the eye of a Pterostichus melanarius adult. Shown are; thin corneal layer (t); corneal
lens (1); subcorneal layer (cl); crystalline cone (c); retinula rhabdom zone (rr); basal retinula zone (br); membrane (bm); secondary
pigment cells (2p); basal pigment cells (bp); axons (a); and lamina ganglionaris (Ig). Scale = 100 /xm. Fig. 52. LM of transverse
section of the eye. Structural component abbreviations as above. Scale = 100 jum. Fig. 53. Same, through the retinula rhabdom zone,
showing retinula cells (rt) and rhabdom (r). Scale == 10 ^xm.
Quaest. Ent., 1979, 15 (3)
332
Kuster
Figures 54 - 57. SEM of the frontal aspect of the head of an Elaphrus americanus adult, showing large bulbous eyes.
Scale = 200 Fig. 55. Same, of a lateral view of the left compound eye, showing hexagonal corneal lenses (I) and ocular sclerite
(os). Vertex positioned at the left. Scale = 100 pm. Fig. 56. Same, of convex distal surfaces of hexagonal corneal lenses (1). Note
cuticular pegs (cp) between some lenses. Scale = 10 /am. Fig. 57. Same of a cuticular peg (cp) of an interfacetal mechanoreceptor.
Scale - 1 ^m.
Compound eyes of Cicindelidae and Carabidae
333
Figures 58 - 60. LM of longitudinal section of the eye of an Elaphrus americanus adult. Shown arc: thin corneal layer (t); corneal lens
(1); subcorneal layer (cl); crystalline cone (c); retinula rhabdom zone (rr); basal retinula zone (br); basement membrane (bm);
secondary pigment cells (2p); basal pigment cells (bp); axons (a); and lamina ganglionaris (Ig).
Scale = 100 ;um. Fig. 59. LM of transverse section of the eye. Structural component abbreviations as above. Scale = 200 yum. Fig. 60.
Same, through the retinula rhabdom zone, showing retinula cells (rt) and rhabdom (r). Scale = 10 pm.
Quaest. Ent., 1979, 15 (3)
Reconstructed Phylogeny of North American Genera of Cicindelidae
Based on Horn, 1926 and Compound Eye Structure and Function
Kuster
334
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INSECT DAMAGE TO OLD OAK BEAMS AT LOWER FORT GARRY, MANITOBA
HORNE R. WONG
Northern Forest Research Centre
Canadian Forestry Service
Environment Canada Quaestiones Entomologicae
Edmonton, Alberta T6H 3S5 1 5: 335-339 1 979
Old oak beams used in construction of ceilings of first and second storeys and lintels of the
limestone warehouse, Lower Fort Garry, Manitoba, were attacked by Dermestes lardarius, Priobium
sericeum, and Pseudohadrotoma sp. The insects were probably brought in on goods or originated
locally.
Les vieilles poutres de chene utilisees dans la contraction des plafonds da premier et da deuxieme etages, et des linteaux des
entrepots en pierre a chaux de Lower Fort Garry, Manitoba,- ont subi I’attaque de Dermestes lardarius, Priobium sericeum et
Pseudohadrotoma sp. Ces insectes furent probablement int rod aits avec des marchandises, on bien etaienl d’origine locale.
INTRODUCTION
Lower Fort Garry was built 32 km north of Winnipeg after a disastrous flood ruined old Fort Garry
(Winnipeg) at the forks of the Red and Assiniboine rivers in 1826 (Goldring, 1970). Construction was
started in 1831 and finished in 1847 with completion of the walls surrounding the buildings. In 1963, an
extensive restoration of the fort began. In the process of restoring the old warehouse building,
H. Van der Putten, Project Manager of Restoration of Lower Fort Garry, observed considerable insect
damage (Fig. 6,7) to the old oak beams. Some of the damage resembled that caused by the death watch
beetle, Xestobium rufovillosum (De Geer), a destructive insect introduced from Europe and known to
attack oak timbers in old buildings.
The purpose of this study was to identify the insect species damaging oak beams and speculate on
their origin.
THE WAREHOUSE BUILDING
The three-storey limestone warehouse building (Fig. 1), constructed between 1835 and 1845
(Van der Putten, pers. comm.), measures 21 m long and 9 m wide. Oak used for the ceilings of the first
and second storeys and lintels over the windows of the warehouse was floated down the Red River to the
fort from the United States and areas around the fort (Van der Putten, pers. comm.). The wood was cut
into beams measuring 9 m long, 17.5-22.5 cm wide, and 20.0-22.5 cm high; the beams were spaced
60-75 cm apart. The lengths of the lintels varied from 1 .8 to 7.2 m.
The building was originally used by the Hudson’s Bay Company to house supplies for the fur trade,
but it also served several subsequent functions: a barracks for the military (1846-48), the first provincial
and later federal penitentiary (1871-77), and Manitoba’s first asylum (1885-86) (Goldring 1970). It
was while the warehouse was used as a penitentary that a lime wash, still evident today (Fig. 2), was
painted over the beams and lintels (Van der Putten, pers. comm.).
336
Wong
METHODS AND MATERIAL
Infested oak beams from the warehouse were cut into small sections and shipped to the Northern
Forest Research Centre in late November 1976. These were split into smaller pieces and examined for
insects within the wood. Adults for identification were obtained by placing those pieces containing living
larvae in 73.12-liter galvanized metal garbage cans, modified as in Fig. 4. The cans were placed in a
rearing room with a temperature of 21 C and relative humidity of 50%. Daily observations were made
for adult emergence.
INSECT SPECIES AND DISCUSSION
Three species of insects were found in the oak beams.
Dermestes lardarius Linnaeus
This was the most abundant species present, observed throughout the beams, although rarely in
lintels or the spruce floor. All larvae, pupae, and adults in the wood were dead (Fig. 6). Their presence in
the oak beams can be attributed to the building having been used to store supplies of the Hudson’s Bay
Company. Full-grown larvae, which feed on furs, hides, and anything of animal origin, are known to
wander about in search of some compact material in which to bore a tunnel to pupate (Hinton 1945).
The compact material near at hand was the oak beams and spruce floor. Instances of the larder beetle
damaging structural wood are common. Hinton (1945) listed a number of such reports, and Craighead
(1950) and Baker (1972) noted that valuable cargos of lumber have been damaged in ship holds where
hides had previously been stored.
It is not known for certain what killed the beetles in the wood. A chemical analysis of the dead larvae
and adults did not disclose any toxic chemicals such as lead, which was present at 140 ppm in the lime
wash. One explanation that may account for the sudden death of numerous larvae, pupae, and adults in
the wood is cold winter temperatures. The warehouse was probably left unheated for at least one year
before the military took over in 1846. If this is so, then dead larvae, pupae, and adults of the larder
beetle must have been entombed in the wood for over a hundred years.
We can only speculate on the possible origin of the larder beetles in the warehouse. Records indicate
that there was little contact between Lower Fort Garry and eastern Canada (Van der Putten, pers.
comm.), but supplies were frequently received from England. It would appear then that this insect
originated from England on or in any number of things listed by Hinton (1945).
Priobium sericeum (Say)
The numerous small exit holes and dustlike powder on the oak beams indicated that the beams had
been attacked by an anobiid beetle. Adults obtained from rearing the larvae in the beams were not death
watch beetles, Xestobium rufovillosum, as expected, but another member of the same family, Priobium
sericeum (Fig. 3). Larvae and exit holes were found at the end of the beams up to 1.2 m from the wall
and in the lintels. Adults of this species emerged in the laboratory on 14 February and 14 March 1977
and 5 January 1978.
This is a native species, which White (1962) recorded from dead branches of oak, wild cherry, and
hickory. According to Simeone (1962), it is distributed from Nova Scotia to Wisconsin in the north,
southwestward to Arizona, and southeastward to Texas and Florida. The Canadian National Collection,
Ottawa, has specimens only from Nova Scotia, New Brunswick, Quebec, and Ontario. This is the first
record of Priobium sericeum in Manitoba.
Insect damage to Lower Fort Garry
337
After the oak was floated down the Red River to Lower Fort Garry, it probably became infested by a
local population of Priobium sericeum, which laid eggs in exposed wood such as knots, cracks, and
crevices. After hatching the larvae entered the wood and riddled it with tunnels, packed with powdery
dust. The small round surface holes mark exits of adults of Priobium sericeum. Populations of this insect
started either outside the warehouse when the oak was stacked ready for use or within the warehouse
from infested oak used in ax handles, farm implements, furniture, or frame-work of Red River carts.
Pseudohadrotoma sp.
Only four adults (Fig. 5) of this species were obtained. Dr. E.C. Becker, Biosystematic Research
Institute, Ottawa, indicated that it is undescribed. Adults emerged on 2 and 14 February and
19 December 1977, and 4 June 1978. In Beal’s key (1967) to species of Pseudohadrotoma, the new
species is close to Pseudohadrotoma falsa (Horn), which Beal indicated as being beaten from Quercus
and which is probably a general scavenger feeding on the remains of other insects in burrows of wood.
This new species probably fed on the remains of larder beetles in the oak beams and did not cause
primary damage to the wood.
Of the three species of insects reared from infested oak beams at Lower Fort Garry, Manitoba, only
Dermestes lardarius and Priobium sericeum caused primary damage. Individuals of the former species,
although most abundant, were all killed, apparently by extreme cold weather when the warehouse was
left unheated for a year. Those of the latter species are still thriving.
ACKNOWLEDGMENTS
Grateful acknowledgment is extended to Henri Van der Putten, Project Manager, Lower Fort Garry
Restoration Project for historical information and material for this study. Thanks are also extended to
M. Tarr, superintendent, and A. Buziak, interpretive officer, Indian and Northern Affairs, Parks
Canada, Selkirk, Manitoba, for their interest; and to V. Hildahl, Canadian Forestry Service, Winnipeg
for his valuable assistance. I am indebted to Dr. Richard E. White, Systematic Entomology Laboratory,
USDA, Washington, D.C. and Dr. E.C. Becker, Biosystematic Research Institute, Ottawa, Ontario, for
the identification of Priobium sericeum (Say) and Pseudohadrotoma sp. respectively; to P.S. Debnam,
Northern Forest Research Centre, Edmonton, Alberta, and Chris Grant, Indian and Northern Affairs,
Parks Canada, Selkirk, Manitoba, for the photographs; and to Y.P. Kalra, Northern Forest Research
Centre, for the chemical analyses in this study.
REFERENCES
Baker, W.L. 1972. Eastern forest insects. U.S. Department of Agriculture Miscellaneous Publication.
No. 1175: 1-642.
Beal, R.S. Jr. 1967. A revisionary study of the North American dermestid beetles formerly included in
the genus Perimegatoma (Coleoptera). Entomological Society of America. Miscellaneous publication
No. 5:279-312.
Craighead, F.C. 1950. Insect enemies of eastern forests. L.S. Department of Agriculture. Miscellaneous
Publication No. 657:1 -679.
Goldring, P. 1970. Lower Fort Garry. Beaver. Summer; 26-28.
Hinton, H.E. 1945. A monograph of the beetles associated with stored products. British Museum.
Natural History Publication. 1: 275-285.
Quaest. Ent. 1979, 15 (3)
338
Wong
Simeone, J.B. 1962. Survey of wood-feeding Anobiidae in north-eastern United States, including a study
of temperature and humidity effects on egg development of Hadrobregmus carinatus Say.
Proceedings of the Xlth International Congress of Entomology 2: 326-335.
White, R.E. 1962. The Anobiidae of Ohio (Coleoptera). Ohio Biological Survey 1(4): 1-58.
Insect damage to Lower Fort Garry
339
Figures 1 - 7. Fig. 1, limestone warehouse, Lower Fort Garry, Manitoba; Fig. 2, oak beams on ceiling showing whitewash and
section of beam restored; Fig. 3, adult of Priobium sericeum just emerged from oak beam; Fig. 4, modified galvanized metal
garbage can used for rearing insects in oak beams; Fig. 5, adult of Pseudohadrotoma sp.; Fig. 6, dead adults of Dermestes
lardarius in oak beam; Fig. 7, holes in oak beam made by Dermestes lardarius and Priobium sericeum.
Quaest. Ent. 1979, 15 (3)
OBSERVATIONS ON THE SNOW SCORPIONFLY BOREUS BRUMALIS FITCH
(BOREIDAE: MECOPTERA) IN SUDBURY, ONTARIO
J.D. SHORTHOUSE
Department of Biology
Laurentian University
Sudbury, Ontario Quaestiones Entomologicae
P3E2C6 15:341-344 1979
Adults of the snow scorpionfly, Boreus brumalis Fitch, are common on the surface of snow from
February to March within the city limits of Sudbury, Ontario when the temperature rises above 0 C.
Ease of locating a mate and dispersal are the suggested reasons for this mid-winter activity.
Les adultes du Mecoptere Boreus brumalis Fitch sont communs sur la neige dans la ville de Sudbury, Ontario, en fevrier et en
mars, quand la temperature s'eleve au-dessus de 0 C. Le fait que la rencontre des partenaires sexuels et la dispersion soient fad les
est suggere comme cause d’une telle activite hivernale.
The occurrence of active insects on the surface of snow in mid-winter is to most biologists an
extraordinary event. Snow scorpionflies of the genus Boreus exhibit this behavior as a regular part of
their life cycle (Crampton, 1940; Byers 1954; Chapman, 1954; Webb et ai, 1975; Penny, 1977) and
although they are not rare insects, they are seldom observed. This paper reviews the biology of Boreus
brumalis Fitch and records the species on snow-covered hills within the city limits of Sudbury, Ontario
(46°30'N, 81°00'W) in February and March of 1977 and 1978. Suggestions also are given as to why
Boreus brumalis emerges onto the surface of the snow.
The Family Boreidae consists of two genera, Boreus and Hesperoboreus (Penny, 1977) with
members of the genus Boreus occurring in Europe, Asia, and North America. Ten species of Boreus are
recorded from North America with only two, Boreus brumalis and Boreus nivoriundus Fitch occurring
east of the Rocky Mountains (Penny, 1977). Boreus brumalis is considered the most common North
American species (Byers 1954). It is usually found in deciduous woodlands and has been recorded from
central Ontario and Quebec south to Tennessee, with the western limits being isolated populations in
Illinois, Wisconsin, and Minnesota (Webb et ai, 1975).
Both male and female Boreus brumalis are small (2.5 mm to 5.0 mm), stout, black insects with biting
mouthparts on the ventrally extended rostrum, a characteristic common to the Order (Richards and
Davies, 1977). The legs and antennae are long and black. Wings of the male are reduced to a pair of
slender vestiges, tapering apically to an acute point (Fig. 1), with coarse black setae found along the
lateral and medial margin. The hind wings are much smaller and lie in a groove in the fore-wing. The
genital claspers are flexed dorsally over the abdomen giving the scorpion-like appearance. Wings of the
female are further reduced to scale-like, oval lobes (Fig. 2). The functional ovipositor is elongated and
consists of modified 9th to 11th abdominal segments (Cooper, 1940). For further details on the
morphology of Boreidae, see Penny, 1977.
Both the immatures and adults of all species live in and feed on mosses. The eggs are laid in late fall
at the base of mosses and the larvae hatch in about ten days (Webb et al., 1975). Crampton (1940)
reported that larvae can be found at all times of the year implying that larvae feed throughout the
winter. The larvae form small chambers in compacted soil and it is thought that they aestivate
throughout part of summer in these chambers (Fraser, 1943; Webb et al.. 1975). Descriptions of Boreus
larvae and pupae are given by Withycombe (1922). Pupation occurs in the chambers (Fraser, 1943) and
the pupal stage lasts from 4 to 8 weeks. Adults of the European species Boreus hyemalis (L) emerge in
342
Shorthouse
late fall and are found crawling around on mosses (Withycombe, 1922; Fraser, 1943). Adults of the
North American species also emerge in the fall with Illinois populations known to crawl onto the surface
of the snow from November to April (Webb et al., 1975). The adults presumably continue feeding on
mosses beneath the snow throughout the winter. The adults feed by thrusting their rostrum into the
young shoots of mosses, chewing off the apex, then eating down into the core (Fraser, 1943).
Figure 1 and 2. Fig. 1, Male Boreus brumalis. Fig. 2, Female Boreas brumalis. Magnification X 15.
The life cycle of Boreus brumalis in Sudbury is likely similar to that of brumalis in Illinois (Webb
et al., 1975). No adults were observed until mid-winter; however it is possible that some adults emerge in
the fall and remain under the snow for several months before crawling to the surface. The first adults
were found on 12 February 1977 and were common until 19 March on days with suitable temperatures.
No adults were seen in 1978 before 4 March. Many copulating pairs were found indicating that
oviposition also takes place in late winter.
Adults come to the surface along passage-ways found next to the stems of shrubs and trunks of trees.
Apparently they only come to the surface when the air temperature approaches 0 C (Wojtusiak, 1950;
Chapman, 1954), although one European species has been reported active at -5.5 C (Fjellberg and
Greve, 1968). On 12 February 1977, the air temperature in Sudbury had risen to 2.3 C and on 19 March
to 2.7 C, both being days that followed periods of subzero temperatures. The same occurred on
11 March 1978 when the temperature was 2.4 C. The adults were always most common and active
aroung 1400 hours; however, they retreated beneath the snow around 1700 hours.
The sex ratio of the 107 specimens collected in Sudbury was 0.57, similar to that recorded for other
species. (Fjellberg and Greve, 1968). Adults are easily spotted against the white snow and often were as
common as 1 per 4m2, The largest populations were found in serai white birch-red maple-red oak forests
on south-facing slopes (Fig. 3). The snow depth on dates of collection was about 30 cm.
The snow scorpionfly Boreus brumalis
343
Figure 3. Typical winter habitat of Boreus brumalis on a hillside dominated by white birch, red maple, and red oak within the
city limits of Sudbury, Ontario.
The mating behavior of Boreus species is peculiar and has been described for several species
(Cockle, 1908; Withycombe, 1926; Cooper, 1940; Crampton, 1940). The Boreus brumalis male upon
finding a female, springs and seizes her body with his modified wings. Once she is securely gripped, the
male uses his hind legs and claspers to manoeuvre her into a position parallel to and above his body. He
then releases his wings allowing the female to flex her rostrum between his coxae, fold her antennae
between her legs, and stretch her legs posteroventrally. Once in this position the male then grips her
fore-legs and rostrum with his wings. This position with the female above the male is maintained
throughout copulation. The male usually runs about during copulation while the female remains
motionless.
The ability to remain active at low temperatures and to use the snow surface as a habitat is a
fascinating adaptation shared by few other arthropods. Besides Boreus, only a few species of
Chironomidae, Heleomyzidae, Tipulidae, Trichoceridae, Plecoptera, Collembola, and spiders have been
reported using the snow surface (Wojtusiak, 1950; Hagvar, 1973; Hagvar, and Ostbye, 1973). Some
spiders even form webs on the surface where they trap collembolans.
Little has been written as to reasons why scorpionflies climb to the surface in mid-winter. The
obvious benefit would be ease of movement since the smooth surface of snow provides an alternative
terrain with few impediments to dispersal. Some individuals were observed hopping a distance of 2 m in
5 minutes, while one was found 70 m from shore on a snow covered lake, obviously far from its site of
emergence. Ease of locating the opposite sex also would be enhanced in this environment while the risk
of attack by predators would be reduced. Some spiders were found on the surface at the same time as the
Sudbury Boreus, but they were not observed feeding.
The temperature of the subnivean air space remains near 0 C with about 30 cm of snow cover,
regardless of the external air temperature (Coulianos and Johnels, 1962) and this is obviously
sufficiently warm for scorpionflies to locate stems and tree trunks and climb to the surface. Once on the
surface their dark coloration would absorb heat thus raising the body temperature higher than it would
Quaest. Ent., 1979, 15 (3)
344
Shorthouse
have been under the snow. This additional heat probably contributes to their improved ability to disperse
on the snow surface. How they get back under the protective snow cover when the air temperature drops
and the cue for their return remain as fascinating problems.
Although members of the genus Boreus are excellent candidates for further studies of cold hardiness,
their greatest value may come from serving educators with another intriguing example of insect
adaptability. Educators and biologists alike should be on the alert for winter insects since they vividly
contradict the common misunderstanding that all in the insect world is inactive from October to April.
I thank M. Derro for his help in making the plates and G.M. Courtin for his comments on the
manuscript. This paper was supported in part by NSERC grant number A0230.
REFERENCES
Byers, G.W. 1954. Notes on North American Mecoptera. Entomological Society of America Annals
47:484-510.
Chapman, J.A. 1954. Observations on snow insects in Western Montana. The Canadian Entomologist
86: 357-363.
Cockle, J.W. 1908. The mating of Boreus californicus. The Canadian Entomologist 40: 101 .
Cooper, K.W. 1940. The genital anatomy and mating behavior of Boreus brumalis Fitch (Mecoptera).
American Midland Naturalist 23: 354-367
Coulianos, C.-C. and A.G. Johnels. 1962. Note on the subnivean environment of small mammals. Arkiv
Fur Zoologi 15: 363-370.
Crampton, G.C. 1940. The mating habits of the winter mecopteron Boreus brumalis Fitch. Psyche
47:125-128.
Fjellberg, A. and L. Greve. 1968. Notes on the genus Boreus in Norway. Norsk Entomologisk Tidsskrift
15: 33-34.
Fraser, F.C. 1943. Ecological and biological notes on Boreus hyemalis (L) (Mecopt. Boreidae). Journal
of the Society for British Entomology 2: 125-129.
Hagvar, S. 1973. Ecological studies on a winter-active spider Bolyphanthes index (Thorell) (Araneida,
Linyphiidae). Norsk Entomologisk Tidsskrift 20: 309-314.
Hagvar, W. and E. Ostbye. 1973. Notes on some winter-active Chironomidae. Norsk Entomologisk
Tidsskrift 20: 253-257.
Penny, N.D. 1977. A systematic study of the Family Boreidae (Mecoptera). The University of Kansas
Science Bulletin 51: 141-217.
Richards, O.W. and R.G. Davies. 1977. Imms’ General Textbook of Entomology. Tenth Edition.
Volume 2: Classification and Biology. Chapman and Hall, London. 421-1354.
Webb, D.W., N.D. Penny and J.C. Marlin. 1975. The Mecoptera, or scorpionflies of Illinois. Illinois
Natural History Survey Bulletin 31: 251-316.
Withycombe, C.L. 1922. On the life-history of Boreus hyemalis L. Entomological Society of London,
Transactions 1921: 312-318.
Withycombe, C.L. 1926. Additional remarks upon Boreus hyemalis L. Entomologist’s Monthly
Magazine 62: 81-83.
Wojtusiak, H. 1950. The temperature preferendum of winter insects of the genus Boreus (Panorpatae)
and Chionea (Diptera). Academie Polanaise des Sciences et des Lettres. Bulletin International.
Series B. 11: 125-143.
NOTES ABOUT CLASSIFICATION OF THE SOUTH AMERICAN PTEROSTICHINI WITH A
KEY FOR DETERMINATION OF SUBTRIBES, GENERA AND SUBGENERA
(COLEOPTERA: CARABIDAE)
S.L. STRANEO
V. Campari 8E
27100 Pavia Quaestiones Entomologicae
ITALY 15:345-356 1979
Treated in the key, which is based on characters of adults, are 37 genera and 20 subgenera, arrayed
in the following seven subtribes: Morionina (two genera); Cratocerina (one genus); Microcephalina (one
genus); Euchroina (four genera); Pterostichina (24 genera, 18 subgenera); Antarctiina (four genera);
and Chaetogenyina (one genus, two subgenera). Excluded from the Pterostichina and ranked as tribes
are the Catapiesini and Agonini.
Tratados en la Have, el cual estd basado en el original de los adultos, se encuentran 37 generos y 20 suh-generos, compiladus en
los siete siguientes sub-tribus: Morionina (dos generos); Cratocerina (un genero); Microcephalina (un generoj; Euchriona (cuatro
genero); Pterostichina (24 generos, 18 sub-generos); Antarctiina (cuatro generos): y Chaetogenyina (un genero, dos sub-generos).
Excluidos del Pterostichini y clasificados como tribus estdn los Catapiesini y Agonini.
INTRODUCTION
In 1976, the late Hans Reichardt (Museu de Zoologia da Universidade de Sao Paulo, Brazil)
suggested to me a revision at the generic level of the South American Pterostichini, which he needed for
his proposed synopsis of the genera of Neotropical Carabidae. This work was published posthumously, in
an incomplete state (Reichardt, 1977). Preparation of my contribution was delayed for a variety of
reasons, and I did not know about plans for completion of the general synopsis, following Dr. Reichardt’s
sudden and untimely death. The synopsis of the South American Pterostichini was published
(Straneo, 1977) in Italian. Some months ago, George E. Ball invited me to write an English translation
of my key for publication in Quaestiones Entomologicae, as a supplement to Reichardt’s work.
The key I offer here is a translation with some modifications in style, as well. 1 have chosen to restrict
the key and preliminary classification to taxa that are either confined to South America, or whose
relatives are in South America. Thus, I do not deal with the northern elements of the Neotropical fauna,
except for Dyschromus, which is the only genus of the Euchroina known from the northern hemisphere.
Only a few references are given here. References to descriptions of taxa can be obtained from Csiki
(1929 and 1930), Blackwelder (1944 and 1957) and Reichardt (1977).
Previously (Straneo, 1977), I presented a list of the generitypes of genera and subgenera of South
American Pterostichini. Generitypes of the following taxa were fixed in that paper: Oribazus Chaudoir;
Meropalpus Tschitscherine; Pseudabarys Chaudoir; Cynthidia Chaudoir; Ogmopleura Tschitscherine;
Trirammatus Chaudoir; Feroniomorpha Tschitscherine; Parhypates Motschulsky; Agraphoderes Bates;
Eutanys Tschitscherine; Adrimus Bates; Feroniola Tschitscherine; and Antarctiola Straneo. Also, the
genus Cephalostichus was described as new.
NOTES ABOUT CLASSIFICATION
Two types of problems were encountered in study of classification of the South American
Pterostichini. First is the matter of inclusion and exclusion of reasonably clearly defined groups of
346
Straneo
genera. Second is the matter of ranking at the genus-group level.
I exclude from the Pterostichini the catapiesines, based on conclusions of Reichardt (1973), though
the exact position of the group remains to be determined. Presently, it is ranked as a tribe, and placed
near the truncatipennian assemblage of tribes. 1 also exclude the agonines, which 1 regard as a group
related to, but at the same rank as, the pterostichines, even though various recent authors (for example
Lindroth 1966: 441-442; Erwin et al., 1977: 4.25-4.32; and Reichardt, 1977: 406) combine these two
groups in a single tribe.
I include as a subtribe of the Pterostichini the morionines because, first, many character states of
adults of these two groups are shared, and second, because morionines are traditionally regarded as a
subgroup of the Pterostichini, and at present carabid specialists are not unanimous about placement and
ranking of morionines. Thus, it seems best to follow tradition.
Ball (in Reichardt, 1977: 408) questioned inclusion of the chaetogenyines in the Pterostichini, but for
the present, I prefer to maintain the status quo.
Inclusion of the Cratocerina, Microcephalina {^Tichoniina Reichardt 1977: 407), Euchroina, and
Antarctiina in the Pterostichini has not been challenged by other recent authors.
Ranking at the level of genus-group (genera and subgenera) is a problem in the Pterostichini. Many
of these taxa were ranked by previous authors as subgenera of Pterostichus (see, for example, Csiki,
1930 and Blackwelder, 1944: 35). Provisionally, at least, I think it best to exclude Pterostichus {sensu
latiore) from the southern hemisphere by removing from it all South American species. Future studies
will have to be made to determine relationships of these taxa to their northern counterparts.
Limits of many of the South American genera are not well defined on the basis of adult character
states, and I feel uncertain about the validity of some of these groups. Some groups ranked by me as
subgenera will probably be treated as genera by other authors. As well, some of my genera will probably
be combined by other authors. In brief, the present arrangement is unstable, and the South American
pterostichine fauna is a rich field for systematic research.
In view of the circumstances outlined above, it may seem presumptuous to offer a key for
determination of the higher taxa of South American Pterostichini. However, a provisional key seems
better than nothing, and I hope that it will be useful both as a stepping stone to development of a better
one, and above all, as a stimulus to investigate the interesting, highly diverse and divergent Neotropical
pterostichine fauna.
Key to adults of subtribes and genera of the South American Pterostichini
1(0) Anterior tibia markedly dilated toward apex. Antennomeres 4-10 moniliform (i.e., each
article thickened, and about as wide as long 2
T Anterior tibia average, not markedly dilated apically. Antennomeres 4-10 filiform,
articles not thickened, and either distinctly longer than wide (most taxa) or as long as
wide 4
2(1) Anterior tibia with terminal angle extended as evident tooth; position of scutellum at base
of elytra normal. Body more or less “scaritoid” and pedunculate
MORIONINA 3
2' Anterior tibia with terminal angle not extended as evident tooth. Scutellum forward in
relation to bases of elytra. Body convex, not pedunculate
CRATOCERINA, Cratocerus Dejean
3(2) Mentum with tooth bilobed. Length of body more than 12 mm Morion Latreille
y Mentum with tooth simple. Length of body less than 12 mm Moriosomus Motschulsky
4 ( L) Labium with apex of ligula extended forward, and with long setae (Fig. 4). Other
Classification of South American Pterostichini
347
mouthparts and basal antennomeres with some long and stiff setae
CHAETOGENYINA, Camptotoma {sensu latiore) Reiche 5
4' Ligula with apex not prominently prolonged forward (Fig. 1,2,3)- Antennomeres and
mouthparts without supernumerary long and stiff setae 6
5(4) Head with single pair of supraorbital setigerous punctures. Pronotum with single pair of
lateral setigerous punctures, anterior pair absent Camptotoma {sensu stricto)
5' Head with two pairs of supraorbital setigerous punctures. Pronotum with two pairs of
lateral setigerous punctures Camptotoma (Chaetogenys) van Emden
6 ( 4') Apex of labial paraglosssa with few setae (Fig. 3) ANTARCTIINA 7
6 Apex of paraglossa without setae (Fig. 1,2) 10
7(6) Penultimate labial palpomere with row of setae (more than two). Pronotum with two
pairs of lateral setigerous punctures in anterior half. Anterior surface (ventral surface, in
repose) of each femur with one or two rows of setigerous punctures, latter more numerous
on posterior femora. Color red-brown, body very convex, short and stout, elytra only
slightly longer than wide. Range - San Ambrosio Island, Chile Kuschelinus Straneo
1' Penultimate labial palpomere bisetose. Body more elongate, more slender, and less
convex 8
8 ( 7") Anterior surface of femur with row of long setae. Abdominal sterna I V-Vl each with
transverse row of setae Antarctiola Straneo
8' Anterior surface of femur with one or two setae only. Abdominal sterna IV-V each with
single pair of setae, only 9
9 ( 8') Penultimate tarsomere of each tarsus markedly bilobed Abropus Waterhouse
9' Penultimate tarsomere not bilobed, average for Pterostichini Metius Curtis
10 ( 6') Sternum VI with setigerous punctures deep and rather large, located near transverse
mid-line (Fig. 6). Abdominal sterna IV-VI each distinctly and transversely bordered
basally. Elytron without scutellar stria. Terminal palpomeres various 1 1
10' Sternum VI with setigerous punctures normal, not foveate; OR subfoveate, but located
along apical margin (Fig. 7). Scutellar stria present or absent. Terminal palpomeres
subcylindrical or fusiform PTEROSTICHINA 1 5
11 (10 )Terminal labial palpomere subcylindrical or fusiform. Elytral interval 3 with one or more
discal setigerous punctures EUCHROINA (in part), Bothynoproctus Tschitscherine
1 r Terminal labial palpomere more or less triangular. Elytral interval 3 without discal
setigerous punctures 12
12 (1 r) Penultimate labial palpomere plurisetose (Fig. 5). Terminal maxillary and labial
palpomeres dilated apically, latter with width at apex subequal to length of medial
margin. Body form like that of an Abax adult. Dorsal surface brilliantly metallic
MICROCEPHALINA, Tichonilla Strand
12' Penultimate labial palpomere bisetose EUCHROINA (in part) 13
13 (12') Body length more than 20 mm. Head and prothorax blue green metallic, elytra coppery
with pronounced metallic luster. Elytron with striae deeply impressed, intervals convex ...
Lobobrachus Sharp
13' Body length less than 20 mm. Color and elytral sculpture various 14
14 (13') Labial mentum with epilobe not extended to apex of lateral lobe; ligula moderately
protruded beyond mental tooth; terminal labial palpomere of male markedly dilated and
triangular, apical and lateral margins subequal; of female, less dilated, with medial and
apical margins subequal. Elytron with striae deeply impressed, intervals convex
Euchroa Brulle
Quaest. Ent., 1979, 15 (3)
348
Straneo
Figures 1 - 7. Fig. 1-4. Outline drawings of apical portion of prementa of various pterostichines. Fig. 1 and 2, paraglossae without
apical setae; Fig. 3, paraglossae setose (Antarctiina); Fig. 4, ligula setose (Chaetogenyina). Fig. 5. Outline drawing of labial
palpomeres 2 and 3 of Microcephalina. Fig. 6 and 7. Outline drawings of ventral aspect of abdominal sterna IV-VI. Fig. 6, sterna
sulcate, margined along base, and with setigerous punctures of sternum VI mediad (Euchroina); Fig. 7, regular or average sterna
for Pterostichini.
Classification of South American Pterostichini
349
14' Mental epilobe extended to apex of lateral lobe; ligula more elongate, markedly projected
beyond apex of mental tooth; terminal labial palpomere less dilated apically (in male,
apical margin much shorter than medial margin; in female, still less dilated). Elytron
with striae less deeply impressed, intervals only slightly convex. Range - mountains of
central Mexico, and island of Hispaniola Dyschromus Chaudoir
15 (10') Antenna short, antennomeres 4-10 each slightly wider than long. Abdominal
sterna IV-VI each sulcate along base, or with vestigial sulcus each side, only
Cephalostichus Straneo
15' Antenna average, elongate, antennomeres 4-10 distinctly longer than wide. Abdominal
sterna sulcate or not 1 6
16 (15')Sterna IV-VI each entirely transversely sulcate and margined basally 17
16' Sterna IV-VI not entirely sulcate and not bordered along basal margin, with or without
short sulcus each side 29
17 (16 ) Elytron with only five deeply impressed striae; intervals markedly convex, catenate
Oribazus Chaudoir
17' Elytron normally striate; intervals uninterrupted throughout length of disc 18
18 ( 17') Elytron with interval 3 impunctate 19
18' Elytral interval 3 with one, two, or three setigerous punctures 22
19 (18 )Alternate intervals of elytron with irregular impressions. Pronotum with single
postero-lateral impression each side Apsaustodon Tschitscherine
19' Intervals of elytron smooth, without irregular impressions 20
20 (19') Dorsal surface of elytra with blue-violaceous metallic luster. Pronotum with two
postero-lateral impressions each side. Body length more than 10 mm. Setigerous
punctures of abdominal sternum VI slightly larger than usual Haplobothynus Tschitscherine
20' Elytra with dorsal surface black, without metallic luster 21
21 (20') Larger, body length 14-15 mm. Body slender, slightly convex. Dorsal surface of elytra
black, glossy, but not iridescent (Middle American specimens of this genus have the
elytra distinctly iridescent [geb]) Ophryogaster Chaudoir
21' Smaller, body length about 8.5 mm. Elytra markedly convex, dorsal surface slightly
iridescent Hybothecus Chaudoir
22 (18 )Tarsal claws pectinate Abaridius Chaudoir
22' Tarsal claws smooth, not pectinate 23
23 (22') Elytron with interval 3 impunctate Pseudabarys Chaudoir
23' Elytron with interval 3 at least bipunctate 24
24 (23') Transverse sulcus of each abdominal sternum with large and deep punctures
Sierrobius (sensu latiore) Straneo 25
24' Transverse sulci of abdominal sterna impunctate 26
25 (24 )Body slender. Apical blade of median lobe of most males asymmetric, right side more
developed than left Sierrobius (sensu stricto)
25' Body stouter. Median lobe either symmetrical, or left side slightly more developed than
right Sierrobius (Pachyabaris) Straneo
26 (24 ) Elytron with scutellar stria more or less developed. (Body form like that of Poecilus
cupreus, or Poecilus lucublandus) Pachythecus Chaudoir
26' Elytron without scutellar stria 27
Quaest. Ent., 1979, 15 (3)
350
Straneo
27 (260Terminal labial palpomere of male triangular, medial margin twice length of apical
margin Meropalpus Tschitscherine
21' Terminal labial palpomere subcylindrical or fusiform 28
28 (27') Abdominal sterna with setigerous punctures slightly larger and deeper than usual.
Smaller (body length 9-10 mm) Eumara Tschitscherine
28' Abdominal sterna with setigerous punctures of average size. Larger, or same size as
above Marsyas Putzeys
29 (16') Tarsal claws pectinate. Head wide; eyes large hemispherical. Body length 4-6 mm.
Elytral interval 3 with single setigerous puncture. Abdominal sterna IV-VI without
transverse sulci laterally Abaris Dejean
29' Tarsal claws smooth, not pectinate. Length more than 6 mm; or eyes small to average in
size 30
30 (29')Sterna IV-VI each with transverse sulcus widely interrupted medially (complete in few
specimens of Blennidus fontainei Tschitscherine) 31
30' Sterna IV-VI without partial transverse sulcus and border; with or without irregular
longitudinal impressions laterally 34
3 1 (30 ) Labrum metallic Cynthidia Chaudoir
31' Labrum not metallic 32
32 (31') Partial sulci of sterna IV-VI each with row of rather wide and deep punctures.
Metepisternum of thorax short or moderately elongate Ogmopleura Tschitscherine
32' Partial sulci of sterna IV-VI smooth, without row of wide and deep punctures.
Metepisternum elongate, with lateral side 1.5 times longer than anterior width Blennidus {sensu
latiore) Motschulsky 33
33 (32') Dorsal surface of body coppery, shiny (similar to Foecilus cupreus; rather flattened
Blennidus (Pseudocynthidia) Straneo
33' Dorsal surface of body black; more convex Blennidus (sensu stricto)
34 (30') Elytral interval 3 impunctate Feroniola Tschitscherine
34' Elytral interval 3 with at least one setigerous puncture 35
35 (34') Elytral intervals 5 and 7 with some setigerous punctures Metoncidus Bates
35' Elytral intervals 5 and 7 impunctate 36
36 (35') Elytral interval 3 with two or more setigerous punctures 37
36' Elytral interval 3 with single setigerous puncture 46
37 (36 )Intercoxal process of prosternum with two setigerous punctures near apex
Parhypates (Argutoridius) Chaudoir
37' Intercoxal process glabrous, without setigerous punctures 38
38 (37') Metepisternum short, anterior and lateral margins subequal Parhypates
(sensu latiore) Motschulsky 39
38' Metepisternum elongate, lateral margin longer than width at anterior margin
Trirammatus (sensu latiore) Chaudoir 43
39 (38 ) Small, body length 7-9 mm. Form stout. Pronotum convex, only slightly narrowed
basally. Range - high altitudes in Andes of Ecuador Parhypates (Agraphoderes) Bates
39' Size generally larger. Form slender. Pronotum only slightly convex, or slightly flattened,
generally evidently narrowed toward base 40
40 (39') Head with two pairs of supraorbital setigerous punctures 41
40' Head with one or three pairs of supraorbital setigerous punctures 42
41 (40 ) Lateral margins of lateral lobes of mentum smooth, not crenulate. Frons with
Classification of South American Pterostichini
351
impressions. Elytron with basal ridge complete. At least hind tarsomere 1 sulcate
laterally. Mandibles shorter Parhypates {sensu stricto)
4T Mentum with lateral margins of lateral lobes crenulate. Frontal impressions indistinct or
nearly so. Elytron with basal ridge complete or rudimentary. Mandibles longer
Parhypates (Eutanys) Tschitscherine
42 (40') Head with single pair of supraorbital setigerous punctures
Parhypates (Antarctobium) Tschitscherine
42' Head with at least three pairs of supraorbital setigerous punctures
Parhypates (Chaetuchenium) Tschitscherine
43 (38') Elytron without scutellar stria Trirammatus (sensu stricto)
43' Elytron with scutellar stria 44
44 (43') Mentum with tooth bilobed. Pronotum with complete anterior submarginal sulcus
Trirammatus (Meraulax) Tschitscherine
45 (44') Prosternum with intercoxal process smooth, not margined apically. Pronotum not
obliquely depressed near postero-lateral angles
Trirammatus (Feroniomorpha) Tschitscherine
45' Intercostal process of prosternum with apex margined. Pronotum obliquely depressed
near postero-lateral angles Trirammatus (Plagioplatys) Tschitscherine
46 (36')Metepimeron with posterior margin obliquely truncate. Metepisternum with lateral
margin only slightly longer than width at anterior margin. Posterior tarsomeres
longitudinally deeply sulcate on outer and inner sides; anterior tarsomeres of male
slightly dilated obliquely, ventral adhesive vestiture confined to inner (medial) half of
tarsomeres 1-3. Pronotum with anterior marginal sulcus complete and deep Adrimus Bates
46' Metepimeron regular in form, posterior margin rounded 47
47 (46') Metepisternum short, width at base and length of lateral margin subequal. Anterior
tarsomeres of male normally, symmetrically dilated Parhypates (Paranortes)
47' Metepisternum elongate (specimens of most taxa), or short. Male with anterior
tarsomeres obliquely dilated 48
48 (47') Pronotum with more than two pairs of lateral setigerous punctures; not pedunculate
Oxycrepis (sensu stricto) Reiche
48' Pronotum with one or two pairs of lateral setigerous punctures; pedunculate or not 49
49 (48') Pronotum not pedunculate Loxandrus LeConte
49' Pronotum pedunculate 50
50 (49') Pronotum as wide as long; lateral margin evidently angulate at level of anterior setigerous
puncture. Elytral striae shallow, very slightly punctulate; intervals flat, interval 3 with
series of nine setigerous punctures near stria 2..... Oxycrepis (Prostolonis) Mateu
50' Pronotum more narrowed anteriorly, wider than long; lateral margin rounded or only
slightly angulate at level of anterior setigerous puncture. Elytral striae deeper and
markedly punctulate; intervals more or less convex, interval 3 with single setigerous
puncture Oxycrepis (Stolonis) Motschulsky
Quaest. Ent., 1979, 15 (3)
352
Straneo
CHECKLIST OF SOUTH AMERICAN PTEROSTICHIN I: SUBTRIBES, GENERA, AND
SUBGENERA
Subtribe MORIONINA
1. Morion Latreille, 1810
2. Moriosomus Motschulsky, 1864
Subtribe CRATOCERINA
3. Cratocerus Dejean, 1829
Subtribe MICROCEPHALINA
4. Tichonilla Strand, 1942
Subtribe EUCHROINA
5. Lobobrachus Sharp, 1885
6. Euchroa Brulle, 1834
{Dyschromus Chaudoir, 1835)
7. Bothynoproctus Tschitscherine, 1900
Subtribe PTEROSTICHIN A
8. Oribazus Chaudoir, 1874
9. Apsaustodon Tschitscherine, 1901
10. Haplobothynus Tschitscherine, 1901
1 1 . Ophryogaster Chaudoir, 1878
12. Hybothecus Chaudoir, 1874
13. Sierrobius {sensu latiore) Straneo, 1951
13.1 Sierrobius (sensu stricto)
13.2 Pachyabaris Straneo, 1951
14. Pachythecus Chaudoir, 1874
15. Meropalpus Tschitscherine, 1900
16. Marsyas Putzeys, 1846
17. Eumara Tschitscherine, 1901
18. Pseudabarys Chaudoir 1873
19. Abaridius Chaudoir 1873
20. Abaris Dejean, 1831
21. Cynthidia Chaudoir, 1873
Classification of South American Pterostichini
353
22. Bletinidus {sensu latiorej Motschulsky, 1865
22.1 Blennidus (sensu strictoj
22.2 Pseudocynthidia Straneo, 1953
23. Tschitscherine, 1898
24. Trirammatus (sensu latiorej Chaudoir, 1838
24.1 Trirammatus (sensu strictoj
24.2 P I agiop lat y s TschiischQunQ, 1900
24.3 Meraulax Tschitschcvine, 1900
24.4 Feroniomorpha Solier, 1 849
25. Parhypates (sensu latiorej Motschulsky 1865
25.1 Parhypates (sensu strictoj
25.2 Agraphoderes Bates, 1891
25.3 Tschitscherine, 1900
25.4 Anarctobium Tschitscherine, 1900
25.5 ChaetaucheniumTsc\\\isc\\Qnr\Q, 1900
25.6 Tschitscherine, 1900
25.7 Argutoridius Chaudoir, 1876
26. Loxandrus LeConte, 1852
27. Oxycrepis (sensu latiorej Reiche, 1843
27.1 Oxycrepis (sensu strictoj
27.2 Prostolonis Mateu, 1976
27.3 Stolonis Motschulsky, 1865
28. AdrimMS Bates, 1872
29. Metoncidus Bates, 1870
30. Feroniola Tschitscherine, 1900
31. Cephalostichus Straneo, 1977
Subtribe ANARCTIINA
32. Kuschelinus Straneo, 1963
33. Metius Curtis 1839
34. Abropus Waterhouse, 1842
35. Antarctiola Straneo 1951
Subtribe CHAETOGEN YINA
36.Camptotoma (sensu latiorej Reiche, 1843
36.1 Camptotoma (sensu strictoj
36.2 Chaetogenys w3Ln Emden, 1958
Quaest. Ent., 1979, 15 (3)
354
Straneo
ACKNOWLEDGEMENTS
Publication costs for this paper were met by a special grant from the Strickland Memorial Trust
Fund. I thank G.E. Ball for making the necessary arrangements. I also thank Maria Figueroa,
Department of Economics, University of Alberta, for preparing a Spanish translation of the abstract.
REFERENCES
Ball, G.E. 1977. [Notes about Chaetogenyina]. In Reichardt, FI. A synopsis of the genera of Neotropical
Carabidae, p. 408.
Blackwelder, R.E. 1944-1957. Checklist of the coleopterous insects of Mexico, Central America, the
West Indies, and South America. Parts 1-6 United States Government Printing Office, Washington,
D.C. United States National Museum, Bulletin 185, 1492 pp.
1944. Part 1,XII + pp. 1-188.
1957. Part 6, vii + pp. 927-1492.
Csiki, E. 1927-1933. Coleoptera, Carabidae. In Junk, W. and S. Schenkling. 1909-1940.
Coleopterorum Catalogus, W. Junk, Berlin and ’s-Gravenhage.
1929. Pars 104, pp. 347-527.
1930. Pars 112, pp. 529-737.
Erwin, T.L., D.R. Whitehead and G.E. Ball. 1977. Family 4. Carabidae, The Ground Beetles,
pp. 4.1-4.68. In Blackwelder, R.W. and R.H. Arnett. Checklist of the beetles of Canada, United
States, Mexico, Central America, and the West Indies. World Digest Publications, Oxycopis Pond,
Wallace Road, Kinderhook, New York, 12106.
Lindroth, C.H. 1966. The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 4.
Opuscula Entomologica, Supplementum 29: 409-648.
Reichardt, H. 1973. Monograph of Catapiesini, a neotropical tribe of Carabidae (Coleoptera). Studia
Entomologica 16: 321-342.
Reichardt, H. 1977. A synopsis of the genera of Neotropical Carabidae. Quaestiones Entomologicae
13(4): 347-485.
Straneo, S.L. 1977. Chiavi per la determinazione dei generi e sottogeneri dei Pterostichini delTAmerica
del Sud. Bollettino della Societa Entomologica Italiana, 109(7-8): 104-1 16.
Classification of South American Pterostichini
355
Index to Names of Taxa
TRIBE AND SUBTRIBES
Agonini, 345
Antarctiina, 345, 346, 347, 355
Catapiesini, 345, 354
Cephalostrichus Straneo, 345
Chaetogenyina, 345, 347, 354, 355
Cratocerina, 345, 346, 352
Euchroina, 345, 346, 347, 352, 355
Microcephalina, 345, 346, 347, 352, 355
Microcephalina ( = Tichoniina), 346
Morionina, 345, 346, 352
Pterostichina, 345, 347
Pterostichini, 345, 346, 347, 355
GENERA AND SUBGENERA
Abaridius Chaudoir, 349, 352
Abaris Dejean, 350, 352
Abropus Waterhouse, 347, 353
Adrimus Bates, 345,351,353
Agraphoderes Bates, 345, 353
Anarctobium Tschitscherine, 353
Antarctiola Straneo, 345, 347, 353
Apsaustodon Tschitscherine, 349, 352
Argutoridius Chaudoir, 353
Blennidus (Pseudocynthidia) Straneo, 350
Blennidus {sensu latiore) Motschulsky, 350,
353
Blennidus (sensu stricto), 350, 353
Bothynoproctus Tschitscherine, 347, 352
Camptotoma (Chaetogenys) van
Emden , 347
Camptotoma (sensu latiore) Reiche, 347, 353
Camptotoma (sensu stricto), 347, 353
Cephalostichus Straneo, 349, 353
Chaetauchenium Tschitscherine, 353
Chaetogenys van Emden, 353
Cratocerus Dejean, 346, 352
Cynthidia Chaudoir, 345, 350, 352
Dyschromus Chaudoir, 349, 352
Euchroa Brulle, 349, 352
Eumara Tschitscherine, 350, 352
Eutanys Tschitscherine, 345, 353
Feroniola Tschitscherine, 345, 350, 353
Feroniomorpha Sober, 353
Feroniomorpha Tschitscherine, 345
Haplobothynus Tschitscherine, 349, 352
Hybothecus Chaudoir, 349, 352
Kuschelinus Straneo, 347, 353
Lobobrachus Sharp, 347, 352
Loxandrus LeConte, 351,353
Marsyas Putzeys, 350, 352
Meraulax Tschitscherine, 353
Meropalpus Tschitscherine, 345, 350, 352
Metius Curtis, 347,353
Metoncidus Bates, 350, 353
Morion Latreille, 346, 352
Moriosomus Motschulsky, 346, 352
Ogmopleura Tschitscherine, 345, 350, 353
Ophryogaster Chaudoir, 349, 352
Oribazus Chaudoir, 345, 349, 352
Oxycrepis (Prostolonis) Mateu, 351
Oxycrepis (sensu latiore) Reiche, 353
Oxycrepis (sensu stricto), 353
Oxycrepis (sensu stricto) Reiche, 351
Oxycrepis (Stolonis) Motschulsky, 351
Pachyabaris Straneo, 352
Pachythecus Chaudoir, 349, 352
Paranortes Tschitscherine, 353
Parhypates (Agraphoderes) Bates, 350
Parhypates (Antarctobium)
Tschitscherine, 351
Parhypates (Argutoridius) Chaudoir, 350
Parhypates (Chaetuchenium)
Tschitscherine, 351
Parhypates (Eutanys) Tschitscherine, 351
Parhypates (Paranortes), 351
Parhypates (sensu latiore)
Motschulsky, 350, 353
Parhypates (sensu stricto), 351, 353
Parhypates Motschulsky, 345
Plagioplatys Tschitscherine, 353
Prostolonis Mateu, 353
Pseudabarys Chaudoir, 345, 349, 352
Pseudocynthidia Straneo, 353
Pterostichus (sensu latiore), 346
Sierrobius (Pachyabaris) Straneo, 349
Sierrobius (sensu latiore) Straneo, 349, 352
Sierrobius (sensu stricto), 349, 352
Quaest. Ent., 1979, 15 (3)
356
Straneo
Stolonis Motschulsky, 353
Tichonilla Strand, 347, 352
Trirammatus (Feroniomorpha)
Tschitscherine, 351
Trirammatus (Meraulax)
Tschitscherine, 351
Trirammatus (Plagioplatys)
Tschitscherine, 351
Trirammatus (sensu latiore) Chaudoir, 350,
353
Trirammatus (sensu stricto), 351, 353
Trirammatus Chaudoir, 345
FOOD OF BLACK FLY LARVAE (DIPTERA: SIMULIIDAE): SEASONAL CHANGES IN GUT
CONTENTS AND SUSPENDED MATERIAL AT SEVERAL SITES IN A SINGLE WATERSHED'
DANIEL C. KURTAK^
Department of Entomology
Cornell University
Ithaca, New York 148353
US.A.
Quaestiones Entomologicae
15: 357-374 1979
Gut contents of black fly larvae and suspended material in water flowing over them in their stream
habitats were studied for three years at five sites in a watershed near Ithaca, New York, U.S.A.
Suspended material was removed from water samples with a centrifuge. Total dry weight varied
from 1-13 mg/l. Organic content {determined by dichromate oxidation) of suspended material varied
from 8-24% of the dry weight. A portion of suspended material was analyzed microscopically.
Particles were classified as diatoms and other algae, mineral fragments (silt), plant fragments, or fine
material {<5 gm).
Quantity and quality of suspended material varied greatly at different sites in the watershed and
with season. Headwater streams, especially in forested areas, contained larger amounts of plant
fragments. In lower reaches there was more total suspended material and a larger proportion of
diatoms. When stream discharges were low and steady, and sunshine was abundant, more diatoms were
produced, while spring floods added more silt.
Gut contents of larvae were also analyzed microscopically, and the proportions of their constituents
were found to agree generally with the proportions in the water. This indicated indiscriminate feeding.
Also there was evidence that habitat preference had an influence on larval food. Diatoms often made up
as much as 50% of some gut contents and were shown to be digested.
Le contenu du tube digestif de larves de mouche noire et le materiel en suspension dans I’eau recueillie autour des larves dans
leurs habitats d’eau courante ont ete etudies pendant trois ans a cinq emplacements situes dans une zone de partage des eaux pres
d'lthaca, dans I’etat de New York (Etats-Unis).
Le materiel en suspension a ete extrait des echantillons d’eau par centrifugation. Le poids sec total varie de I a 1 3 mg/l. La
fraction organique (determinee par oxydation an dichromatej du materiel en suspension constitue de 8 a 24% du poids sec. Une
partie du materiel en suspension a ete analysee au microscope. Les particules ont ete classifiees soit comme diatomees et autres
algues, fragments mineraux (argile), fragments vegetaux, ou comme materiaux fins (<5 ur>9-
La quantite et la qualite du materiel en suspension varient beaucoup suivant les differents emplacements dans la zone de partage
des eaux et suivant les saisons. Les ruisseaux de tete de cours, particulierement dans les zones forestieres. contiennent de grandes
quantites de fragments vegetaux. Dans les zones basses du bassin hydrographique, le materiel en suspension est plus abondant et la
proportion de diatomees, plus elevee. Dans les cours d’eau abondamment ensoleilles et a ecoulement lent et regulier, les diatomees
sont produites en plus grands nombres, tandis les inondations printani'eres causent un apport accru d’argile.
Le contenu du tube digestif des larves a egalement ete analyse au microscope. Les proportions des constituants presents sont
generalement semblables d celles trouvees dans Lean, indiquant que les larves s’alimentent au hasard. 1 1 semble aussi que le choix de
I'habitat ait une influence sur la nourriture des larves. Souvent, les diatomees constituent jusqu’a 50% du contenu de certains tubes
digestifs et I’analyse montre qu’elles sont digerees.
'Part of a thesis accepted by the Graduate Faculty of Cornell University in partial fulfillment of
the requirements for the degree of Doctor of Philosophy.
^Present address: Rt. 2 Box 74R, Bishop, California 93514 U.S.A.
358
Kurtak
INTRODUCTION
Black fly larvae are filter feeders in practically all types of lotic waters, ranging from tiny streamlets
to large rivers. Larvae of some species can filter material down to colloidal size (Wotton, 1976), whereas
others “graze” on filamentous algae (Burton, 1973). On the basis of gut analyses, larvae feed rather
indiscriminately on particulate material as shown by the studies of Anderson and Dicke (1960), Carlsson
et aL, (1977), Chance (1970), Davies and Syme (1958), Emery (1913), Naumann (1925), Pacaud
(1942), Puri (1925), Williams (1961), and Wotton (1977).
This study was designed to measure seasonal changes in the quantity and quality of suspended
material at various points along the length of a stream in Tompkins County, New York, U.S.A. The gut
contents of black fly larvae at each site were also examined to determine the degree of selectivity and
extent to which habitat specificity influenced type of food material ingested.
MATERIALS AND METHODS
The study area
Pinkovsky (1970) and Naumann (1965) give a good ecological background for the area in central
New York State (U.S.A.) where the work was done. Situated at the northern edge of the Appalachian
Plateau, the area is characterized by rolling hills to the south and plains to the north. Natural vegetation
is beech-maple woodland. Such vegetation exists in the more rugged areas, and on poorer soils. About
40% of Tompkins County is cultivated.
Variations in topography and vegetation produce a wide variety of stream habitats. Temperature,
flow regime, discharge, substrate, and type of watershed all vary widely.
The climate is continental humid and there are large seasonal variations in stream temperature and
flow. In summer months small streams dry up and large streams have low, steady, base flows.
Occasionally a thunderstorm produces a freshet. In fall, winter, and spring, streams have higher and less
regular flows. Snowmelt produces spring floods, with peak discharges 20-40 times the base flow (United
States Geological Survey, Water Resources Data, 1965). Maximum daily water temperatures vary
throughout the year and from 0 C to over 26 C for an unshaded, slow-moving stream in summer. Ice
may be present for 2-3 months. Intradiel variations in temperature may be high in the summer,
exceeding 10 C. There are more than 20 species of black flies present in the county (Pinkovsky, 1970, p.
156).
The sites were located along Sixmile Creek and its tributaries in Tompkins County, New York. This
stream is 33 km long and originates in the uplands east of Ithaca, New York at an elevation of 520 m
(Fig. 1). Flowing first for approximately 5 km through mixed wood and farm land, it then passes
through a gorge and over a waterfall before travelling through 7 km of farm land and a second gorge.
From there it traverses a series of waterfalls, dams and reservoirs before emptying into Cayuga Lake
(elevation 1 17 m) at Ithaca.
Sampling sites and times
The stream was sampled at Sites 1-4 (Fig. 1), chosen as representatives of different aspects of the
stream (e.g. headv/aters and lower reaches) and as known locations of black fly larvae. Site 1 is on one of
the headwaters of Sixmile Creek where the stream is less than 1 m wide, temporary, and running
through an open marshy meadow. Site 2 (Hurd Road Stream) is a small stream in a forested area,
typical of many small branches of Sixmile Creek. This particular branch enters a swamp which feeds
Seasonal changes in larval black fly food
359
Figure 1. Sixmile Creek, Tompkins County, New York, USA. (Plan above and profile below) showing collecting sites:
1 = Sixmile Creek Headwaters (open headwater stream), 2 = Hurd Road Stream (forested headwater stream), 3 = Sixmile
Creek at Sixhundred Road (forested lower reach), 4 = Sixmile Creek at Brooktondale (open lower reach), 5 = Sixmile Creek
at Burns Road (dam spillway).
Quaest. Ent., 1979, 15 (3)
360
Kurtak
both Sixmile Creek and Cascadilla Creek. At Site 3, Sixmile Creek (at Sixhundred Road) has become
large and permanent and flows in a rocky gorge through hilly, forested land. At Site 4 (at
Brooktondale), the stream is in a shallow gorge, surrounded by farm land. Between Sites 3 and 4, there
are long reaches of stream with fine-textured substrates (sand and fine gravel) which support a few
larvae. A few samples were also taken at Site 5 (at Burns Road) where the stream passes over a dam
spillway before entering a long gorge with forested sides. Detailed descriptions of these collection sites
are given in Kurtak (1973).
Sites were visited monthly, or at least once in each of the four seasons in 1970 and 1971, and some
again in 1972. Samples were taken during the daylight hours, but not at specific times.
Samples from different parts of the watershed were usually taken within a few days of each other, but
not commonly on the same day, due to lack of time. If rainfall occurred in the interval between samples,
a significant change in suspended material was possible.
In one instance, a special effort was made to sample the sites in a rapid sequence. Sites 1, 3, 4 and 5
were visited within a few hours.
Sampling procedure
Environmental parameters were measured at each site. These included surface velocity (measured by
timing a float passing along a calibrated cord held in the current); temperature (measured to the nearest
degree centigrade with an alcohol thermometer); and pH (measured with a Helige comparator using
bromophenol red indicator). Notes were taken on the dimensions of the streams , the nature of the
substrate, bank vegetation, and any other factors that seemed relevant.
Characteristics of the sites are summarized in Table 1 and 2. Two significant trends down the stream
were increases in velocity and water temperature (in summer). Also, pH increased downstream in
general, but decreased again at Site 5. Site 4 was densely populated in summer with up to 50 larvae/cm-
of Simulium pictipes. Note that Sites 1 and 2 were both dry in the summer and early fall.
Black fly larvae were collected and returned to the laboratory alive for identification and dissection.
They were identified according to Peterson (1970), Pinkovsky (1970), Stone (1964), and Stone and
Jamnback (1955). Voucher specimens have been deposited in the Cornell University Insect Collection,
Ithaca, New York (Lot. No. 1038). Whenever possible, identifications were made before removing the
gut; otherwise, specimens were determined on the basis of preserved head capsules and skins. At least
three individuals of each major species found at each site were selected and their guts removed and
smeared on individual slides in Hoyer’s medium. The composition of the gut content was determined in
the same way as composition of samples of suspended material (see below).
The majority of the larvae collected were of the following species:
Cnephia mutata (Malloch)
Prosimulium fontanum Syme and Davies
Prosimulium fuscum Syme and Davies
Prosimulium magnum Dyar and Shannon
Simulium decorum Walker
Simulium parnassum Malloch
Simulium pictipes Hagen
Simulium tuberosum (Lunstrom)
Simulium venustum Say
Simulium verecundum Stone and Jamnback (species complex)
Simulium vittatum Zetterstedt
Table 1. Characteristics of stream sites. All sites are in Tompkins County, New York, USA.
Seasonal changes in larval black fly food
Quaest. Ent., 1979, 15 (3)
Average dimensions through a 10-m reach.
Surface velocity.
362
Kurtak
Seven liters of water were collected at each site for analysis of suspended material. The sample was
dipped out as close as possible to the attachment sites of larvae when present, taking care not to stir up
the substrate. Samples were stored at 4 C and usually processed within two days. They were centrifuged
in a Foerst® continuous centrifuge, permitting removal of 98% of the plankton-sized material and
25-50% of the bacteria (Welch, 1948). The centrifugate was subsampled by scraping the wall of the
centrifuge bowl in several vertical bands with a 4 mm wide blade. Each subsample (two per water
sample) was then spread thinly on a slide in a drop of Hoyer’s medium. These slides were observed at a
magnification of 300 diameters using a phase contrast microscope. At least five fields laid out in an “X”
pattern were observed on each slide. Areas where particles were not in a single layer were avoided.
Within each field, a 50 jum x 300 nm strip was divided into 12.5 )um x 12.5 jam squares using an ocular
grid. Each of these units was examined and the category of particle which filled or dominated (>50%) it
was determined.
Table 2. Additional characteristics of stream sites.
Particles were classified as (1) mineral fragments (identifiable by their sharp-fractured edges and
including fine sand, very fine sand, and silt^, (2) organic fragments (mainly pieces of plant tissue), (3)
diatoms and other algae, and (4) “fine material” less than 5 ij,m in diameter whose nature was
undeterminable (including particles the size of fine silt and clay O- Polarized light was sometimes used to
identify mineral grains by their crystalline nature. Other categories, such as filamentous algae, were
employed as needed. No effort was made to count fungi, bacteria, or colloid particles. The size range of
the particles was not determined in detail.
In the examination process, empty squares were not counted. Areas of large irregular particles were
estimated as though the particles were smoothed to rectilinear shapes. Groups of particles less than
12.5 fim in length were assumed to be crowded together so that the space between them was negligible.
Then the area (number of squares) occupied by the group was estimated. Finally, the percentage of each
category in relation to the total number of squares counted was calculated for the five fields, and the
mean of the two subsamples was determined. This permitted estimates of the composition of the samples
on basis of area occupied by various types of particles in a thin smear.
^See United States Department of Agriculture (1951) for definitions of the sizes of soil
particles.
Seasonal changes in larval black fly food
363
The remaining 95% of the centrifugate was transferred to an aluminum weighing dish and dried at
60 C for 24 hours. The residue was then weighed to the nearest 0.01 mg to determine mg/1 total
suspended material.
Selected samples were also analyzed for oxidizable organic matter by the micro method of Maciolek
(1962).
Digestion of diatoms
Although this study did not concern itself with what portion of the ingested suspended material was
actually digested, a simple experiment was conducted with diatoms. Fresh, living diatoms
{Cymbella sp.) were fed larvae of Simulium jenningsi, Simulium pictipes and Si mu Hum tuberosum in
the laboratory. The larvae were given no additional food, so the diatoms were retained for several hours.
Every five minutes for the first 30 minutes after ingestion and at one-hour intervals for several hours
thereafter, a sample of the larvae was taken. The guts were immediately removed and squashed on a
slide and observed by phase contrast microscopy.
RESULTS
Suspended material and gut contents
The data from the stream water samples and gut-content analyses are presented in Tables 3 - 7. In
these tables, each value for “water” represents the mean for 1-10 samples taken at that site in that
month during the three years of the study. Each “gut content” value represents a mean for at least three
larvae of each of the main species present when the water was sampled. The data for different species
were combined if there were no significant differences among them. Coefficients of variation for groups
of subsamples of a given water sample and groups of larvae collected at the same time were 10-20%. The
stream stage (percent of the highest level of flow observed) and water temperature are also shown in the
tables.
The dry weights of suspended material normally ranged from 1-13 mg/1 (x = 4.6 mg/1). During
flash flood periods, however, large amounts of silt and coarser mineral particles briefly raised the
weights to several hundred mg/1. Such samples are not included in the means.
Usually at least 50% of the suspended material was mineral fragments or material less than 5 gm in
diameter. This largely inorganic portion (see “Organic Content” below) accounted for most of the
increase in total suspended material during high water periods (e.g. during March at Site 4, Table 6).
The organic portion consisted largely of plant fragments and diatoms. Fragilaria, Meridion,
Gomphonema, and Cymbella were the main diatom genera seen. Visible diatom coverings on the rocks
in the streams tended to be more common in the late spring and early fall than in summer or winter.
In addition to the types of materials indicated in the tables, other materials were found occasionally
in the gut contents. These included fragments of filamentous algae, aquatic mites, and fragments from
other arthropods, including black fly larvae. Pollen, probably of Pinus strobus L., was important at
Site 2 on one occasion, in both guts and water.
In terms of size, mineral particles generally ranged from 5-50 pm, diatoms from 20-100 pm, and
organic particles varied from five to several hundred pm in diameter. Particles larger than 300-400 pm
were rarely seen in the guts. Particles less than 5 pm, not separated as to type, often made up a large
portion of the samples.
Proportions of various components of suspended material varied greatly with time and place. Water
in upper reaches (Sites 1-3) contained fewer diatoms than that from lower reaches (Site 4). This trend is
also clear from samples taken on one day (Table 7). The forested headwater stream at Site 2 contained
Quaest. Ent., 1979, 15 (3)
364
Kurtak
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Seasonal changes in larval black fly food
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Seasonal changes in larval black fly food
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368
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more organic particles and fewer diatoms at Site 3 than at Site 4, even though the substrates were very
similar. In general, the lower reaches had more total suspended material.
Maximum quantities of suspended material occurred in spring and consisted mainly of mineral
fragments or material less than 5 iim. In summer and fall, diatoms made up a large portion of total
suspended material in the lower reaches, sometimes up to 50% (e.g. November at Site 4). Plant
fragments were numerous at forested headwater site (Site 2) in fall, and at Site 3 in spring.
Proportions of material in guts tended to correspond to proportions of materials carried in the stream,
except that larvae commonly contained a higher percentage of diatoms. Usually, gut contents of
different species at the same site were quite similar. At Site 1 in January, however, there was
considerable difference between those of Cnephia mutata and Prosimulium fontanum.
Differences between the years were not analyzed, since duration of this study was short.
Organic content of the suspended material
Results of the analyses for organic content are shown in Table 8. Total organic content varied from
0.35-1.11 mg/1 (7.9-24%). The highest percentage of organic matter in the suspended material did not
coincide with the highest total amount of organic matter per liter.
The highest percentages of organic matter were found where organic fragments were common (e.g.
Site 2 XI-4-70). Unfortunately, none of the water samples from Sixmile Creek chosen for analysis had a
high proportion of diatoms. To show that large quantities of diatoms also increased the organic content,
a result from a site similar to Site 4 (Taughannock Creek) is also shown. Low percentages were
associated with large proportions of mineral fragments and/or material less than 5 pm (e.g.
Site 2 IV-6-71, Site 3 1-26-71, Site 4 1-26-71, and 11-25-71).
Digestion of diatoms
Diatoms examined 15 and 30 minutes after ingestion showed no visible change in appearance other
than loss of mobility. After 40-60 minutes, the diatoms began to lose their golden-yellow color and
became more transparent, revealing the striations on the frustules and the internal organelles. The
clearing occurred first in diatoms near the gut wall. Within 3.5 hours, frustules became completely clear.
This is of course much longer than the usual retention time (see Discussion). Non-ingested fresh diatoms
which were killed in hot water and stored at room temperature did not clear for a least 48 hours. The
clearing occurred at similar rates for Cymbella sp. in guts of larval Simulium pictipes, Simulium
tuberosum and Simulium jenningsi.
DISCUSSION
Reliability of data
Reliability of data gathered for this work is limited by the method of estimating proportions of
various materials in samples. Applied consistently, estimation of composition by area should give good
comparative information. However, for absolute measurement of suspended material, estimation by
microscopical measurement of area is less accurate than volumetric measurement or direct chemical
analyses, but is more accurate than counting (Sladeckova, 1962). Neither area or volume estimates
allow for differences in density. In Table 8, there is only a general agreement between the microscopical
and chemical analyses.
Also, the particulate material examined was only a small fraction of the total material collected in
water samples, which in turn were a very small percentage of flow of the stream at a particular time.
Another aspeet of the sampling problem was daily variability in stream discharge arid concurrent
Quaest. Ent., 1979, 15 (3)
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Seasonal changes in larval black fly food
371
variability in content of suspended material. Sampling was done in periods of stable base flow when
possible. Another approach would be to use automatic equipment capable of continuous sampling.
Interpretation of the stream samples
Factors such as type of terrain in the watershed, the nature of the substrate, degree of shading by
streamside vegetation, and water depth and velocity can be used to explain the observed temporal and
spatial variations in the quantity and composition of suspended material in the stream, as well as
distribution of black fly larvae.
In forested headwater streams such as Site 2, fallen leaves are the main part of the organic material.
Coarse leaf fragments (>300 /xm) are especially important in the fall. These streams provide practically
no diatoms because of heavy shading. In open headwater streams, such as Site 1 (which is on a
south-facing slope), diatoms grew abundantly on trailing grass even in January. Both types of
headwaters can produce large numbers of larvae, but of different species (i.e. Prosimulium magnum at
Site 2; Prosimulium fontanum, Cnephia mutata, and several Simulium spp. at Site 1 ).
Changes seen at downstream sites in this study can be explained by a description of changes along a
stream similar to that shown by Cummins (1977). The material from headwater streams is comminuted
and diluted as it is carried downstream. Additional organic particles enter the stream from the
vegetation bordering the stream, but this contribution becomes less important in relation to the volume
of the stream as it coalesces and increases in discharge. Diatoms and other algae washed off of the
substrate contribute more to organic content at this stage. Amount of algal growth varies with suitability
of substrate and amount of sunlight. Sand and silt are continuously eroded from banks. Higher water
velocities downstream tend to keep more material suspended.
Consistent with this picture of downstream changes, diatoms were more important at Site 3 and at
Site 2, but organic particles still made up over 50% of the organic fraction. The large (>300 pm) leaf
fragments common at Site 2 were not seen at Site 3. The bedrock substrate at Site 3 would certainly be
suitable for algal growth, but diatoms may have been limited by shade from the forested banks. At
Site 4, the substrate was bedrock, but was exposed to the sun with only a thin layer of water flowing over
it. The suspended material there exceeded 10 mg/1 and was often nearly 50% diatoms, especially during
later summer months when discharge was low, illumination high, and floodwaters did not frequently
scour the streambed. In early spring, with high water, more silt was present.
The data taken all on the same day (Table 7) support the idea of a trend toward fewer large particles
and more diatoms downstream. Below Site 4, however, the diatoms were mostly empty frustules.
The presence of diatoms and organic particles increases the percentage of organic matter, but does
not necessarily increase the total amount per liter. However, a small amount of organic matter in a pure
form may be more beneficial than a larger amount heavily diluted with silt.
Interpretation of the gut contents
Proportions of materials in gut contents generally reflected proportions in the water, with little
difference among larvae of different black fly species present at a given site. This indicates that feeding
is indiscriminate. It was not uncommon, however, to find larvae that contained a higher proportion of
diatoms than that found in the associated water. In the case of Simulium pictipes (Site 4 in April), this
could be the result of more efficient feeding on the diatoms, which are larger than the silt particles
(Kurtak, 1978). Also other fractions of gut contents may be digested, leaving behind only the insoluble
diatom frustules and falsely inflating the percentage of diatoms.
At Site 1 in January, larvae of Cnephia mutata contained a much larger proportion of diatoms than
did the water or larvae of Prosimulium fontanum at the same location. This can be explained by the
Quaest. Ent., 1979, 15 (3)
372
Kurtak
observation that larvae of Cnephia mutata often spend considerable time scraping material from the
substrate rather than filter feeding (Kurtak, 1973).
No attempt was made to determine which components of gut contents are important in nutrition of
black fly larvae, except to show that diatoms are in fact digested. The use of radioactive tracers would be
one way to determine what portions of the gut content are assimilated.
It is clear that diatoms can be at least partly digested in the normal retention time, which is about one
hour (Kurtak, 1978). Diatoms from larvae collected in natural habitats always appeared dead and clear
since the larvae were held alive several hours before dissection.
Black fly larvae are known to ingest bacteria (Snoddy and Chipley, 1971) and there is evidence that
bacteria may be an important food source in large rivers (Fredeen, 1960,1964). No bacteria could be
found in Sixmile Creek, but coliform bacteria are certainly present where the stream banks are densely
populated. Truly rheophilic bacteria may also be present. These, however, are usually few in number and
are difficult to culture (Hynes, 1970). Considerable numbers of bacteria may be associated with
decomposing plant fragments in the water.
Black fly larvae also reportedly ingest particles as small as colloids (Wotton, 1976), but these
particles require electron microscopical techniques to detect.
From the data it appears that the choice of a specific habitat by a black fly species has a strong
influence on what kind of food the larvae will be exposed to. Larvae of Simulium pictipes, for example,
which were characteristically found on bedrock substrates near waterfalls, usually contained large
numbers of diatoms. The larvae of Prosimulium magnum on the other hand, usually contained large
amounts of leaf fragments obtained in the small, shaded, forest streams where they lived. Another
instance of direct relationship between habitat and food is reported by Carlsson et al., (1977), who
reported that larvae occurring below a lake in Sweden received nutrients in the form of fine particulate
organic matter from the lake bottom. In the region of the present study, however, there were other
species which occurred in a very wide range of habitats, such as Simulium tuberosum which Pinkovsky
(1970) found at 85% of the sites he sampled in Tomkins County. Larvae of such a species would be
exposed to a wide range of food.
ACKNOWLEDGEMENTS
Thanks are due to Dr. B.V. Travis, Dr. C.O. Berg, and Dr. J.M. Kingsbury of Cornell University who
served as the author’s Ph.D. thesis committee and guided this study.
This research was supported in part by a National Science Foundation Fellowship and by a Cornell
Fellowship. Further support came from Hatch Act funds and a grant from the Grace Griswold Fund of
the Cornell Department of Entomology helped to defray publication costs.
Dr. Gary R. Mullen of Auburn University and Dr. Richard W. Merritt of Michigan State University
read the manuscript and made valuable suggestions.
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Seasonal changes in larval black fly food
373
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64: 1467-1468.
Stone, A. 1964. Guide to the insects of Connecticut. Part VI. The Diptera or true flies of Connecticut.
Fascicle 9. Simuliidae and Thaumaleidae. Bulletin of the Connecticut State Geological and Natural
History Survey No. 97.
Stone, A., and H.A. Jamnback. 1955. The black flies of New York State (Diptera: Simuliidae). Bulletin
of the New York State Museum No. 349; 1-144.
United States Department of Agriculture. 1951. Soil Survey Manual (United States Department of
Agriculture Handbook No. 18) United States Government Printing Office, Washington, D.C.
503 pp.
United States Geological Survey. 1967. Water resources data for New York (1965). Part 1. Surface
water records. Water Resources Division of the United States Geological Survey, Albany, New York.
490 pp.
Quaest. Ent., 1979, 15 (3)
374
Kurtak
Welch, P.S. 1948. Limnological methods. McGraw-Hill Book Company, New York, 381 pp.
Williams, T.R. 1961. Size of particles ingested by Simulium larvae. Nature, London. 189: 78.
Wotton, R.S. 1976. Evidence that black fly larvae can feed on particles of colloidal size. Nature, London
261(5562): 697.
Wotton, R.S. 1977. The size of particles ingested by moorland stream blackfly larvae (Simuliidae).
Oikos 29: 332-335.
1
I
s.
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
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QL
m
QI3
Ent
Quaest
lones
Entomoloqicae
A periodical record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.
VOLUME 15
NUMBER 4
OCTOBER 1979
QUAESTIONES ENTOMOLOGICAE
ISSN 0033-5037
A periodical record of entomological investigation published at the Department of Entomology,
University of Alberta, Edmonton, Alberta.
Volume 15 Number 4 October 1979
CONTENTS
Editor’s Acknowledgements 375
Bell and Bell - Rhysodini of the world Part II. Revisions of the smaller genera (Coleoptera:
Carabidae or Rhysodidae) 377
Wheeler - Revision and cladistics of the Middle American genus
Creagrophorus Matthews (Coleoptera: Leiodidae) 447
Corrigenda - Volume 15(3) 48 1
ACKNOWLEDGEMENTS
Production of Volume 15 of Quaestiones Entomologicae was overseen by two Editors, acting
consecutively. Douglas A. Craig served as editor while I enjoyed the delights of a year’s sabbatical leave,
returning just in time to attend to the last details in production of the fourth issue. I thank Dr. Craig not
only for his work on most of this volume, but also for his contributions to publication of issues 3 and 4 of
Volume 14.
During Douglas Craig’s term as Editor, we transferred from using a Selectric Composer to use of an
electronic computer for setting up the issues. Although electronic computers may be expected to
ultimately ease the task of production of copy for printing, initially a lot of work is involved, both by
editor and operator. So, I am pleased to express my deep appreciation not only to Douglas, but also to
Twyla Gibson, who acquired the necessary expertise to employ a computer for our work.
On behalf of Dr. Craig, I record appreciation of the efforts of the following persons, who acted at his
request as referees of papers published in this volume:
J.A. Downes, Biosystematics Research Institute, Ottawa, Ontario;
J.L. Eaton, Department of Entomology, Virginia Polytechnic and State University,
Blacksburg, Virginia;
R.W. Merritt, Department of Entomology, Michigan State University, Lansing,
Michigan;
G. R. Mullen, Department of Zoology-Entomology, Auburn University, Auburn,
Alabama;
A.P. Nimmo, Department of Entomology, University of Alberta;
H. G. Philip, Entomology Laboratory, Plant Industry Laboratory, Agriculture Alberta.
Maria Figueroa, Department of Economics, University of Alberta, prepared the Spanish translation
of the abstract for one of the papers. Henri Goulet, Biosystematics Research Institute, Ottawa and
Jean-Francois Landry, a graduate student in my Department, provided French translations of abstracts
of several papers.
Again, I take pleasure in thanking staff members of Printing Services, Universty of Alberta, for their
excellent work in production of this volume.
George E. Ball
RHYSODINI OF THE WORLD PART II. REVISIONS OF THE SMALLER GENERA
(COLEOPTERA: CARABIDAE OR RHYSODIDAE)
ROSS T. BELL
Department of Zoology
University of Vermont
Burlington, VERMONT 05401
JOYCE R. BELL
24 East Terrace Quaestiones Entomologicae
South Burlington, VERMONT 05401 1 5: 377-446 1 979
This paper is the second of a series which will constitute a revision of the Rhysodini of the world
and consists of revisions of all genera except for the four largest ones. Treated are: Leoglymmius Bell
and Bell', Tangaroa Bell and Bell, Dhysores Grouvelle', Neodhysores Bell and Bell; Rhysodes Dalman;
Kupea Bell and Bell; Kaveinga Bell and Bell; Grouvellina Bell and Bell; Xhosores Bell and Bell;
Yamatosa Bell; Shyrodes Grouvelle; Srimara Bell and Bell; Arrowina Bell and Bell; and
Plesioglymmius Bell and Bell. Some of the genera have been redescribed, incorporating new
information. Yamatosa NEW NAME replaces Yamatoa Bell 1977 which is preoccupied by Yamatoa
Kiriaknff 1967
In Kaveinga Bell and Bell, four subgenera are recognized: Angekiva, NEW SUBGENUS,
ry^pe-Kaveinga frontalis (Grouvelle), one sp., Australia: Ingevaka, NEW SUBGENUS, r>^/?e-Kaveinga
orbitosa (Broun), one sp.. New Zealand; Vakeinga, NEW SUBGENUS, two spp.. New Zealand, New
Caledonia: Kaveinga (sensu stricto), 14 spp.. New Guinea, Bismark Archipelago, Solomon Islands,
Santa Cruz Island, Moluccas, and Mindanao.
In Plesioglymmius Bell and Bell, three subgenera are recognized: Plesioglymmius sensu stricto two
spp., Sumatra, Borneo, Mindanao; Ameroglymmius NEW SUBGENUS, /j^pe-Plesioglymmius
meridionalis (Grouvelle), three spp.. South America, Cuba; Juxtaglymmius NEW SUBGENUS,
r>^/7^-Plesioglymmius jugatus NEW SPECIES, one sp., Java.
Rhysodes parvus Grouvelle, accidentally omitted from Part I, is assigned to Kaveinga (sensu
stricto).
The following new species are described (and type localities indicated): Dhysores pan (ZAIRE,
Kivu, Itombwe, 2300 m, Terr. Uvira, Nyalengwe); Dhysores liber (LIBERIA, crest of Mt. Nimba,
1400 m): Kaveinga fibulata (NEW BRITAIN, Rabaul): Kaveinga pignoris (SOLOMON IS.,
Bougainville, Kokure, 690 m) Kaveinga nudicornis (SOLOMON IS., Russell Is., Yandina); Kaveinga
kukum (SOLOMON IS., Guadalcanal, Kukum): Kaveinga ulteria (SOLOMON IS., Santa Cruz Group,
Reef Is.); Kaveinga lupata (NEW GUINEA, Mt. Kaindi, 16 km SW Wau); Kaveinga oka pa (NEW
GUINEA, Kamira, Eastern Highlands); Kaveinga marifuanga (NEW GUINEA, Marifuanga,
Asaro-Chimbu Divide): Kaveinga histrio (PHILIPPINE IS., Mindanao, e. slope Mt. McKinley, 3300',
Davao Prov.); Kaveinga strigiceps (Buru); Grouvellina gigas (MADAGASCAR, Chutes de la Mort);
Grouvellina cuneata (MADAGASCAR, Tamatave and Foret Alahakato); Grouvellina hova
(MADAGASCAR, Tsaramainiandro); Grouvellina ranavalona, Grouvellina edentata, Grouvellina
cooperi, Grouvellina montana (MADAGASCAR, only); Grouvellina radama (MADAGASCAR,
Mahatsinjo); Grouvellina descarpentriesi (MADAGASCAR, Annanarivo); Grouvellina divergens
(MADAGASCAR, Mt. d’Ambre); Grouvellina dentipes (MADAGASCAR, Fizono); Grouvellina cinerea
(MADAGASCAR, Perinet); Arrowina pygmaea (SRI LANKA, Dikoya); Plesioglymmius silus
(PHILIPPINE IS., Mindanao, Zamboanga, Kabasalan); Plesioglymmius reichardti (VENEZUELA,
378
Bell and Bell
Suapure, Caura R.}; Plesioglymmius compactus (CUBA); and Plesioglymmius jugatus {JAVA).
Cet article est la deuxieme d’une serie qui constitueront une revue taxonomique des Rhysodini du
monde. La deuxieme partie est composee des revisions taxonomiques de tous les genres sauf les quatre
les plus grands. Les genres discutes sont: Leoglymmius Bell et Bell; Tangaroa Bell et Bell; Dhysores
Grouvelle; Neodhysores Bell et Bell; Rhysodes Dalman; Kupea Bell et Bell; Kaveinga Bell et Bell;
Grouvellina Bell et Bell; Xhosores Bell et Bell; Yamatosa Bell; Shyrodes Grouvelle; Srimara Bell et
Bell; Arrowina Bell et Bell; et Plesioglymmius Bell et Bell.
Nous decrivons certains genres de nouveau, comprendant des renseignements nouveaux. Yamatosa
NOUVEAU NOM remplace Yamatoa Bell 1977, car il est preoccupee de Yamatoa Kiriakoff 1967.
Nous divisons le genre Kaveinga Bell et Bell parmi quatre sous-genres: Angekiva NOUVEAUX
SOUS-GENRE, r^^pe-Kaveinga frontalis (Grouvelle), une sp., Australie; Ingevaka NOUVEAU
SOUS-GENRE, ?j;pe-Kaveinga orbitosa (Broun), une sp., Nouvelle Zelande; Vakeinga NOUVEAU
SOUS-GENRE, />^p^-Kaveinga setosa (Grouvelle), deux spp., Nouvelle Zelande et Nouveau Caledonie;
Kaveinga (sensu stricto) 14 spp., Nouvelle Guinee, Archipel de Bismarque, lies de Solomon, lies de
Saint Croix, lies Moluques, et Mindanao.
Nous divisons le genre Plesioglymmius parmi trois sous-genres: Plesioglymmius sensu stricto, deux
spp., Sumatra, Borneo, et Mindanao; Ameroglymmius NOUVEAU SOUS-GENRE,
r>^p^-Plesioglymmius meridionalis (Grouvelle), trois spp., Amerique du Sud et Cuba; Juxtaglymmius,
NOUVEAU SOUS-GENRE, fj;p^-Plesioglymmius jugatus NOUVELLE ESPECE, une sp., Java.
Par hazard, nous omimes Rhysodes parvus Grouvelle de la premiere partie. C’appartient au genre
Kaveinga, sous-genre Kaveinga (sensu stricto)
On decrit les especes nouvelles que void, en indiquant pour chacune la localite du specimen type:
Dhysores pan (ZAIRE, Kivu, Itombwe, 2300 m, Terr. Uvira, Nyalengwe); Dhysores liber (LIBERIA,
crde du mont Nimba, 1400 m); Kaveinga fibulata (NOUVELLE BRETANGE, RABAUL): Kaveinga
pignoris (ILES DE SOLOMON, Bougainville, Kokure, 690 m) Kaevinga nudicornis (ILLS DE
SOLOMON, He Russell, Yandina); Kaveinga kukum (ILES DE SOLOMON, Guadalcanal, Kukum):
Kaveinga ulteria (ILES DE SOLOMON, groupe Santa Cruz, He Reef); Kaveinga lupata (NOUVELLE
GUINEE, Mt. Kaindi, 16 km S.O. Wau); Kaveinga okapa (NOUVELLE GUINEE, Kamira, Highlands
de TEst); Kaveinga marifuanga (NOUVELLE GUINEE, Marifuanga, Asro-Chimbu Divide); Kaveinga
histrio (ILES PHILIPPINES IS., Mindanao, versant est du mont McKinley, 3300', province du
Davao); Kaveinga strigiceps (INDONESIE, Buru); Grouvellina gigas (MADAGASCAR, Chutes de la
Mort); Grouvellina cuneata (MADAGASCAR, Tamatave et Ford Alahakato); Grouvellina hova
(MADAGASCAR, Tsaramainiandro); Grouvellina ranavalona, Grouvellina edentata, Grouvellina
cooperi, Grouvellina montana (MADAGASCAR, sans localites specifics); Grouvellina radama
(MADAGASCAR, Mahatsinjo); Grouvellina descarpentriesi (MADAGASCAR, Annanarivo);
Grouvellina divergens (MADAGASCAR, mont d[Ambre); Grouvellina dentipes (MADAGASCAR,
Fizono); Grouvellina cinerea (MADAGASCAR, PERINET); Arrowina pygmaea (SRI LANKA,
Dikoya); Plesioglymmius silus (ILES PHILIPPINES IS., Mindanao, Zamboanga, Kabasalan);
Plesioglymmius reichardti (VENEZUELA, Suapure, Riv. Caura); Plesioglymmius compactus (CUBA);
et Plesioglymmius jugatus (JAVA).
Classification of Rhysodini
379
TABLE OF CONTENTS
Introduction 379
Sources of Material 379
Subtribe Leoglymmiina 380
Subtribe Dhysorina 381
Fig. 1-12 383
Subtribe Rhysodina 386
Fig. 13-24 391
Fig. 25-39 392
Fig. 40-51 393
Subtribe Clinidiina 408
Fig. 52-63 414
Fig. 64-74 415
Subtribe Omoglymmiina 423
Fig. 75-87 426
Fig. 88 - 102 433
Fig. 103- 114 440
Acknowledgements 443
Supplementary References 443
Indices 444
INTRODUCTION
This paper, the second of a series of five, includes revisions of all the genera of Rhysodidae except
Clinidium Kirby, Rhyzodiastes Fairmaire, Omoglymmius Ganglbauer, and Pyxiglymmius Bell and
Bell. Some of the genera have been redescribed, incorporating new information. Otherwise, subtribal
and generic descriptions have not been repeated; instead, the reader is referred to the appropriate pages
in Part I (Bell and Bell 1978). A short list of additional references is added to supplement the extensive
list in Part I.
SOURCES OF MATERIAL
The following abbreviations designate collections cited in this paper. The names in parentheses are
the curators of the respective institutions.
AIM Auckland Institute and Museum, New Zealand (K.A.J. Wise) ;
AMS Instituut voor Taxonomische Zoologie, Amsterdam, Netherlands (J. Duffels);
BMNH British Museum, Natural History, London (R. Pope);
BPBM Bernice P. Bishop Museum, Honolulu (G Samuelson);
Quaest. Ent., 1979, 15 (4)
380
Bell and Bell
BSL Naturhistorisches Museum, Basel, Switzerland (W. Wittmer);
CAS California Academy of Sciences, San Francisco, CA (D. Kavanaugh);
CMP Carnegie Museum of Natural History, Pittsburgh, PA (G. Wallace);
CNHM Field Museum of Natural History, Chicago, IL (H. Dybas);
DSIR Department of Scientific and Industrial Research, Auckland, N.Z. (J. Watt);
DY Daniel K. Young, Michigan State University, E. Lansing;
LCC Lincoln College, Canterbury, N.Z. (R.M. Emberson);
LEI Rijksmuseum von Natuurlijke Historie, Leiden, Netherlands (J. Krikken);
MCZ Museum of Comparative Zoology, Cambridge, MA (J. Lawrence);
MNHB Museum fiir Naturkunde der Humboldt-Universtat, Berlin, DDR (F. Hieke);
MNHN Museum National d’Histoire Naturelle, Paris, France (A. Descarpentries);
MRAC Musee Royal de I’Afrique Centrale, Tervuren, Belgium (P. Basilewsky);
MZSP Museu de Zoologia da Universidade Sao Paulo, Brazil (U.R. Martins);
NMNH United States National Museum of Natural History, Washington, D.C. (P. Hurd);
NMNZ National Museum of New Zealand, Wellington (R.G. Ordish);
SATO Masatako Sato, Nagoya, Japan.
SUBTRIBE LEOGLYMMIINA
Description. - Part I, 53. Only the one genus is known.
Leoglymmius Bell and Bell 1978
Type species. - Rhysodes lignarius Olliff 1885: 471.
Description. — Large, stout rhysodine of somewhat caraboid appearance; with the characters of the subtribe; in addition, with
the following : labrum with one pair of setae; one or two minute temporal setae, in isolated punctures posteromedial to eye; temporal
lobe otherwise impunctate.
Pronotum slightly longer than wide, its sides curved, widest point anterior to middle; sides strongly convergent to apex; more
moderately convergent to base; each side slightly sinuate anterior to hind angle; latter obtuse; median and paramedian grooves narrow,
nearly linear, coarsely punctate, closed anteriorly, but widely open posteriorly; marginal groove single, slightly dilated, punctate; one
or two marginal setae present; prosternum and propleura densely punctate; prosternum densely punctate, hairy.
Elytra relatively short and broad for a rhysodine; humeral tubercle absent; basal scarp transverse, indistinctly defined; base of
Stria VI continued anteromedially from humerus to base of scarp; elytral striae finely punctate, intervals nearly flat; Striae I-VII
joined posteriorly in broad densely microsculptured depression; apical tubercle absent; two striae ventral to marginal Stria; upper one
(Stria VllI) discontinuous, basal portion opposite metasternum, and apical portion opposite apical half of Sternum VI of abdomen;
lower one (Stria IX) on border of epipleural rim, entire; metasternum punctate at sides, with one or more irregular longitudinal rows
of punctures near midline; abdominal sterna coarsely punctate; female with densely microsculptured lateral areas on Sterna Ill-V,
those of IV and V in form of shallow lateral pits.
Legs relatively long for a rhysodine; setae of tarsi fewer, coarser than in other genera; male with ventral tooth on anterior femur;
middle calcar small, acute, hind calcar shorter and less acute than middle one; female with acute inner process on middle tibia,
simulating a calcar.
This genus is a phylogenetic relict. In a number of respects it is more like a normal carabid than are
other rhysodines. These include: a suture separating gena and mentum; broad bands of minor setae on
the outer antennal segments; the absence of an apical tubercle; and the indistinctness of the basal scarp
of the elytron.
We have assigned two additional species to the genus. L. blackburni (Grouvelle 1903) and L.
trichosternus (Lea 1904). The only type specimens studied by us are a series of three L. blackburni in
the Grouvelle collections (MNHN), all labelled as types. We hereby designate one of these types as
Classification of Rhysodini
381
LECTOTYPE. This is a male, labelled “Victoria, Australia”. PARALECTOTYPES: one female, same
data as lectotype; one male, labelled “doit provenir d”Australie”.
The three nominal species were not compared with one another in the original descriptions, and we
have not investigated whether they are really separate species or merely three names for the same
species. B.P. Moore will deal with the question in his forthcoming revision of the Australian rhysodines.
SUBTRIBE DHYSORINA
Description. - Part I, 53.
Key to Genera. - Part I, 54.
Genus Tangaroa Bell and Bell 1978
Type species. - Rhysodes pensus Broun 1880.
Description. - Part I, 54. Only one species is known.
Tangaroa pensus (Broun) 1880)
Rhysodes pensus Broun 1880: 215.
Type material. - HOLOTYPE labelled: “TYPE, Mt. Mangaia, Broun Coll.” (BMNH)
Another specimen, in the same collection, labelled COTYPE, is without locality data. In the original
description, the type locality is given as Whangarei Harbour, and mention is made of an additional
specimen from the Wairoa. In the main collection of the BMNH there is an additional TYPE specimen
labelled “NEW ZEALAND:Broun”. We have not had the opportunity to determine which, if any, of the
above specimens represent the genuine type specimen.
Description. — Length 7-9 mm. Minor setae on Segments Vi-X; antenna with basal setae on Segments V or Vl-X; anterior
tentorial pits large, V-shaped; labrum with two setae; orbital groove absent; temporal setae absent; marginal groove of pronotum
single; precoxal carina absent; elytral Striae l-V impressed, punctate; Striae VI and Vil effaced in anterior third; Stria VI not
impressed, represented only by row of fine punctures; Stria VII with apical fifth impressed, remainder represented by fine punctures;
metasternum relatively short, impunctate; abdominal sterna with scattered punctures; lateral pits not present in either sex; inner and
outer spurs nearly equal on middle and hind tibiae; male with only slight development of calcars, each represented by a swelling on
anteromedial portion of apex of tibia; no other obvious secondary sexual characters.
This species is a large, rather stout rhysodine, with a big pronotum and short, rather broad elytra.
The absence of frontal grooves easily separates it from other New Zealand rhysodines.
Range. - This species is confined to the North Island of New Zealand where it seems to be restricted
to the northern half of the island. In addition to the type material, we have seen the following specimens:
one female, Coroglen, 13-1-64, coll. P. Johns, “mixed podocarp & broadleaf forest” (LCC); one female,
Kopu Rd., Coromandel Range, 30 Sept., 1967, coll. B. May (DSIR); one male. Little Barrier Island,
coll. H. Swale (BMNH); one male, Parakao, Whangarei, 23-IX-1956, coll. R.A. Crowson (CAS); one
female, Tamaha, North Auckland, no date, coll. C.E. Clarke (AIM); one male, Titirangi, 18-9-1915
(BMNH); one male, Whangarei, Mar. 18-20, 1931, coll. E.S. Gourley (DSIR).
Quaest. Ent., 1979, 15 (4)
382
Bell and Bell
Genus Dhy sores Grouvelle 1 903
(Fig. 1-8)
Type species. - Rhysodes thoreyi Grouvelle 1903.
Description. - Part I, 54.
Type species. - D. thoreyi of South Africa occurs from sea level to an elevation of 6000 feet (Brinck
1965). All the remaining species live in tropical Africa, where they are confined to montane forests.
They are allopatric, and, with the possible exception of D. quadriimpressus, each is limited to a single
mountain range. Adults of all the species are much alike externally, but males show clear-cut differences
in genitalia. The tropical species do not form clear-cut groups among themselves, suggesting that each of
them may have evolved from a relict population of D. thoreyi, assuming that the latter species was once
able to spread northward during a period of cooler and moister climate.
KEY TO SPECIES
1 Pronotum with discal striole plus basal impression 87% of length of pronotum;
prosternum with well-developed precoxal carinae Dhysores pan new species, p. 382
V Discal striole plus basal impression 50-80% of length of pronotum; precoxal carinae
rudimentary or absent 2
2 (F) Anterior tentorial pits and prefrontal pits both large and deep, in form of prominent
rectangle Dhysores quadriimpressus (Grouvelle), p. 384
2' Anterior tentorial pits small; prefrontal pits varied from large to minute 3
3 (2') Pronotum narrow - sides nearly parallel, base and apex only slightly narrowed; small,
narrow beetles, length 4. 6-5.0 mm Dhysores liber new species, p. 384
3' Pronotum broadened at middle - sides strongly curved, base and apex distinctly
narrowed; larger, broader beetles, length 5. 2-6. 5 mm 4
4 (30 Discal striole plus basal impression 50% of length of pronotum; humeral tubercle reduced
(Fig. 8) Dhysores thoreyi (Grouvelle), p. 384
4' Discal striole 60-80% of length of pronotum; humeral tubercle larger, more prominent
(Fig. 7) 5
5 (40 Prefrontal pit deep, larger than anterior tentorial p\i.. Dhysores rhodesianus (Brinck), p. 385
5' Prefrontal pit shallow, equal to or smaller than anterior tentorial pit
Dhysores basilewskyi (Brinck), p. 385
Dhysores pan new species
(Fig. 1)
Type material. - HOLOTYPE male, labelled: “Kivu: Itombwe; 2300 m., Terr. Uvira, Nyalengwe,
N. Leleup, XI, 1959” (MRAC). The locality is in eastern Zaire, west of Lake Kivu.
Description. — Length 5.9 mm. Anterior tentorial pits small, indistinct; prefrontal pits large, oval, not at all oblique; frontal
grooves scarcely visible; two pairs of postlabial setae; pronotum relatively elongate, its margin not sinuate anterior to hind angle, latter
obtuse; base of pronotum conspicuously sinuate medial to hind angle; hind angle obtuse; discal striole (including basal impression)
about 87% of length of pronotum, rather sharply divided into basal impression 25% of length of pronotum, and long and sinuate discal
striole proper; strioles convergent to anterior fourth of pronotum, slightly divergent more anteriorly; one marginal seta; prosternum
with long precoxal carina, extended over half the distance between coxal and anterior margin; Striae VI and VII nearly complete, but
faded near humeral tubercle.
This species is easily recognized by the long discal striole and well-developed precoxal carinae.
Classification of Rhysodini
383
Figures 1-12. Fig. 1-8, Genus Dhysores\ Fig. 1-6, Head and pronotum, dorsal aspect; Fig. 1, D. pan new species; Fig. 2,
D. quadriimpressus (Grouvelle); Fig. 3, D. thoreyi (Grouvelle); Fig. 4, D. liber new species; Fig. 5, D. rhodesianus (Brinck);
Fig. 6, D. basilewskyi (Brinck); Fig. 7,8, Base of right elytron, dorsal aspect; Fig. 7, D. basilewskyi (Brinck); Fig. 8, D. thoreyi
(Grouvelle); Fig. 9-12, Genus Rhysodes\ Fig. 9,10, Head and pronotum, dorsal aspect; Fig. 9, R. sulcatus (Fabricius); Fig. 10,
R. comes (Lewis); Fig. 11,12, Hind tibia, male, apical portion. Fig. 11,/?. sulcatus (Fabricius); Fig. 12, R. comes (Lewis).
Quaest. Ent., 1979, 15 (4)
384
Bell and Bell
Dhysores quadriimpressus (Grouvelle)
(Fig. 2)
Rhysodes quadriimpressus Grouvelle 1910: 325.
Type material. - LECTOTYPE (here designated) male, labelled: “Usumbara, Nguelo” (MNHN).
The locality is at present in BURUNDI, formerly German East Africa. Arrow (1942) and Hincks
(1950) erroneously attributed it to Tanganyika. PARALECTOTYPE one female, same data as
lectotype (MNHN).
Description. — Length 6. 0-6. 8 mm. Anterior tentorial pits large, round, equal in size to prefrontal pits, the four in form of a
conspicuous square; frontal grooves relatively distinct, though very shallow; two pairs of postlabial setae; pronotum relatively narrow
anteriorly, basal impressions uniformly deep, about 70% of length of pronotum, narrowed uniformly anteriorly; hind angles
denticulate; one marginal seta; Stria VI reduced to fine row of punctures, its anterior fourth effaced; Stria VII impressed, its anterior
fourth effaced.
This species is recognized by the enlarged anterior tentorial pits. In addition to the type material, a
specimen from Tanzania possibly belongs to this species. It is a female labelled “Tanzanie:
Mts. Uluguru, Kinola, for. transition, alt. 1500-1750 m., arbres morts 6-13-VI-71, coll. L. Berger, N.
Leleup, J. Debecker” (MR AC). This specimen differs from the type material only in lacking the
marginal seta of the pronotum. It may not be conspecific with the specimens from Burundi, but a final
decision will have to await study of genitalia of males from both localities.
Dhysores liber new species
(Fig. 4)
Type material. - HOLOTYPE male, labelled: “LIBERIA, Crest of Mount Nimba 1400 meters,
15-VIII-1966. coll. E.S. Ross, K. Lorenzen” (CAS). PARATYPE female, same data (CAS).
Description. — Length 4. 6-5.0 mm. Narrower and more cylindrical than other members of the genus; anterior tentorial pits
small, punctiform; prefrontal pits shallow, small, elongate oval; frontal grooves distinct though very shallow; one pair of postlabial
setae; pronotum with margin nearly parallel, slightly sinuate anterior to scarcely denticulate hind angles; discal stride (including basal
impression) about 70% of length of pronotum; basal impression shallow; discal striole almost straight; marginal seta of pronotum
absent; precoxal carina short; Stria VI represented by fine punctures, extended almost to humerus; Stria VII very fine, shallow,
represented anteriorly by row of fine punctures.
This is the only species of Dhysores known from West Africa. It can be recognized by its small size,
narrow form, and nearly parallel-sided pronotum.
The paratype differs from all other Dhysores in that the minor setae begin on antennal segment IV.
Dhysores thoreyi (Grouvelle 1903)
(Fig. 3,8)
Rhysodes thoreyi Grouvelle 1903: 125.
Type material. - not examined by us. According to Grouvelle (1903) the type series is in the Cape
Town Museum. The species is represented by abundant material (BMNH and other museums).
Description. — Length 5. 0-6.0 mm. Microsculpture better developed than in related species, body less shining; anterior
tentorial pits relatively distinct, oblique; prefrontal pits shallow, small, punctiform, in some specimens virtually obliterated; frontal
groove very shallow to almost absent; two pairs of postlabial setae; pronotal margin with short sinuation anterior to hind angle, latter
rectangular or nearly so, minutely denticulate; base markedly sinuate medial to hind angle; discal striole, including basal impression,
about 50% of length of pronotum; discal striole anterior to basal impression narrow, shallow; marginal seta present; precoxal carina
absent; elytra more narrowed at base than in other Dhysores, humeral tubercle smaller (Fig. 8); Stria VI reduced to row of minute
punctures not extended to humerus; Stria VII effaced in anterior third and interrupted opposite apical striole, gap with several large
setigerous punctures; penis with tip tapered to narrow point on left side.
Classification of Rhysodini
385
This species is easily recognized by the reduced humeral tubercles, narrow, oblique anterior tentorial
pits, markedly reduced or obsolete prefrontal pits, and short discal striae. Dhysores thoreyi is known
only from the Union of South Africa, where it is found in Natal Province and the eastern Cape Province,
from sea level to 6000 feet in altitude (Brinck 1965).
Brinck (1965) provides a map of the localities.
Dhysores rhodesianus (Brinck 1965)
(Fig. 5)
Rhysodes (Dhysores) rhodesianus Brinck 1965: 467-468.
Type material. - According to Brinck, the type series includes many adults in the Transvaal
Museum, from RHODESIA: Mt. Selinda Forest, 9-17.4, 1956, leg. G. Van Son and L. Vari. We have
not seen this material, but have studied a PARATYPE female, labelled: “RHODESIA: Chirinda
Forest, Dec., 1901, leg. G.A.K. Marshall” (BMNH).
Description. — Length 5. 2-6. 6 mm. Anterior tentorial pits punctiform, small but distinct; prefrontal pits large, oval, somewhat
oblique; frontal grooves evident though very shallow; two pairs of postlabial setae; pronotal margin sinuate anterior to minutely
denticulate hind angle, base markedly obliquely sinuate medial to hind angle; discal striole including basal impression about 80% of
length of pronotum; anterior portion of discal striole very shallow; strioles convergent anteriorly except extreme anterior ends parallel
to one another; marginal seta present; prosternum without trace of precoxal carina; Stria VI and VII almost entire, effaced only in
anterior tenth of elytron; penis with apex triangular, in form of short, obliquely truncated apical lobe.
The elongate discal strioles give this species a resemblance to Dhysores pan, from which it is easily
separated by the sinuation anterior to the hind angles and absence of a precoxal carina. The
well-developed obliquely oval prefrontal pits are unique in Dhysores, and recall those of Neodhysores.
In addition to the paratype we have seen one male specimen in MRAC labelled “Chriinda
(misspelled?) F., XII-52, vSon”.
Dhysores basilewskyi (Brinck 1965)
(Fig. 6,7)
Rhysodes (Dhysores) basilewskyi Brinck 1965: 468-469.
Type material. - HOLOTYPE male, labelled: “RUANDA, Astrida, 1750 m., 22-2-1953, coll.
P. Basilewsky” (MRAC). PARATYPE female, same data as type (MRAC).
Description. — Length 5. 0-6.5 mm. Anterior tentorial pits scarcely evident; prefrontal pits medium-sized, oval, moderately
deep, not oblique; mentum with one pair of postlabial setae; lateral margins of pronotum distinctly sinuate anterior to denticulate hind
angles; sinuation medial to hind angle shorter than in related species; discal striole, including basal impression 60-75% of length of
pronotum; discal strioles convergent anteriorly, very shallow near apex; marginal seta of pronotum absent from most specimens
(present unilaterally in one specimen); Stria VI and VII completely effaced in anterior third of elytron; apex of penis much more
obtuse than in Dhysores thoreyi and Dhysores rhodesianus males.
The denticulate hind angles, shorter discal strioles, and the absence of precoxal carinae separate this
species from Dhysores pan. The reduced anterior tentorial pits separate it from Dhysores
quadriimpressus, while smaller prefrontal pits, the shorter discal striole and the more extensive
abbreviation of Striae VI and VII are the most obvious differences from Dhysores rhodesianus.
In addition to the type material, a series of six specimens from Zaire appear to belong to this species.
These are labelled “B. CONGO, 42 mi. N. of Kapona, 1520 m., 1-13-58, coll. E.S. Ross, R.E. Leech”
(CAS).
Quaest. Ent., 1979, 15 (4)
386
Bell and Bell
Genus Neodhy sores Bell and Bell 1978
Type species. - Neodhy sores seximpressus Bell and Bell 1978
Description. - Part I, 56.
KEY TO SPECIES
1 Pronotum relatively short; length/greatest width 1.28; antennal Segment XI as wide as
long Neodhysores seximpressus Bell and Bell, p. 386
T Pronotum elongate, length/greatest width 1.43; antennal Segment XI much longer than
wide Neodhysores schreiberi (Vulcano and Pereira), p. 386
Neodhysores seximpressus Bell and Bell 1978
Type material. - Part I, 56. The type locality is Nova Teutonia, Santa Catarina State, Brazil.
Description. - Part I, 56.
Neodhysores schreiberi (Vulcano and Pereira) 1975a)
Rhysodes schreiberi Vulcano and Pereira 1975a: 219.
Type material. - One type and one paratype in the collection of the describers; we have not studied
the types. The type locality is Belo Horizonte, Minas Gerais State, Brazil.
Description. - Vulcano and Pereira 1975a: 219.
SUBTRIBE RHYSODINA
Description. - Part I, 56.
Key to Genera. - Part I, 57.
Genus Rhysodes Dalman 1823
Fig. 9-12
Type species. - Cucujus sulcatus Fabricius 17^1 Rhysodes exaratus Dalman 1823, by monotypy)
Synonym. - Epiglymmius Lewis 1888: 79 (Type species Cucujus sulcatus Fabricius 1787,
designated by Hincks, 1950)
Description. — Part I, 58. Prosternum without precoxal carina.
KEY TO SPECIES
1
Pronotum relatively short, length/greatest width 1.1; discal striole of pronotum well
separated from anterior margin Rhysodes sulcatus (Fabricius), p. 387
Classification of Rhysodini
387
Pronotum more elongate, length/greatest width 1.2; discal striole almost reaching
anterior margin of pronotum Rhysodes comes (Lewis), p. 388
Rhysodes sulcatus (Fabricius 1787)
(Fig. 9,11)
Cucujus sulcatus Fabricius 1787: 165.
Rhysodes exaratus Dalman 1823: 93.
Rhizodes europaeus Ahrens 1814: 1.
Type material. - Not located.
Description. — Length 6. 8-8. 3 mm. Antennal Segment XI slightly less than twice as long as wide; stylet more than 0.33 of
length of cone; clypeus slightly emarginate in midline anteriorly; narrow oblique bridge connecting clypeal rim to median lobe,
interrupting clypeal groove anterior to each anterior tentorial pit; frontal grooves oblique, sinuate; median lobe distinctly narrowed
between openings of frontal pit, medial emargination of temporal lobe relatively wide and deep; anterolateral projection of temporal
lobe relatively approximate to antennal rim; latter with distinct narrow brace directed posteromedially from its posterior end, opposite
distinct emargination in temporal lobe; orbital groove relatively dilated; base of mentum in form of distinctly projecting lobe in lateral
view; mentum with numerous short setae in form of “beard” in male, only scattered setae in female; gular grooves with anterior halves
indistinct.
Pronotum relatively short and broad, length/greatest width 1.1, both apex and base more narrowed than in R. comes\ discal striole
plus basal impression 80% of length of pronotum; basal impression about 33% of length of pronotum; hind angles acute; lateral
margins parallel in basal fifth, divergent Just anterior to hind angles; lateral margin deeply sinuate anterior to hind angles; precoxal
rim with short stub representing rudimentary precoxal carina; punctures of prosternum each with prominent seta.
Elytral intervals flatter than in R. comes-. Stria Vll with five to seven setae near apex; apical striole with three to four setae.
Anterior and middle tibiae hairy beneath, more densely so in male; all tibiae hairy medially in both sexes; middle tibia of male
with conspicuous lateral setae, that of female with few setae; hind calcar large, pointed, only slightly dorsad to spur ( Fig. 11).
Range. - Widely distributed in Europe, but now confined to small, scattered relict areas, probably
because centuries of intensive use of wood have restricted the development of overmature trees. Dajoz
(1975) gives a detailed account of the recent and nineteenth century distributions. The westernmost
localities are in the Pyrenees of southern France. The species also occurs in Italy, Germany, Poland,
Ukrainia, Yugoslavia, Rumania, and Sweden. We have seen specimens (NMNH) from Asiatic Turkey,
from the Taurus Mountains near Adana. Dajoz also lists it from the Caucasus. It does not now occur in
Britain, but is known from subfossil remains (Kelley and Osborne, 1965; Buckland and Kenward, 1972)
the most recent remains being from about 1000 B.C.
Bionomics. - Despite its rarity, Rhysodes sulcatus is better known ecologically than any other species
of Rhysodini. Dajoz (1975) and Burakowsky (1975) have summarized previous work and have added
their personal observations about the species. Tiberghien (1969) gives a detailed account of R. sulcatus
in the Pyrenees. Considering the accounts of these authors together, it is evident that Rhysodes sulcatus
depends for its survival on the presence of very large, old trees. It is seemingly indifferent to the species
of tree. It has been reported from fir {Abies alba Mill.), spruce {Picea excelsa (Law.) Lk., beech {Fagus
silvatica L.), aspen {Populus tremula L.), and other species of poplar {Populus spp.). It is confined to
dead wood, but apparently is not particular about the state of the wood, occurring in badly decayed logs,
as well as dry, hard ones and even charred stumps in a burned area. It is known from both standing and
fallen trunks. Waga (1841, quoted by Grouvelle, 1903) collected specimens from the roots of poplar
{Populus) at depths down to two meters.
Tiberghien (1969) found Rhysodes sulcatus specimens associated with myxomycetes in burnt stumps
of fir {Abies), while Dajoz (1975) collected them in stumps of beech {Fagus) which were infested with
oyster fungus {Pleurotus ostreatus). Burakowsky (1975) states that Trella (1926, 1939) found numbers
of adults hibernating in galleries of the larvae of Ceruchus chrysomelinus Hochw. (Coleoptera,
Lueanidae) in a rotten fir {Abies) trunk.
Quaest. Ent., 1979, 15 (4)
388
Bell and Bell
Burakowsky (1975) has described and illustrated larvae and pupae as well as illustrating many
details of structure of adults. He believes that two years are required to complete metamorphosis, as
shown by the presence of two size classes of larvae during the summer. Larvae, unlike adults, live in
well-defined galleries. These are in the moister, more decayed parts of wood. The older parts of the
gallery, behind the larvae, are tightly packed with wood fragments.
Rhysodes comes (Lewis 1888)
(Fig. 10,12)
Epiglymmius comes Lewis 1888: 79.
Type material. - LECTOTYPE (here designated) male, labelled: “JAPAN: Nikko, Aug. 10-18,
1881, coll. G. Lewis” (BMNH). PARALECTOTYPES, two females, JAPAN: Sapporo, Aug. 5-6,
1880, coll. G. Lewis (BMNH).
Description. — Length 6. 5-8.0 mm. Antennal Segment XI more than twice as long as wide; cone elongate, stylet less than 33%
as long as cone; clypeus truncate anteriorly, not emarginate; bridge from median lobe to margin of clypeus incomplete or lacking;
frontal grooves almost longitudinal, in form of acute angle with antennal grooves; median lobe narrowed opposite frontal pits; medial
emargination of temporal lobe smaller and shallower than in Rhysodes sulcatus; temporal lobe without distinct anterolateral
projection; antennal rim without posteromedial brace; temporal lobe scarcely emarginate opposite antennal rim; antennal groove
nearly straight; orbital groove relatively narrow, in contact with antennal groove at slight angle anterior to eye; mentum with “beard”
in male, scattered setae in female; base of mentum less prominent in lateral view than in Rhysodes sulcatus-, gular groove complete.
Pronotum longer and narrower than in Rhysodes sulcatus, length/greatest width about 1.20; base and apex not as strongly
narrowed as in Rhysodes sulcatus-, basal impession less than 0.25 of total length of pronotum; discal strioles elongate, more than twice
as long as basal impressions, extending almost to anterior margin of pronotum; hind angles slightly obtuse; sides of pronotum not
parallel anterior to hind angles; lateral sinuation much shallower than in Rhysodes sulcatus-, precoxal carina absent; punctures of
prosternum without evident setae.
Elytral intervals more convex than in Rhysodes sulcatus-, elytron with one seta at tip of Stria IV, three to six at tip of Stria VII;
and with or without one in apical striole.
Male with about four ventral setae on anterior femur, and with or without one ventral seta on middle femur; female without
ventral setae on femora; tibiae with only a few medial setae; middle tibia without lateral setae; calcar of hind tibia very small, very
obtusely angled, distinctly dorsad to spur (Fig. 12).
Range. - Japan and Siberia. In Japan, known from the following islands; HONSHU: Nikko (the
lectotype); Sanno-Toge, Oku-Nikko (DY); Yumato, Iwase (DY); Ooda lima-Gnu (Mie Pref.) (SATO);
KYUSHU: Wakamatsu (DY); HOKKAIDO: the paralectotypes. The Siberian specimen, the first
record from the U.S.S.R., is from Kongaus, Siberia, coll. Cockerell, August, 1923. The locality is on
Nakhodka Bay, about 75 miles east of Vladivostok.
Genus Kupea Bell and Bell 1978
Type species. - Clinidium arcuatum Chevrolat 1873a.
Description. - Part I, 58. Only one species is known.
Kupea arcuatus (Chevrolat 1873a)
Clinidium arcuatum Chevrolat 1873a: 216.
Rhysodes aterrimus Broun 1880: 214 (nec Chevrolat 1873a)
Rhysodes brouni Lewis 1888: 79.
Classification of Rhysodini
389
Type material. - Rhysodes arcuatus: HOLOTYPE male, no locality, labelled: ^"Rhyz. antarcticus ”,
as far as we can determine an unpublished name (MNHN); R. aterrimus Broun; LECTOTYPE - sex
not determined, labelled: “NEW ZEALAND: Tairua” (BMNH-Broun Collection).
PARALECTOTYPES two specimens, sex not determined (BMNH-1 in general collection, the other in
Broun Collection). Rhysodes brouni Lewis was a substitute name for Rhysodes aterrimus Broun,
preoccupied by Rhysodes aterrimus Chevrolat 1873a.
Description. — with characters of genus; length 5. 0-8. 0mm. Pronotum with angular seta but without marginal setae;
prothorax with several postcoxal setae; precoxal carina absent; Stria 11 with one to two setae near tip; Stria IV with one seta near base
and another near tip; Striole with one or more setae; Stria VI 1 with about five setae in apical fifth; middle tibia with prominent acute
calcar, directed ventroanteriorly; calcar of hind tibia smaller than that of middle tibia, acute, located entirely dorsad to apex of tibia.
This species is easily recognized among New Zealand Rhysodini by the form of the head, with the
narrow, parallel-sided median lobe separated on either side from the temporal lobe by a linear frontal
groove.
Range. - The North Island of New Zealand and nearby coastal islands. The locality records indicate
that it occurs throughout the North Island. In addition to the type material, we have seen specimens
from the following localities: one female. Boatmans Reefton (BMNH); one female, Hunua, coll. Drury (BMNH); three
males, two females, Kawau I., 6-1902, coll. J.J. Walker (BMNH); one male, three females. Little Barrier 1., coll. H. Swale, 1913
(BMNH); one female, Mararainui, Bay of Plenty, May, 1928, coll. A.E. Brookes (DSIR); two males, Maud 1., 1-6-3-53, coll.
E.S. Gourley (DSIR); two males, one female, Mayar 1., Dira Bay, 11 Nov. 55, coll. J.C. Watt (DSIR); one female, Mayar 1., Te
Ohineitid, 14 Nov. 55, coll. J.C. Watt (DSIR); one female, Okauai, Maramara, Waikato, 1-1 1-1931, coll. A.E. Brookes (DSIR); one
female, Oruru, 6-9-10, coll. A.E Brookes (DSIR); one male, Papakura, Auckland, 26-12, 1926, coll. A.E. Brookes (DSIR); one male,
Paparoa, coll. A.E. Brookes (DSIR); one female, Plummerton (MCZ); one male, one female, Pollok, Auckland, 7-1-64, coll. P. & M
Johns (LCC); one female, Ratanihipiki, Taranaki (BMNH); one female. Springs Junct. Br., 5-V-1977, coll. R.M. Emberson (LCC);
three males, one female, Titirangi, 21-3-1913 (Broun Colin.) (BMNH), same locality, 1913-14, coll. T. Broun (DSIR), same locality,
5-10-27, coll. E.S. Gourley (DSIR); one male, Waikanae, lX-1930, coll. G.V. Hudson (BMNH), one male, same locality. Sept. 5,
1938, “GVH” (DSIR); two females, Waipoua, 4-12-63, Kauri bark, coll. P.M. Johns (LCC); one female, Waitakeri, 31-12-14, coll.
A.E. Brookes (DSIR); six males, four females, Wellington (may refer either to province or city) (BMNH); one male, Whangarei,
Western Hills, 13-1X-1956, coll. R.A. Crowson (CAS); one male, Whangarei, 18-20,3,31, coll. E.S. Gourley (DSIR); one male,
Whangarei Dist., Maunga Karamea, 13-2-1926, coll. E. Fairburn (DSIR); two males, two females, Wilton’s Bush, 2-1 1-44, coll. G.V.
Hudson (BMNH), same locality, 6-1 1-20 (DSIR).
Genus Kaveinga Bell and Bell 1978
(Fig. 13-51)
Type species. - Rhysodes abbreviatus Lea 1904.
Description. — Antennal stylet absent except in one undescribed Australian species; antennal Segment V without minor setae,
latter on Segments Vl-X, but confined to more distal articles in specimens of a few species; labrum with two setae; medial margin of
temporal lobe curved or oblique, not closely parallel to margin of median lobe; temporal and medial lobes entirely separated or else in
contact for short distance posteriorly; orbital groove, if present, terminated near posterior margin of eye; pronotum in most species
with complete, entire paramedian grooves (in Kaveinga orbitosa) paramedian groove represented by row of very coarse punctures);
precoxal carinae present on prosternum; humeral tubercle well developed; middle and hind tibiae each with one spur.
In many species, the tip of the median lobe is supported ventrally by a slender pillar. This feature is
easily seen in species such as K. histrio, in which the median lobe is markedly raised above the pronotum
but would require dissection to verify in those species in which the head is low and wide. One specimen of
K. abbreviata had the tip of the median lobe broken off, revealing a pillar. Perhaps this feature will
prove to be an invariable characteristic of Kaveinga. The pillar is entirely posterior to the frontal pit, so
that the latter forms a transverse passageway beneath the median lobe (Fig. 37,47).
This markedly diverse and divergent genus differs from its relatives in having only two labral setae,
and in having precoxal carinae. It differs from Rhysodes in having distinct humeral tubercles and in
having the orbital groove ending at or anterior to the posterior margin of the eye. It differs from Kupea
in having the paramedian grooves complete or nearly so, and in not having the margins of the medial
lobe and temporal lobes parallel and close to one another along the entire length of the head.
Quaest. Ent., 1979, 15 (4)
390
Bell and Bell
Kaveinga ranges from New Zealand, New Caledonia and the Santa Cruz Islands through Australia
and New Guinea to Buru in the Moluccas and Mindanao in the Philippines.
Phytogeny. - The described species of Kaveinga are included in four subgenera. The subgenera can
be grouped into pairs which appear to represent major phyletic lines. Within each pair, the two genera
are allopatric. In Vakeinga and Kaveinga (sensu stricto), the paramedian grooves are broad, deep,
entire, and impunctate. Vakeinga occurs in New Zealand and New Caledonia, while Kaveinga (sensu
stricto) occupies the remainder of the range of the genus. The second pair of subgenera appear less
certainly related to one another, though they are linked together by having the pronotal grooves coarsely
punctate. Angekiva, confined to Australia, have the paramedian grooves entire, curved, and nearly
linear. Ingevaka, confined to New Zealand, have the paramedian grooves represented by a series of very
coarse punctures.
There is an undescribed species from Australia which does not fit into the above subgenera, and may
require erection of a fifth subgenus, although it may prove to be an aberrant species of Angekiva. This
species will be described by Dr. Moore in his monograph of the Australian Rhysodini. This species was
mistakenly regarded as Rhysodes lignarius Olliff by Grouvelle. (The true R. lignarius belongs to
Leoglymmius.) There are two specimens of the undescribed species in the MNHN, one labelled
“Richmond R., N.S. Wales”, the other without specific locality data. We have also seen two specimens
in the MCZ collection, one from the McPherson Range, Queensland, and the other from rain forest
north of Dunoon, New South Wales.
The undescribed species differs from all other Kaveinga in the presence of an antennal stylet. Both
sexes have a ventral tooth on the anterior femur, though that of the female is very small. The pronotum
is like that of Angekiva, except that the paramedian grooves are scarcely punctate.
KEY TO SUBGENERA
Paramedian grooves entire, open both anteriorly and posteriorly 2
Paramedian groove incomplete, closed both anteriorly and posteriorly, represented by
row of very coarse punctures Ingevaka new subgenus, p. 394
Paramedian grooves linear, markedly curved, punctate; pollinosity, if present, limited to
punctures Angekiva new subgenus, p. 390
Paramedian grooves not linear, not markedly curved, without punctures, pollinosity
continuous from base to apex 3
Temporal setae numerous; marginal setae of pronotum numerous, in form of continuous
row Vakeinga new subgenus, p. 395
Temporal setae 1-3; marginal setae 0-3, when present, limited to anterior and posterior
ends of marginal groove Kaveinga (sensu stricto), p. 397
Subgenus Angekiva new subgenus
Type species. - Rhysodes frontalis Grouvelle 1903.
Description. — Antennal Segment Xi without stylet; antennal segments not pollinose; median groove of pronotum coarsely
punctate; paramedian grooves curved, linear except at base; complete, coarsely to sparsely punctate; pollinosity restricted to punctures,
or else absent; angular seta present or absent; marginal setae absent; prosternum coarsely punctate, punctures either generally
distributed or else confined to margins of precoxal carinae; elytral striae glabrous; femora glabrous; femur of anterior leg without
ventral tooth in either sex (in species where both sexes are known); middle tibia without lateral serrulation, and without lateral setae.
1
V
2 (1)
1'
3 (20
3'
Classification of Rhysodini
391
Figures 13-24. Fig. 13-51, Genus Kaveinga; Fig. 13, Flead and pronotum, dorsal aspect, K. (Angekiva) frontalis (Grouvelle);
Fig. 14-16, K. (Ingevaka) orbitosa (Broun); Fig. 14, Head and pronotum, dorsal aspect; Fig. 15, Head and pronotum, lateral
aspect; Fig. 16, Head, anterior aspect, antennae omitted; Fig. 17-19, K. (Vakeinga) lusca (Chevrolat); Fig. 17, Head and
pronotum, dorsal aspect; Fig. 18, Middle tibia; Fig. 19, Head and pronotum, lateral aspect; Fig. 20,21, K. (V.) selosa
(Grouvelle); Fig. 20, Head and pronotum, dorsal aspect; Fig. 21, Head and pronotum, lateral aspect; Fig. 22-27; Head and
pronotum, dorsal aspect; Fig. 22; K. {Kaveinga} abbreviata (Lea); Fig. 23, K. (K.) fibulata, new species; Fig. 24, K. (K.) kukum
new species.
Quaest. Ent., 1979, 15 (4)
392
Bell and Bell
Figures 25 - 39. Fig. 25, K. (K.) pignoris new species; Fig. 26, K. (K.) ulteria new species; Fig. 27, K. (K.) nudicornis new
species; Fig. 28,29, Antennal Segments IX-XI; Fig. 28, K. (K.) pignoris new species; Fig. 29, K. (K.) nudicornis new species;
Fig. 30,31, Hind tibia, male, apical portion; Fig. 30, K. (K.) pignoris new species; Fig. 31, X. (K.) ulteria new species; Fig. 32-35
K. (K.) cylindrica (Arrow); Fig. 32, Head and pronotum, dorsal aspect; Fig. 33, Head and pronotum, lateral aspect; Fig. 34,
Head, anterior aspect, antennae omitted; Fig. 35, Base of right elytron, dorsal aspect; Fig. 36-39, K. (K.) lupata new species;
Fig. 36, Base of right elytron dorsal aspect; Fig. 37, head, posterolateral aspect; Fig. 38, Head and pronotum, dorsal aspect;
Fig. 39, Head and pronotum, lateral aspect.
Classification of Rhysodini
393
Figures 40 - 51. Fig. 40, K. (K.) okapa new species; Fig. 41, A^. {K.} marifuanga new species; Fig. 40,41 Flead and pronotum,
dorsal aspect; Fig. 42, Middle tibia, K. (K.) marifuanga new species; Fig. 43,44, Head and pronotum, dorsal aspect; Fig. 43,
K. (K.) occipitalis (Grouvelle); Fig. 44, K. (K.) parva (Grouvelle); Fig. 45-48, K. (K.j histrio new species; Fig. 45, Head and
pronotum, dorsal aspect; Fig. 46, Head and pronotum, lateral aspect; Fig. 47, Head, posterolateral aspect; Fig. 48, Head,
anterior aspect, antennae omitted; Fig. 49-51, K. (K.) strigiceps new species; Fig. 49, Head and pronotum, dorsal aspect; Fig. 50,
head and pronotum, lateral aspect; Fig. 51, head, anterior aspect, antennae omitted.
Quaest. Ent., 1979, 15 (4)
394
Bell and Bell
The curved, linear, punctate paramedian grooves are diagnostic of this subgenus. The type species is
the only described species. However, we have also studied two unnamed species which will be described
by Dr. Moore. One species has large eyes which are deeper than long, a complete orbital groove, and a
very shallow, glabrous postantennal groove. This species is known from Malanda, north Queensland
(MCZ), and from Mt. Kairi, Queensland (CAS).
The second species resembles K. frontalis in having the eye reduced, the orbital groove absent, and
the postantennal groove deep. It differs in having the median lobe of the head very slender, strongly
constricted near the middle, the median pit of the clypeus very small, and the eye circular. This species is
represented by a specimen from Mt. Spurgeon, northern Queensland (MCZ).
Kaveinga (Angekiva) frontalis (Grouvelle 1903) NEW COMBINATION
(Fig. 13)
Rhysodes frontalis Gxo\x\qWq 1903: 104-105.
Type material - HOLOTYPE male, labelled: “TASMANIA” (MNHN).
Description. — Length 7 mm. Antennal Segment XI obtuse, 1.5 times longer than wide; minor setae on Segments VI-X; basal
setae on Segments VI-X. Head as wide as long; clypeal setae absent; clypeus with enlarged median pit; clypeal grooves dilated;
postclypeal grooves incomplete, so median lobe is connected to parafrontal bosses; median lobe relatively broad, about 0.25 of width of
head; median lobe only slightly constricted anteriorly; postantennal grooves deep, pollinose; one temporal seta; orbital groove reduced
to minute vestige; eye deeper than long, apparently slightly reduced (in holotype very heavily pigmented so that it is difficult to see its
outlines).
Pronotum longer than wide, oval, widest near middle, its base and apex both strongly narrowed; sides strongly curved; angular seta
absent; prosternum and propleura coarsely punctate.
Elytral setae restricted to apex of Stria VII; metasternum coarsely punctate; abdominal Sterna entirely coarsely punctate; male
with very small ventral tooth on anterior femur; female unknown.
Subgenus Ingevaka new subgenus
Type species. - Rhysodes orbitosus Broun 1880
Description. — Antennal Segment XI without stylet; antennal segments not pollinose; median groove of pronotum coarsely
punctate; paramedian grooves nearly straight, coarsely punctate; impressed posteriorly, reduced to row of punctures anteriorly; closed
both anteriorly and posteriorly; angular seta present; marginal seta varyingly developed, most often absent; anterior lateral pits absent;
elytral striae shallow, glabrous; anterior femur of male with ventral tooth in some specimens, absent from others; females without
ventral tooth on anterior femur; tibia of middle leg without lateral serration or lateral setae.
The incomplete paramedian grooves are the mark of this subgenus. The short, high, transverse head
resembles those of certain species of Kaveinga (sensu stricto), surely as a result of convergent evolution.
Subgenus Ingevaka contains only one species, which is confined to the North Island of New Zealand.
Kaveinga (Ingevaka) orbitosa (Broun 1880) NEW COMBINATION
(Fig 14-16)
Rhysodes orbitosus Broun 1880: 215-216.
Rhysodes luscus auct., nec Chevrolat 1875.
Rhysodes eminens auct., nec Broun 1880
Lewis (1888) wrongly supposed this species to be the male of R. eminens Broun {Rhysodes luscus
Chevrolat).
Type material. - We have studied a specimen labelled as cotype in BMNH. It is without locality
data. We have not located any other type material. According to the original description the type
material was collected at Whangarei Harbour.
Classification of Rhysodini
395
Description. — Length 4-5 mm. Body very short and transverse for a rhysodine; head short, strongly transverse, about twice as
wide as long; clypeal setae present; clypeus separated from median lobe by pollinose impression; clypeus with three conspicuous pits;
clypeus continuous laterally with antennal lobes; each antennal lobe with two medial braces; median lobe short, high, in form of raised
crest (Fig. 16), its anterior end acutely pointed, its posterior end rounded; temporal lobes markedly transverse, their medial margins
fitted into cavities on sides of median lobe; three to four temporal setae, each inserted in conspicuous pit (in many specimens pits fused
in form of one elongate depression; in other specimens pits separate, in one specimen pits fused on left lobe and separate on right lobe);
eye markedly reduced, about twice as long as wide, with about sixty ommatidia; cornea thickened, without facets.
Pronotum subquadrate, widest anterior to middle; sides slightly curved and distinctly sinuate well anterior to posterior angles;
marginal groove distinct; prosternum with wide groove medial to each precoxal carina; prosternum impunctate; propleuron with a few
very coarse punctures.
Elytra very short, convex, inflated; strial punctures very coarse; Stria 1 with two setae in apex; Stria II and IV with setae both at
apex and at base; in many specimens with continuous series of setae in either II or IV, in a few with continuous series in both striae;
apical striole with several setae; apex of Stria VII with several setae; metasternum concave in midline, but not truly suleate;
metasternum with median line of coarse punctures, a cluster of punctures anterior to each hind coxa, and with five to six large lateral
punctures; abdominal sterna Ill-V each with narrow transverse, almost uniseriate band of punctures; calcar of male small and pointed.
Variation. - Seven of the nine males examined had well-developed ventral teeth on the anterior
femora. Two males differ strikingly in lacking the teeth. One of these is from Little Barrier Island
(BMNH), and the other (NMNH) is without locality data. One female from Waitakiri (DSIR) has
numerous marginal setae on the pronotum, while in all other specimens the marginal setae are restricted
to one near the angular setae or are entirely absent. The variations listed above are quite comparable to
specific differences seen in other groups of Rhysodini, so it is possible that future work will show that
there are several species of Ingevaka in New Zealand.
Range. - We have seen specimens with the following locality data: AUCKLAND: one male. Little Barrier i.,
coll. H. Swale, 1913-117 (BMNH); one male, Okauia, Matamata, Waikato, A.E. Brookes Colin., 11-3-1923 (DSIR); one female,
Omahuta, Kauri foret, Jan. 1972, coll. G.W. Ramsay (DSIR); one male, Waimatenui, Hobson Co., C.E. Clarke Colin. (AIM); one
male, one female, Wainui Bay, saddle, Kaea Northland, 4-Nov-67, coll. J.l. Townsend (DSIR); one female, Waipqua, 15 June 66,
coll. J.L Townsend (DSIR); one male, one female, Whangerei, Heads, C.E. Clarke colln. (AIM); one male, Warkworth, The Dome,
21 Oct. 1962, coll. J.C. Watt (DSIR); one female, Whangarei, Tiraki, A.E. Brookes colln, 8-9-23 (DSIR); one male, Whangarei,
Western Hills, 13-1X-1956, coll. R.A. Crowson (CAS); HAWKES BAY(?): two females, Waipaua S.F., VI-9, 13-66, coll. J.C. Watt
(DSIR); one female, Waitakiri, 26-10-14(380), T. Broun A.E. Brookes colln. (DSIR).
Subgenus Vakeinga new subgenus
Type species. - Rhysodes setosus Grouvelle 1903
Description. — Antennal Segment XI without stylet; antennal Segment I finely pollinose above; remaining antennal segments
without pollinosity; minor setae on Segments Vl-X; head with distinct parafrontal bosses; four to five temporal setae; orbital groove
absent or vestigial; grooves of pronotum deep, pollinose, impunctate; paramedian grooves relatively broad, open anteriorly and
posteriorly; inner carinae not abbreviated anteriorly; marginal groove complete; angular seta present; five to nine marginal setae;
prosternum, propleura impunctate; several postcoxal setae on prothorax; elytral striae without pollinosity; elytron with several setae in
apices of Striae 1 and Vll, and complete series in striae II and IV (setae sparse or absent from middle third of Striae II and IV in some
specimens; female with small lateral pits in Sterna IV-V; femora glabrous; anterior femur without ventral tooth in either sex; middle
tibia with row of setae on lateral aspect, but without lateral serrulation.
This subgenus is similar to the allopatric Kaveinga (sensu stricto) in the structure of the prothorax. It
differs from the latter in having many lateral setae, and in lacking lateral serrulation on the middle tibia
(Fig. 18). The deep, entire, pollinose, impunctate paramedian gooves are the best means of separating it
from the sympatric Ingevaka. Vakeinga has one species in New Zealand, and one in New Caledonia.
KEY TO SPECIES
1 Eye large, round, its diameter about half depth of head; pronotum without setae at bases
of inner carinae Kaveinga setosa (Grouvelle), p. 396
Quaest. Ent., 1979,15 (4)
396
Bell and Bell
r Eye very small, oblique, its diameter less than 0.2 depth of head; one seta on each inner
Carina near its posterior end Kaveinga lusca (Chevrolat), p. 396
Kaveinga (Vakeingaj setosa (Grouvelle 1903) NEW COMBINATION
(Fig. 20,21)
Rhysodes setosus Grouvelle 1903: 108.
Type material - According to Grouvelle, the type was in the Oberthur collection. We could not
locate it in MNHN. However, the description and the locality are sufficient to recognize the species.
Description. — Length 4. 8-5.0 mm. Minor setae well developed on antennal Segments VLX; basal setae absent; head slightly
longerthanwide;medianlobenarrowedbetweeneyes,itsposteriorendspatulate,pointed;anteromedialmarginoftemporallobestrongly
angulate midway between eye and medial angle; orbital groove vestigial; eye large, nearly round, its diameter over half depth of head;
cornea facetted, unpigmented.
Pronotum moderately elongate, length/greatest width 1.20, widest near middle, sides strongly curved; inner pronotal carina
without seta near base; five to six marginal setae; prosternum with pair of pits between precoxal carinae; elytra narrow, cylindrical;
striae coarsely punctate.
This species is easily distinguished from the next by: fully developed eyes, absence of seta from the
base of the inner carina, broader median lobe, shorter head and pronotum, and absence of basal setae
from the antennae.
Range. - New Caledonia. The only specimens we have seen are two males and one female labelled:
“NEW CALEDONIA, Foret de Thi, Hanna, VII-16-1958, leg. B. Malkin” (CNHM).
Kaveinga (Vakeingaj lusca (Chevrolat 1875) NEW COMBINATION
(Fig. 17-19)
Rhysodes luscus Chevrolat 1875: 183.
Rhysodes eminens Broun 1880; 215.
Type material. - R. luscus: not located by us. R. eminens: “COTYPE”, sex not recorded, without
specific locality data in Broun Collection (BMNH). Broun states that the original specimens were
collected at Whangarei Harbour. We do not know whether or not there are additional type specimens.
Grouvelle (1903) states that he had studied types of R. luscus and had concluded that R. eminens and
R. orbitusus were conspecific, but this is not so. The original description of R. luscus agrees with that of
R. eminens and not R. orbitosus.
Description. — Length 4. 7-6. 6 mm. Minor setae sparse on Segment VI, well developed on Segments Vll-Xl; basal setae on
Segments VI-X, head distinctly longer than wide, median lobe narrower than in K. setosa, convex, constricted between eyes;
anteromedian margin less distinctly angulate than in K. setosa; orbital groove absent; eye markedly reduced and modified; cornea
oblique, twice as long as deep; cornea not facetted, clear in some (presumably younger) specimens, markedly pigmented in other
specimens; ommatidia about twenty, grouped in round central disc much smaller than cornea.
Pronotum distinctly elongate, length/greatest width about 1.25; sides distinctly curved; inner carina with seta near base; seven to
nine marginal setae; precoxal carinae markedly developed, extended to anterior margin of prosternum; each precoxal carina bounded
medially by wide groove; these grooves closely approximated at middle of length, separated by median carina; no pits between precoxal
carinae; elytra moderately broad, somewhat flattened; striae less coarsely punctate than in K. setosa.
The reduced, oblique eyes distinguish this species from K. setosa. K. lusca differs markedly from K.
(Ingevakaj orbitosa, with which it was formerly considered conspecific. K. lusca has complete
paramedian grooves, a low, elongate head and temporal lobes which are not in close contact with the
median lobe.
Classification of Rhysodini
397
Range. - Confined to the North Island of New Zealand, where it is widespread. We have seen
specimens from the following localities: one male, one female, Herekino, Mangonui Co., G.E. Clarke Colin. (AIM); one
female, Mt. Hikurangi, 4000, 1 7-1-64, coll. P. Johns (LCC); one male, Motu R., 1 2-1 1-28, A.E. Brookes Colin. (DSlR); one male, two
females, Mt. Maungapohatu, 914-1219 m., 3-Mar-71, coll. J.I. Townsend (DSlR); one male, Parkanae Opononi, 17-XI-1968,
Hokianga Co., coll. K.A.J. Wise (AIM) one male, Pukerui Hills, Whangarei 21-1 1, coll. G. Given (DSlR); two males, Tepaki Coastal
Park, North Cape, 7 Feb 1975, coll. J.C. Watt (DSlR); three males, five females, Tepaki Coastal Park, S. Pandora, 7 Feb 1975, coll.
S.E. Nichols (DSlR); one male, Unuwhao trig.. Spirits Bay, 20-28-V111-1957, coll. J.C. Watt (DSlR); three males, Waimatenui,
Hobson Co., C.E. Clarke Colin. (AIM); two females, Wainui Bay, saddle, Kaea, 4 Nov 62, coll. J.I. Townsend (DSlR); two males,
three females, Waipoua, VI-66-, X-67, coll. J.C. Watt and Vl-66, coll. J.I. Townsend (DSlR); one male, one Whanaki North
Whangarei Co., 15-X1-1968, coll. K.A.J. Wise (AIM); one male, one female, Whangarei, 8-9-23, A.E. Brookes Colin. (DSlR); one
female, Whangarei, Western Hills, 13-X-56, coll. R.A. Crowson (CAS).
Subgenus Kaveinga sensu stricto Bell and Bell 1978
Type species. - Rhysodes abbreviatus Lea 1904
Description. — Antennal Segment XI without stylet; antennal Segment I pollinose above; in adults of most species pollinose
bands or spots on more distal segments; in adults of few species, pollinosity confined to Segment 1; minor setae on Segments Vl-X;
parafrontal boss either distinct or else fused to median lobe; one to three temporal setae; pronotum with grooves deep, entire, pollinose;
paramedian grooves broad, open anteriorly and posteriorly; marginal groove complete; angular seta present or absent; 0-three
marginal setae, when present, located near anterior and/or posterior angles of pronotum; prosternum, propleura impunctate; postcoxal
setae absent; elytral pollinosity and setae variably developed; abdominal sterna of female without enlarged lateral pits; anterior femur
without ventral tooth in either sex; femora in most species with dorsal and ventral pollinose strips (in a few species reduced or absent);
middle tibia with row of lateral setae, with a minute tooth between each pair of setae, the teeth in form of serrate lateral margin in
anterior or posterior aspect (Fig. 42); serration indistinctly developed in K. strigiceps and some specimens of K. abbreviata.
Members of this markedly diverse and divergent subgenus differ from those of all other subgenera in
having the lateral margin of the middle tibia serrulate. In addition, females differ from those of
Vakeinga in lacking lateral pits from the abdomen of the female, and in having fewer setae and better
developed pollinosity. Kaveinga {sensu stricto) occupies most of the range of the genus, but is absent
from New Caledonia and New Zealand. In Australia it is limited to northern Queensland.
Phytogeny. - The species Kaveinga (sensu stricto) comprise three distinct groups. The members of
Group I have deep, pollinose elytral striae and subcarinate to carinate intervals. The abdominal sterna
are transversely sulcate. This group includes K. abbreviata of Australia, K.fibulata of New Britain and
the K. pignoris complex of the Solomon Islands and Santa Cruz Islands. In this group, all species except
K. abbreviata have the pronotum elongate.
The species in Group II have the striae shallow or not impressed and without pollinosity. The
abdominal sterna each have a transverse row of punctures with an interruption at the midline. This
group includes K. parva, K. cylindrica, K. lupata, K. okapa, and K. marifuanga, all from New Guinea.
In this group, all species except K. parva have the pronotum scarcely elongate.
The species of Group III have the head very short and broad, and the margin the pronotum raised.
They show marked convergence in head structure with subgenus Ingevaka of New Zealand. The elytral
striae are deep and pollinose, and the intervals are narrow and convex to carinate. The abdominal sterna
are transversely sulcate, and the pronotum is not elongate. This group includes K. occipitalis of New
Guinea, K. histrio of Mindanao, and K. strigiceps of Buru.
An important question is whether each of these groups is an independent phyletic line or whether
either Group I or Group II is simply a collection of primitive (plesiomorphic) species. To answer this
question, it is necessary to decide the character states of the elytral striae and intervals in the common
ancestor of the subgenus. The other subgenera of genus Kaveinga have the elytral striae deep but not
pollinose, and the elytral intervals convex but not carinate. In other words, they are intermediate
between those of Groups I and II. Of all the species of Kaveinga (sensu stricto)., only K. parva of
Group II approaches the other subgenera in elytral structure.
Quaest. Ent., 1979, 15 (4)
398
Bell and Bell
One possible hypothesis would be that K. parva is primitive in the structure of its elytra, and that the
remaining species have departed from the ancestral condition in two directions. In the remainder of
Group II, the striae have become more shallow and the punctures finer, while in Groups I and III the
striae have become deepened and pollinose, and the intervals have become subcarinate to carinate.
A second possible hypothesis is that pollinose striae and subcarinate intervals characterized the
ancestral species, and that members of Group II have undergone a secondary loss of pollinosity and a
reduction of the depth of the striae. Several lines of evidence seem to us to point to the second hypothesis.
Significant are the geographical distributions of the two character states, evidence supplied by
pollinosity of the femora, and that of sculpture of the abdominal sterna. The species characterized by
shallow, non-pollinose striae and flat elytral intervals form a compact group in New Guinea, while those
characterized by carinate intervals and pollinose striae are much more widely distributed and much
more varied in structure. This suggests that Group II represents a local radiation within New Guinea.
Reduced striation is a conspicuous feature of the species of Omoglymmius {sensu stricto) and
Omoglymmius {Nitiglymmius} in New Guinea also, suggesting that it is in some way an adaptation to
local conditions.
Most species of Kaveinga (sensu stricto) have conspicuous pollinose bands on the femora. Some
species of Group II are characterized by traces of these bands, while in others, the bands are entirely
absent. This suggests that the ancestor of the subgenus had such bands, and that they are in the process
of being lost in the species of Group II, an idea consistent with the theory that the common ancestor of
the subgenus was more like Group I than Group II.
Members of subgenera other than Kaveinga (sensu stricto) have the abdominal sterna with the
punctures in broad bands. Members of Kaveinga (sensu stricto) either have the punctures of each
sternum in a uniseriate transverse row (Group II) or else have a transverse sulcus on each sternum
(Groups I, III). Transverse sulci may contain punctures, though they are in many specimens concealed
by the pollinosity. We postulate that the development of sulci resulted in alignment of the punctures in
uniseriate rows, and that a subsequent reduction in sculpture of the body in Group II resulted in the
disappearance of the sulci, leaving behind the uniseriate rows of punctures.
If it is accepted that deep, pollinose striae, subcarinate intervals, partly pilose femora, and sulcate
abdominal sterna were features of the ancestral species of Kaveinga (sensu stricto), then the possibility
exists that Group I is merely a collection of primitive (plesiomorphic) species, and is not a single phyletic
line. K.fibulata and the K. pignoris complex are small, narrow species which appear to be closely related
to one another. K. abbreviata, however, differs from the preceding species in its shorter, broader form,
and in the tendency for the abdominal sulci to be interrupted. It may really be as close to the species in
Groups II and III as it is to the remainder of Group I.
In Group II, the most isolated species is K. parva, in which the striae are distinctly impressed and the
pronotum is elongate. If deep pollinose striae and carinate intervals are regarded as primitive within the
subgenus, then this species is the least modified in the group. The four remaining species have the striae
reduced to rows of punctures, or, at most, with a few of the inner striae slightly impressed. They also
have the pronotum, at most, slightly longer than wide. They form two pairs of closely related species. K.
cylindrica and K. lupata inhabit the middle altitudes of the mountains of the extreme eastern part of
New Guinea. K. okapa and K. marifuanga form a similar pair in the east central mountains. The
distribution is markedly similar to that of the New Guinea species of Omoglymmius subgenus
Nitiglymmius. It suggests for each genus that the range of an ancestral species was split, and two
daughter species evolved, one in the east central mountains, and the other in the eastern mountains. This
was followed by a further fragmentation of range, in which each daughter species became divided into
two species. Still later, local barriers disappeared, leading to sympatry within each mountain region. In
Classification of Rhysodini
399
Group II of Kaveinga, the more specialized pair of species inhabits the east central mountains, while in
Nitiglymmius, the reverse is true, with the less specialized pair on the east central mountains.
Group III has one species in New Guinea and two species in the islands further west. K. histrio of
Mindanao and K. strigiceps of Burn are much more specialized than K. occipitalis of New Guinea, and
are obviously related to one another. K. occipitalis stands out from other New Guinea Kaveinga in
having carinate elytral intervals and a transversely sulcate abdomen. Its resemblances are to the
members of Group I, especially K. abbreviata, rather than to Group II. Its presence on New Guinea
requires confirmation, as it is known only from the holotype, collected over a century ago, though there is
no concrete reason to doubt the authenticity of the label.
KEY TO SPECIES
1
V
2
2'
3
3'
4
4'
5
5'
6
6'
1
r
8
8'
9
9'
10
Head distinctly longer than deep, at least slightly longer than wide 2
Head deeper than long, much wider than long 12
( 1) Elytral striae deeply impressed, continuously pollinose; elytral intervals markedly convex,
at least the outer ones carinate 3
Elytral striae shallowly impressed or not impressed; striae not pollinose; interval flat’or
nearly so 8
( 2 ) Pronotum relatively short, broad, length/greatest width 1.1 or less
Kaveinga abbreviata (Lea), p. 400
Pronotum elongate, length/greatest width 1.2-1. 3 4
( 3') Hind angle of pronotum rounded; parafrontal boss well developed
Kaveinga fibulata new species, p. 401
hind angle of pronotum obtuse; parafrontal boss reduced or absent 5
( 4') Basal setae of antennal Segments VI-X well developed, forming transverse row on each
segment (Fig. 28); median lobe shallowly sinuate on either side 6
Basal setae of antennal Segments VI-X reduced, each segment with setae limited to
lateral margins (Fig. 29); median lobe deeply sinuate on either side
Kaveinga nudicornis new species, p. 403
( 5 ) Strial punctures very coarse, each puncture about 0.5 as wide as interval; hind calcar of
male relatively longer and less obtuse (Fig. 31) Kaveinga ulteria new species, p. 402
Strial punctures less coarse, each puncture about 0.3 as wide as an interval; hind calcar of
male shorter, obtuse (Fig. 30) 7
( 6') Outer carina of pronotum without basal knob; pronotum shorter, its lateral margins more
curved Kaveinga pignoris new species, p. 402
Outer Carina with distinct basal knob; pronotum more elongate, its sides more nearly
straight Kaveinga kukum new species, p. 402
( T) Pronotum elongate, length/greatest width about 1.25; striae distinctly impressed
Kaveinga parva (Grouvelle), p. 403
Pronotum not elongate; length/greatest width 1.0 to 1.1; striae not impressed or only
inner striae impressed 9
( 8') Parafrontal boss separate from median lobe; lateral margins of median lobe shallowly
emarginate 10
Parafrontal boss partly fused to median lobe; lateral margins of median lobe straight 1 1
( 9 ) Humeral tubercle not exserted (Fig. 35); elytra narrow, cylindrical
Quaest. Ent., 1979, 15 (4)
400
Bell and Bell
10'
11 (90
11'
12 (10
12'
13 (120
13'
Kaveinga cylindrica (Arrow), p.
Humeral tubercle exserted (Fig. 36); elytra broader, somewhat flattened
Kaveinga lupata new species, p.
Suborbital tubercle prominent; median lobe partly or entirely separated from clypeus by
pollinose area; transverse rows of punctures on abdominal sterna III-V not interrupted...
Kaveinga okapa new species, p.
Suborbital tubercle very small; median lobe broadly continuous with clypeus; transverse
rows of punctures on abdominal sterna III-V broadly interrupted medially
Kaveinga marifuanga new species, p.
Temporal lobe transverse posteriorly; humeral tubercle exserted
Kaveinga occipitalis (Grouvelle), p.
Temporal lobe with posterior margin strongly oblique in dorsal view; humeral tubercle
not exserted
Orbital grooved extended posterior to eye; median lobe smooth; temporal lobe with few
rugae Kaveinga histrio new species, p.
Orbital groove ending opposite eye; median and temporal lobes rugose
Kaveinga strigiceps new species, p.
403
404
405
406
406
13
407
407
Kaveinga (sensu stricto) abbreviata (Lea 1904)
(Fig. 22)
Rhysodes abbreviatus Lea 1904: 79-80
Type material. - Lea cites “Cairns, Q.” as the type locality and the type depository as the Macleay
Collection. We have not been able to see the type specimen but have seen a long series from the type
locality with the following data: 11 males, 10 females, Queensland, Upper Little Mulgrave river,
S.W. Cairns, 23,27- VII and 3-VIII-69, coll. J.E. Tobler; one female, Cairns, 1952, coll. J. Sedlacek (all
CAS).
Description. — Length 5. 2-6. 7 mm. Antennal Segment I pollinose dorsally; Segments II-V each with narrow pollinose band;
basal setae reduced, well developed only on Segments VIII-X or IX-X, in some specimens with few on more proximal segments; head
slightly longer than wide; clypeus broadly separated from median lobe by band of pollinosity; parafrontal boss triangular, as wide as
long, widely separated from median lobe; sides of median lobe broadly sinuate; orbital groove as long as eye, slightly dilated; temporal
lobes slightly wider than long, oblique, anteromedial margins converging posteriorly; medial angle rounded, scarcely overlapped by
median lobe; one temporal seta; pollinosity of postorbit well developed, extended to eye ventrally; temporal lobe with slight overhang in
lateral view; suborbital tubercle and gular ridge absent.
Pronotum relatively short, broad, length/greatest width 1.10; widest near middle, sides markedly curved, convergent to narrow
apex; sides slightly curved in posterior half, side distinctly sinuate anterior to hind angle; latter obtuse, distinct emargination between
hind angle and basal knob; latter small, depressed, pollinose; paramedian grooves deep, pollinose, width at middle more than half that
of outer Carina; anterior ends of inner carinae pollinose, so the glabrous areas of these carinae appear strongly abbreviated anteriorly;
marginal grooves broad; marginal setae absent; angular seta present; prosternum with shallow transverse groove between precoxal
carinae; latter not extended to anterior margin of prosternum.
Elytra broad, slightly flattened; humeral tubercles slightly exserted; striae deep, pollinose; intervals narrow, convex, outer ones
subcarinate; striole punctures coarse, each puncture about 0.5 as wide as one interval; Stria II with one basal and one apical seta;
Stria IV with about six setae; striole without setae; several setae in apex of Stria VII; abdominal Sterna Ill-V each with coarsely
punctate transverse sulcus, in most specimens interrupted at midline; femora with pollinose bands; serrulation of middle tibia less
developed than in other members of subgenus; hind calcar of male acute.
The genitalia were figured (Part I, 57; Fig. 26).
This species, from the base of the Cape York Peninsula, is the only known representative of the
subgenus in Australia. The subcarinate elytral intervals and unspecialized head put it in Group I. The
short, rather broad pronotum separates it from other members of the group.
Classification of Rhysodini
401
Kaveinga (sensu stricto} fibulata new species
(Fig. 23)
Type material. - HOLOTYPE male, labelled: “NEW BRITAIN: Rabaul, 29-X, 1940,
J.L. Froggatt, in dead stump, C-2408, pres, by Imp. Inst. Ent., BM 1946-35” (BMNH).
Description. — Length 4.8 mm. Antennal Segment 1 pollinose dorsally; Segments II-V each with narrow dorsal pollinose band;
basal setae sparse on Segment VI, well developed on Segments Vll-X; head longer than wide; clypeus with margins pollinose with
small central glabrous area widely separated from median lobe by pollinose depression; parafrontal boss large, oval, longer than wide,
widely separated from median lobe; sides of median lobe broadly sinuate; orbital groove dilated, not extended to posterior margin of
eye; temporal lobes slightly longer than wide, their anteromedial margins oblique, convergent posteriorly; medial angle rounded, not
overlapped by median lobe; one temporal seta; postorbit entirely pollinose; temporal lobe without overhang in lateral view; suborbital
tubercle and gular ridge absent.
Pronotum rather narrow, elongate, length/greatest width 1.20; widest near middle, sides strongly curved to apex and to base;
latter only slightly wider than apex; sides not at all sinuate anterior to hind angles; latter rounded; margin not at all sinuate
posteromedial to hind angles; paramedian grooves deep, pollinose, broad, at middle nearly equal to outer carinae; anterior ends of inner
carinae pollinose, glabrous areas apparently abbreviated anteriorly; marginal groove rather narrow; marginal setae absent; angular
seta present; no transverse groove or punctures between precoxal carinae; latter not extended to anterior margin of prosternum.
Elytra narrow, cylindrical; humeral tubercles not exserted; striae deep, pollinose; intervals narrow, convex; outer intervals carinate;
strial punctures large, each about 0.33 as wide as an interval; Stria II with one basal and two subapical setae; Stria IV with five setae;
Striole without a seta; Stria Vll with several setae in apex; abdominal Sterna III-V each with transverse sulcus, latter not interrupted
medially; femora with pollinose bands; hind calcar of male small, acute.
Adults are small and narrow, belonging to Group I, but differing from other members of the group in
having the hind angles rounded. The members of the pignoris complex differ in having the hind angles
distinct and the parafrontal bosses markedly reduced or absent. K. parva, of Group II, is superficially
similar, but has shallow striae which are not continuously pollinose, and has the clypeus continuous with
the median lobe.
The pignoris complex
The subgenus Kaveinga {sensu stricto) is known from three of the Solomon Islands and one island of
the Santa Cruz group. Although these beetles are very similar to one another, they do not seem, on the
basis of very limited material, to be identical. These four forms could be treated as subspecies of a single
species. Since Rhysodini from continental landmasses rarely, if ever, have morphologically distinct
subspecies, we prefer to regard the Solomon Island’s forms as distinct species. It is nevertheless
convenient to describe the complex as a whole before describing the individual species.
Description. — Length 4. 9-6. 4 mm. Antennal Segment 1 pollinose dorsally; Segments 11-IV each with narrow pollinose band;
basal setae various in development; head longer than wide; clypeus with margins pollinose, with small central area glabrous, latter
widely separated from median lobe by pollinose depression; parafrontal boss reduced or absent; orbital groove extended to posterior
margin of eye or nearly so, dilated anteriorly, tapered posteriorly; temporal lobes as wide as long, anteromedial margin of temporal
lobe curved, margins convergent posteriorly; median angle rounded, scarcely overlapped by median lobe; one temporal seta in most
specimens (holotype of K. pignoris with three temporal setae on one temporal lobe and one on the other); postorbit entirely pollinose;
temporal lobe without overhang in lateral view; suborbital tubercle and gular ridge absent.
Pronotum elongate, narrow; hind angles distinct, obtuse; margin distinct posteromedial to hind angle; paramedian grooves deep,
pollinose, almost as wide as outer carina at middle of length; anterior end of inner carina pollinose, glabrous part of inner carina
apparently abbreviated anteriorly; marginal groove well developed; marginal setae absent; angular seta present; no pits or grooves
between precoxal carinae.
Elytra narrow, cylindrical; humeral tubercle not exserted; striae deep, pollinose; intervals narrow, convex, outer ones carinate;
strial punctures large; Stria II with setae confined to apex or to base and apex; Stria IV with continuous series of approximately six
setae; apical striole without setae; several setae in apex of Stria VII; abdominal Sterna III-VI each with complete transverse sulcus;
femora with pollinose bands.
Members of this complex are small and narrow with carinate elytral intervals. They differ from K.
fibulata in having the hind angles distinct, and separated from the base by an emargination.
Quaest. Ent., 1979, 15 (4)
402
Bell and Bell
Kaevinga (sensu stricto) pignoris new species
(Fig. 25,28,30)
Type material. - HOLOTYPE male, labelled “SOLOMON ISLANDS, Bougainville (S). Kokure,
690 m., June 13, 1956, E.J. Ford, Jr.” (BPBM).
Description. — With the characters of the pignoris complex; length 5.0 mm; basal setae sparse on antennal Segment VI, well
developed on Segments Vll-X (Fig. 28); parafrontal boss entirely absent; sides of median lobe broadly, rather shallowly emarginate;
temporal lobe evenly rounded posteriorly; pronotum moderately elongate, length/greatest width 1.20; pronotum widest near middle,
sides feebly curved except strongly curved near apex; basal knob absent; strial punctures about 0.33 as wide as an interval; Stria II
with one seta at base and two to three near apex; hind calcar small, narrow, its apex truncate, its width about 0.6 of apical width of
hind tibia.
This species differs from K. kukum in having the pronotum shorter, its sides more curved, in having
the temporal lobe evenly rounded posteriorly, and in having the hind calcar longer and more distinctly
truncate.
Kaveinga (sensu stricto) kukum new species
(Fig. 24)
Type material. - HOLOTYPE male, labelled; “SOLOMON ISLANDS, Guadalcanal, Kukum;
lljA, 1963, P. Greenslade 5109, B.M. 1966-477” (BMNH). PARATYPE female, same data as
holotype (mounted on same pin).
Description. — With the characters of the pignoris complex; length 5. 2-6. 4 mm; basal setae sparse on Antennal Segment VI,
well developed on Segments VIl-X; parafrontal boss entirely absent; sides of median lobe broadly, rather shallowly emarginate;
temporal lobe more distinctly sinuate medial to occipital angles than in K. pignoris-, pronotum more elongate than in K. pignoris
length/greatest width 1.30; pronotum with sides nearly parallel and straight except at base and apex; small detached basal knob
posterior to outer carina; strial punctures about 0.33 as wide as an interval; Stria 11 with one seta at base and one or two near apex;
hind calcar much smaller and less distinctly truncate than in K. pignoris, its width about 0.33 of apical width of tibia.
This species is close to the preceding, but differs in the form of the hind calcar, the more elongate
pronotum with parallel sides, and the presence of a basal knob on the outer carina.
The paralectotype shows an anomaly which we have not seen in any other rhysodine: the anterior half
of the median groove is absent, so that the two inner carinae fuse anteriorly.
Kaveinga (sensu stricto) ulteria new species
(Fig. 26,31)
Type material. - HOLOYPE male, labelled: “SOLOMON ISLANDS; Santa Cruz Group; Reef Is.
25/11, 1964, 11813 P. Greenslade BM 1966-477” (BMNH); PARATYPE female, same data as male,
mounted on same pin, head and prothorax missing.
Description. — With characters of pignoris complex; length 5.0 mm; basal setae on Segments Vll-X well developed;
parafrontal boss absent; sides of median lobe broadly, rather shallowly emarginate; temporal lobe markedly sinuate medial to occipital
angle; pronotum rather elongate; length/greatest width 1.28; pronotum with sides parallel except at base and apex; sides markedly
curved and narrowed to apex; pronotum with distinct basal knob posteromedial to base of outer carina; strial punctures very coarse,
about 0.5 as wide as an interval; hind calcar more elongate, more narrowly truncate than in K. pignoris (Fig. 31 ).
This species differs from both K. pignoris and K. kukum in having coarser punctures in the elytral
striae. The presence of a basal knob on the pronotum is a similarity to K. kukum, but the hind calcars of
the two species are quite different.
Classification of Rhysodini
403
Kaveinga (sensu stricto) nudicornis new species
(Fig. 27,29)
Type material. - HOLOTYPE sex unknown (hind legs missing); labelled; “SOLOMON IS. Russell
Islands; Yandina, in logs, 22-24/11/1967, P.J.M. Greenslade, 13463, log 7, B.M. 1966-477” (BMNH).
Description. — With the characters of the pignoris complex; length 4.9 mm; basal setae of Antennal Segments Vl-X markedly
reduced, represented on each segment by one or two lateral setae only (Fig. 29); parafrontal boss represented by small, oblique
glabrous area; sides of median lobe more narrowly and deeply emarginate than in related species, thus median lobe strongly
constricted at middle; hind margin of temporal lobe not sinuate medial to occipital angle; pronotum elongate, length/greatest width
1.30; pronotum widest at apical fourth, sides strongly curved from there to apex; sides straight, very slightly convergent from widest
point to hind angles; basal knob absent from base of outer carina; strial punctures coarse, each puncture about 0.33 as wide as an
interval; Stria 11 without a seta at base, with two setae near apex.
Reduction of the basal setae of the antennae and the sharp constriction of the median lobe of the head
differentiate this species from its relatives.
Kaveinga {sensu stricto} parva (Grouvelle 1895) NEW COMBINATION
(This species was accidentally omitted from Part I, 59)
(Fig. 44)
Rhysodes parvus Grouvelle 1895a: 157.
Type material. - HOLOTYPE female, labelled: “Nouv. Guinee, Bale de Geelvink, Raffray &
Maindron- 78” (MNHN). Grouvelle, in his written description, cited “Dorey” as the type locality.
Geelfinkk Bay, now called Teluk Sarera, is located on the north side of the western (Indonesian) part of
New Guinea. The data is too imprecise to determine whether this species, like its relatives, is a montane
one.
Description. — Length 4.8 mm. Antenna without pollinosity; basal setae well developed on Segments Vl-X; head longer than
wide; clypeus continuous with median lobe; parafrontal boss large, triangular, widely separated from median lobe; sides of median lobe
broadly sinuate, orbital groove fine, not extended to posterior margin of eye; temporal lobe oblique, longer than wide; anteromedial
margin oblique, nearly straight; margins converging posteriorly; medial angle rounded, well separated from median lobe; one temporal
seta; postorbit finely pollinose; temporal lobe with slight overhang in lateral view; suborbital tubercle and gular ridge absent.
Pronotum elongate, narrow; length/greatest width 1.25; widest anterior to middle; sides markedly curved to apex; sides oblique,
nearly straight, slightly convergent from widest point to hind angles; latter obtuse; margin shallowly sinuate posteromedial to hind
angle; paramedian groove deep, broad, pollinose, at middle nearly equal to outer carinae; inner carinae not at all abbreviated
anteriorly, glabrous areas extended anteriorly as far as outer carinae extended; marginal groove deep; marginal setae absent; angular
seta present.
Elytra narrow, cylindrical; humeral tubercles not exserted; striae shallow, not pollinose, intervals nearly flat; each strial puncture
about 0.25 as wide as an interval, punctures thus coarser than in other members of Group II; Stria II with one basal and one apical
seta; Stria IV with seven setae; striole without setae; several setae near tip of Stria VII.
This is a small, narrow species, superficially like K. fibulata and the K. pignoris complex, but with
the clypeus continuous with the median lobe and the elytral striae not continuously pollinose. It differs
from the other species from New Guinea in having the pronotum elongate and narrow.
Kaveinga (sensu stricto) cylindrica (Arrow)
(Fig. 32-35)
Rhysodes cylindricus Arrow, 1942: 178.
Type material. - LECTOTYPE male, labelled “PAPUA: Mt. Tafa, 8,500 ft.. Ill- 1934,
L.E. Cheesman, BM 1934-244” (BMNH). Six PARALECTOTYPES, one male, one female, same data
as lectotype; two males, two females, same data as lectotype 11-1934 (all (BMNH).
Quaest. Ent., 1979, 15 (4)
404
Bell and Bell
Description. — Length 5. 5-7.0 mm. Antennal Segment I pollinose dorsally; more distal segments not pollinose; basal setae well
developed on Segments VI-X; head slightly longer than wide; clypeus with pollinose margins and isolated central glabrous area, latter
separated from median lobe by slightly depressed pollinose area; parafrontal boss twice as long as wide, broadly separated from
median lobe; median lobe broad, sides of median lobe shallowly sinuate; orbital groove narrow, not quite as long as eye; temporal lobe
oblique, wider than long, its anteromedial margin oblique, lobes convergent posteriorly; medial angle rounded, slightly overlapped by
median lobe; one to two temporal setae; occipital angles prominent; postorbit glabrous; temporal lobe without overhang in lateral view;
gular ridge and suborbital tubercle absent; postorbital tubercle very small or indistinct.
Pronotum as long as wide or slightly longer than wide; pronotum widest near middle, sides slightly curved both anteriorly and
posteriorly, sides strongly curved medially near apex; margin sinuate anterior to rectangular hind angle, margin distinctly sinuate
posteromedial to hind angle; paramedian grooves deep, pollinose, strongly narrowed in middle, less than 0.33 as wide as outer carina at
middle; anterior ends of inner carinae pollinose, so that glabrous areas appear abbreviated anteriorly; posterior end of outer carina
depressed, pollinose; basal knob of outer carina pollinose; marginal groove rather narrow; one to three marginal setae present; angular
seta present; precoxal carinae reaching anterior margin of prosternum; a transverse groove present between precoxal carinae.
Elytra narrow, cylindrical; humeral tubercle narrow, not exserted (Fig. 35); Striae I, 11 slightly impressed; remaining striae
represented by rows of punctures; intervals flat; strial punctures small, less than 0.2 times as wide as one interval; Stria 11 with one
basal seta, without apical seta; Stria IV with four to five setae; Striole without setae; Stria Vll with several setae near apex; abdominal
Sterna 111-V each with uniseriate transverse row of punctures, latter not interrupted at midline; anterior femora with fine pollinose
bands, other femora with pollinosity reduced or absent; hind calcar of male small, rather obtuse.
This species is broader and has a shorter pronotum than does K. parva, and is narrower and more
cylindrical than are the remaining members of Group II. The most similar species is K. lupata, which
has broader elytra with the humeral tubercle strongly exserted.
Range. - K. cylindrica is confined to the most eastern mountains of New Guinea, from Mt. Tafa to
the vicinity of Wau. In addition to the type material, we have seen the following specimens: one male, 6 km
W of Wau, Nami Creek, 1700 m, lO-Vl-1962, coll. J. Sedlacek; one male, Mt. Missim, 1500-2000., 22-30-IV-1968, coll. J.L. Gressitt,
R.C.A. Rice & J. Sedlacek; one female, Mt. Kaindi, 2250 m., lO-V-1968, coll. J.L. Gressitt, J. Sedlacek (all BPBM).
Kaveinga (sensu stricto) lupata new species
(Fig. 36-39)
Type material. - HOLOTYPE male, labelled; “New Guinea: (NE) Mt. Kaindi, 16 km SW of
Wau,2300 m., 8-9-VI-1962, coll. J. Sedlacek” (BPBM). 17 PARATYPES (all BPBM); five males,
same data as holotype; one female, same locality and collector as holotype, 10-1-1962, one male, two
females, “N. Guinea (NE) Wau, 2400 m., 9-12-1, 1962, coll. J. Sedlacek, C. Monteith & native”; two
males, “N. Guinea (NE) Wau, Morobe Dist. 2400 m., 9-12-1-1962, coll. J.H. & M. Sedlacek
C. Monteith & native”; one male, “N. Guinea (NE) Wau, Morobe Dist. 1300 m., 28-1-1963, coll.
J. Sedlacek”; one female, “N. Guinea (NE) Wau, Morobe Dist., 1700-1800 m., 7-X-1962, coll.
J.&M. Sedlacek”; one male, “N. Guinea (NE) 6 km W of Wau, Nami, Creek, 1700 m., lO-VI-1962,
coll. J. Sedlacek”; one female, same locality, collector, 12-VI-1962; one female, “N. Guinea (NE) Wau,
Morobe Dist., Nami Creek, 1600-1650 m., 24-11-1963, coll. J. Sedlacek”; one male, “N. Guinea (NE),
lalibu, 2900 m., 8-14-1968, coll. Gressitt & Maa”.
Description. — Length 5.0-6. 2 mm. Antennal Segment I pollinose dorsally, more distal segments not pollinose; basal setae
sparse on Segment VI, well developed on Segments VII-X; head slightly longer than wide; clypeus with pollinose margins, and
glabrous central area; latter narrowly separated from median lobe by shallow pollinose depression in some specimens, clypeus and
median lobe narrowly joined at midline in other specimens; parafrontal boss twice as long as wide, broadly separated from median
lobe; median lobe broad, sides of median lobe shallowly sinuate; orbital groove slightly dilated, ended opposite middle of eye; temporal
lobe oblique, slightly wider than long, its anteromedial margin oblique; the two lobes convergent posteriorly; medial angle rounded,
slightly overlapped by median lobe; one temporal seta inserted in enlarged puncture; temporal lobe deeper than that of K. cylindrica,
distinctly overhanging occiput in lateral view; distinct gular ridge lateral to each gular groove; each ridge ended posteriorly in
prominent suborbital tubercle (Fig. 39); postorbital tubercle absent.
Pronotum as wide as long, widest near base, sides almost parallel except strongly convergent near anterior margin; margin not
sinuate anterior to hind angle, latter rectangular; margin scarcely emarginate posteromedial to hind angle; paramedian grooves deep,
entire, moderately wide, about 0.5 as wide as outer carina at middle; anterior ends of inner carinae pollinose, so that glabrous areas
appear abbreviated anteriorly; posterior end of outer carina depressed, pollinose; basal knob of outer carina pollinose; marginal groove
narrow; marginal setae absent from most specimens, a few specimens with one marginal seta anterior to the angular seta; precoxal
Classification of Rhysodini
405
carinae not extended to anterior margin of prosternum; transverse row of two pits between precoxal carinae.
Elytra somewhat flattened, broader than in K. cylindrica-, humeral tubercle prominently exserted (Fig. 36); elytral striae not
impressed, represented only by rows of punctures; intervals flat; strial punctures small, less than 0.2 of width of one interval;
Stria II with one basal seta, and in most specimens with one apical seta; in some specimens apical seta absent from Stria II, but
apical seta in Stria 1; Stria IV with four to five setae; Stride without a seta; Stria Vll with five to ten setae near apex; abdominal
sterna 111-V each with uniseriate transverse row of punctures, the row not interrupted in midline; anterior femora with fine
pollinose bands; remaining femora without pollinosity; hind calcar of male very small.
This species is sympatric with K. cylindrica, but is readily separated by the exserted humeral
tubercles, the wider paramedian and marginal grooves, and the well-developed gular ridges and
suborbital tubercles.
With this species we provisionally place one specimen from Mt. Missim, New Guinea, 1650 m
(BPBM). It matches the description of K. lupata in most respects, but differs in having the suborbital
tubercles almost absent; the parafrontal bosses large, quadrate, and the total length 6.8 mm. Since we
have doubts about the inclusion of this species, we have not designated it as a paratype.
Kaveinga (sensu stricto) okapa new species
(Fig. 40)
Type material. - HOLOTYPE male, labelled: “New Guinea; Okapa, Kamira, Eastern Highlands,
2-9-1964, coll. R. Hornabrook (NMNZ). Seven PARATYPES (NEW GUINEA) one female, same
data as holotype; one male, Okapa, 12-11-1964; one female, Okapa, Okosa, 12-1-1965; one female,
Daulo Pass, Asato-Chimbu Divide, 8-4-1972 and one male, 16-9-72; one female, 18-10-72 and one male,
Feb., 1971, Lufa, Mt. Michael. All specimens collected by R. Hornabrook and deposited at NMNZ.
Description. — Length 5. 0-6. 3 mm. Antennal Segment I pollinose dorsally, more distal segments not pollinose; basal setae
sparse on Segment VI, well developed on Segments Vll-X; head slightly longer than wide; clypeus with pollinose margins, glabrous
central area; latter narrowly separated from median lobe by shallow pollinose depression in some specimens, clypeus and median lobe
narrowly joined at midline in other specimens; parafrontal boss connected narrowly to median lobe; median lobe broad, its lateral
margins nearly parallel; orbital groove slightly dilated, short, ended well anterior to posterior margin of eye; temporal lobe oblique,
slightly longer than wide, its anteromedial margin oblique; lobes convergent posteriorly; median angle rounded, slightly overlapped by
median lobe; one temporal seta, inserted in prominent puncture; occipital angle prominent, somewhat lobate; postorbit extensively
glabrous; pollinosity not extended to eye; temporal lobe with slight overhang in lateral view; gular ridge prominent, partly to entirely
pollinose; posterior end of gular ridge in form of prominent suborbital tubercle.
Pronotum varied in size and proportions, in most specimens as wide as long, greatest width at base; in few specimens slightly
longer than wide, with widest point near middle, and base slightly narrowed; lateral margin not sinuate anterior to hind angles, latter
rectangular; paramedian grooves deep, markedly narrowed at middle, their outer margins straight, elevated, sharply defined; their
inner margins ill-defined, sloped gradually from inner carinae; pollinosity of paramedian groove reduced to narrow strip along outer
margin; anterior end of inner carina pollinose, inner carina apparently abbreviated anteriorly; basal knob pollinose, depressed below
level of outer carina; marginal groove fine; marginal setae absent; angular seta present; precoxal carinae ending close to anterior
margin of prosternum; a transverse row of two pits between precoxal carinae.
Elytra broad, slightly flattened; humeral tubercle slightly exserted; elytral striae not at all impressed, represented only by rows of
fine punctures; intervals flat; strial punctures small, less than 0.2 as wide as an interval; Stria II with basal seta and apical seta or one
or both of these absent; Stria IV with one or two setae near base and one at apex, but without setae in middle third; Striole asetose;
Stria VII with seven to ten setae near apex; abdominal Sterna IIl-V each with uninterrupted transverse row of punctures; femora with
pollinose bands absent or represented by small vestiges; hind calcar of male very short, obtuse.
This species and K. marifuanga differ from all other members of the subgenus in having the
parafrontal bosses joined to the median lobe. This species is distinguished by having large suborbital
tubercles, parafrontal bosses only narrowly joined to the median lobe, and the latter, at most narrowly in
contact with the clypeus. In most specimens of K. okapa, the pronotum is much less elongate than in K.
marifuanga, and is not at all narrowed at the base. In a few specimens, which we provisionally interpret
as variants of K. Okapa, the pronotum is slightly elongate and slightly narrowed at the base,
approaching that of K. marifuanga. Possibly we are including more than one species in our concept of K.
okapa. The point cannot be settled until many more specimens are available.
Quaest. Ent., 1979,15 (4)
406
Bell and Bell
Kaveinga (sensu stricto) marifuanga new species
(Fig. 41, 42)
Type material. - HOLOTYPE male, labelled: “NEW GUINEA, Marifuanga, Asaro-Chimbu
Divide, 1-6-72, coll. R. Hornabrook” (NMNZ). Three PARATYPES: one male, same data as holotype;
two females, Daulo Pass, Asato-Chimbu Divide, 8-4-72 and 4-1-75, coll. R. Hornabrook (all NMNZ).
Description. — Length 5. 4-7.0 mm. Antennal Segment 1 pollinose dorsally, more distal segments not pollinose; basal setae
sparse on Segment VI, well developed on Segments VII-X; head slightly longer than wide, clypeus with pollinose anterior and lateral
margins, broadly joined to median lobe posteriorly; parafrontal boss broadly fused to median lobe; median lobe broad, its lateral
margins nearly parallel; orbital groove slightly dilated, short, ending anterior to middle of eye; temporal lobe oblique, slightly longer
than wide, its anteromedial margin oblique, straight; temporal lobes convergent posteriorly; medial angle rounded, slightly overlapped
by median lobe; one temporal seta, inserted in prominent puncture; occipital angle prominent, in form of small lobe; postorbit glabrous;
temporal lobe not overhanging occiput in lateral view; gular ridge low, glabrous, its posterior end in form of small suborbital tubercle
(suborbital tubercles smaller and closer together than in K. okapa). Pronotum distinctly longer than wide, length/greatest width
averages 1.16; widest near middle, sides curved to apex, nearly straight, slightly convergent posteriorly; margin distinctly sinuate
anterior to hind angles; latter rectangular; paramedian grooves shallower than in K. okapa, markedly narrowed at middle, their outer
margins straight, elevated, sharply defined; their inner margins ill-defined, sloped gradually from inner carinae; pollinosity of
paramedian groove reduced to narrow strip along outer margin; anterior end of inner carina not pollinose nor apparently abbreviated;
basal knob of outer carina glabrous, appearing as continuation of outer carina; marginal groove fine; marginal setae absent from most
specimens, but one present unilaterally near anterior angle on one specimen; angular seta present; prosternum with two pits in
transverse row between precoxal carinae.
Elytra relatively narrow, cylindrical; humeral tubercles not exserted: elytral striae not impressed, represented by rows of
punctures; intervals flat; strial punctures very small, less than 0.15 as wide as an interval; Stria 1 in some specimens with seta near
apex, in other specimens this seta absent; Stria II without setae; Stria IV with one seta near base and two near apex; Striole without
setae; Stria VII with approximately nine setae near apex; Sternites IIl-V each with transverse row of coarse punctures, latter broadly
interrupted in midline; femora with pollinose bands absent; hind calcar obtuse, but more prominent than in K. okapa.
This species is similiar to K. okapa, but differs in having the suborbital tubercles much smaller, the
clypeus broadly joined to the median lobe; the parafrontal bosses more broadly joined to the median
lobe, and the transverse rows of punctures of the abdomen broadly interrupted in the midline. The
pronotum is very different in shape from most specimens of K. okapa, but a few specimens which we
interpret as variants of the latter species approach the shape seen in K. marifuanga.
Kaveinga (sensu stricto) occipitalis (Grouvelle 1903)
(Fig. 43)
Rhysodes occipitalis GvouwqWq 1903: 105-106.
Type material. - HOLOTYPE female, labelled; “NUOVA GUINEA, Fly River, L.M. D’Albertis
1876-1877” (MNHN).
Description. — Length 7.0 mm. Basal setae sparse on Segment VI, well developed on Segments Vll-X; head distinctly wider
than long; clypeus with glabrous central area, narrow pollinose anterior margin and widely pollinose lateral margins, separated from
median lobe by deep pollinose transverse impression; clypeus not constricted posteriorly; median lobe connected to each antennal rim
by oblique, pollinose carina, but true parafrontal bosses absent; sides of median lobe shallowly sinuate; orbital groove very broadly
dilated, extended beyond posterior margin of eye; temporal lobe transverse, its anterior margin rounded, its medial angle closely fitted
to and overlapped by median lobe; median and temporal lobes not rugose; posterior margin of temporal lobe transverse, its most
posterior point midway between midline and lateral margin of head; one temporal seta, inserted in prominent puncture; postorbit
pollinose; temporal lobe without overhang in lateral view; gular ridge and suborbital tubercle absent.
Pronotum as wide as long, widest at base; sides nearly straight, slightly convergent almost to apex, where they are strongly curved
medially, lateral margin not sinuate anterior to hind angle; latter rectangular; anterior margin of pronotum deeply emarginate
medially; paramedian grooves deep, rather markedly narrowed at middle, width at middle about 0.5 width of outer carina; anterior
end of inner carina pollinose, latter much shorter than outer one, appearing oval; posterior end of outer carina not depressed; basal
knob absent; marginal groove deep, holotype with one marginal seta on left side, just anterior to angular seta, but without marginal
seta on right side; angular seta present; precoxal carina not extended to anterior margin of prosternum; no pits or transverse grooves
between precoxal carinae.
Elytra broad, somewhat flattened; humeral tubercle exserted; striae deep, pollinose, intervals carinate; Intervals II, III more
elevated than others at base, slighty prominent, in form of slight anterior prominence; strial punctures oval, longer than wide, about
Classification of Rhysodini
407
0.33 as wide as interval; Stria II with one basal and one apical seta; Stria IV with one basal and two apical setae; apical striole
without setae; Stria VII with about ten setae near apex; abdominal Sterna III-V each with complete transverse sulcus; Sternum VI
with complete basal transverse sulcus and group of coarse punctures near apex; femora with pollinose bands.
This is the least specialized member of Group III, the “short-faced” species. It differs from the other
members of the group in having the temporal lobes transverse posteriorly. The broad head and the deep
pollinose elytral striae will separate it from all other known Kaveinga species from New Guinea, all of
which belong to Group II.
Kaveinga (sensu stricto) histrio new species
(Fig. 45-48)
Type material. - HOLOTYPE male, labelled; “Mindanao, E. Slope Mt. McKinley, 3300 ft., Davao
Prov., X-1-46, H. Hoogstraal (CNHM Philippines Zoo. Exped. 1946-57)” (CNHM). Two
PARATYPES, one female, same data as type; one female, same locality as type, IX-28-1946,
F.G. Werner (both CNHM).
Description. — Length 5. 5-7.0 mm. Antennal Segment 1 pollinose dorsally; Segments II-V each with narrow pollinose band;
basal setae well developed on Segments VI-X; head twice as wide as long, much deeper than long; clypeus entirely pollinose,
constricted at base, its sides angulate, its anterior and lateral margins raised; clypeus separated from median lobe by deep pollinose
impression; anterior tentorial pits enlarged, close together; parafrontal boss absent; median lobe with lateral margins straight; median
lobe sloped abruptly from postclypeal depression, markedly convex in lateral view (Fig. 46); posterior end of median lobe curved
ventrally over occiput; orbital groove markedly dilated; extended beyond posterior margin of eye; two temporal setae, each inserted in
prominent pit, one or both of latter confluent laterally with orbital groove in some specimens; temporal lobe much wider than long,
curved, its medial margin closely fitted to median lobe but not countersunk in it or extended above it; median lobe smooth, temporal
lobe with few longitudinal rugae near border with median lobe; temporal lobe markedly divergent posteriorly; posterior margin of
temporal lobe oblique, sinuate, most posterior point much closer to lateral margin than to midline; postorbit pollinose; temporal lobe
without overhang in lateral view; gular ridge, suborbital tubercle absent.
Pronotum slightly longer than wide; length/greatest width 1.1; widest slightly anterior to middle; sides curved; base and apex both
narrowed; lateral margin shallowly sinuate anterior to hind angle, latter nearly rectangular; anterior margin of pronotum shallowly
emarginate medially; paramedian grooves wide, deep, as wide at middle as are outer carinae; both inner and outer carinae pollinose
and apparently abbreviated anteriorly; inner carinae more abbreviated than outer one; basal knob of outer carina glabrous, prominent,
separated from rest of carina by pollinose depression; marginal groove broad, deep; one marginal seta near anterior angle (unilaterally
absent in one specimen); one to two marginal setae near angular seta; angular seta present; precoxal carinae prominent, but not
extended to anterior margin of prosternum; transverse groove between precoxal carinae; anterior part of prosternum and precoxal
carinae pollinose; anterior part of pleuron pollinose, pollinosity extending posteriorly along notopleural suture.
Elytra narrow, cylindrical; humeral tubercle pilose at base, not exserted; striae deep, very narrowly pollinose; intervals convex but
not carinate; strial punctures very large, about 0.33 as wide as an interval; Stria II with one seta at base and two near apex; Stria IV
with four to six setae; striole with one seta; Stria VII with one seta at humerus and six near apex; Sternum II of abdomen with two
transverse rows of punctures; Sterna III-V each with complete transverse sulcus; Sternum VI with basal sulcus which is interrupted in
midline and with group of coarse punctures near apex; femora with pollinose band; hind calcar of male small, obtuse.
In this species and the next one, the head is an extraordinary, masklike structure, rising high above
the pronotum, and largely hollowed posteriorly, exposing the slender pillar which supports the median
lobe (Fig. 47). In dorsal view, the diverging temporal lobes suggest the head of the hammer-head shark
{Sphyrna). The long orbital grooves, the smooth median lobe, and the abbreviated pronotal carinae
provide convenient separation of this species from K. strigiceps.
Kaveinga (sensu stricto} strigiceps new species
(Fig. 49-51)
Type material. - HOLOTYPE male, labelled; “Burn, Station 12, 4-7 Febr. 1922, L.J. Toxopeus”
(AMS).
Description. — Length 5.4 mm. Antennal Segment I pollinose dorsally; Segments II-V each with narrow pollinose band; basal
setae well developed on Segments Vl-X; head almost twice as wide as long, twice as deep as long; clypeus glabrous in middle, its
anterior and lateral margins pollinose; clypeus constricted at base, its anterior margin curved; margins of clypeus reflexed; clypeus
Quaest. Ent., 1979, 15 (4)
408
Bell and Bell
separated from median lobe by deep pollinose depression; anterior tentorial pits greatly enlarged, close together; parafrontal boss
absent; median lobe with lateral margins straight; median lobe sloped almost vertically from postclypeal depression, markedly
convex in lateral view; posterior end of median lobe curved ventrally over occiput (Fig. 50); anterior half of median lobe
longitudinally rugose, posterior half with few scattered rugae, but otherwise smooth; orbital groove dilated, not extended posterior
to eye; one temporal seta far from orbital groove; temporal lobes oblique, divergent posteriorly, posterior margin of temporal lobe
sinuate; temporal lobe densely, longitudinally rugose (Fig. 51); temporal lobes very convex, sloped above and partly overlapping
median lobe; postorbit entirely pollinose, gular ridge, suborbital tubercle absent.
Pronotum as wide as long, nearly quadrate; margins straight and parallel except near apex, where they curve medially; lateral
margin not sinuate anterior to hind angle, latter rectangular; anterior margin of pronotum shallowly emarginate medially;
paramedian grooves deep, narrower than in K. histrio, at middle equal in width to outer carina; neither outer nor inner carina
abbreviated or pollinose anteriorly; basal knob of outer carina small, depressed, pollinose; marginal groove deep, broad; marginal
setae absent; angular seta present; inner carina with basal seta; precoxal carinae not extended to anterior margin of prosternum; no
transverse groove or pits between precoxal carinae; anterior part of prosternum, precoxal carinae, and propleura pollinose;
pollinosity of pleuron extended posteriorly along notopleural suture.
Elytra narrow, cylindrical; humeral tubercle not exserted; striae deep, narrowly pollinose; intervals convex, narrow, but not
carinate; strial punctures very small, less than 0.2 of width of one interval; Stria II without setae; Stria IV with one apical seta;
Striole without seta; Stria VII with few setae near apex; Sternum II of abdomen with two transverse rows of punctures;
Sterna III-V each with complete transverse sulcus; Sternum VI with transverse basal sulcus interrupted at midline, and group of
coarse punctures near apex; femora with pollinose bands; lateral margin of middle tibia scarcely serrulate; hind calcar of male
small, acute.
The high, domed head with conspicuous rugae on both median and temporal lobes easily separate this
species from all others.
SUBTRIBE CLINIDIINA
Description. - Part I, 59.
Key to Genera. - Part I, 59
Genus Grouvellina Bell and Bell, 1978
(Fig. 52-74)
Type species. - Rhysodes tubericeps Fairmaire 1868.
Description. — Antennal stylet well developed; antennal segments with both basal and apical series of setae well developed;
minor setae of antennal segments forming a ventral tuft on each segment, in most species beginning on Segment V; dorsal side of
antennal segments with bands or spots of pollinosity.
Eye large, deeper than long, with about 150 ommatidia; antennal lobe separated from temporal lobe by deep postantennal groove;
most species with elevated glabrous parafrontal boss lateral to median lobe and anterior to postantennal groove; head extensively
pilose, in most species with well-developed orbital band of pilosity in place of orbital groove; longer pilosity in frontal pit; pilosity of
occipital region in form of “rufF’ anterior to glabrous neck condyle; labrum with one or two pairs of setae; clypeus with one pair; two to
six temporal setae; postlabials one to many pairs.
Pronotum with complete paramedian grooves; marginal groove single, complete in most species (but abbreviated posteriorly in G.
grouvellei)-, prothorax with “collar” of long pilosity around the anterior margin; angular seta present; 0 to six marginal setae.
Elytra! striation complete; Stria VII marginal; apical striole absent; pilosity well developed in elytral striae (in some species also
invading intervals); hind wings well developed in those species checked. Abdominal sterna transversely sulcate.
Anterior tibia with cleaning organ entirely or largely distad to base of tarsus; middle of cleaning organ with short, stiff spinose
setae, in form of “comb”, in most species sharply contrasted with longer, flexible setae at either end of the row, in form of pair of
“brushes”; in most species both proximal and distal spurs distinct, opposite the “brushes” (in a few species, proximal spur absent);
distal spur apparently secondary point on preapical tooth; proximal spur inserted just distad of intermediate tooth.
Apex of middle tibia alike in both sexes, with one spur and a medial process (in some species also with a lateral process); male
without middle calcar; hind tibia in female with two spurs; in male with one spur and a calcar; all legs with a complex pattern of pilose
and glabrous areas.
The genus is unique among Clinidiina in possessing large eyes, functional hind wings, and complete
elytral striation. Grouvellina is known only from Madagascar, but Rhysodes planifrons Fairmaire 1893,
described from Mayotte, in the Comoro Islands, will probably prove to belong to the genus. We have
Classification of Rhysodini
409
been unable to locate the type of this species or that of Rhysodes canaliculatus Castelnau 1836. The
description of the latter species is consistent with membership in Grouvellina, but is not detailed enough
to indicate to which, if any, of the species described below it applies. The size range is consistent with it
being G. hova, G. cuneata, or G. gigas.
Phytogeny. - The radiation of this remarkable genus must have taken place entirely within
Madagascar. Deciphering the phylogeny has proven difficult, as many of the characters occur in
bewildering combinations. It is not made easier by the fact that six of the species are known from one sex
only, that four species are not known from any specific locality, and that six of the remaining species are
recorded from one locality each. The above figures suggest that we know only a fraction of the actual
species.
We present a hypothetical phylogenetic diagram of the genus (Diagram 1). Species 1, the
hypothetical common ancestor of the genus, is presumed to have had the following characteristics:
antenna with tufts of minor setae on Segments V-X; antennal Segments II-X each with complete dorsal
apical and basal bands of pollinosity; head slightly longer than broad; labrum with two pairs of setae;
median lobe glabrous; parafrontal boss present; orbital groove represented by a complete band of
pollinosity medial to the eye; all pronotal carinae extended the full length of the pronotum, and broadly
glabrous; all pronotal grooves complete and deep; pronotum with several marginal setae; precoxal carina
well developed; elytral intervals equal, subcarinate; Stria I with several setae near the apex*; Striae II
and IV with complete series of setae; metasternum coarsely punctate on disc and margins; anterior
femur with a ventral tooth in both sexes; anterior tibia of male with proximal tooth (absent in female).
Of these characters, perhaps the most important are the number of setae on the labrum, the presence
of the ventral femoral and proximal tibial teeth, and the precoxal carina. Four is the number of labral
setae in the more primitive genera of Rhysodini, and it is closer to the number characteristics of most
other Carabidae (six), therefore it is reasonable to conclude that having only two setae is a derived
characteristic. Femoral and proximal tibial teeth are present in a number of otherwise very dissimilar
Grouvellina; therefore it seems likely that they were present in the common ancestor, and have been lost
independently several times. The same line of reasoning leads us to hypothesize the presence of the
precoxal carina in Species I. Grouvellina tubericeps is the one member of this genus which has all the
hypothetical ancestral characteristics.
With the exception of G. tubericeps, all Grouvellina can be assigned to two major lines, descended
respectively from Species 2 and 3. In Species 2, the basal bands of pollinosity were lost from antennal
Segments III-X; while in Species 3, the lateral setae of the labrum were lost.
Species 2 was the ancestor of two lines, represented by Species 4 and 5. Species 4 lost the precoxal
carina but was otherwise not modified. Species 5 retained the precoxal carina, but lost the femoral and
proximal tibial teeth, and the basal pollinose band on Segment II of the antennae, and the anterior setae
of Stria II.
Species 5 was the ancestor of G. edentata and G. cooperi. G. edentata lost the anterior setae from
elytral Stria II, but otherwise remain unmodified. G. cooperi retained one anterior setae in Stria II, but
had the posterior setae of that stria reduced to one, and lost the parafrontal boss and the marginal setae
of the pronotum.
Species 4 gave rise to Species 6 and 7. In Species 6, the metasternal punctures were reduced in size
and the anterior setae of Stria II were lost. In Species 7, the metasternal punctures remained large and
the anterior setae in Stria II remained, but the head became short and broad, the body increased in size,
and the intervals became more distinctly carinate.
Species 6 is the ancestor of G. montana and G. ranavalona. The former species lost the anterior part
of the orbital pilose band, while the metasternal (discal) punctures disappeared entirely, the marginal
Quaest. Ent., 1979, 15 (4)
410
Bell and Bell
Phylogenetic Diagram I. Reconstructed Phylogeny of species of Grouvellina.
Classification of Rhysodini
411
setae of the pronotum were reduced in number, the body size diminished, and the elytra intervals
became flattened. The femoral and proximal tibial teeth were retained. In G. ranavalona the femoral
tooth was lost, while the size remained large and the metasternal punctures remained distinct, though
fine. The male of this species is unknown, but it seems likely that it will be shown to lack the proximal
tibial tooth. (All of the known males of Grouvellina which do not have a femoral tooth lack the proximal
tibial tooth as well.)
Species 7 was the ancestor of Species 8 and of G. cuneata. In Species 8, Stria I lost its setae, while in
G. cuneata these setae were retained, but the anterior setae of Stria II were lost, the pronotum became
strongly narrowed anteriorly, and the striae and intervals became strongly unequal.
The two descendants of Species 8 are G. gigas and G. hova. The former species remains essentially
like Species 8, while the latter species lost the parafrontal boss.
Species 3 (the common ancestor to the species with two labral setae) was ancestral to Species 9 and
Species 10. In Species 9, the basal pollinose bands of the outer antennal segments became broken into
isolated pilose spots, and the femoral and proximal tibial teeth were lost. Species 10 retained the
pollinose bands intact, and retained the femoral and proximal tibial teeth, but became highly specialized
in many features, including the abbreviation of the outer pronotal carina posteriorly, the development of
pollinosity on the median lobe, the loss of all but the most anterior of the pronotal marginal setae, and
the loss of the parafrontal boss. In addition, the head became shortened and conspicuous tufts of
pollinosity developed around each temporal seta.
Species 9 gave rise to G. descarpentriesi and to Species 1 1 . The former species lost the discal
punctures of the metasternum, the precoxal carina, the setae of Stria I and the setae of the anterior half
of Stria II. The apical pollinose rings of the outer remained intact. Species 1 1 had the apical band
broken into isolated pilose spots, but it retained the metasternal punctures, the precoxal carinae and the
setae of Stria I and the anterior half of Stria II.
Species 1 1 gave rise to G. radama and Species 1 2 G. radama remained close to Species 1 1 in
structure, but the median lobe became very narrow and the postlabial setae underwent multiplication.
Species 12 lost the tuft of minor setae and the basal setae from antennal Segment V, while the marginal
carina became abbreviated posteriorly, the precoxal carina was lost and the metasternal punctures
became fine.
Species 12 gave rise to G. divergens and G. grouvellei. The former species remains close to Species 12
in structure, but has lost the marginal setae of the pronotum and the anterior setae from Stria II. G.
grouvellei retains these setae, while it has become highly specialized in having the pollinosity greatly
increased so that it covers almost the entire dorsal surface, while the pronotal carinae and elytral
intervals have become much less convex.
Species 10 gave rise to G. dentipes and G. cinerea. G. dentipes retained the femoral and proximal
tibial teeth, while G. cinerea lost them.
The above phylogeny is by necessity somewhat speculative, and doubtless will require some
modification when more is known about Grouvellina. Perhaps the most questionable points are the
existence of Species 6 (a common ancestor for G. montana and G. ranavalona), and the question of
whether G. radama is really more closely related to Species 1 2 than is G. descarpentriesi.
KEY TO SPECIES
1 Outer carina of pronotum complete, glabrous, extended from anterior margin to vicinity
of hind angle; marginal carina glabrous, ended distinctly anterior to hind angle 2
Quaest. Ent., 1979, 15 (4)
412
Bell and Bell
V
2
2'
3
3'
4
4'
5
5'
6
6'
1
r
8
8'
9
9'
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ir
12
12'
Outer Carina not both complete and glabrous, either abbreviated clearly anterior to hind
angle or else entirely pollinose and coalescent with marginal carina posteriorly
( 1 ) Femur of anterior leg with ventral tooth in both sexes; male with proximal tooth on
anterior tibia (Fig. 74)
Femur of anterior leg without ventral tooth; male (where known) without proximal tooth
on anterior tibia
( 2 ) Orbital groove complete, pilose, eye and temporal lobe separated; elytral intervals more
or less carinate
Orbital groove incomplete, eye in contact with glabrous area of temporal lobe; elytral
intervals not carinate Grouvellina montana new species, p.
( 3 ) Head with large distinct parafrontal boss
Parafrontal boss absent, entire superantennal area pollinose or pilose
Grouvellina hova new species, p.
( 4 ) Lateral margins of pronotum markedly convergent anteriorly, width at middle less than
width at base; inner striae narrower than outer ones
Grouvellina cuneata new species, p.
Lateral margins of pronotum not markedly convergent, width at middle equal to or
greater than width at base; inner and outer striae of equal width
( 5') Prosternum with short precoxal carina; body length 6.0-8.0 mm; postmentum glabrous
(Fig. 60); dorsal basal pollinose bands on antennal Segments II-VIII (Fig. 59); median
lobe long, narrow Grouvellina tubericeps (Fairmaire), p.
Precoxal carina absent; body length 8.0-10.5 mm; postmentum pollinose (Fig. 63); dorsal
basal pollinose bands only on antennal Segment II (Fig. 62); median lobe short, broad,
broadly rounded posteriorly Grouvellina gigas new species, p.
( 2') Labrum with four setae
Labrum with two setae
( 7 ) Precoxal carina absent; elytral intervals carinate; base of elytral interval II not elevated;
length 9.0 mm or more Grouvellina ranavalona new species, p.
Precoxal carina present; elytral intervals not carinate; base of elytral interval II elevated;
length 5.0-7.0 mm
( 8') Parafrontal boss present Grouvellina edentata new species, p.
Parafrontal boss absent Grouvellina cooperi new species, p.
( 7') Median lobe of head long, narrow, parallel-sided; carinae of pronotum narrow, equal to
or slightly wider than pronotal grooves; precoxal carina present
Grouvellina radama new species, p.
Median lobe of head wider; carinae of pronotum wider than pronotal grooves; precoxal
carina absent
(10') Medial angle of temporal lobe rounded; antennal stylet elongate, acute; discal
metasternal punctures absent; pronotum with 4-5 marginal
setae Grouvellina descarpentriesi new species, p.
Medial angle of temporal lobe more pointed; antennal stylet obtuse; discal metasternal
punctures present; marginal setae absent Grouvellina divergens new species, p.
( 1') Outer carina of pronotum entirely glabrous, abbreviated posteriorly; marginal groove of
pronotum complete; each temporal seta concealed within dense tuft of pilosity, temporal
lobe otherwise glabrous
Outer carina completely pollinose or with small glabrous areas; outer carina not
12
3
7
4
419
5
417
416
6
413
413
8
10
417
9
418
418
419
11
420
421
13
Classification of Rhysodini
413
abbreviated posteriorly, but coalescent with marginal carina, marginal groove
incomplete; temporal lobe largely pilose, no tufts around setae
Grouvellina grouvellei (Fairmaire), p. 421
13 (12 ) Medial angle of temporal lobe more distinct; anterior femur with ventral tooth; male with
proximal tooth on anterior tibia Grouvellina dentipes new species, p. 422
13' Medial angle of temporal lobe more rounded; anterior femur without ventral tooth; male
without proximal tooth on anterior tibia Grouvellina cinerea new species, p. 423
Grouvellina tubericeps (Fairmare, 1868)
(Fig. 58-60)
Rhysodes tubericeps Fairmaire, 1868: 782.
Type material. - HOLOTYPE male, labelled; “Madagascar” (MNHN); labelled as Fairmaire type.
According to the original description, the specimen was collected by Charles Coquerel. The holotype is
in poor condition with hind legs missing and perhaps some setae broken off.
Description. — Length6.0-8.0 mm. Antennal Segment XI as long as wide; apical stylet short, conical; tufts of minor setae on
Segments V-X; antennal Segment I extensively pollinose on dorsal aspect; Segments II-IX each with two transverse pollinose bands;
Segment X with basal band only (Fig. 59). Head slightly elongate; frontal and postantennal grooves deep, relatively wide; median lobe
relatively narrow, obtusely pointed posteriorly; parafrontal bosses well developed; temporal lobe slightly longer than wide; sinuate
anterior to medial angles, latter narrowly separated, obtusely pointed; two to three temporal setae; four labral setae; orbital groove
complete, broadly pilose; three to four pairs of postlabial setae; mentum pollinose, postmentum contrastingly glabrous (Fig. 60).
Pronotum elongate, narrow, length/greatest width about 1.35; lateral margins nearly parallel, width at middle equal to that at
base, apex only slightly narrowed; lateral margin not sinuate anterior to hind angle; outer carina narrowed and more or less bent
outward at base; pronotum with about six marginal setae in most specimens (but only two in holotype); prosternum with
well-developed precoxal carina extended at least halfway to anterior margin; carina bounded on either side by longitudinal
impressions.
Elytral striae deep, coarsely punctate; intervals narrow, carinate; humerus prominent, quadrangular, with conspicuous patch of
golden pilosity; Stria I with two setae near apex; Stria 11 with five setae; metasternum with many discal punctures coarser than
marginal ones; margins of metasternum pilose.
Anterior femur with ventral tooth in both sexes; anterior tibia slender, with proximal tooth present in male, absent from female;
cleaning organ with proximal spur small but distinct; as small triangular tooth on distal side of intermediate tooth; distal spur
flattened; comb teeth short, slightly overlapping brushes. Male with hind calcar tapered, its tip truncate.
This species is similar to G. gigas, from which it differs in its smaller size, somewhat narrower body,
and especially in having a well-developed precoxal carina.
Range. - The holotype, which is in poor condition, and without exact locality data, appears to us to be
conspecific with a series of specimens from the northern tip of Madagascar, representing the following
localities: Cap d’Ambre, coll. F. Schneider, one male (MNHN); Diego Suarez, collector not indicated,
two females (MNHN); Mont Ambre, coll. Sicard, 1930, two males one female (MNHN).
Grouvellina gigas new species
(Fig. 61-63)
Type material. - HOLOTYPE male, labelled: “MADAGASCAR, Chutes de la Mort, XI- 10- 1959,
coll. E.S. Ross” (CAS). Thirteen PARATYPES (all from Madagascar) one female same data as type
(CAS); one male, 1883, coll. Humblot, ex coll. Oberthur 1904-175 (BMNH); four males, six females,
Ambodivoangy, 1959-1961, coll. J. Vadon (MRAC); one male, Madagascar Sud, 1901, coll. Alluaud
(MNHN).
Description. — Length 8.0-10.5 mm, females averaging larger than males. Antennal Segment XI distinctly longer than wide;
stylet prominent, conical, acute; tufts of minor hairs on Segments V-X; antennal Segment 1 with broad dorsal band of pollinosity;
Segment II with two dorsal bands; Segments Ill-X with narrow apical pollinose bands, but without basal bands (Fig. 62).
Quaest. Ent., 1979, 15 (4)
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Bell and Bell
Figures 52 - 63. Fig. 52-74, Genus Grouvellina\ Fig. 52-58, Head and pronotum, dorsal aspect; Fig. 52, G. montana new
species; Fig. 53, G. hova new species; Fig. 54, G. cuneata new species; Fig. 55, G. radama new species; Fig. 56, G. divergens
new species; Fig. 57, G. ranavalona new species; Fig. 58-60, G. tubericeps (Fairmaire); Fig. 58, Head and pronotum, dorsal
aspect; Fig. 59, Antennal Segments I-Vl; Fig. 60, Head, ventral aspect; Fig. 61-63, G. gigas, new species; Fig. 61, Head and
pronotum, dorsal aspect; Fig. 62, Antennal Segments I-Vl; Fig. 63, Head, ventral aspect.
Classification of Rhysodini
415
Figures 64 - 74. Fig. 64, Flead and pronotum, dorsal aspect, G. edentata, new species; Fig. 65-67, G. descarpentriesi, new
species; Fig. 65, Head and pronotum, dorsal aspect; Fig. 66, Head, dorsal aspect; Fig. 67, Metasternum, metacoxae; Fig. 68-70,
G. cooperi new species; Fig. 68, Head and pronotum, dorsal aspect; Fig. 69, Prothorax, ventrolateral aspect; Fig. 70,
Metasternum, metacoxae; Fig. 71-73, Head and pronotum, dorsal aspect; Fig. 71, G. grouvellei (Fairmaire); Fig. 72, G. cinerea,
new species; Fig. 73, G. dentipes, new species; Fig. 74. Femur, tibia, male, G. dentipes, new species.
Quaest. Ent., 1979, 15 (4)
416
Bell and Bell
Head distinctly wider than long; labrum with four setae; frontal and postantennal grooves broad, pollinose; median lobe relatively
short, broad, obtusely pointed posteriorly; frontal groove shallow, parafrontal bosses distinct; temporal lobes slightly wider than long,
sinuate anterior to medial angles, latter apparently acute, because of pollinosity on the posterior surface; three to four temporal setae;
orbital groove marked by broad, complete band of pollinosity; occiput and frontal pit with conspicuous reddish pilosity; three to four
pairs of postlabial setae; postmentum pollinose, separating the glabrous shining genae (Fig. 63).
Pronotum elongate, length/geatest width 1.33; sides nearly parallel, widest point slightly posterior to middle; slightly narrowed at
base, more distinctly at apex; lateral margin not at all sinuate anterior to hind angle; pronotal carinae convex, rather narrow; outer
Carina relatively broadly truncate at base, at most slightly bent outwards there; inner carina broadened or rounded at base; four to six
marginal setae; precoxal carina absent.
Elytral striae deep, pollinose, coarsely punctured; intervals narrow, carinate; humerus prominent, quadrangular, with
conspicuous patch of golden pilosity; base of second interval elevated; Stria 1 without setae; Stria II with five to seven setae; Stria IV
with six to eight setae; Stria VII with four to five setae near apex; metasternum with many discal punctures, latter slightly coarser
than marginals; metasternum with narrow lateral pilose strip.
Anterior femur with ventral tooth in both sexes; anterior tibia with proximal tooth in male, absent from female; cleaning organ
with wide comb, strongly overlapped by brushes, comb teeth blunt; proximal spur stout, conical; distal spur small, hind calcar small, its
tip blunt, spur clearly removed from calcar, small, acute.
Females of this species are among the largest of Rhysodini. G. gigas differs from G. tubericeps in its
larger size and stouter form, in the absence of a precoxal carina, in the presence of the postmental
pilosity and the absence of the dorsal basal pilose bands of antennal Segments III-X. It differs from G.
cuneata in the shape of the pronotum and from G. hova in the presence of parafrontal bosses.
Variation. - Of the eighteen specimens we have studied, the following characters exhibit variation:
shape of medial angle of temporal lobe, width of base of the inner carina, depth of the frontal grooves,
extent of glabrous area of the parafrontal boss, elevation of bases of the third and fifth elytral intervals
in addition to the second and degree of sinuation of the lateral margin of the humeral tubercle. However,
there has been no consistent pattern of characters that would suggest another species in this group.
In addition, there is a male (MNHN) labelled: “Madagascar, Est, Presqu’Ile de Masoala, Hiaraka,
XII- 1968, coll. Vadon/Peyrieras” which resembles the type series except for reduction of orbital pilosity.
The latter is very narrow, and is almost discontinuous anteriorly, leaving the bases of the temporal setae
partly isolated in round clumps of pilosity. These, however, are in narrow contact with the pilosity of the
orbital groove, and are not completely isolated as in G. dentipes and G. cinerea.
Final judgement of these aberrant forms will have to be held until there are more specimens available
and better locality data recorded.
Grouvellina cuneata new species
(Fig. 54)
Type material. - HOLOTYPE male, labelled: “MADAGASCAR: Tamatave et Foret Alahakato,
1/VII, 1888, ex. coll. Breuning” (MRAC).
Description. - Length 7.2 mm. Antennal Segment XI distinctly longer than wide; stylet prominent, conical, 0.33 as long as
segment; tufts of minor setae on Segments V-X; Segment 1 with broad dorsal band of pollinosity; Segment II with two transverse
bands of pollinosity; Segments III-X each with apical pollinose band, but without basal one.
Head about as wide as long; labrum with four setae; frontal and postantennal grooves broad, pollinose; median lobe short, broad,
obtusely rounded posteriorly; parafrontal bosses distinct; temporal lobes slightly longer than wide, sinuate anterior to medial angles,
latter narrowly obtuse and narrowly separated; three to four temporal setae; orbital groove marked by broad, complete band of
pollinosity; occiput and frontal pit with reddish pilosity; four pairs of postlabial setae; postmentum and most of mentum glabrous,
shining, these areas separated by narrow transverse pollinose area.
Pronotum elongate, distinctly wedge-shaped; broadest just anterior to base, sides convergent anteriorly, apex narrow; length of
pronotum/greatest width 1.33; pronotal carinae rather narrow; convex; inner carina broadest at base, pointed anteriorly; marginal
carina narrower than in G. tubericeps and G. gigas\ outer carina obliquely truncate at base; three marginal setae; precoxal carina
absent.
Elytral striae conspicuously unequal; l-V impressed; I the narrowest, others successively wider to V, latter prominent and widely
pilose; Stria VI reduced to row of isolated punctures anteriorly, punctures narrowly connected in posterior half; Stria VII irregularly
curved, somewhat zigzag; intervals broader and less distinctly carinate than in G. tubericeps and G. gigas, humerus prominent,
subrectangular, conspicuously pilose; Stria I with one seta in apical half; Striae II with three setae in apical half; Stria IV with six to
eight setae; Stria VII with about five setae in apical fifth; metasternum with many coarse distal punctures, but without lateral pilosity.
Classification of Rhysodini
417
Femur of anterior leg relatively slender, with ventral tooth; male with proximal tooth on anterior tibia; hind calcar very small,
obtusely pointed, in rear view, doubly curved, so that its outline resembles the toe of a boot.
The most conspicuous marks of this species are the wedge-shaped pronotum, and the conspicuously
unequal elytral striae and intervals.
Grouvellina hova new species
(Fig. 53)
Type material. - HOLOTYPE male, labelled: “MADAGASCAR: Tsaramainiandro, X-1950, coll.
J. Vadon” (MNHN). Three PARATYPES two males, same data as type (MNHN); one male labelled:
“Madagascar” coll. Humblot 1885 (MNHN).
Description. — Length 8. 7-9.0 mm (only males are known; if the females are conspicuously larger than males, as in G. gigas.
they may be the largest member of the genus). Antennal Segment XI distinctly longer than wide, stylet conical, acute; tufts of minor
setae on Segments V-X; antennal Segment I with broad dorsal band of pollinosity; Segment 11 with two transverse pollinose bands;
Segments III-X with apical but not basal bands.
Head short, clearly wider than long; labrum with four setae; frontal, postantennal grooves broad, pollinose; median lobe short,
broadly rounded posteriorly; parafrontal bosses absent, superantennal area entirely pollinose; temporal lobes transverse, wider than
long, markedly sinuate anterior to medial angle, latter apparently acute, because of pollinosity on posterior surface; five to six temporal
setae; orbital groove marked by very broad band of pollinosity; occiput and frontal groove with conspicuous pilosity; three pairs of
postlabial setae; postmentum pollinose. *
Pronotum shorter than in G. gigas, length/greatest width 1.25; greatest width near middle, sides curved; apex strongly narrowed,
base slightly narrowed; lateral margin slightly sinuate anterior to hind angles; pronotal carinae convex, rather narrow; outer carina
rather broadly, obliquely truncate at base; marginal carina narrow, nearly vertical; five to six marginal setae; precoxal carinae absent.
Elytral striae deep, coarsely punctate; intervals narrow, carinate; humerus prominent, subrectangular, conspicuously pollinose;
Stria 1 without setae; Stria II with about ten setae; Stria IV with about nine setae; Stria VII with about five in its apical fifth;
metasternum with many discal punctures which are as coarse as lateral punctures; metasternum without lateral pollinosity.
Anterior femur with ventral tooth; male with proximal tooth on anterior tibia (female unknown); cleaning organ with comb hairs
in line with brush hairs, indistinctly differentiated from them and not overlapping them; both spurs of anterior tibia present, small,
acute; calcar of hind leg truncate, flattened.
This very large species resembles G. gigas, but is more stoutly built, and is easily distinguished by
complete absence of the parafrontal bosses. G. cooperi and most specimens of G. descarpentriesi also
lack the bosses, but differ from G. hova in being much smaller, in lacking the femoral tooth, and in
having the head elongate.
Grouvellina ranavalona new species
(Fig. 57)
Type material. - HOLOTYPE female, labelled: “MADAGASCAR, cn. Fairmaire” (MNHN).
PARATYPE female, labelled; “MADAGASCAR 1901-267” (BMNH).
Description. — Length 9.7-10.0 mm. Antennal Segment XI longer than wide; stylet large, conical, acute; tufts of minor setae
present on Segments V-X; antennal Segment I with broad dorsal band of pollinosity; Segment II with two transverse pollinose bands;
Segments III-X with apical pollinose bands but without basal ones.
Head slightly wider than long; labrum with four setae; frontal and postantennal grooves broadly pollinose; median lobe short,
broadly rounded posteriorly; parafrontal bosses distinct; temporal lobes slightly longer than wide; medial angles obtuse, narrow; two to
three temporal setae; orbital groove marked by broad, complete band of pollinosity; occiput and frontal pit with conspicuous
pollinosity; three to four pairs of postlabial setae; mentum nearly impunctate, glabrous except for narrow posterior trans\*erse strip of
pollinosity; postmentum also glabrous.
Pronotum relatively shorter than in G. gigas, length/greatest width 1.26: pronotum widest near middle, sides curved; apex strongly
narrowed, base slightly so; margin not sinuate anterior to hind angles; pronotal carinae convex, rather narrow; outer carinae narrowed
and abruptly divergent at base; five to six marginal setae; precoxal carina absent.
Elytral striae deep, coarsely punctured; intervals narrow, carinate, humeral tubercle prominent, conspicuously pilose; Stria I with
one seta near apex; Stria 11 with four setae in apical half; Stria IV with seven to eight setae; Stria VII with about nine setae in apical
fourth; metasternum with many discal punctures, latter finer than marginal ones; metasternum with some lateral pollinosity.
Anterior femur of female without ventral tooth, but with rounded swelling; male unknown.
Quaest. Ent., 1979, 15 (4)
418
Bell and Bell
This is a very large, broad-headed species similar to G. gigas and G. hova, but differing from both in
the absence of a ventral tooth on the anterior femur.
Grouvellina edentata new species
(Fig. 64)
Type material. - HOLOTYPE male, labelled: “MADAGASCAR, 1885, coll. Humblot no.
85/6044” (MNHN). PARATYPES three males, same data (MNHN).
Description. — Length 5. 0-6.0 mm. Antennal Segment XI slightly longer than wide; stylet short, conical; tufts of minor setae
on Segments V-X; Segment 1 with broad dorsal pollinose band; Segments II-X with narrow apical bands but no basal ones.
Head slightly longer than wide; four labral setae; frontal and postantennal grooves deep; median lobe broadly rounded at apex;
parafrontal bosses distinct, narrow, elongate; temporal lobes slightly longer than wide, rather shallowly sinuate anterior to medial
angles; latter obtuse; two temporal setae; orbital groove marked by narrow but complete band of pollinosity; two to three pairs of
postlabial setae.
Pronotum elongate, length/greatest width 1.40; widest near middle, sides slightly curved; base and apex only moderately
narrowed; lateral margin not sinuate anterior to hind angle; pronotal carinae rather broad, convex; inner carina widest near middle,
base and apex nearly equally tapered; outer carina obliquely truncate at base, but not divergent; three marginal setae; precoxal carina
present.
Elytral striae fine, shallow, very coarsely punctured; intervals convex but not carinate; base of Interval II elevated in form of small,
indistinct tooth; humeral tubercle rather small; Stria I with two to four setae in apical fifth; Stria II with two to three sete in apical
fourth; Stria IV with four to six setae; Stria VII with about five setae near apex; metasternum with scattered discal punctures, latter
finer than lateral ones; metasternum not pilose laterally.
Anterior femur without ventral tooth; anterior tibia without proximal tooth (male); cleaning organ with comb teeth not flattened;
distal spur of front tibia large; proximal one absent; male with calcar of hind leg acuminate, with narrow base, very close to spur, latter
large. Female unknown.
This is the smallest species in the genus. It has a complete, though narrow, orbital groove. It is most
similar to G. cooperi, from which it differs in having a distinct parafrontal boss, and in having the inner
pronotal carinae less narrowed anteriorly.
Grouvellina cooperi new species
(Fig. 68-70)
Type material. - HOLOTYPE female, labelled: “MADAGASCAR, coll. Rogez” (MCZ).
Description. — Length 6.9 mm. Antennal Segment XI slightly longer than wide; stylet conical, acuminate; tufts of minor setae
on Segments V-X; Segments 1,11 broadly pollinose dorsally; Segments I II-X with apical pollinose bands but no basal ones.
Head longer than wide; labrum with four setae; frontal and postantennal grooves deep; median lobe tapered anteriorly, widest near
broadly rounded apex, parafrontal bosses absent, superantennal area entirely pollinose; temporal lobes distinctly longer than wide,
rather shallowly sinuate anterior to medial angles; latter obtuse; three temporal setae; orbital groove marked by narrow but complete
band of pollinosity; three to four pairs of postlabial setae.
Pronotum narrow, elongate, length/greatest width about 1.33; pronotum widest distinctly anterior to middle; sides curved,
moderately narrowed at apex, slightly less so at base; lateral margin slightly sinuate anterior to hind angle; pronotal carinae broad,
rather convex; inner carina widest in basal fourth, apex strongly tapered; outer carina with extreme base markedly narrowed, but not
divergent; marginal setae 0, or one, located near angular seta; precoxal carina well developed, nearly extended to anterior margin (Fig.
69); postcoxal tooth large, blunt.
Flytral striae deep, narrow, coarsely punctate, punctures wider than striae; inner intervals nearly flat; outer ones more convex;
base of Interval II in form of distinct tooth; humeral tubercle slightly prominent, pollinose; Stria I without setae; Stria II with one seta
near base and another near apex; Striae IV and VII without setae; metasternum coarsely punctate, discal punctures equal in size to
marginal ones; metasternum with lateral pilose strips (Fig. 70).
Anterior femur without ventral tooth; both spurs distinct on anterior tibia; anterior and middle femora and all tibiae with many
more setae than in other members of genus. Male unknown.
This species is a small one with a complete, though narrow orbital groove. It differs from G. edentata
in lacking the parafrontal boss. In the latter feature, it resembles G. hova, but the latter species is much
larger, has carinate elytral intervals, and a ventral tooth on the anterior femur, and lacks a precoxal
carina. Most specimens of G. descarpentriesi also lack a parafrontal boss. These can be distinguished
Classification of Rhysodini
419
from G. cooperi by the absence of the outer labral setae and the precoxal carina.
It is a pleasure to dedicate this species to Dr. Kenneth Cooper, whose contributions to entomology
have been multiple in both place and time.
Grouvellina montana new species
(Fig. 52)
Type material. - HOLOTYPE male, labelled: “MADAGASCAR; coll. C. Schauf’ (MNHN),
written in Grouvelle’s hand “/?. montanus, type”. This name was not published. PARATYPES three
females, labelled: “MADAG.” (MNHN).
Description. — Length 6. 2-7.0 mm. Antennal Segment XI twice as long as wide; stylet acute; tufts of minor setae on
Segments V-X; Segment I with broad dorsal pollinose band; Segment II with basal and apical band, separated by narrow glabrous
band; Segments 111-X with apical bands but without basal ones.
Head slightly longer than wide; labrum with four setae; frontal and postantennal grooves deep; median lobe nearly parallel-sided,
its apex slightly broadened and rounded; parafrontal bosses distinct, triangular; temporal lobe slightly longer than wide, medial margin
shallowly sinuate anterior to obtuse medial angle; two temporal setae; orbital groove incomplete; pollinosity absent anterior to anterior
temporal seta, with glabrous area of temporal lobe extended to eye; five pairs of postlabial setae in pilose transverse band which
separates glabrous submentum from mentum.
Pronotum elongate, length/greatest width 1.35, widest anterior to middle, moderately narrowed both to base and apex; side not
sinuate anterior to hind angle; pronotal carinae broad, flat; extreme base of outer carina narrowed and divergent at hind angle; two to
three marginal setae, most anterior one near anterior angle; precoxal carina absent.
Elytral striae shallow, narrower than intervals, rather finely punctate; intervals slightly convex near base, otherwise nearly flat;
Stria VI not impressed, marked only by a row of punctures; humeral tubercle narrow but prominent; Stria I with one seta near apex;
Stria II with four in apical fifth; Stria IV with four to five setae; Stria VII with about six setae in apex; metasternum without discal
punctures, without pollinosity except in each marginal puncture.
Anterior femur with ventral tooth; that of male large, that of female small and partly concealed by pollinosity; male with large
proximal tooth on anterior tibia; female without such tooth; cleaning organ with comb comprised of short distinctly flattened teeth;
proximal and distal spurs both rather large; male with acutely pointed hind calcar; female with spurs of middle and hind tibiae very
unequal, anterior ones much reduced.
This is a distinctive small species with an incomplete orbital groove, a feature distinguishing it from
all other species of Grouvellina. Absence of a precoxal carina and presence of a femoral tooth distinguish
it from G. edentata and G. cooperi.
Grouvellina radama new species
(Fig. 55)
Type material. - HOLOTYPE male, labelled: “MADAGASCAR: Mahatsinjo pres Tananarive”
(MNHN). Six PARATYPES one male, same data as type (MNHN); one male, labelled:
“MADAGASCAR: Antsianaka, 1923, acq. Le Moult” (AMS); one female, labelled:
“MADAGASCAR, foret de Fito, ex. coll. Dr. Breuning” (MRAC); one male, two females, labelled:
“MADAGASCAR, col. Le Moult” (MRAC).
Description. — Length 7. 5-8. 9 mm. Antennal Segment XI slightly longer than wide; stylet short, conical; tufts of minor setae
on Segments V-X; Segments 1, II broadly pilose dorsally; Segments III, IV with broad apical pollinose bands; Segments V, VI with
narrow ones; Segments Vll-X each with two transverse rows of pollinose spots.
Head distinctly longer than wide, preocular portion elongate; labrum with two setae; frontal and postantennal grooves deep;
median lobe very narrow, its sides parallel, its tip rounded; parafrontal bosses distinct; temporal lobe longer than wide; margin scarcely
sinuate anterior to obtuse medial angle; five temporal setae; orbital groove represented by complete pollinose band; many pairs of
postlabial setae, of two distinct sizes; postmental area pollinose; mentum glabrous with scattered punctures.
Pronotum elongate, length/greatest width 1.35; widest near middle, sides almost parallel, base and apex only slightly narrowed;
lateral margin slightly sinuate anterior to hind angles; pronotal carinae very narrow, scarcely wider than the grooves, convex; inner
carinae straight, pointed at both ends; outer carina obtuse at apex, its base narrow, and divergent; two to three marginal setae;
precoxal carina fine, extended about halfway to anterior margin; prosternum with very deep medial pit; one specimen with pair of
prosternal setae near anterior ends of carinae.
Quaest. Ent., 1979, 15 (4)
420
Bell and Bell
Elytra! striae very broad, very coarsely punctured, expanded opposite base of each puncture; intervals narrow, less than half as
wide as striae, scarcely carinate, expanded punctures making the intervals appear segmented, slightly zigzag; humeral tubercle
small; Stria I with one seta at apex; Stria II with five to six apical setae; Stria IV with eight setae; apical tubercle with two setae;
Stria VII with about six setae in apical fifth; metasternum with lateral pollinose strips and with coarse discal punctures.
Anterior femur without ventral tooth; male without proximal tooth on anterior tibia; cleaning organ with stiff comb teeth near
middle gradually graded to long brush hairs on either side (not graded abruptly as in other species); distal spur well developed;
proximal spur vestige at base of intermediate tooth; hind calcar rather long, its dorsal and ventral margins nearly parallel, its tip
truncate.
This large species has a distinctive long, narrow, parallel-sided median lobe, duplicated only in the
otherwise very different G. grouvellei. The pronotal carinae are narrower than in other species. The most
similar species is G. divergens, which has a similarly elongated head, but which has a wide median lobe,
and lacks the precoxal carinae.
Grouvellina descarpentriesi new species
(Fig. 65-67)
Rhysodes tubericeps auct. nec. Fairmaire. Both Grouvelle (1903) and Arrow (1942) interpreted this
species as R. tubericeps.
Type material. - HOLOTYPE male, labelled: “MADAGASCAR, Annanarivo (Sikora)” (MNHN).
18 PARATYPES (all from Madagascar) as follows; one female, same data as type; one male, two
females, Antananal; one female, Antananarivo; one male, three females (on same pin), Madagascar,
Sikora; four males, one female, “Madagas” (all MNHN); one female, “Madagas”. Fry Colin. 1905-100
(BMNH); one female, “Madagas”. 79.18 (BMNH); one female, foret de Fito, ex. coll. Dr. Breuning
(MRAC); one female, “Madagas., ontv. 6 April 08, Ant. Grouvelle” (LEI).
Description. — Length 6. 1-8.0 mm. Antennal Segment XI almost twice as long as wide; stylet long, acute, about 0.33 total
length of segment; tufts of minor setae on Segments V-X; Segment I and II with broad dorsal pollinose band; Segments Ill-V with
narrow apical pollinose bands; Segments Vl-X with apical pollinose bands and basal transverse rows of pollinose spots.
Head longer than wide; preocular portion elongate; labrum with two setae; frontal grooves deep, rather broad, postantennal
grooves deep; median lobe moderately broad, slightly broadened posteriorly, tip broadly rounded; parafrontal boss variable, in most
specimens absent, in few specimens suggested by small, irregular glabrous areas; temporal lobe longer than broad; margin shallowly
sinuate anterior to medial angles, latter obtusely rounded, margin not emarginate posterior to medial angle; two temporal setae; orbital
groove marked by rather narrow band of pilosity; two pairs of postlabial setae; mentum with lateral margins pilose, remainder
glabrous; postmentum glabrous, separated from genae and mentum by narrow pollinose band.
Pronotum elongate, narrow, length/greatest width 1.35; widest point anterior to middle, apex scarcely narrower than base; margin
shallowly sinuate anterior to hind angles; pronotal carinae rather narrow, convex, only slightly broader than paramedian grooves; outer
Carina narrowed and abruptly divergent at base; three marginal setae, most anterior one near anterior angle; precoxal carina
represented by small rudiment between two precoxal pits.
Elytral striae moderately broad, slightly wider than intervals, coarsely, closely punctate; striae dilated around each puncture,
intervals of irregular width; intervals convex but not carinate; base of Interval II raised in form of small “tooth”; humeral tubercle
moderately prominent; Stria I without setae; Stria 11 with two near apex; Stria IV with five setae; Stria Vll with five setae in apical
fifth; metasternum with lateral punctures but without discal ones or lateral pilosity (Fig. 67).
Anterior femur without ventral tooth; male without proximal tooth on anterior tibia; cleaning organ with central comb sharply
distinct from overlapping brushes on hairs at either end; distal spur well developed; proximal spur absent; hind calcar of male very
small, blunt; hind femur of male sinuate ventrally.
This is a moderate-sized species with an elongate head. It is most similar to G. radama, but the latter
is larger, has a narrower, more elongate median lobe and much narrower pronotal carina, as well as
having a well-developed precoxal carina. G. divergens is also similar to G. descarpentriesi, but differs in
the shape of the medial angle of the temporal lobe, in lacking a tuft of minor setae on antennal
Segment V, and in having the marginal carina of the pronotum abbreviated posteriorly.
Of the 20 specimens studied, two have a distinct parafrontal boss. They are labelled as follows; one
male, “Madagascar, Centre, Pays Betsileo, Route du Sud, km. 292, 1700m., 14/15-11-1974, P. Viette et
A. Peyrieras” (MNHN) (Fig. 66); one female, “Madagascar, int. austr. Hildebrandt S.” (MNHB). We
Classification of Rhysodini
421
have tentatively identified these as G. descarpentriesi based on the other species characters, but have not
made them paratypes. More specimens are needed to determine whether or not this is a distinct species.
It is a pleasure to dedicate this species to Dr. A. Descarpentries of the Museum National d’Histoire
Naturelle, Paris, whose generous aid during our study of the Grouvelle Collection was vital to the
completion of this work.
Grouvellina divergens new species
(Fig. 56)
Type material. - HOLOTYPE male, labelled; “MADAGASCAR, Mt. d’Ambre, 1930, coll. Sicard”
(MNHN). Nine PARATYPES one female, same data as type (MNHN); one female, same locality as
type, Dec. 1900 (MNHN); four females, Mt. d’Ambre, no date (MNHN); one male, two females
labelled: “MADAGASCAR, Diego-Suarez, 1893, coll. C. Alluaud’’ (MNHN).
Description. — Length 6. 0-8. 4 mm. Antennal Segment XI slightly longer than wide; stylet short, conical; tufts of minor setae
present on Segments VI-X (absent from Segment V); basal setae absent from Segments V and VI; Segments 1-11 broadly pilose
dorsally; Segments lll-V with broad apical pollinose band; Segments Vll-X each with two transverse rows.of pollinose spots.
Head slightly longer than wide; preocular part narrow; slightly elongate; labrum with two setae; frontal and postantennal grooves
deep; median lobe broad, dilated, its tip very broadly rounded; parafrontal bosses distinct; temporal lobe slightly longer than wide, its
margin rather deeply sinuate anterior to medial angles, latter apparently acute, partly because of occipital pilosity; margin posterior to
medial angles distinctly emarginate; two to three temporal setae; orbital groove represented by rather narrow band of pollinosity, its
medial margin irregular; two pairs of postlabial setae; sides of mentum pilose; submentum pilose in most specimens, small glabrous
median area present in others.
Pronotum moderately elongate; length/greatest width 1.31; widest anterior to middle, base moderately narrowed, apex rather
markedly so; lateral margin sinuate anterior to hind angle; pronotal carinae moderately narrow, convex; outer carina narrowed and
abruptly divergent at base; marginal carina abbreviated posteriorly; marginal setae absent, angular seta present; precoxal carina
absent.
Elytral striae broad, slightly wider than intervals; latter convex, subcarinate; borders of intervals slightly irregular; humeral
tubercle rather small. Stria I with three setae in apex; Stria 11 with six setae in apical half; Stria IV with six setae; Stria Vll with
about six in apical fifth; metasternum with discal punctures much finer than marginal ones; metasternum without lateral pollinose
strips.
Anterior femur without ventral tooth; male without proximal tooth on anterior tibia; cleaning organ with comb teeth conical, not
flattened, graded gradually to brush hairs on either side, rows not overlapping; both spurs of anterior tibia well developed; hind calcar
small, more or less pointed, only slightly larger than spur.
This is a moderate-sized species, superficially like G. radama and G. descarpentriesi, but differing
from them in the absence of minor setae from antennal Segment V. In the latter feature it resembles
only G. grouvellei.
Grouvellina grouvellei (Fairmaire)
(Fig. 71)
Rhysodes grouvellei Fairmaire, p. 10
Type material. - LECTOTYPE (here designated) male, labelled: “MADAGASCAR” (MNHN).
PARALECTOTYPE one female, labelled: “MADAGASCAR, Fry Coll. 1905-100”, labelled as
“cotype” (BMNH).
Description. — Length 7.0-8. 8 mm. Antennal Segment XI longer than wide; stylet conical, short; tufts of minor setae on
Segments VI-X (absent from Segment V); basal setae absent from Segment V; Segment 1 with most of dorsal surface pilose; Segment
II with pollinose band; Segments lll-V with narrow apical pollinose band; Segment VI with apical band and basal row of pilose spots;
Segments Vll-X with two rows of pilose spots.
Head distinctly longer than wide, preocular portion narrow, elongate; labrum with two setae; frontal and postantennal grooves
deep; median lobe very long, narrow, its sides parallel, its tip obtusely rounded; parafrontal boss absent from lectotype but well
developed in paralectotype; temporal lobe longer than broad, its medial angles obtuse; glabrous area of temporal lobe very small, less
than 0.33 of total width of temporal lobe; three to four temporal setae; two to three pairs of postlabial setae; sides of mentum and
entire postmentum pollinose; mentum with scattered pilose punctures.
Quaest. Ent., 1979, 15 (4)
422
Bell and Bell
Pronotum rather narrow, elongate, length/greatest width 1.36; widest anterior to middle; base moderately narrow, apex more
strongly so; margin with long shallow sinuation anterior to hind angle; median and paramedian grooves deep, wide; marginal groove
incomplete, closed both anteriorly and posteriorly, confined to middle third of pronotum; pronotal carinae extensively pollinose,
inner carina with narrow glabrous space extended most of length; outer carina with traces of interrupted linear glabrous area;
marginal carina with small glabrous area near middle of length; four to five marginal setae; rudimentary precoxal carina present;
postcoxal tooth small.
Elytral striae shallow, with very coarse deep punctures; intervals nearly flat, almost entirely pollinose, except for narrow strip
along suture and narrow, irregular discontinuous strips on Intervals II and 111 and on apical tubercle and traces in some of the
outer intervals; some of the strial punctures also glabrous; humeral tubercle small; Stria 1 with two setae near apex; Striae II with
about twelve setae (some displaced laterad to stria); Stria IV with about ten setae (some displaced laterad to stria); Stria VI with
about twelve setae; about six setae on apical tubercle; about seven in apical fifth of Stria VII; metasternum with discal punctures
finer than marginal ones; metasternum with lateral pollinose strip.
Anterior femur without ventral tooth; proximal tooth absent from anterior tibia of male; all femora unusually long and slender
for genus; cleaning organ with central row of comb teeth overlapped at either end by row of slender brush hairs; distal spur well
developed; proximal one absent; hind calcar slender, tapered, its tip blunt.
This species is easily recognized by the abbreviation of the marginal groove and by the extensive
development of the pollinosity, which covers almost the entire dorsal surface.
Variation. - As aforementioned, the parafrontal boss is absent from the lectotype, but well developed
in the paralectotype. As for G. descarpentriesi, two species may be represented, but more specimens will
be needed to make certain. In addition to the type series, we have seen two other specimens, a male
labelled: “MADAG.” (MNHN), and another male, labelled: “MADAGASCAR: Annanarivo (Sikora)”
(MNHN). Both of these specimens lack the parafrontal boss.
Grouvellina dentipes new species
(Fig. 73, 74)
Type material. - HOLOTYPE male, labelled: “MADAG ASCAR:Fizono; IX, 1959, coll. J. Vadon”
(MRAC).
Description. — Length 7.8 mm. Antennal Segment XI slightly longer than wide; stylet conical, large, rather blunt; tufts of
minor setae present on Segments V-X; Segment I with broad dorsal pollinose band; Segment II with partly confluent basal and apical
pollinose bands; Segments Ill-V with both basal and apical bands; Segment VI with basal band and apical row of pilose spots;
Segments VlI-VIll similar, but with apical spots much reduced; Segments IX, X with basal band but no apical spots or bands.
Head wider than long; labrum with two setae; postantennal and frontal grooves deep; median lobe short, broad, parallel-sided,
abruptly truncate posteriorly; anterior third of median lobe finely pollinose; parafrontal boss absent, superantennal area entirely
pollinose; temporal lobe broader than long; margin deeply sinuate anterior to narrow, distinct medial angle; orbital groove represented
by narrow but complete strip of pilosity; row of five prominent round tufts medial to it, each more or less concealing a temporal seta;
two pairs of postlabial setae.
Pronotum very short for genus, length/ widest width 1.20; pronotum widest anterior to middle, moderately narrowed to base,
markedly narrowed to apex; margin distinctly sinuate anterior to hind angle; pronotal carinae convex, narrow, their margins pollinose,
centers glabrous; inner carina entire; outer carina abbreviated at basal fifth of pronotum; marginal groove narrow, confluent with
paramedian groove at base of outer carina; marginal carina entire, broader than outer carina; pronotum with angular seta, and one
marginal on each side, near anterior angle; precoxal carina represented by small rudiment, between two precoxal pits.
Elytral striae broad, deep, coarsely punctate; intervals narrow, convex, subcarinate less than half width of striae; humeral tubercle
small; Stria I with four setae in apical fifth; Stria 11 with two near apex; Stria IV with five; Stria VII with four in apical fifth;
metasternum with discal punctures coarser than marginal ones; metasternum with lateral pilose strip; Sternite V with deep large
lateral pits.
Anterior femur with large ventral tooth; male with large proximal tooth on anterior tibia (Fig. 74) (female unknown); hind calcar
of male tapered, prominent, its tip truncate.
The abbreviated outer pronotal carina and the conspicuous tufts on the temporal lobe separate this
species from all others excepting G. cinerea. From the latter it is easily distinguished by the large ventral
tooth on the anterior femur.
Classification of Rhysodini
423
Grouvellina cinerea new species
(Fig. 72)
Type material. - HOLOTYPE male, labelled: “MADAGASCAR: Perinet, sur Tavolo, 16-1-64,
Rec. Brunck, coll. CTFT No. Ill” (BMNH).
Description. — Length 9.2 mm. Antennal Segments X-XI missing from holotype; tufts of minor setae begin on Segment V;
Segment I with broad band of pollinosity; Segment 11 with broad basal band; Segments 111-V each with broad basal and apical bands,
separated by narrow glabrous bands; Segment VI with basal band and trace of apical band; Segments VII-IX with basal band but
without apical band or spots.
Head wider than long; labrum with two setae; postantennal and frontal grooves deep; median lobe short, broad, parallel-sided, its
middle third pollinose, its tip truncate; parafrontal boss absent; superantennal area pollinose anteriorly, more pilose posteriorly;
temporal lobe broader than long; margin deeply sinuate anterior to medial angle; latter blunt, broader than in G. dentipes; orbital
groove represented by narrow strip of pollinosity, isolated tufts medial to it, one or two anterior tufts, and three to four tufts in form of
transverse row posteriorly, each tuft more or less concealing a temporal seta; four pairs of postlabial setae.
Pronotum rather short, length/greatest width is 1.27; widest anterior to middle, moderately narrowed to apex and to base; margin
slightly sinuate anterior to hind angle; pronotal carinae convex, narrow, glabrous at center, both inner and outer margins pollinose;
inner carinae entire; outer carina abbreviated at basal fifth of pronotum; marginal groove narrow, confluent with paramedian groove
at base of outer carina; marginal carina entire, broader than outer carina; pronotum with angular seta and one marginal located near
anterior angle; precoxal carina represented by rudiment between two precoxal pits.
Elytral striae broad, deep, coarsely punctate; intervals narrow, convex, subcarinate, less than half width of striae; intervals
distinctly unequal II, IV, V, and especially 111, dilated at base; II very narrow posteriorly, its glabrous area more or less interrupted;
Interval IV abbreviated posteriorly, distinctly shorter than 111; humeral tubercle small; Stria I with two setae near apex; Stria II
withone near apex; Stria IV with five setae; Stria VI 1 with about five in apical fifth; metasternum with coarse discal punctures;
metasternum with lateral pilose strip; Sternite V with large deep lateral pits.
Anterior femur without ventral tooth; male without proximal tooth on anterior tibia (female unknown); hind calcar of male very
small, acutely pointed.
This species is similar in most respects to G. dentipes, but differs in the absence of the anterior
femoral and proximal tibial teeth.
SUBTRIBE OMOGLYMMIINA
Description. - Part I, 66.
Key to Genera. - Part I, 66-67.
Gtnu?, Xhosores Bell and Bell 1978
Type species. - Rhysodes figuratus Germar 1840
Description. - Part I, 67. Only one species is known.
Xhosores figuratus (Germar 1840)
Rhysodes figuratus Germar 1840: 352.
Type material. - We have studied a male specimen from MNHB believed to be the type. It is
labelled “CAP, 43699”.
Ouaest. Ent., 1979, 15 (4)
424
Bell and Bell
Genus Yamatosa NEW NAME
(Fig. 75-85)
Yamatoa Bell 1977, nec Kiriakoff 1967. We are grateful to Dr. Takehiko Nakane for pointing out this
synonym.
Type species. - Rhysodes niponensis Lewis 1888.
Description. — Antennal Segment XI with stylet minute or absent; base of antennal Segment III more or less elongate;
Segments V-X wider than Segments ll-IV; minor setae in form of subapical rings on Segments V-X; basal setae absent; labrum with
two setae; frontal, antennal grooves narrow; orbital grooves absent; clypeal setae one pair; temporal setae absent; postlabial setae one
or two pairs.
Pronotum with basal impression on each side, preceded by discal striole; basal impression with tubercle; disc of pronotum
continued laterad to basal impression, in form of basilateral lobe; basal impression partly closed posteriorly by “brace” laterad to inner
lobe of base; marginal groove of pronotum single, fine, in most species ended anteriorly near anterior margin of pronotum (but reduced
in Y. reitteri)\ pronotal setae absent.
Elytral striae coarsely punctured; basal scarp of elytron transverse medially; humeral tubercle present; elytral setae absent from
Stria H, confined to apex of Stria IV or else absent; metasternum with marginal punctures; abdominal sterna with transverse bands of
punctures; lateral pits present in both sexes, in most species confined to Sternum IV, in a few species also present on Sternum V;
Sternum VI without setae.
Middle and hind tibiae each with two spurs; calcar of middle leg of male acute, directed distomedially; hind calcar blunt.
The genitalia of Y. reitteri have been described and figured (Part I, 68-69).
Lack of paramedian grooves separates this genus from most others in Subtribe Omoglymmiina.
Shyrodes and Srimara appear to be related to Yamatosa, or perhaps represent highly specialized
offshoots of it. Both Shyrodes and Srimara have ocelliform compound eyes, and have the basal scarp of
the elytron modified. In Yamatosa the eyes may be fully developed or reduced, but are not ocelliform,
and the basal scarp is transverse. Plesioglymmius is another genus which does not have complete
paramedian grooves, but differs from Yamatosa in having only one spur on each middle and hind tibia.
In Plesioglymmius, all species except P. jugatus have the discal strioles dilated, and not linear.
This genus is confined to the Oriental Region, from the Himalaya to Japan, Taiwan, and Java.
Phylogeny. - With the exception of Y. arrowi, the species of Yamatosa are grouped clearly into an
“Eastern Line” and a “Western Line”. In the Eastern Line (Y. niponensis, Y. longior, and Y.
peninsularis), the antenna has a distinct apical stylet, there is a beard of setae on the mentum of the
male, and there are distinct lateral pits on abdominal Sternum IV, but not on Sternum V. In the
Western Line, the antennal stylet is absent, the beard is absent, and there are lateral pits on both
abdominal Sternum IV and V. Y. boy si, Y. draco and Y. reitteri belong to the Western Line, known only
from the Himalaya. Y. arrowi, despite its being a Himalayan species, clearly belongs to the Eastern
Line.
Among the species of the Eastern Line, Y. longior and Y. peninsularis are clearly closely related,
both having a precoxal carina, short discal strioles, and, in the male, the anterior tibia grooved medially.
In Y. longior, both sexes have a bearded mentum, while the female of Y. peninsularis is unknown. In
Y. niponensis, the precoxal carina is absent, the discal striole is long, the beard is present in the male but
absent from the female, and the anterior tibia of the male is not grooved. The relationships of Y. arrowi
are somewhat problematical, partly because the male is unknown. The absence of a precoxal carina and
the long discal striole seem to link it to Y. niponensis, while the presence of a beard in the female is a
character shared with the two preceding species. It also differs from all other members of the genus in
several characters, including large size, broad-based pronotum, dilated frontal grooves, and enlarged
anterior tentorial pits.
Among the species of the Western Line, Y. boy si and Y. draco are clearly closely related to one
another. Both species have reduced eyes, and probably both are flightless (F. boy si has vestigial hind
wings, while Y. draco has not yet been checked for this character). Both species have one seta each in the
apex of Striae II and IV of most specimens, while such setae are not found in other species of Yamatosa.
Classification of Rhysodini
425
V. reitteri is a more isolated species. Unlike the two preceding species, it has full-sized eyes and hind
wings. It has two specializations not seen in any other member of the genus: the marginal groove of the
pronotum has been reduced to a basal remnant, while the male has lost the ventral tooth of the anterior
femur.
KEY TO SPECIES
1 Segment XI of antenna with distinct apical stylet; both eye and marginal groove of
pronotum fully developed 2
V Segment XI of antenna obtuse, without apical stylet; either eye reduced or else marginal
groove of pronotum reduced 5
2 (1 ) Prosternum with distinct precoxal carinae; discal stride ended at or posterior to middle of
pronotum 3
1' Prosternum without precoxal carinae; discal stride ended at or anterior to apical third of
pronotum 4
3 (2 ) Precoxal carina extended more than 75% of distance from coxa to anterior margin of
prosternum; discal stride 50% pronotal length Yamatosa longior (Grouvelle), p. 425
y Precoxal carina, extended about 33% of distance from coxa to anterior margin of
prosternum; discal stiole 33% pronotal length Yamatosa peninsularis (Arrow), p. 427
4 (2') Frontal and antennal grooves narrow, equal in width to posterior part of clypeal grooves;
posterior margins of frontal and antennal grooves sharply defined; discal stride ended at
anterior third of pronotum Yamatosa niponensis (Lewis), p. 427
4' Frontal and antennal grooves dilated, much wider than posterior part of clypeal grooves;
posterior margins of frontal and antennal grooves not sharply defined; discal stride
extended nearly to anterior margin of pronotum Yamatosa arrowi (Grouvelle), p. 428
5 (L) Marginal groove of pronotum absent except in basal fourth of pronotum; eyes large,
much deeper than long Yamatosa reitteri (Bell), p. 429
5' Marginal groove of pronotum nearly complete, ended short distance from anterior
margin of pronotum; eyes more or less reduced 6
6 (50 Head evenly rounded posteriorly, widest point just posterior to eye; eye only moderately
reduced, with about 100 ommatidia, deeper than long (Fig. 77)
Yamatosa draco (Bell), p. 429
6' Head broadened posteriorly, widest point far posterior to eye; eye markedly reduced, with
about 50 ommatidia, longer than deep (Fig. 85) Yamatosa boysi (Arrow), p. 430
Yamatosa longior (Grouvelle 1903) NEW COMBINATION
Fig. 78-80)
Rhysodes niponensis var. longior GtouvqWq 1903: 107.
Yamatoa longior (Bell and Bell) 1978.
Type material. - LECTOTYPE (here designated) female, labelled: “JAVA:Tougou, 3-96, coll.
J.D. Pasteur” (MNHN). Two PARALECTOTYPES: one female, same data as lectotype (MNHN);
one female, same data as lectotype, except that locality is spelled “Toegoe”, in Dutch orthography
(LEI).
Quaest. Ent., 1979, 15 (4)
426
Bell and Bell
Figures 75 - 87. Fig. 75-85, Genus Yamatosa\ Fig. 75-76, Head and pronotum, dorsal aspect; Fig. 75, Y. peninsularis (Arrow);
Fig. 76, Y. draco Bell; Fig. 77, Head, lateral aspect, Y. draco Bell, Fig. 78-80, Y. longior (Grouvelle); Fig. 78. Head and
pronotum, dorsal aspect; Fig. 79, Prothorax, ventrolateral aspect; Fig. 80. Head, ventral aspect; Fig. 81-84, Head and pronotum,
dorsal aspect; Fig. 81, K niponensis (Lewis); Fig. 82, Y. arrowi (Grouvelle); Fig. 83, Y. reitteri Bell; Fig. 84, Y. boysi (Arrow);
Fig. 85, Head, lateral aspect, Y. boysi (Arrow); Fig. 86,87, Head and pronotum, dorsal aspect; Fig. 86, Shyrodes dohertyi
Grouvelle; Fig. 87. Srimara planicollis Bell and Bell.
Classification of Rhysodini
427
Description. — Length 5. 3-6. 9 mm. Antennal Segment XI with apical stylet distinct; head cordate, anterior tentorial pits
small, punctiform; frontal grooves narrow, well defined; eye large, deeper than long; mentum bearded in both sexes (Fig. 80); one pair
of postlabial setae in most specimens.
Pronotum of medium length for the genus, length/greatest width about 1.33; base moderately narrowed; apex more strongly so;
discal striole extended approximately to middle of pronotum; marginal groove complete; precoxal carina markedly developed, extended
almost to anterior margin of prosternum (Fig. 79).
Strial punctures relatively fine; intervals relatively flat; Striae II, III abbreviated at base; Stria V effaced in basal sixth, only
slightly impressed; Stria VI not impressed, absent in basal third, represented by row of minute punctures in apical two-thirds;
Stria VII effaced in basal half, represented by minute punctures in apical half; elytral setae confined to apex of Stria VII.
Abdominal Sternum IV with very small lateral pits; Sternum V without pits; male with shallow oval median impression extended
from middle of metasternum to abdominal Sternum III.
Anterior femur with ventral tooth in both sexes; anterior tibia of male with medial groove, latter bounded both anteriorly and
posteriorly near base by pair of flanges; spurs of middle and hind tibiae nearly equal; hind calcar with tip just above level of spurs.
Although this species was described as a “variety” or subspecies of Y. niponensis, it is actually a
distinct species, more like Y. peninsularis than like Y. niponensis. Contrary to Grouvelle’s key, it is not
more elongate than Y. niponensis. The complete precoxal carinae separates Y. longior from all other
members of the genus.
Range. - Java and Sumatra. In addition to the type material, we have seen the following specimens:
JAVA: one female, Kopeng, 1600 m., 14-5-1933, coll. P.H.v. Doesburg (LEI); two males, Malang
(MNHN); one female, Mt. Tengge (MNHN); one male, one female, G. Papandajan, 3-1916, coll.
Drescher (AMS); one male, one female. Preanger, coll. P.F. Sijthoff (LEI); SUMATRA: one male, two
females, Pajakombo (MNHN).
Yamatosa peninsularis (Arrow 1942) NEW COMBINATION
(Fig. 75)
Rhysodes peninsularis Arrow 1942: 178.
Yamatoa peninsularis (Bell and Bell) 1978.
Type material. - HOLOTYPE male, labelled: “FEDERATED MALAY STATES: PAHANG:
Tanak Rata, 4800 ft., Cameron’s Highlands, May 28, 1931, coll. H.M. Pendlebury” (BMNH).
Description. — Length 5.2 mm. Antennal Segment XI with distinct stylet; head cordate; anterior tentorial pits small,
punctiform; frontal grooves narrow, well defined; eyes large, deeper than long; mentum bearded in male, beard les extensive than in
Y. longior, status in female unknown; one pair of postlabial setae.
Pronotum longer and narrower than in Y. longior, length/greatest width 1.40; base moderately narrowed; apex markedly so; discal
striole short, not extending beyond basal third of pronotum; marginal groove complete; precoxal carina present, shorter than in
Y. longior, extended less than half the distance from coxa to anterior margin of pronotum.
Strial punctures relatively fine; intervals relatively flat; Striae 11, 111 abbreviated at base; Stria V effaced in basal sixth, only
slightly impressed; Stria VI not impressed, absent in basal third, represented by row of minute punctures in apical two-thirds;
Stria VII effaced in basal half, represented by minute punctures in apical half; elytral setae confined to apex of Stria VII.
Abdominal Sternum IV of male with lateral pit larger than Y. longior (female unknown); Sternum V without lateral pits; holotype
with raised tubercle in middle of metasternum (possibly an abnormality), metasternum not as definitely impressed at middle as in
Y. longior.
Anterior leg of male as in Y. longior, spurs of middle and hind tibiae nearly equal; hind calcar larger than in Y. longior, with
upper oblique margin longer.
This species is easily identified by the half-length precoxal carina. It also differs from Y. longior in
having a more elongate pronotum with notably small basal impressions and shorter discal strides.
Yamatosa niponensis (Lewis 1888) NEW COMBINATION
(Fig. 81)
Rhysodes niponensis Lewis 1888: 82.
Yamatoa niponensis Bell and Bell 1978
Quaest. Ent., 1979, 15 (4)
428
Bell and Bell
Type material. - LECTOTYPE (here designated) male, labelled; “JAPAN: Honshu, Hakone
District, coll. G. Lewis” (BMNH). Six PARALECTOTYPES: one male, two females, same data as
lectotype (BMNH); one female, JAPAN: Subushri (BMNH); one male, one female (no specific
locality), coll. G. Lewis, 1910-320, C.F. Baker Colin. (1927) (NMNH). The male is labelled as a
“co-type”.
Description. — Length 5. 4-7.0 mm. Antennal Segment XI with apical stylet distinct, though small; head cordate, anterior
tentorial pits small, punctiform; frontal grooves narrow, well-defined; eye large, deeper than long; mentum conspicuously punctate and
bearded in male; in female beard greatly reduced to absent, punctures very fine; one pair of postlabial setae.
Pronotum moderately elongate for genus, length/greatest width about 1.40; base moderately narrowed; apex more strongly
narrowed; discal striole extended 0.66 length of pronotum; marginal groove complete; precoxal carina absent; a group of many
punctures anterior to each front coxa.
Strial punctures fine, intervals relatively convex; Striae 11, 111 scarcely abbreviated at base; Stria V slightly abbreviated at base,
only shallowly impressed, coarsely punctate; Stria VI not impressed, represented by row of fine punctures; Stria VII represented by
minute punctures in anterior half; those of posterior half coarser; only apex of Stria Vll impressed; elytral setae confined to apex of
Stria Vll.
Abdominal Sternum IV with lateral pits; Sternum V without pits; metasternum shallowly impressed medially in male.
Anterior femur with ventral tooth in both sexes; anterior tibia of male slightly swollen near base, with trace of groove on medial
surface (much less distinct than in preceding species); spurs of middle and hind tibiae nearly equal; hind calcar of male small, forming
subacute angle immediately above spurs.
This species resembles Y. peninsularis in form, but differs from the latter in the complete absence of
a precoxal carina, and in the much less distinct medial groove on the anterior tibia.
Range. - Japan and Taiwan. A series in MNHN, labelled: “Sydney, Australia”, is surely mislabelled,
as the genus is otherwise unknown east of Java. In addition to the type material, we have seen the
following specimens: JAPAN: (HOKKAIDO) one female, Wakoto, 5-V11-1958, coll. F. Takechi (SATO); (HONSHU) one
female, Hida, Vll, 1957, coll. H. Torigai (SATO); one female, Hida, Futatsuya, VI-1954, coll. H. Torigai (SATO); one female, Hida,
Idani, Kawai, Vll-1953, coll. H. Torigai (SATO); Ikenoshima, 900' (Higashimata, Nikko District) (sex and number not recorded)
(BMNH); two males, Tsuta-yu (Towada), Vll-25-1954, coll. J. Aoki (DY); (KYUSHU) one male, one female, Mt. Homan, Fukuoka,
7-1V-1965, coll. M.T. Chuyo (MNHB); Nagasaki, Chiuzenji, Ichiuchi, Oyayama District (numbers and sexes not recorded)
(BMNH); (SHIKOKU) one male, one female, Mt. Kohtsu (Tokushima), 31-X-1965, coll. M. Sakai (SATO); one female, Omogo,
lO-Vll-1956, coll. M. Sato (SATO). TAIWAN (FORMOSA): one male, Taiheizan, 6-V-1932, coll. L. Gressitt (CAS).
According to Lewis, this species is found under the bark of firs {Finns) (now in the genus Abies).
Yamatosa arrowi (Grouvelle 1908) NEW COMBINATION
(Fig. 82)
Rhysodes arrowi Grouvelle 1908: 318.
Yamatoa arrowi Bell and Bell 1978.
Type material. - HOLOTYPE female, labelled: “SIKKIM” (MNHN).
Description. — Length 9.5 mm. Antennal Segment XI with distinct stubby stylet; head cordate; anterior tentorial pits large,
rounded; frontal and antennal grooves dilated, their posterior margins indistinctly defined; eye large, deeper than long; mentum
punctate and bearded in female (male unknown); two pairs of postlabial setae.
Pronotum moderately elongate; length/greatest width 1.38; apex markedly narrowed; base much less narrowed than in other
members of the genus; discal striole 80% of length of pronotum; marginal groove complete; precoxal carina and precoxal punctures
absent.
Strial punctures relatively coarse; elytral setae limited to apex of Stria Vll; abdominal Sternum IV with distinct lateral pits;
Sternum V without lateral pits.
Anterior femur with large acute ventral tooth in female (male unknown); spurs of middle and hind tibiae nearly equal.
Enlarged anterior tentorial pits, dilated frontal grooves, broad base of the pronotum, and very long
discal strides are diagnostic of this species, which is also much larger than other members of the genus.
Classification of Rhysodini
429
Yamatosa reitteri (Bell 1977) NEW COMBINATION
(Fig. 83)
Rhysodes boysi auct., nec. Arrow
Yamatoa reitteri (Bell), 1977: 152.
Type material. - HOLOTYPE male, labelled; “INDIA, U. Gumti Val. W. Almora Dn., Apr. 19,
coll. H. Gower Champion” (BMNH). 14 PARATYPES, listed in Bell (1977), 7 (BMNH), 4(MNHN),
3(BSL).
Description. — Length 6. 2-7. 3 mm. Antennal Segment XI obtuse, stylet absent; head cordate, sides evenly rounded; anterior
tentorial pits, rounded; frontal grooves narrow, well defined; eye large, deep, with more than 150 ommatidia; mentum nearly smooth,
without beard; one pair of postlabial setae.
Pronotum moderately long for the genus, length/greatest width 1.37; base moderately narrowed, apex more strongly narrowed;
discal striole ending anteriorly at basal third of pronotum; marginal groove largely effaced, distinct only in basal fourth of pronotum;
precoxal carina absent; precoxal punctures absent.
Elytra relatively elongate, narrow; humeral tubercle relatively prominent in form of small, laterally-directed tooth; strial punctures
relatively coarse; intervals moderately convex; Striae II, 111 abbreviated at base; Stria V slightly impressed, its anterior end slightly
posterior to humeral tubercle; Stria VI only slightly abbreviated anteriorly; elytral setae confined to apex of Stria VII; hind wings fully
developed.
Male with neither metasternum nor abdomen concave; Sternite IV (both sexes), with conspicuous lateral pits, shallower in male;
Sternite V with smaller, shallower lateral pits.
Anterior femur without ventral tooth in either sex; anterior tibia of male without medial groove or swelling; anterior spur
conspicuously smaller than posterior one on both middle and hind tibiae; hind calcar of male in form of small acute angle immediately
above level of spurs.
The male genitalia were figured in Part 1, 68.
This species differs from all others in the genus in having the marginal grooves of the pronotum
largely effaced.
Y. reitteri is confined to the middle and eastern Himalaya, from Kumaon District to Bhutan. It is not
known from Himachal Pradesh or Kashmir. Bell (1977) gives a list of localities.
Yamatosa draco (Bell 1977) NEW COMBINATION
(Fig. 76,77)
Yamatoa draco Bell 1977: 155
Type material. - HOLOTYPE male, labelled; “BHUTAN: Tango, 12 kilometers north of Thimphu,
2500-2900 m. June 30, 1972, coll. Basel Natural History Museum Expedition” (BSL).
Description. — Length 6.7 mm. Antennal Segment XI obtuse; stylet absent; head cordate, sides evenly rounded, widest point
opposite middle of temporal lobe, just posterior to eyes; anterior tentorial pits small, oblique; frontal grooves very narrow, clearly
defined; eye reduced (Fig. 77), deeper than long, entirely anterior to middle of temporal lobe, with about 100 ommatidia; cornea
distinctly facetted; mentum nearly smooth, without beard; one pair of postlabial setae.
Pronotum relatively short for genus, length/greatest width 1.33; base distinctly narrowed; apex more strongly narrowed; discal
striole ended anteriorly at basal third of pronotum; marginal groove nearly complete, effaced only near anterior end of pronotum;
precoxal carina absent; precoxal punctures absent.
Elytra relatively short, broad for genus; humeral tubercle small; strial punctures small but deep; Striae II, HI slightly abbreviated
at base; Stria V represented only by row of punctures in anterior half; slightly impressed in posterior half; Stria V with origin slightly
posterior to humerus; Stria VI only slightly abbreviated anteriorly; Striae II and IV each with one seta near apex; several setae also in
apex of Stria VII; status of hind wings not investigated.
Male with metasternum broadly concave, abdominal sterna not concave; Sternum IV with deep lateral pits; Sternum V with
shallow ones; lateral pits suggested on Sternum 111.
Anterior femur of male with ventral tooth (female unknown); anterior tibia of male without medial groove or swelling; spurs of
middle tibiae unequal, anterior ones smaller; spurs of hind tibiae equal; calcar small, obtusely angled.
This species is closely related to the allopatric Y. boysi, with which it agrees in having reduced eyes,
oblique anterior tentorial pits, and setae in elytral Striae II and IV. This species has the eyes much less
reduced and the shape of the head less modified than in Y. boysi. Size and shape are similar to those of
Quaest. Ent., 1979, 15 (4)
430
Bell and Bell
the sympatric Y. reitteri, but the latter species has reduced marginal grooves on the pronotum, and eyes
which are not reduced.
Yamatosa boy si (Arrow 1901) NEW COMBINATION
(Fig. 84, 85)
Rhysodes boy si Arrow 1901: 87.
Rhysodes kaschmirensis Reitter 1922: 57.
Yamatoa boy si Bell and Bell 1978.
Type material. - R. boysv. LECTOTYPE (here designated) female, labelled: “INDIA: 1901-58, coll.
Capt. Boys” (BMNH). PARALECTOTYPE: the original description states that there were two females
in the type series. We were unable to locate the second female. If it still exists, it constitutes a
paralectotype.
R. kaschmirensis: A series of four specimens in Paris (MMHN) are labelled: “Kaschmir, Reitter”.
One of these bears a determination in the same handwriting “R. kaschmirensis Reitter”. It is our belief
that these specimens are cotypes.
Description. — Length 6. 0-7. 2 mm. Antennal Segment XI obtuse; stylet absent; head inflated, distinctly broader than long,
widest point at basal fourth, far behind eyes; anterior tentorial pits small, oblique; frontal grooves narrow, well defined; eye markedly
reduced (Fig. 85), longer than wide, rounded anteriorly, angled posteriorly, extended only 0.33 of distance from antennal base to rear
of temporal lobe, only 0.25 of depth of head; eye flat, cornea not facetted; about 50 ommatidia; mentum nearly smooth, without beard;
one pair of postlabial setae.
Pronotum moderately short for genus, length/greatest width about 1.38; base distinctly narrowed; apex more markedly narrowed;
discal striole ended anteriorly at basal third of pronotum; marginal groove nearly complete, effaced only near anterior end of
pronotum; precoxal carina absent; precoxal punctures absent.
Elytra short and broad for genus; humeral tubercle small; strial punctures coarse; Striae II, III slightly abbreviated at base;
Stria V originating slightly posterior to humerus, its anterior half represented by a row of punctures, its posterior half slightly
impressed; Stria VI only slightly abbreviated anteriorly; Striae II and IV each with apical seta (one or the other missing in some
specimens); several setae also present in apex of Stria VII; hind wings reduced to small vestiges.
Male with metasternum broadly, shallowly concave; abdominal sterna not concave; metasternum of female evenly convex;
Sternite IV with deep lateral pits; Sternite V with shallow ones.
Anterior femur of male with ventral tooth; that of female without tooth; anterior tibia of male without medial groove or swelling;
spurs of middle and posterior tibiae nearly equal; hind calcar of male shaped as in Y. draco, but distinctly larger. The male genitalia
have been figured by Saha, Mukherjee and Sengupta (1978).
Inflated, somewhat wedge-shaped head and strongly reduced eyes give this species a distinctive
appearance. It is the only species of Yamatosa in which the sexes are known to differ in development of
the femoral tooth. (In Y. draco, however, the female is unknown.)
Range. - As presently known, almost entirely to the west of that of Y. reitteri, the two species
occurring together only in Kumaon. The range of Y. boysi extends westward along the Himalaya to the
Indus River in the Pakistani part of Kashmir. We have seen specimens from the following localities:
INDIA; two males, Chakrata Div. U.P., 7000 ft., l-VII-1932, coll. H.G. Champion (BMNH); four males, two females, Chapal,
Himachal Pradesh, 2400-2750 m, 7-5-1977, coll. W. Wittmer, Brancucci (BSL); one male, Dudhatoli, Garhwal, 9000 ft., June 1920,
coll. H.G. Champion (BMNH); one female, Gori Valley, Kumaon, no date, 7000 ft., coll. H.G. Champion (BMNH); one female,
Parbatti V., Kulu, Punjab, 6000-8000 ft., no date, coll. H.G. Champion (BMNH); KASHMIR: one female, Chhangla Gali, 7000 ft.
Hazara, 12-V1-1974, coll. C. Baroni Urban! (BSL); one male, three females, Gulmarg, 16-VI1-31, Fletcher colln. (BMNH); one male,
one female, Yusmarg, 2300-2400 m., 5-7-1976, coll. W. Wittmer (BSL).
In addition, nine other specimens which we have not seen have been reported by Saha, Mukherjee and Sengupta (1978) from
Himachal Pradesh as follows: three males, four females, Bagi, 2518 m., 21-VI-I975; one male, one female, Narkanda, 2725 m.,
20-VI-1975, all collected by T. Sengupta.
Classification of Rhysodini
431
Genus Shyrodes Grouvelle 1903
Type species. - Rhysodes dohertyi Grouvelle 1903.
Description. - Part I, 69. Only one species is known.
Shyrodes dohertyi (Grouvelle 1903)
(Fig. 86)
Rhysodes dohertyi Grouvelle 1903: 126.
Description. - Part I, 69.
Type material. - Part I, 69.
The name of the country of origin, BURMA, was inadvertently omitted from Part I. “Ruby Mines”,
the type locality, refers to the vicinity of Mogok, about 140 km north of Mandalay. A Doherty specimen
in the MCZ collection has additional collecting data of 5000-7000 feet.
Genus Srimara Bell & Bell 1978
Type species. - Srimara planicollis Bell & Bell 1978.
Description. - Part I, 70.
Type material. - Part I, 70. Only one species is known.
Srimara planicollis Bell & Bell 1978
(Fig. 87)
Srimara planicollis Bell & Bell 1978: 70.
Description. - Part I, 70.
Type material. - Part I, 70.
Genus Plesioglymmius Bell and Bell 1978
(Fig. 88-102)
Type species. - Rhysodes elegans Grouvelle 1903.
Description. — Segment IX of antenna varied from sharply pointed to obtuse; minor setae on Segments V-X; basal setae
present or absent; antennal bases more or less displaced dorsomedially; clypeus separated from median lobe by transverse impression,
latter ill-defined or very deep and distinct; or else clypeus and median lobe broadly separated by junction of antennal sclerites in
midline; clypeal setae present or absent; eyes fully developed, with distinct ommatidia.
Pronotum with paramedian grooves incomplete anteriorly, in most species paramedian grooves broad, in form of paramedian
grooves characteristic of Omoglymmius except not extended to reach anterior margin; in subgenus Juxtaglymmius, anterior portions
of paramedian grooves in form of discal strioles as in Yamatosa; marginal grooves fine, complete.
Elytral setae virtually constant within genus; Stria II with one to four in apex; Stria IV with many setae in form of complete series;
one seta on apical tubercle; one to three setae in apical striole; several setae in tip of Stria VII; hind wings long; middle and hind tibiae
each with one spur.
The above description has been modified from the original one (Part I, 70-71) to accommodate
Plesioglymmius (Juxtaglymmius) jugatus. The latter species has the paramedian grooves reduced to
discal strioles, much as in Yamatosa. Nevertheless, loss of inner spurs of the middle and hind tibiae and
tendency for the antennal bases to approach one another seem to indicate that P. jugatus is a modified
Quaest. Ent., 1979, 15 (4)
432
Bell and Bell
Plesioglymmius and not a relative of Yamatosa.
Plesioglymmius is found in two widely disjunct areas: the Greater Sunda Islands (Sumatra, Borneo,
Java and Mindanao), and Latin America (Brazil, Venezuela, and perhaps Cuba). The three subgenera
are clearly distinct, and there is no decisive evidence to indicate that any two of them are more closely
related than either is to the third.
Although there are six distinct species, the genus is known from less than 20 specimens. This makes it
excessively rare, even among Rhysodini, and suggests that it is limited to some special and inaccessible
habitat.
KEY TO SUBGENERA
1 Antennal sclerites separated from one another (Fig. 88); paramedian groove not linear,
extending about 90% of distance from base to apex of pronotum 2
L Antennal sclerites joined as median suture between clypeus and median lobe, latter thus
separated (Fig. 89); paramedian groove linear, in form of discal striole, ended near
middle of pronotum Juxtaglymmius new subgenus, p. 437
2 (1) Paramedian grooves coarsely punctate, glabrous except for pollinosity in punctures;
medial angles of temporal lobes distinct; median lobe of head rhomboid
Ameroglymmius new subgenus, p. 435
1' Paramedian grooves entirely pilose, without visible punctures; medial margin of temporal
lobe truncate, without distinct medial angle; median head lobe oval
Plesioglymmius (sensu stricto) Bell and Bell, p. 432
Subgenus Plesioglymmius {sensu stricto) Bell and Bell 1978
Type species. - Rhysodes elegans Grouvelle 1903
Description. — Antennal Segment XI longer than wide; stylet short; Segments I, II pollinose dorsally; Segments 1 1 1-1 V each
with narrow subapical dorsal pollinose band; basal setae present on Segments V-X.
Clypeus concave, with pair of setae; median lobe of head oval, isolated from clypeus by deep transverse pilose impression, latter
bounded anteriorly by raised, pollinose posterior margin of clypeus; antennal sclerites widely separated from one another; medial
margin of temporal lobes truncate, the two temporal lobes thus separated by narrow linear space; one or more coarse punctures near
medial margin of each temporal lobe; orbital groove represented by strip of pollinosity; temporal setae present or absent; postorbit
evenly convex; postorbital tubercle absent; suborbital tubercles prominent, in form of posterior ends of distinct genal ridges; much of
ventral surface of head, including genal ridges pilose; two to four pairs of postlabial setae.
Paramedian grooves deep, straight, about 95% as long as pronotum; both median and paramedian grooves coarsely punctate, but
with punctures almost completely concealed by coarse pilosity; pleural regions impunctate.
Base of elytron opposite Interval I extended anteriorly in form of angulate tooth; elytral striae deep; strial punctures coarse; apical
depression of elytron including tips of Striae I, IV; medial margin of apical tubercle distinctly sinuate.
The truncate medial margin of the temporal lobe is unique to this subgenus. It also differs from other
subgenera of Plesioglymmius in having the paramedian grooves straight, and entirely pollinose.
The subgenus Plesioglymmius (sensu stricto) is known from Sumatra, Borneo, and Mindanao.
KEY TO SPECIES
1
Temporal setae absent; orbital groove represented by narrow strip of pollinosity; postorbit
without a seta Plesioglymmius elegans (Grouvelle), p. 434
433
Classification of Rhysodini
Figures 88 - 102. Genus Plesioglymmius. Fig. 88,89, Flead, anterior aspect, semi-diagrammatic, antennal segments, black;
antennal sclerites, stippled; Fig. 88, P. (Plesioglymmius} silus new species; Fig. 89, P. (Juxtaglymmius) jugatus new species;
Fig. 90, Head and pronotum, dorsal aspect, P. (P.) silus, new species; Fig. 91-93, P. (P.) elegans (Grouvelle); Fig. 91, Head and
pronotum, dorsal aspect; Fig. 92, Metasternum and abdomen, ventral aspect, male; Fig. 93, Head, male, lateral aspect;
Fig. 94-98, Head lateral aspect; Fig. 94, P. (P.) silus new species; Fig. 95, P. (Ameroglymmius) meridionalis (Grouvelle);
Fig. 96, P. (J.) jugatus new species Fig. 97, P. (A.) reichardti new species; Fig. 98, P. (A.) compactus new species; Fig. 99-102,
Head and pronotum, dorsal aspect; Fig. 99, P. (AJ meridionalis (Grouvelle); Fig. 100, P. (A.) reichardti new species; Fig. 101,
P. (J.) jugatus new species; Fig. 102, P. (A.) compactus new species.
Quaest. Ent., 1979, 15 (4)
434
Bell and Bell
r Four to five temporal setae present; orbital groove represented by broad strip of
pollinosity; postorbit with a seta Plesioglymmius situs new species, p. 434
Plesioglymmius (sensu stricto) elegans (Grouvelle 1903)
(Fig. 91-93)
Rhysodes elegans Grouvelle 1903: 117-118
Type material. - LECTOTYPE (here designated) male, labelled; “SUMATRA, Palembang”
(MNHN). Three PARALECTOTYPES: two males, one female, same data as lectotype (MNHN).
Description. — Length 4. 3-5. 5 mm. Head slightly longer than broad; antennal stylet short, acute; clypeus less constricted
posteriorly than in P. situs, its base only moderately raised; temporal lobe with one to two punctures near medial margin; temporal
setae absent; orbital groove represented by very narrow pollinose strip, latter with small tufts of pollinosity; postorbit pollinose,
impunctate; postorbital seta absent (Fig. 93); mentum entirely pollinose except anterior margin in male, pollinosity limited to medial
area of mentum and genae in female; one to two pairs of postlabial setae.
Pronotum elongate, length/greatest width about 1.45; pronotum hexagonal, sides straight, nearly parallel, apex and base both
abruptly narrow, each forming a distinct angle with remainder of lateral margin; median groove narrow, linear near middle;
paramedian grooves relatively narrow; pronotal carinae impunctate; in male, prosternum pilose except for small area anterior to each
coxa; in female, prosternum finely pollinose in anterior fourth, remainder coarsely punctate.
Each elytral stria with fine strip of pollinosity connecting punctures; metasternum of male with median pilose area extended from
slightly posterior to middle coxal cavities to posterior margin; sides of metasternum coarsely punctate; in female, metasternum
glabrous, median area with few scattered punctures, lateral regions coarsely punctate; pollinosity of male extended onto medial portion
of hind coxae and abdominal Sterna 1, II, ending abruptly at anterior transverse row of punctures on Sternum 111 (Fig. 92); female
without pilosity on hind coxae or abdominal Sterna I, 11; in both sexes. Sterna 11-V each with two transverse rows of punctures, those
of anterior row very coarse, surrounded by pollinosity, more or less coalescent into transverse pollinose strip; punctures of posterior row
much finer, not surrounded by pollinosity; both sexes with distinct lateral pits in Sternum IV; male with ventral tooth on anterior
femur; middle and hind calcars both small, acute.
Absence of temporal setae, and hexagonal form of the pronotum are among several striking
differences between this species and P. situs.
Range. - Sumatra and Borneo. In addition to the type material, we have seen one male, labelled:
“BORNEO: S. East; German Mission 59773, Fry Collection 1905. 100” (BMNH).
Plesioglymmius {sensu stricto) situs new species
(Fig. 88,90,94)
Type material. - HOLOTYPE female, labelled; “PHILIPPINES: Kabasalan, Zamboanga,
Mindanao, VIII- 1932, coll. H.G. Muzzell” (CAS). PARATYPES two females, same data as type
(CAS).
Description. — Length 6. 0-6. 9 mm. Head shorter and broader than in P. elegans, clearly broader than long; antennal stylet
short, blunt; clypeus more markedly constricted at base than in P. elegans, its base strongly raised; temporal lobe with cluster of two to
five coarse punctures near medial margin; four to five prominent temporal setae present, inserted within broad strip of pollinosity
which represents orbital groove; postorbit finely pollinose, with distinct coarse punctures; postorbit with one prominent seta opposite
upper fourth of eye (Fig. 94); mentum largely glabrous, contrasting with pollinose genal ridges; four pairs of postlabial setae.
Pronotum shorter than in P. elegans, length/greatest width is 1.36; widest point anterior to middle, sides curved; apex markedly
narrowed; base moderately narrowed; median groove broader than in P. elegans; paramedian grooves slightly broader than in
P. elegans; outer carina with few coarse punctures near lateral margin, and many very fine, scattered punctures; prosternum of female
extensively pollinose anteriorly, pollinosity extended to coxal cavities laterally; posterior half of prosternum glabrous, coarsely punctate
(male unknown).
Elytra with strial punctures not connected by pollinosity; metasternum of female entirely coarsely punctate, not pollinose (male
unknown); abdominal Sterna 1 11-V each with two transverse rows of punctures; anterior rows coarse, surrounded by pollinosity, latter
more or less coalescent in form of transverse pollinose bands; posterior rows fine, without pollinosity; female with distinct lateral pits in
abdominal Sternum IV (male unknown).
Classification of Rhysodini
435
This species has a shorter, broader head than does P. elegans, and is easily recognized by the presence
of temporal setae.
Subgenus Ameroglymmius new subgenus
Type species. - Rhysodes meridionalis Grouvelle 1903
Description. — Antennal Segment Xi either elongated, conical or shorter, obtuse; basal setae present on Segments V-X, or
limited to Segments Vlll-X, or entirely absent; Segment I pollinose dorsally; remaining segments either with small pollinose spots at
bases of basal setae, of else entirely without pollinosity; clypeus flat, without setae; median lobe of head rhomboidal; transverse groove
between clypeus and median lobe either shallow and indefinite or deep and distinct; antennal sclerites widely separated; medial margin
of temporal lobe angulate, the medial angles contiguous or narrowly separated; orbital groove and temporal setae absent; postorbital
tubercle present; suborbital tubercle and genal ridges absent; lower surface of head glabrous except for gular grooves; one pair of
postlabial setae.
Paramedian grooves deep, broad, gradually tapered anteriorly, extended about 95% of length of pronotum; paramedian grooves at
least slightly sinuate, their anterior ends slightly divergent; paramedian grooves coarsely punctate, without pollinosity except within
punctures.
Base of elytron opposite Interval 1 either in form of rounded prominence or else not projected; elytral striae shallow or not
impressed; strial punctures coarse; no pollinosity between punctures; apical depression limited to apices of Striae I and 11; apical
tubercle not sinuate medially; metasternum coarsely punctate; abdominal punctures not in rows, but irregularly distributed; lateral
pits on Sterum IV of female, absent from male.
The coarsely punctate paramedian grooves and the rhomboidal median lobe are diagnostic of this
subgenus. It is found in eastern South America, and possibly in Cuba.
Phylogeny. - P. meridionalis stands apart from the other two species in having a relatively long head,
lateral eyes, antennal bases far apart, and frontal grooves relatively broad. In all these features, it is the
least specialized species of Plesioglymmius. P. reichardti and P. compact us have the head shortened,
eyes directed anterolaterally, frontal grooves linear, and antennal bases relatively close together;
antennal apex obtuse, and basal setae of the antenna reduced.
KEY TO SPECIES
1 Antennal Segment XI acutely pointed; frontal grooves relatively broad; outer carina of
pronotum coarsely punctate Plesioglymmius meridionalis (Grouvelle), p. 435
V Antennal Segment XI obtuse; frontal grooves linear; outer carina impunctate or with few
small punctures 2
2 (1) Median lobe truncate posteriorly at level of anterior margin of eye; marginal setae
absent; anterior and posterior median pits of pronotum deep, expanded
Plesioglymmius reichardti new species, p. 436
2' Median lobe obtusely pointed posteriorly at level of posterior margin of eye; marginal
setae present; anterior and posterior median pits of pronotum shallow
Plesioglymmius compact us new species, p. 437
Plesioglymmius (Ameroglymmius) meridionalis (Grouvelle) NEW COMBINATION
(Fig. 95, 99)
Rhysodes meridionalis GvomwqWq 1903: 108-109
Quaest. Ent., 1979, 15 (4)
436
Bell and Bell
Type material. - LECTOTYPE (here designated) male, labelled: “Brasilia” (MNHN).
PARALECTOTYPES one male, labelled: “Brazilia, co-type. Fry Coll. 1905-100” (BMNH); one
female, labelled: “Rio San°, co-type. Fry. Coll. 1905-100” (BMNH).
Description. — Length 5. 2-6. 7 mm. Antennal Segment XI longer than wide, its apex bluntly pointed, suggesting vestigial
stylet; basal setae on Segments V-X; Segment 1 pollinose dorsally with deeper pilose punctures; Segments Il-Vi with shallow pollinose
punctures.
Head slightly longer than broad; eyes completely lateral; antennal sclerites relatively widely separated; transverse groove between
clypeus and median lobe shallow, incompletely pollinose, in some specimens apparently interrupted in midline; median lobe small,
obtuse to nearly acute; median lobe with apex opposite middle of eye; frontal grooves broad; medial angles obtuse, slightly separated;
temporal lobes coarsely, sparsely, irregularly punctate; postorbit with two irregular vertical rows of coarse punctures posterior to eye,
very finely pollinose posterior to the punctures; postorbital tubercle distinct, but postorbit very shallowly emarginate dorsal to it
(Fig. 95); mentum coarsely punctate, pollinosity of gular grooves incomplete.
Pronotum narrow, rather elongate, widest at posterior third; sides curved, base distinctly narrowed, apex more strongly so;
paramedian groove coarsely punctate; marginal groove more dilated than in related species; marginal setae absent; inner carina
impunctate; outer carina coarsely, irregularly punctate; epipleura and prosternum coarsely punctate.
Basal scarp of elytron transverse except for rounded prominence opposite base of Stria i; both sexes with prominent ventral tooth
on anterior femur, and obtuse one on hind femur. Male with middle calcar acute, triangular; hind calcar obtuse.
The coarsely punctate outer carinae and temporal lobes distinguish this species. It can be separated
from P. reichardti also by the pointed median lobe and the more elongate head.
Distribution. - Southern Brazil. In addition to the type material, we have seen the following
specimens: one male, one female, Cantaneira (San Paulo State), 20-M-1958, coll. K. Lenko (MZSP); one female. Nova Teutonia
(Santa Catarina State), XI-1940, 300-500 m., coll. Fritz Plaumann (MZSP); one female, Brazil, Parana, 1901-31, coll. G. Lewis
(BMNH); one female, Represa do Cabeca, Corcovado (Rio, D.F.), 27-7-1967, coll. Wygodzinsky (MZSP).
Vulcano and Pereira (1975a) also list this species from Agua Funda, Alto da Serra and Parana,
Curitiba. We have not studied the specimens from these localities. Their record from Suapure,
Venezuela actually pertains to P. reichardti. These specimens of P. meridionalis in the Carnegie
Museum (CMP) are labelled: “Denver, Colorado, H. Klages coll.” C.M. acc. 11414. According to
G. Wallace, E.A. Klages, a brother of H. Klages, collected extensively in South America, and these are
no doubt mislabelled Brazilian specimens.
Plesioglymmius (Ameroglymmius) reichardti new species
(Fig. 97,100)
Type material. - HOLOTYPE male, labelled: “VENEZUELA: Suapure, Caura R., Apr. 25, 1899,
coll. E.A. Klages” (MZSP). PARATYPE female, labelled: “BRAZIL: Rio Madeira, Mann & Baker,
W.M. Mann Coll. 1954” (MZSP).
Description. — Length 5. 0-5. 3 mm. Antennal Segment XI somewhat compressed, longer than wide, obtusely rounded at apex,
stylet absent; basal setae absent from Segments V-VIl, but present on Vlll-X; antenna without pollinosity except for dorsal spot on
Segment 1.
Head wider than long; eyes directed slightly anteriorly and dorsally; antennal sclerites relatively narrowly separated; clypeus and
median lobe separated by a deep, pollinose transverse groove; median lobe relatively short, broad, pentagonal, abruptly truncate at
level of anterior margin of eye; frontal grooves linear; medial angles narrowly separated; temporal lobes apparently impunctate except
at high magnification; postorbit pollinose, impunctate; postorbital tubercle distinct (Fig. 97); gular grooves occupied by broad,
continuous bands of pollinosity.
Pronotum narrow, rather elongate, widest at posterior third, sides slightly curved to base, latter slightly narrowed, markedly
curved to apex, latter markedly narrowed; paramedian grooves coarsely, rather densely punctate; basal impressions with rather long
pilosity; marginal groove fine, shallow; marginal setae absent; both inner and outer carinae entirely impunctate; epipleura and
propleuron impunctate; prosternum coarsely punctate.
Basal scarp of elytron transverse except for rounded prominence opposite Stria I; male with ventral tooth on anterior and posterior
femora; female without femoral teeth; calcar of middle tibia acute; hind calcar small, narrow, its tip slightly upturned.
The markedly truncate median lobe is the recognition mark of P. reichardti. It also differs
conspicuously from P. meridionalis in absence of punctures from the outer pronotal carinae and
temporal lobes. This species is dedicated to the memory of Dr. Hans Reichardt, whose untimely death in
Classification of Rhysodini
437
1976 was a great loss for the science of entomology. This species appears to be restricted to the Orinoco
and Amazon Basins, and to be allopatric to P. meridionalis.
Plesioglymmius (Ameroglymmius) compactus new species
(Fig. 98,102)
Type material - HOLOTYPE female, labelled: “CUBA” (MCZ).
Description. — Length 4.8 mm. Antennal Segment XI slightly longer than wide, cone tapered, its tip obtusely rounded, stylet
absent; basal setae absent; Segment I pollinose above; remaining segments devoid of pollinosity.
Head wider than long; eyes directed slightly dorsally and anteriorly; antennal sclerites relatively close together; clypeus and
median lobe separated by deep, pollinose transverse groove; clypeal setae absent; antennal sclerites relatively close together, median
lobe rhomboidal, its tip obtusely pointed at level of posterior margin of eye; temporal lobes short, rather inflated; frontal grooves
sublinear; medial angles obtusely rounded, contiguous; temporal lobes impunctate; postorbit glabrous, convex; postorbital tubercle
minute, pollinose (Fig. 98); mentum with few median punctures; gular grooves occupied by broad, continuous bands of pollinosity.
Pronotum elongate, narrow, broadest clearly anterior to middle; sides almost straight, slightly convergent posteriorly; base and
apex distinctly narrowed; paramedian grooves relatively narrowed; the punctures of each groove in form of a single irregular line;
marginal groove fine, shallow; marginal setae present (some setae missing on holotype, but the punctures indicate two setae near apex
and one near base on each side); both inner and outer carina impunctate; prosternum with broad, irregular band of coarse punctures
on either side of midline, otherwise impunctate; proepipleura and propleuron both impunctate.
Elytral humerus narrow, triangular; basal scarp of elytron angled opposite base of Stria IV; no prominence opposite base of
Stria I; striae not impressed, represented by rows of fine punctures; Striae II, III slightly abbreviated at base; female with distinct
lateral pit on abdominal Sternum IV; male unknown.
This species is the only member of the genus to have marginal setae on the pronotum. The
arrangement of the punctures of the paramedian groove in a single line is also unique. The holotype has
the tip of one wing protruding between the elytra, as is often seen in Carabidae collected at lights. At
first glance, the eyes of this species appear reduced, but this is an optical illusion caused by the
shortening and inflation of the temporal lobes.
According to Dr. P.J. Darlington, Jr. (personal communication), this specimen was from the Blaisdell
collection which contains many erroneous locality records. So perhaps, the specimen is not really from
Cuba. If so, it would probably prove to be a South American species. Bell (1970) discussed but did not
name this species.
Subgenus Juxtaglymmius new subgenus
Type species. - Plesioglymmius (Juxtaglymmius) jugatus new species
Description. — Antennal Segment XI longer than wide, acutely pointed in form of vestigial stylet or none; basal setae sparse,
on Segments V-X; Segment I pollinose dorsally; Segment II with basal pollinose band, remaining segments without pollinosity;
antennal sclerites meeting in contact in form of median suture, clypeus thus widely separated from median lobe; clypeus flat; clypeal
setae absent; median lobe small, oval, markedly convex; medial angles of temporal lobes obtuse, narrowly separated; orbital groove and
temporal setae absent; postorbital tubercle prominent, separated from temporal lobe by deep notch; suborbital tubercles, genal ridges
absent; lower surface of head nearly glabrous; nearly impunctate; one pair of postlabial setae.
Paramedian groove represented by basal impression and short discal striole, latter extended approximately to middle of pronotum,
and without pollinosity or punctures.
Basal scarp of elytron transverse, without prominence opposite base of Stria I; striae shallow, finely punctured; pollinosity limited
to punctures; apical depression small, limited to apices of Striae I, II; apical tubercle not sinuate medially; metasternum with one
median and two lateral longitudinal bands of punctures, otherwise impunctate; punctures of abdominal Sterna lll-V in form of two
irregular transverse rows (but less regular in arrangement than in Plesioglymmius (sensu stricto)): posterior row of each sternum as
coarse as anterior row; female with lateral pits on Sterum IV and V, V slightly deeper; male with slight suggestion of these pits.
Male with ventral tooth on anterior femur, but without one on posterior femur; female without ventral tooth on either femur.
Only one species is known.
Quaest. Ent., 1979, 15 (4)
438
Bell and Bell
Plesioglymmius (Juxtaglymmius) jugatus new species
(Fig. 89,96,101)
Type material. - HOLOTYPE male, labelled: “JAVA” (MNHN). PARATYPE female, same data
as type (MNHN).
Description. — Length 5. 6-6.0 mm. As described for subgenus; head slightly wider than long; clypeus longer than wide, its
sides parallel; eyes directed anterolaterally; temporal lobe rounded posteriorly and laterally; postorbital tubercle and posterior margin
of temporal lobe pilose, notch between them glabrous (Fig. 96).
Pronotum relatively short, widest at middle, sides strongly curved to base and apex; disc of pronotum impunctate; marginal groove
fine; prosternum, propleuron, and epipleura impunctate; pollinosity of prosternum limited to anterior margin and intercoxal pit.
Calcar of middle tibia small, acute; that of hind tibia broadly triangular, angle acute, well above level of tibial spur.
The tendency for the antennal bases to approach one another is seen throughout the genus. In this
species it reaches its limit, with the antennal sclerites actually meeting in the midline. The bases of the
antennae are separated by only slightly more than the width of one basal condyle so that it is difficult to
see how they avoid interfering with one another.
This singular species has as much in common with Ameroglymmius of Brazil as it does with
Plesioglymmius (sensu stricto) from nearby islands, and it may prove that the locality labels are
erroneous.
Genus Arrowina Bell and Bell 1978
(Fig. 103-114)
Type species. - Rhysodes taprobanae Fairmaire 1873
Description. - Part 1,71.
Phylogeny. - With the exception of A. taprobanae and A. pygmaea, the members of this genus are all
very different from one another. The most aberrant species is the Japanese A. rostrata, the only species
which has an elongate rostrum and a femoral tooth. All the remaining species are confined to southern
India and Ceylon. Although A. anguliceps and A. nilgiriensis are very different from one another, they
share some common characters, such as a dilated orbital groove, that suggest that they are more closely
related to one another than either is to A. taprobanae and A. pygmaea.
KEY TO SPECIES
1 Head almost twice as long as wide; anterior femur of male with ventral tooth (female
unknown) Arrowina rostrata (Lewis), p.
V Head only slightly longer than wide; anterior femur of male without ventral tooth
2 (1 ) Orbital groove absent; lateral margin of inner pronotal carina sloped gradually into
paramedian groove
T Orbital groove complete, somewhat dilated; lateral margin of inner pronotal carina
vertical, sharply defined
3 (2 ) Metasternum with punctures confined to row along each margin; length 5.0 mm or more .
Arrowina taprobanae (Fairmaire), p.
y Metasternum with punctures in middle of disc as well as along margins; length 4.3 mm or
less Arrowina pygmaea new species, p.
4 (2') Eyes large, deeper than long; posteromedial margin of temporal lobe sinuate posterior to
medial angles Arrowina nilgiriensis (Arrow), p.
439
2
3
4
439
441
441
Classification of Rhysodini
439
4' Eyes reduced, longer than deep; posteromedial margin of temporal lobe not emarginate....
Arrowina anguliceps (Arrow), p. 442
Arrowina rostrata (Lewis 1888)
(Fig. 108,109)
Rhysodes rostratus Lewis 1888: 81.
Type material. - LECTOTYPE (here designated) male, labelled: “JAPAN: Oyayama, Higo
District, G. Lewis, 1910-320 (Kyushu Island)” (BMNH). PARALECTOTYPE: According to the
original description, there were two specimens, both males. The second specimen is not in the BMNH. If
it is still extant, it is a paralectotype.
Description. — Length 5.5 mm. Antennal Segment XI short, almost spherical; stylet short, acuminate, larger than in other
members of genus; head almost twice as long as wide, with rostrum distinctly elongate; postclypeal grooves sublinear; median lobe
relatively broad, its posterior tip obtusely pointed, projected slightly above frontal pit; antennal groove fine, nearly linear; medial
angles obtuse, distinctly separated; posteromedial margin emarginate between medial and occipital angles; orbital grooves obsolete;
eye large, deeper than long; postorbital tubercle entirely absent (Fig. 109).
Pronotum narrow, elongate, length/greatest width about 1.40; widest anterior to middle; sides slightly curved, slightly convergent
from widest point to base; markedly curved from widest point to apex; median groove of nearly even width, slightly expanded at
posterior median pit, latter partly divided by constriction; paramedian grooves narrow, much narrower than inner carinae, and only
slightly wider than median groove; lateral margin of inner carina vertical, sharply defined; marginal groove fine; prosternum and
propleura impunctate.
Elytra elongate, lateral margins parallel in middle third; humeral tubercle well developed; elytral striae very shallowly impressed;
strial punctures much finer than in other members of genus; elytral setae restricted to apex of Stria VII; metasternum impunctate;
male with acute ventral tooth on anterior femur (female unknown); spurs of middle and hind tibiae nearly equal; calcar of hind tibia
obtuse, raised slightly above level of tibial spurs.
The elongate rostrum, very narrow paramedian grooves, and absence of metasternal punctures are all
diagnostic of this species.
Arrowina taprobanae (Fairmaire 1873)
(Fig. 110-112)
Rhyzodes taprobanae Fairmaire 1873; 389.
Type material. - HOLOTYPE female, labelled: “CEYLON” (MNHN).
Description. — Length 5.0-6. 2 mm. Antennal Segment XI slightly longer than wide; stylet very small but evident; head about
1.20 longer than wide; rostrum not elongate; postclypeal grooves dilated; median lobe otusely pointed posteriorly, its tip barely
overhanging frontal pit; antennal groove narrow; medial angles nearly rectangular, almost contiguous, each angle slightly closer to
occiput than to tip of median lobe; posteromedial margin very shallowly emarginate between medial and occipital angles; orbital
groove absent; postorbital tubercle absent (Fig. 111).
Pronotum slightly longer than wide, length/greatest width 1.17; widest at middle, sides curved, apex markedly narrowed, base
moderately so; inner carina wider than outer one; lateral margin of inner carina ill-defined, sloping gradually into paramedian groove;
pronotum with one to two precoxal punctures on each side.
Elytra with lateral margins parallel in middle third; humeral tubercle well-developed, visible in dorsal view; elytral striae shallow,
intervals nearly flat; strial punctures of inner striae moderate, those of outer striae coarse; basal puncture of Stria I enlarged; elytron
with one seta in apex of Stria IV, one in apical striole, and several in apex of Stria Vll; metasternum with row of coarse punctures
along each margin, but without discal punctures (Fig. 1 1 2); female with deep lateral pits on abdominal Sternum V.
Middle and hind tibiae with spurs nearly equal, those of female much larger than those of male; ventral tooth absent from anterior
femur; male with calcars well developed; hind calcar obtuse, distinctly raised above level of spurs.
This species and A. pygmaea differ from other members of the genus in having the lateral margin of
the inner carina slope gradually into the paramedian groove. In all other species of Arrowina, the carina
and the groove are separated by a sharply-defined, nearly vertical surface. Absence of discal punctures
on the metasternum separates this species from A. pygmaea.
Quaest. Ent., 1979, 15 (4)
440
Bell and Bell
Figures 103 - 114. Genus Arrowina. Fig. 103-105, A. anguliceps (Arrow); Fig. 103, Habitus, dorsal aspect; Fig. 104, Head and
pronotum, dorsal aspect; Fig. 105, Head, lateral aspect; Fig. 106,107, A. nilgiriensis (Arrow); Fig. 106, Head and pronotum,
dorsal aspect; Fig. 107, Head, lateral aspect; Fig. 108,109, A. rostrata (Arrow); Fig. 108, Head and pronotum, dorsal aspect;
Fig. 109, Head, lateral aspect; Fig. 110-112, A. taprobanae (Fairmaire); Fig. 1 10, Head and pronotum, dorsal aspect; Fig. Ill,
Head, lateral aspect; Fig. 112, Metasternum, metacoxae; Fig. 113,114, A. pygmaea new species; Fig. 113, Head and pronotum,
dorsal aspect; Fig. 1 14, Metasternum, metacoxae.
Classification of Rhysodini
441
Distribution. - Known with certainty only from Ceylon. Many specimens in MNHN, BMNH, AMS
are labelled only with the name of the country. The only specimens we have seen with specific locality
data are as follows: Three females, Dikoya, 3800-4200 feet; 6-XII-81 to 16-1-82, coll. G. Lewis
(BMNH); one male, Maturata, no date, coll. Simon (MNHN); one female, Pundalaya, IV, 1888, coll.
H.P. Green (BMNH). One male (probably from southern India rather than Ceylon), “Madura, Ind.
Or.” from Coding, 1935 (NMNH). This almost certainly means the District of Madura (also spelled
“Madurai”), and not the island of the same name near Java. The location is in the Tamil Nadu State,
and is well to the east of the known ranges of other south Indian Arrowina. A specimen in MNHN,
labelled: “Sydney, Austral”, surely has incorrect locality data. A record for Formosa by Miwa (1931), is
probably a misidentification.
Arrowina pygmaea new species
(Fig. 113,114)
Type material. - HOLOTYPE male, labelled: “CEYLON: Dikoya, coll. G. Lewis, 1910-320 (under
label, 17-12-81)” (BMNH). Three PARATYPES one male, labelled: “CEYLON, Nuwara Eliya,
6234-8000 ft., 8-11-II-82, coll. G. Lewis 1910-320” (BMNH); one female, labelled; “CEYLON, coll.
G. Lewis (no date). Sharp Colin. 1905-313” (BMNH); one male labelled: “CEYLON” (MNHN).
Description. — Length 4. 0-4.4 mm. Very similar to A. taprobanae, with appearance of dwarf form of latter, but differing in
following: head and pronotum relatively longer; elytra proportionally shorter; frontal grooves narrower, more linear; median lobe of
head longer, its tip acute, clearly overhanging frontal pit; median angles of temporal lobes obtuse, distinctly separated; posterolateral
end of antennal groove expanded, in form of rudimentary orbital groove; pronotum narrower and more elongate, sightly more
narrowed at base; hind angle more nearly rectangular; elytral striae deeper, intervals slightly convex; elytral setae more numerous;
three setae in Stria IV (one at base, one in middle, one at apex); one seta at apex of Stria VI; one on apical striole; several in apex of
Stria Vll; metasternum with coarse marginal punctures and several coarse punctures in middle of disc (in most specimens in form of
median line (Fig. 1 14); in one specimen limited to group near anterior margin and few scattered on disc).
The coarse discal punctures on the metasternum and the much smaller size separate this species from
A. taprobanae.
Arrowina nilgiriensis (Arrow 1942)
(Fig. 106, 107)
Rhysodes nilgiriensis Know 1942: 179.
Type material. - LECTOTYPE male, labelled: “INDIA: Nilgiri Hills, coll. Andrewes” (BMNH).
Three PARALECTOTYPES two females (on same pin), same data as lectotype (BMNH); one female,
same data as lectotype (MNHN).
Description. — Length 5.2-6. l mm. Antennal Segment XI as wide as long; stylet minute, blunt, scarcely evident; head 1.25
times longer than wide; rostrum not elongate; postclypeal grooves widely dilated; median lobe obtusely pointed posteriorly,
overhanging frontal pit; antennal groove dilated, short; frontal pit relatively broad; medial angles of temporal lobes nearly rectangular,
contiguous; medial angles much closer to occipital angles than to tip of median lobe; posteromedial margin distinctly emarginate
between medial and occipital margins; margin of temporal lobe oblique between eye and occipital angle, latter prominent; orbital
groove dilated, extended to posterior margin of eye; eye large, deeper than long; postorbital tubercle large, deep, separated from
temporal lobe by broad emargination in lateral view (Fig. 107).
Pronotum relatively short, length/greatest width is 1.12; widest at middle; sides curved; base moderately narrowed; apex markedly
narrowed; paramedian grooves broad, almost as wide as inner or outer carinae; inner carina slightly wider than outer one at middle of
pronotum; both margins of inner carinae sharply defined, separated from adjacent grooves by nearly vertical surfaces; about six
precoxal punctures on each side of prosternum.
Elytra moderately elongate, sides nearly parallel in middle third; humeral tubercle well developed; elytral striae distinctly
impressed; intervals convex; strial punctures coarse, those of inner striae elongate, those of outer striae rounded; basal puncture of
Stria 1 not enlarged; Stria IV with four evenly spaced setae; one seta on medial surface of apical tubercle; about seven setae in apex of
Stria Vll; no setae in apical striole; metasternum with coarse marginal punctures, but without discal punctures; female with distinct
lateral pits in abdominal Sterna IV and V, those of Sternum V deeper than those of Sternum IV; male with vestiges of lateral pits
Quaest. Ent., 1979, 15 (4)
442
Bell and Bell
in IV and V.
Male without ventral tooth on anterior femur; spurs of middle and hind tibiae strikingly unequal, anterior one of each pair less
than 0.5 as long as posterior one; hind calcar very obtuse, raised high above spurs, its upper margin small notch about 0.33 of
distance from spurs to base of tibia.
The prominent occipital angles of the head give this species much the appearance of Omoglymmius
subgenus Orthoglymmius. It differs strikingly from the latter in the presence of two tibial spurs. The
narrow pronotal carinae differentiate it from all Orthoglymmius except from O. carinatus (Grouvelle)
and O. hexagonus Grouvelle. The latter two species differ from A. nilgiriensis in having conspicuously
unequal elytral intervals.
Range. - Southern India, apparently confined to the Cardamum Mountains which form the
boundary between Kerala and Tamil Nadu States. In addition to the type series, known from the
following localities: one female, Chambagaganor, Madura (MMHN); one female, Cinchana, Anamalai
Hills, 1050 m., IV-1957, coll. P.S. Nathan (BPBM), one female Shimbaganor (MNHN).
Arrowina anguliceps (Arrow 1901)
(Fig. 103-105)
Rhysodes anguliceps Arrow 1901: 89.
Type material. - LECTOTYPE (here designated) male, labelled: “S. INDIA, Pascoe Coll. 93-60”
(BMNH). PARALECTOTYPES: one female, same data as male (BMNH). According to the original
description, there are also two specimens in the Hope Department at Oxford. If these specimens still
exist, they are also paralectotypes.
Description. — Length 5. 8-6. 9 mm. Antennal Segment XI longer than wide; stylet minute, scarcely evident; head 1.25 times
longer than wide; rostrum not elongate; postclypeal grooves broadly dilated; median lobe short, its tip anterior to eye. its apex obtuse,
not overhanging frontal pit; antennal groove dilated, short; frontal pit small and narrow; medial angles much closer to tip of median
lobe than to occipital angles; medial angles obtuse, distinctly separated; posteromedial margin oblique between medial and occipital
angles; margin of head rounded between eye and occipital angle; orbital groove dilated, extending beyond posterior margin of eye; its
medial margin indistinctly defined; eye strongly reduced, lying entirely anterior to widest part of head, cornea flat, with only traces of
facetting; eye longer than deep, with about 80 ommatida; postorbital tubercle small, but more dorsal than in A. nilgiriensis, prominent
in dorsal view (Fig. 105).
Pronotum rather elongate, length/greatest width is 1.29; widest point posterior to middle; sides markedly curved, base moderately
narrowed, apex strongly narrowed; paramedian grooves broad, but somewhat narrower than outer carinae; both margins of inner
carinae sharply defined, separated from adjacent grooves by nearly vertical surfaces; inner carina distinctly broader than outer
carinae; marginal groove slightly dilated; about 12 precoxal punctures anterior to each coxal cavity and V-shaped line of punctures
present on anterior part of prosternum.
Elytra short, oval, inflated; sides not parallel; humerous prominent, but humeral tubercle not visible in dorsal view; elytral striae
slightly impressed; strial punctures very coarse, those of inner striae elongate oval, those of outer striae nearly round; punctures wider
than intervals; latter convex; some punctures irregularly fused together, especially in humeral region, and near apical tubercle; basal
puncture of Stria 1 not enlarged, but displaced laterally; Stria IV with up to four setae, when four are present they are evenly spaced,
when setae are reduced, more anterior ones are missing; one seta on apical tubercle; none on apical striole; four to five in apex of
Stria Vll; hind wings reduced to small vestiges; metasternum with coarse marginal punctures, and with few discal ones anterior to
coxae; female with shallow lateral pits in abdominal Sternum IV and deep ones in Sternum V; male with vestiges of pits in both
Sterna IV and V.
Male without ventral tooth on anterior femur; all femora coarsely punctate on dorsal surface; spurs of middle and hind tibiae
strikingly unequal, anterior one of each pair about 0.25 as long as posterior one; hind calcar of male obtusely rounded, its top distinctly
proximad to tibiae, but not so proximad as in A. nilgiriensis. Penis illustrated in Part 1, 68.
This is one of the most distinctive species in the tribe. The elytra are shorter and more convex than in
other Rhysodini, suggesting those of Dyschirius (Scaritini). The reduced eye is unique within Arrowina.,
but is similar to that of Tangaroa pensus.
Range. - The Cardamum Mountains of southern India. The only definite locality record is a series of
12 specimens from Dodabetta Peak, 2475 m, III- 1 1-62, coll. E.S. Ross, D.Q. Cavagnaro (CAS).
Classification of Rhysodini
443
ACKNOWLEDGEMENTS
As in the previous part of this work, we acknowledge our indebtedness to all the curators and
collectors who have helped us in this study. We are especially grateful to Mr. M. Bacchus and
Dr. R.B. Madge at the British Musem of Natural History, London, and Dr. A. Descarpentries,
Mme. A. Bons and Dr. J.J. Menier at the Laboratoire d’Entomologie, Museum National d’Histoire
Naturelle, Paris, all of whom were very helpful during our second visit to Europe in May of 1978. We
are also indebted to Dr. Barry P. Moore for further information about Australian Rhysodini, and to Pere
Andre Larochelle and Mr. Paul Choate for calling additional references to our attention.
As before, we are grateful to Mrs. Joyce Murray for her skillful typing of the final manuscript.
SUPPLEMENTARY REFERENCES ‘
Ahrens, A. 1814. Fauna Insecta Europa VI: 1.
Bell, R.T. and J.R. Bell. 1978. Rhysodini of the world. Part I. A new classification of the tribe, and a
synopsis of Omoglymmius subgenus Nitiglymmius new subgenus. (Coleoptera; Carabidae or
Rhysodidae). Quaestiones Entomologicae 14(1): 43-88.
Buckland, P.C. and H.K. Kenward. 1972. Thorne Moor: a palaeo-ecological study of a bronze age site.
Nature 241: 405-406.
Kelley, M. and P.J. Osborne. 1965. Two faunas and floras from the alluvium at Shustoke,
Warwickshire. Proceedings of the Linnean Society of London 176(1): 37-65.
Miwa, Y. 1931. A systematic catalogue of Formosan Coleoptera. Reprinted from Report 55.
Department of Agriculture, Government Research Institute, Taihoku, Formosa. Contribution 32: 21.
Saha, S.K., Mukherjee, A.K. and T. Sengupta. 1978. On the Family Rhysodidae (Coleoptera:
Adephaga) with description of male Rhysodes boysi Arrow from Himachal Pradesh, India. Oriental
Insects 12(1): 115-118.
Trella, T. 1926. Wykaz chrzaszcdw okolic Przemysla. Cicindelidae, Carabidae, Rhysodidae. Polski
Pismo Entomologiczne, Lwow 5: 68-73.
Waga, A. 1841. Rhyzodes europaeus aux environs de Varsovie. Annales de la Societe Entomologique de
France 10: 5.
‘This supplements the complete list of references found in Part I.
Quaest. Ent., 1979, 15 (4)
444
Bell and Bell
Index to Names of Taxa
TRIBE AND SUBTRIBES
Clinidiina, 408
Dhysorina, 381
Leoglymmiina, 380
Omoglymmiina, 423
Rhysodina, 386
Rhysodini, 387, 389, 390, 395, 416, 442
GENERA AND SUBGENERA
Ameroglymmius new subgenus, 377, 432, 435
Angekiva new subgenus, 377, 390
Arrowina Bell and Bell, 377, 438
Clinidium Kirby, 379
Dhysores Grouvelle, 377, 382
Dyschirius (Scaritini), 442
Epiglymmius Lewis, 386
Grouvellina Bell and Bell, 377, 408
Ingevaka new subgenus, 377, 390, 394
Juxtaglymmius new subgenus, 377, 432, 437
Kaveinga (sensu latiore) Bell and Bell, 377,
389
Kaveinga (sensu stricto) Bell and Bell,
390, 394, 395, 397
Kupea Bell and Bell, 377, 388
Leoglymmius Bell and Bell, 377, 380
Neodhysores Bell and Bell, 377, 386
Omoglymmius (Nitiglymmius) , 398
Omoglymmius (sensu latiore}
Ganglbauer, 379
Omoglymmius (sensu stricto), 398
Plesioglymmius (sensu latiore) Bell and
Bell, 377,431
Plesioglymmius (sensu stricto) Bell and
Bell, 432, 438
Pyxiglymmius Bell and Bell, 379
Rhysodes Dalman, 377, 386
Rhyzodiastes Fairmaire, 379
Shyrodes Grouvelle, 377,431
Srimara Bell and Bell, 377,431
Tangaroa Bell and Bell, 377, 381
Vakeinga new subgenus, 377, 390, 395
Xhosores Bell and Bell, 377, 423
Yamatoa Bell, 377
Yamatoa Kiriakoff, 377
Yamatosa Bell and Bell, 377, 424
SPECIES AND SUBSPECIES
abbreviata (Lea), Kaveinga (sensu
stricto), 399, 400
abbreviatus Lea, Rhysodes, 389, 397, 400
anguliceps (Arrow), Arrowina, 439
anguliceps Arrow, Rhysodes, 442
arcuatum Chevrolat, Clinidium, 388
arcuatus (Chevrolat), Kupea, 388
arrow! (Grouvelle), Yamatosa, 425, 428
arrow! Bell and Bell, Yamatoa, 428
arrow! Grouvelle, Rhysodes, 428
aterrimus Broun, Rhysodes, 388, 389
aterrimus Chevrolat, Rhysodes, 389
basilewskyi (Brinck), Dhysores, 382, 385
boysi (Arrow), Yamatosa, 425, 430
boysi Arrow, Rhysodes, 430, 443
boysi auct. nec Arrow, Rhysodes, 429
boysi Bell and Bell, Yamatoa, 430
brouni Lewis, Rhysodes, 388, 389
canaliculatus Castelnau, Rhysodes, 409
cinerea new species, Grouvellina, 413
comes (Lewis), Rhysodes, 387, 388
comes Lewis, Epiglymmius, 388
compactus new species,
Plesioglymmius (Ameroglymmius), 435,
437
cooper! new species, Grouvellina, 412,418
cuneata new species, Grouvellina, 412, 416
cylindrica (Arrow), Kaveinga, 400
cylindrica (Arrow), Rhysodes, 403
cylindricus Arrow, Rhysodes, 403
dentipes new species, Grouvellina, 413, 422
descarpentriesi new species,
Grouvellina, 412,420
divergens new species, Grouvellina, 412, 421
dohertyi (Grouvelle), Shyrodes, 431
dohertyi Grouvelle, Rhysodes, 43 1
draco (Bell), Yamatosa, 425 429
draco Bell, Yamatoa, 429
edentata new species, Grouvellina, 412
elegans (Grouvelle), Plesioglymmius, 432
elegans Grouvelle, Rhysodes, 43 1 , 432, 434
eminens auct., Rhysodes, 394
eminens Broun, Rhysodes, 396
europaeus Ahrens, Rhizodes, 387
exaratus Dalman, Rhysodes, 386, 387
Classification of Rhysodini
445
fibulata new species, Kaveinga (sensu
stricto), 401
figuratus (Germar), Xhosores, 423
figuratus Germar, Rhysodes, 423
frontalis (Grouvelle), Kaveinga
(Angekiva), 377, 394
frontalis Grouvelle, Rhysodes, 390, 394
gigas new species, Grouvellina, 412,413
grouvelle! (Fairmaire), Grouvellina, 413, 421
grouvelle! Fairmaire, Rhysodes, 421
histrio new species, Kaveinga, 400
hova new species, Grouvellina, 412,417
jugatus , Plesioglymmius, 377
jugatus new species, Plesioglymmius
(Juxtaglymmius), 378,431,437,438
kaschmirensis Reitter, Rhysodes, 430
kukum new species, Kaveinga (sensu
stricto), 399, 402
liber new species, Dhysores, 382, 384
lignarius Olliff, Rhysodes, 380, 390
longior (Bell and Bell), Yamatoa, 425
longior (Grouvelle), Yamatosa, 425
lupata new species, Kaveinga, 400
lusca (Chevrolat), Kaveinga
(Vakeinga), 396
luscus auct., Rhysodes, 394
luscus Chevrolat, Rhysodes, 394, 396
marifuanga new species, Kaveinga
(sensu stricto), 400, 406
meridionalis (Grouvelle),
Plesioglymmius, 435
meridionalis (Grouvelle),
Plesioglymmius (Ameroglymmius), 377,
435
meridionalis Grouvelle, Rhysodes, 435
montana new species, Grouvellina, 412, 419
nilgiriensis (Arrow), Arrowina, 438, 441
nilgiriensis Arrow, Rhysodes, 441
niponensis (Lewis), Yamatosa, 425, 427
niponensis Bell and Bell, Yamatoa, 427
niponensis Lewis, Rhysodes, 424, 427
niponensis var. longior Grouvelle,
Rhysodes, 425
nudicornis new species, Kaveinga
(sensu stricto), 399, 403
occipitalis (Grouvelle), Kaveinga
(sensu stricto), 406
occipitalis Grouvelle, Rhysodes, 406
okapa new species, Kaveinga (sensu
stricto), 400, 405
orbitosa (Broun), Kaveinga
(Ingevaka), 377, 394
orbitosus Broun, Rhysodes, 394
pan new species, Dhysores, 382, 385
parva (Grouvelle), Kaveinga, 399
parvus Grouvelle, Rhysodes, 403
peninsularis (Arrow), Yamatosa, 425, 427
peninsularis (Bell and Bell), Yamatoa, 427
peninsularis Arrow, Rhysodes, 427
pensus (Broun), Tangaroa, 381, 442
pensus Broun, Rhysodes, 381
pignoris new species, Kaveinga (sensu
stricto), 399, 402
planicollis Bell & Bell, Srimara, 431
planifrons Fairmaire, Rhysodes, 408
pygmaea new species, Arrowina, 438, 441
quadriimpressus (Grouvelle),
Dhysores, 382, 384
quadriimpressus Grouvelle, Rhysodes, 384
radama new species, Grouvellina, 412,419
ranavalona new species, Grouvellina, 412,
417
reichardti new species, Plesioglymmius, 435
reichardti new species, Plesioglymmius
(Ameroglymmius), 436
reitteri (Bell), Yamatoa, 429
reitteri (Bell), Yamatosa, 425, 429
rhodesianus (Brinck), Dhysores, 382, 385
rostrata (Lewis), Arrowina, 438, 439
rostratus Lewis, Rhysodes, 439
schreiberi (Vulcano and Pereira),
Neodhysores, 386
schreiberi Vulcano and Pereira,
Rhysodes, 386
setosa (Grouvelle), Kaveinga
(Vakeinga), 395
setosus Grouvelle, Rhysodes, 395, 396
seximpressus Bell and Bell,
Neodhysores, 386
silus new species, Plesioglymmius
(sensu stricto), 434
strigiceps new species, Kaveinga, 400
sulcatus (Fabricius), Rhysodes, 386
Quaest. Ent., 1979, 15 (4)
446
Bell and Bell
sulcatus Fabricius, Cucujus, 386, 387
taprobanae (Fairmaire), Arrowina, 438, 439
taprobanae Fairmaire, Rhysodes, 438
taprobanae Fairmaire, Rhyzodes, 439
thoreyi (Grouvelle), Dhysores, 382, 384
thoreyi Grouvelle, Rhysodes, 382, 384
tubericeps (Fairmaire), Grouvellina, 409,
412,413
tubericeps auct. nec Fairmaire,
Rhysodes, 420
tubericeps Fairmaire, Rhysodes, 408, 413
ulteria new species, Kaveinga (sensu
stricto), 399, 402
KEYS
Key to species of Ameroglymmius, 435
Key to species of Arrowina, 438
Key to species of Dhysores, 382
Key to species of Grouvellina, 411
Key to subgenera of Kaveinga, 390
Key to species of Kaveinga
{sensu stricto), 399
Key to species of Neodhy sores, 386
Key to subgenera of Plesioglymmius, 432
Key to species of Plesioglymmius
(sensu stricto), 432
Key to species of Rhysodes, 386
Key to species of Vakeinga, 395
Key to species of Yamatosa, 425
REVISION AND CLADISTICS OF THE MIDDLE AMERICAN GENUS CREAGROPHORUS
MATTHEWS (COLEOPTERA: LEIODIDAE)
QUENTIN D. WHEELER
Department of Entomology
The Ohio State University
Columbus, OHIO 43210 Quaestiones Entomologicae
US. A. 15:447-479 1979
Creagrophorus is a compact genus of four Middle American species whose members eat fruiting
bodies of puffballs (Gasteromycetes); Creagrophorus hamatus Matthews {Mexico, Panama),
Creagrophorus bihamatus Matthews (Panama), Creagrophorous jamaicensis Peck (Jamaica), and
Creagrophorus spinaculeus new species (Panama). A cladistic analysis led to the following conclusions:
Creagrophorus is supported as a monophyletic group by apotypic broad labial palpus segment HI with
taeniaform seta, spines and flagellum of endophallus, absence of epicranial lines, and fimbriae of
galeae and presence of four setae on segment I of the urogomphus of the larvae, transverse coxites and
flattened, sclerotized styli of female genitalia, male femoral hooks (denticles), expanded middle tibia,
and puffball ecological relationships; the Aglyptinus association is a monophyletic lineage consisting of
Creagrophorus, Aglyptinus, and Scotocryptini, and defined synapotypically by 3-3-3 tarsi and oblique
lines on abdominal sternum HI; Aglyptinus is the sister group of CvtdLgvoyTaovm-Scotocryptini;
Scotocryptini is the sister group of Creagrophorus; C. hamatus is the sister to the
jamaicensis-bihamatus-spinaculeus lineage, and C. bihamatus sister species to C. spinaculeus.
Creagrophorus is hypothesized to be an endemic Middle American genus, which arose after its ancestor
migrated from South America where the sister group differentiated in body structure and ecological
relationships (Scotocryptini); the hamatus lineage was isolated north of the Isthmus of Tehuantepec;
the jamaicensis lineage was isolated in nuclear Middle America (and dispersed to Jamaica); and
bihamatus/spinaculeus became north/south vicariants due to volcanism in the Chiriqui region.
Creagrophorus beetles are continental in differentiation pattern, and as such are implied to have wide
geographic ranges and require relatively long periods of time for speciation. The Aglyptinus
association and Creagrophorus are defined, species of Creagrophorus described, and the following
presented; keys to genera of Aglyptinus association and species of Creagrophorus, habitus drawings of
adult and larval beetles, illustrations of important character states and geographic distribution,
designation of lectotypes for Matthews' species, and discussions about ecological, cladistic, and
zoogeographic relationships.
Creagrophorus est un petit genre de Leiodidae d’Amerique Centrale comprenant quatre especes qui se nourrissent de
vesses-de-loup (champignons Gasteromycetes); ce sont: C. hamatus Matthews (Mexique et Panama), C. bihamatus Matthews
(Panama, C. jamaicensis Peck (Jamaique) et C. spinaculeus Wheeler, une espece nouvelle (Panama). Une analyse cladistique permet
de tirer les conclusions suivantes; les Creagrophorus forment un taxon monophyleetique, a cause de la presence des caracteres
apotypiques suivants: troisieme segment des palpes labiaux large et portant une sole taeniaforme, phallobase avec epines et flagelle,
absence de lignes epicraniales et de fimbriae sur les galeas, presence de quatre soies sur le premier segment de I'urogomphe chez les
larves, coxas transversaux et aplatis, stylets de I'armature genitale des femelles sclerifies, femurs des males portant des crochets
(denticules), tibias intermediaires elargis, et rapports ecologiques avec les vesses-de-loup; le groupe Aglytinus constitue une lignee
monophyletique et comprend les Creagrophorus, les Aglyptinus et les Scotocryptini. Ce groupe est defini par les synapomorphies
suivantes: formule tarsale 3-3-3, et presence de lignes obliques sur le troisieme sternite abdominal. Le genre Aglyptinus est
monophyletique et est apparente aux Creagrophoms-Scotocryptini. C. hamatus est apparente a la lignee
jamaicensis-bihamatus-spinaculeus et C. jamaicensis est apparente a la lignee bihamatus-spinaculeus. On pose comme hypothese que
Creagrophorus est un genre endemique a I’Amerique Centrale et qu’il est apparu a la suite de la migration vers le nord de son ancetre
de I’Amerique du Sud, bu le groupe apparente (Scotocryptini) s’est differencie morphologiquement et ecologiquement. La lignee
hamatus a ete isolee au nord de I’isthme de Tehauntepec tandis que la lignee jamaicensis a ete isoleee an coeur de I’Amerique
Centrale (et s’est disperee jusqu’en Jamaique). Finalement, C. bihamatus et C. spinaculeus sont devenus des vicariants, le premier au
nord, le second au sud, en raison de I’activite volcanique dans la region de Chiriqui. Etant donne que les Creagrophorus ont un patron
de differenciation continental. Us devraient avoir une distribution couvrant de vastes regions geographiques et leur processus de
speciation devrait s’etendre sur une periode de temps relativement longue. On definit le groupe Aglyptinus et les Creagrophorus. et on
448
Wheeler
decrit les especes de Creagrophorus. On presente aussi une clef d’identification pour chacun des genres faisant partie du groupe
Aglyptinus et pour les especes de Creagrophorus, ainsi que des illustraions de inspect general des larves et des adultes, des
illustrations des principaux caracteres et des cartes montrant la distribution geographique. On designe des lectotypes pour les
especes decrites par Matthews, et enfin on discute des rapports ecologiques, cladistiques et biogeographiques.
TABLE OF CONTENTS
Introduction 448
Materials and Methods 449
Functional Morphology 451
Aglyptinus Association of Genera 452
Key to Genera 453
Genus Creagrophorus Matthew 453
Key to Adults of Species 456
The Species 456
Larvae 465
Ecological Relationships 468
Cladistics 469
Character Polarity 469
Cladistic Relationships 471
Zoogeography 471
Conclusions 476
Acknowledgements 476
References 476
Index 479
INTRODUCTION
Small, convex, shiny beetles (Fig. 1,2) which comprise the leiodid genus Creagrophorus Matthews
are distributed in Middle America, from central Mexico south to Panama, and in Jamaica in the
Antilles (Peck, 1972). Their minute size and obscure ecological habits have made these beetles
exceedingly rare in museum collections. During recent field investigations on tropical, lowland,
terrestrial carabids for Terry L. Erwin (National Museum of Natural History) in Panama, I collected a
large series of adults and larvae of an undescribed species and, perhaps more significantly, information
about the ecological relationships of Creagrophorus.
Taxonomy of Creagrophorus has remained largely neglected since the generic description by
Matthews in 1888. Peck (1977a) pointed out inconsistencies between the tarsal formula of his Jamaican
species and the formula recorded by Matthews for his species. Following Peck’s suggestion, I have
re-examined Matthews’ types and found that they, as well as Peck’s species, have 3-3-3 tarsi.
Creagrophorus, together with Aglyptinus and the Scotocryptini apparently form a monophyletic
group which I define below as the taxonomically informal Aglyptinus association of genera. Members of
this lineage were used as the out-group for determination of character polarity in Creagrophorus. Lack
of specimens has made decisions about cladistic and zoogeographic relationships difficult. However,
sister group relationships are hypothesized using existing information about structure and ecology of
Creagrophorus and related genera, and a few cursory observations are made about Creagrophorus
zoogeography, based on the cladistic conclusions.
Revision and cladistics of Creagrophorus
449
Figures 1 - 2. Creagrophorus spinaculeus, habitus: Fig. 1, dorsal aspect; Fig. 2, lateral aspect.
The purposes of this study are to define the genus Creagrophorus, revise known species, discuss
ecological, cladistic (intrageneric and suprageneric), and zoogeographic relationships, and describe the
larvae of a member of Creagrophorus.
MATERIALS AND METHODS
Materials
This study is based on about 65 adult and larval Creagrophorus, including type-specimens for three
previously described species. Specimens of the following species were studied as out-group taxa, and
voucher specimens placed in the National Museum of Natural History (USNM, listed below):
Scotocryptus sp., Aglyptinus laevis (LeConte), Aglyptinus matthewsi Champion, Aglyptinus sp.
(Panama), and a new species of Aglyptinus to be described elsewhere. The following acronyms represent
collections from which material was borrowed, or into which type material is deposited:
BMNH British Museum (Natural History), London;
CNCI Canadian National Collection of Insects, Ottawa;
MCZC Musem of Comparative Zoology, Cambridge;
OSUC Ohio State University, Columbus;
QDWC Author’s private collection, Columbus;
USNM National Museum of Natural History, Washington.
Quaest. Ent. 1979, 15 (4)
450
Wheeler
Methods
I have generally followed those methods discussed in a previous leiodid study (Wheeler, 1979),
including both philosophical and procedural methods. Exceptions, modifications, and additions to those
methods are discussed below.
Species criteria
I use both the theoretical and practical criteria for defining and recognizing species here as in my
study of Anisotoma (Wheeler, 1979). Because so few specimens of Creagrophorus were available,
delimitation of the range of intraspecific variation was impossible. I have used the structure of the
aedeagus, particularly the endophallus, as a source of structures on which decisions regarding separation
of species were made. Eversion of the endophallus was only possible in one species C. spinaculeus, since
so few males were seen. Consequently, drawings of endophallus structures were prepared by observation
through the median lobe. While these are only approximations of the actual detailed structure of
endophallus armature, they do show distinct differences in size, number, and configuration of spines
present.
Cladistic methods
My philosophical approach is essentially that of Hennig (1965,1966), as discussed previously
(Wheeler, 1979). Criteria for determination of character polarity were discussed in detail by Munroe
(1974) and Ekis (1977), and cladistics generally outlined by Griffiths (1972), and need not be pursued
here. I have based my decisions about character polarity on out-group comparisons with taxa of the
Aglyptinus association of genera.
Classification methods and formal ranking
My classifications of species and genera are phyletic sequence classifications as formulated and
applied by Nelson (1972, 1973), Cracraft (1974), Schuh (1976), and Wheeler (1979). No formal rank is
assigned to the monophyletic lineage formed by Creagrophorus and related genera. Rather, the informal
Aglyptinus association of genera is proposed and defined. This allows present statements about cladistic
relationships, and avoids premature reranking of Leiodini taxa before global studies are made, as
suggested by Erwin (1975) as a general taxonomic rule. Similarly, I refrain from reranking the
Scotocryptini now, but do state my views as to their phylogenetic significance.
Examination of larvae
Larvae were collected and stored in 70% ethanol until studied. For examination, they were cleared in
Nesbitt’s solution at room temperature and mounted directly into Hoyer’s medium (Krantz, 1978).
Magnifications up to X450 were used for observation and drawing.
Disarticulation
In addition to use of standardized dissecting methods for male and female genitalia (Wheeler, 1979),
it is important to make disarticulations of entire beetles whenever possible. Inaccurate interpretations of
detailed structure (e.g. tarsomere numbers in Creagrophorus and Aglyptinus, etc.) have plagued leiodid
classification, but can be avoided by use of high magnification with disarticulated specimens as
comparison of structures is enhanced and many additional characters are available for cladistic analysis.
After disarticulation of relaxed and partially cleared specimens, parts are further cleared in dilute KOH,
rinsed in water, and stored and studied in a mixture of glycerin and glycerin jelly as applied elsewhere by
Wheeler (1979) and Triplehorn and Wheeler (1979). Thus parts may be oriented for view from any
Revision and cladistics of Creagrophorus
451
angle with the use of gentle heat, yet are held firmly enough for preparation of accurate drawings. I
susbscribe to the views of Watrous (In press), who discusses both practical and philosophical reasons for
preparing disarticulations in relation to his work with Lathrobium staphylinid beetles.
Descriptive format
All of the species could not be studied in the same detail as C spinaculeus. Therefore, C. spinaculeus
is described in considerable detail (including larvae), the genus diagnosed and presumed to share
characters of that species except as noted, and other species diagnosed and briefly described. Where
appropriate, the following are given for each species: type data, diagnostic combination, description,
geographic relationships, cladistic relationships, material examined, and illustrations. Measurements
were made as discussed previously (Wheeler, 1979), and are only approximations due to varying
contractability of individual specimens.
COMMENTS ON FUNCTIONAL MORPHOLOGY
Aside from my observations of adult and larval C. spinaculeus in Panama, nothing is known about
Creagrophorus habits. During the present study, I have developed ideas about possible functions of some
structures of adult beetles. I present these ideas as possible explanations for the structures. Certainly
other explanations are not difficult to conceive, but at least my ideas may stimulate further field
observations which can test them and suggest realistic alternatives.
Female genitalia (Fig. 33)
Flattened, sclerotized styli of the female genitalia contrast so sharply with the plesiotypic form (i.e.,
lightly sclerotized, digitiform) seen in the sister group and Leiodinae in general that some specialized
function is strongly implied. I hypothesize that it is an adaptation for piercing the hardened outer wall of
host puffball fruiting bodies making oviposition possible, or perhaps for cutting through tough outer
walls of immature fruiting bodies before their emergence above ground (see discussion under “Legs”
below). Oviposition preceding apical pore formation by the host is supported by my observations in the
field, although there is no reason why older fruiting bodies could not be used at times, beetles gaining
entry through the apical pore.
Legs. (Fig. 16-19, 21, 22-26)
Enlargement of tibiae, presence of spines, and compaction of tarsomeres all may indicate fossorial
habits. Most puffballs develop under ground until maturity, and it is conceivable that Creagrophorus
beetles seek them out. “Primitive” Leiodini (e.g., Leiodes spp.) feed on hypogeous fungi (e.g., truffels:
see Arzone 1970, 1971), and Creagrophorus, like other leiodines, retain the antennal vesicles (Crowson,
1967; Peck, 1977b), which are presumably chemoreceptive. Subterranean activities aside, adult beetles
must move within the fruiting bodies in order to emerge, and these consist internally of little more than a
large mass of spores. Male femoral hooks are another matter (Fig. 11-14). I do not believe they function
directly in copulatory activities (males have spatulate adhesive setae on tarsomeres), but rather in some
male/male aggression or male/female courtship behavior.
Mouthparts
Distinct molar teeth are not present on adult (Fig. 8,9,28) or larval (Fig. 6,64) mandibles. Such teeth
were proposed as spore-crushing devices in Anisotoma beetles which feed on slime mold spores
(Wheeler, 1979). Crenulations (Fig. 9) of adult mandibles may be homologus with these teeth, and the
Quaest. Ent. 1979, 15 (4)
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Wheeler
molar region probably still functions in spore crushing to some extent. Setae and spines of the maxillae
in adults and larvae (Fig. 29,62) probably help to rake spores into the mouth. The strangest modification
of the mouthparts is the taeniaform seta of the labial palpus (Fig. 3,30), for which I do not hazard a
functional explanation.
The Aglyptinus association of genera
Creagrophorus, Aglyptinus, and Scotocryptini appear to consitute a monophyletic lineage.
Synapotypic character states which support this arrangement are discussed more fully under
“Cladistics”, but include the 3-3-3 tarsomere configuration and oblique lines of the first visible
abdominal sternum (sternum III). I have not made a study of Scotocryptini gcncraiScotocryptus,
Parabystus, Synaristus, and Scotocryptodes), and I am therefore not in a position to resolve specific
problems in that taxonomic group. The current classification is based on what I perceive as weak
taxonomic characters (Portevin, 1907,1937; Hatch, 1929a), and all members of Scotocryptini need to be
re-examined in detail. For the purposes of this study of Creagrophorus, I have made the following
assumptions about Scotocryptini: first, that Scotocryptini is a monophyletic group, and second, that the
single species of Scotocryptus which I have studied is representative of this monophyletic lineage. An
inevitable conclusion, if the above taxonomic arrangement is correct, is that Scotocryptini should not be
alloted tribal rank. Until a study of these beetles can be made, however, I conservatively refrain from
re-ranking the Scotocryptini. Nonetheless, I am convinced that it is a phylogenetically indefensible
taxonomic rank.
Table 1. Phyletic sequence classification of Aglyptinus association genera and Creagrophorus species,
and geographic distributions of taxa in North America (NAM), nuclear Middle America
(nMA), the Antilles (ANT), southern Middle America (sMA), and South America (SAM).
Numbers beside supraspecific taxa indicate number of species in area.
Revision and cladistics of Creagrophorus
453
KEY TO GENERA OF THE AGLYPTINUS ASSOCIATION
Adults
1 Tarsi 3-3-3; abdominal sternum III with oblique lines (Fig. 34), Aglyptinus associaton.. 2
Tarsal formula greater than 3-3-3; sternum III without oblique lines
(other Leiodini genera, not in key)
2 Antennal club compact; tarsi compact, tarsomeres subcylindrical 3
Antennal club loosely articulated; tarsi very long, thin; usually associated with fungi
Aglyptinus Cockerell
3 Eyes present; body glabrous; labial palpus with taeniaform seta (Fig. 30); stylus of
female genitalia flattened, heavily sclerotized (Fig. 33); middle tibia wider than hind
tibia
Creagrophorus Matthews
- Eyes absent; body pubescent; labial palpus without taeniaform seta; stylus long,
digitiform; middle tibia not wider than hind tibia Scotocryptini
(Includes following genera: Scotocryptus Girard, type species Scotocryptus meliponae
Girard; Parabystus Portevin, type species Scotocryptus inquilinus Matthews; Synaristus
Portevin, type species Synaristus pilosus Portevin; and Scotocryptodes Portevin, type
species Scotocryptodes germaini Portevin. Key in Hatch, 1929a).
Genus Creagrophorus Matthews
Type species. - Creagrophorus hamatus Matthews, 1888, by subsequent designation (Hatch,
1929b).
Diagnostic combination. - Staphylinoidae; Leiodidae; Leiodinae; Leiodini; ADULTS (Fig. 1,2) -
Tarsi 3-3-3; abdominal sternum III with oblique lines (Fig. 34); spermathecal capsule short, broad,
cylindrical, with long distal process (Fig. 35); stylus of female genitalia flattened, markedly sclerotized
(Fig. 33); aedeagus ventrally curved, endophallus with large annulate flagellum and various spination
(Fig. 36-40,50); middle tibia wider than hind tibia; antennal club of five antennomeres, compact
(Fig. 5), antennomere VIII smaller than VII or IX; tarsi compact, subcylindrical (Fig. 24); labial palpus
with taeniaform seta apically (Fig. 30); LARVAE (Fig. 53) - Galea not fimbriate (Fig. 62); epicranial
lines absent (Fig. 54); urogomphal article I quadrisetose, one seta very long (Fig. 58), segment II long,
crenulate; terga of abdomen each with four forked dorsal setae and one pointed lateral pair (Fig. 55);
sterna with simple setae only (Fig. 56); ECOLOGICAL RELATIONSHIPS - Adults and larvae feed
on fruiting bodies of puffballs (Gasteromycetes).
Cladistic relationships. - The sister group appears to be Scotocryptini, as discussed under
‘Cladistics’.
Geographic relationships. - The sister group is primarily South American with a northern
subtraction pattern; Creagrophorus is entirely Middle American including political divisions Mexico,
Panama, and Jamaica. Relationships discussed in detail under ‘Zoogeography’ section.
Species classification. - Table 1 represents a phyletic sequence classification of the species of
Creagrophorus, and members of the Aglyptinus association. It is based on relationships concluded from
the cladistic analysis.
Semaphoronts. - Key and descriptions of adult beetles are followed by a discussion and description of
larvae.
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Wheeler
Figures 3 - 5. C. spinaculeus: Fig. 3, head, dorsal aspect; Fig. 4, head, ventral aspect; Fig. 5, antenna.
Revision and cladistics of Creagrophorus
455
Figures 6 - 9. C. spinaculeus: Fig. 6, mola, larval mandible; Fig. 7, prostheca, adult mandible; 8-9, mola, adult mandible.
Quaest. Ent. 1979,15 (4)
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Wheeler
KEY TO ADULT CREAGROPHORUS
1 Elytral punctures distinct; hind femur with single hook (Fig. 1 2); middle femur with hook
(Fig. 11); endophallus with small spines near middle (Fig. 36); Mexico, Panama
Creagrophorus hamatus Matthews, p. 458
Elytral punctures indistinct or absent; hind femur with two hooks (Fig. 14); middle femur
without hook (Fig. 10); endophallus with more extensive spination; Panama or Jamaica ...
2
2 Color black; endophallus with patches of very large spines near apex (Fig. 39,40); median
lobe dorsally curved near apex (Fig. 41); Panama
Creagrophorus spinaculeus new species, p. 456
Color reddish-brown; endophallus with patches of smaller spines (Fig. 38) or small spines
only (Fig. 37), lacking patches of large spines * 3
3 Endophallus with small spines only (Fig. 37), lacking patches of large spines; female
middle tiba about as wide as in male; Jamaica Creagrophorus jamaicensis Peck, p. 461
Endophallus with two longitudinal patches of larger spines (Fig. 38); female middle tibia
much narrower than in male; Panama
Creagrophorus bihamatus Matthews, p. 458
Creagrophorus spinaculeus new species
(Fig. 1-10,13,15-19,22,23,27-35,39-43,69)
Holotype. - Male, USNM.
Type locality. - PANAMA, Canal Zone, Barro Colorado Island.
Paratypes. - 20, same locality as holotype; BMNH (2 males, 2 females), CNCI (2 males, 2 females),
MCZC (1 male, 2 females), USNM (1 male, 7 females), QDWC (1 male).
Diagnostic combination. - Color black; shining; male hind femur bidentate (Fig. 13); female middle
tibia similar to that of male; endophallus with patches of large spines near apex (Fig. 39,40).
Description. — Form subhemispherical (Fig. 1,2); length about 1 mm.
Color. Black, mouthparts, legs, and venter paler.
Microsculpture. Micropunctules distinct, moderately dense on head, increasingly sparse and obscure on pronotum and elytra.
Luster. Shining.
Head. Obovate, transverse (Fig. 3). Eyes large, conspicuous not protruding (Fig. 3). Postocular tempora not developed. Antennal
grooves ventral (Fig. 4). Gular sutures subparallel at middle, divergent anteriorly and posteriorly. Antennae short, club compact,
antennomeres as illustrated (Fig. 5).
Mouthparts. Labrum (Fig. 31) with six pairs apical setae, setae I-V increasing in length, VI shorter. Mandibles (Fig. 28) short,
robust; apical dens large, pointed, grooved on inner margin; prostheca large, with mesal, forked cuticular processes (Fig. 7); mola with
transverse grooves lined with crenulations (Fig. 8,9). Maxilla: three palpomeres, palpomere 111 aciculate (Fig. 29); lacinia and galea
with dense tufts of fine setae. Labium: three palpomeres. 111 broad, curved, with several long setae medially and broad taeniaform seta
apically (Fig. 30), taeniaform seta about length of palpus.
Thorax. Pronotum wider than long (Fig. 1); anterior margin medially excised; disc broadly curved. Prosternum short, medially
carinate, carina forming ventrally-directed, bidentate, laminiform process. Procoxal cavities closed anteriorly and posteriorly.
Mesosternum carinate; posterior intercoxal process declivous, forming transverse laminiform process. Hind wings long. Elytra broad,
sutural stria complete, epipleural fold very wide basally, gradually narrowed posteriorly (Fig. 27).
Legs. Tarsal formula 3-3-3, tarsomeres compact, subclyindrical (Fig. 17); basal tarsomere of male front and middle legs with
spatulate setae (Fig. 16,18). Front tibia narrow (Fig. 16). Middle tibia greatly enlarged (Fig. 17,18,19), much wider than hind tibia
(Fig. 22,23), with many stout spines. Hind tibia gradually widened. Middle femur wide. Hind femur wide (Fig. 13,15), bidentate in
male (Fig. 1 3).
Abdomen. Sterna Ill-Vlll visible, lll-Vll each with a single, transverse row of fine setae, VIII with medial and apical rows.
Sternum III with oblique lines (Fig. 34).
Revision and cladistics of Creagrophorus
457
Male genitalia. Median lobe widest subapically, narrowed at apex (Fig. 42); generally curved ventrally, apex curved dorsally
(Fig. 41). Endophallus with large annulated flagellum, and apical patches of large setae (Fig. 39,40). Parameres as illustrated
(Fig. 43).
Female genitalia. Coxites broad, transversely oriented. Styli lateraly curved, flattened, sclerotized, each with single stout seta
and finer mesal setae (Fig. 33).
Spermatheca. Capsule short, broad, cylindrical, with tubular distal process, short tubular basal process, long, tubular,
sclerotized duct leading to longer membranous duct (Fig. 35). Spermathecal gland small, membranous, inserted basally at side
(Fig. 35).
Figures 10 - 15. Creagrophorus spp., femora; Fig. 10, C. spinaculeus, male, middle leg; 11, C hamatus, male, middle leg;
Fig. 12, C. hamatus, male, hind leg; Fig. 13, C. spinaculeus, male, hind leg; Fig. 14, C bihamatus, male, hind leg; Fig. 15, C
spinaculeus, female, hind leg.
Quaest. Ent. 1979, 15 (4)
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Geographic relationships. - This species is known only from the Canal Zone in Panama. Both C.
hamatus and C. bihamatus seem to occur in Chiriqui Province, Panama. Full ranges of all three species
are undetermined.
Cladistic relationships. - C. spinaculeus is the sister species of C. bihamatus, based on synapotypic
occurrence of patches of large spines on the endophallus (Fig. 38,39,40).
Material examined. - In addition to specimens designated as types, one male and one female were
disarticulated and are in my collection (QDWC). Illustrations were made primarily from these
specimens.
Larvae. - The larvae are described under a section following descriptions of adults.
Creagrophorus hamatus Matthews
(Fig. 1 1,12,20,25,26,36,50-52,69)
Creagrophorus hamatus Matthews, 1888: 83.
Lectotype. - Male, BMNH, here designated (examined).
Type locality. - MEXICO, Puebla.
Diagnositc combination. - Elytral punctation distinct; male middle femur with hook (Fig. 11), hind
femur with single hook (Fig. 12); reddish-brown in color; area of endophallus near middle with small
spines (Fig. 36).
Description. — Length about 2.3 mm.
Color. Reddish-brown, appendages more yellowish.
Microsculpture. Sparse, fine punctules on head and pronotum, slightly larger, irregularly distributed punctules on elytra.
Luster. Shining.
Legs. Male middle tibia clavate apically (Fig. 26); middle femur with single hook (Fig. 1 1); hind tibia slightly clavate (Fig. 25);
hind femur with single hook (Fig. 1 2).
Male genitalia. Median lobe ventrally curved (Fig. 50); narrowed apically (Fig. 51). Paramere fused to median lobe, in part
(Fig. 52). Endophallus with small spines confined to area near middle (Fig. 36).
Female. - 1 have seen no females of this species.
Geographic relationships. - This is the only species of Creagrophorus known to occur as far north as
Mexico. Matthews (1888) also records the species from Chiriqui where C. bihamatus also occurs.
Cladistic relationships. - C. hamatus is the sister of remaining species, the
jamaicensis-bihamatus-spinaculeus lineage. The latter species share synapotypic character states
including bidenticulate male hind femora, and more extensive spination of the endophallus.
Material examined. - The lectotype specimen only, labelled: SYN-TYPE (circular, ringed with
blue)/TYPE (circular, ringed with red)/Puebla (hand-written), Mexico. Salle coll. /hamatus
(hand-wwritten)/B.C.A., Col. ,11, 1. /Creagrophorus hamatus Matt. <3 (hand written) det Hlisnikovsky
1961 /Creagrophorus hamatus Matth., M.E. Bacchus det 197, SYNTYPE/and my labels.
Creagrophorus bihamatus Matthews
(Fig. 14,21,24,38,47-49,69)
Creagrophorus bihamatus Matthews, 1888: 84.
Lectotype. - Male, BMNH, here designated (examined).
Type locality. - PANAMA, Chiriqui Province, Volcan de Chiriqui.
Diagnostic combination. - Color reddish-brown; male middle femur without hook, hind femur with
two hooks (Fig. 14); endophallus with two longitudinal patches of large spines (Fig. 38).
Description. — Length about 2.1 mm.
Revision and cladistics of Creagrophorus
459
Figures 16 - 21. Creagrophorus spp., legs and antenna: Fig. 16-19, C. spinaculeus. Fig. 16, male, front. Fig. 17, female,
middle (mesal surface). Fig. 18, male, middle. Fig. 19, female, middle (outer surface); Fig. 20, C. haniatus, antenna; Fig. 21, C.
bihamatus, male, middle.
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460
Figures 22 - 26. Creagrophorus spp., legs: Fig. 22-23, C. spinaculeus. Fig. 22, male, middle. Fig. 23, female, middle; Fig. 24, C.
bihamatus, male hind; Fig. 25-26, C hamatus, male. Fig. 25, hind. Fig. 26, middle.
Revision and cladistics of Creagrophorus
461
Color. Dark reddish brown, appendages paler.
Microsculpture. Obscure punctules on head, pronotum, and elytra.
Luster. Shining.
Legs. Male middle femur edentate, tibia broad (Fig. 21); hind femur bidentate (Fig. 14), tibia slightly wider apically
(Fig. 24).
Male genitalia. Median lobe ventrally curved (Fig. 47), narrowed apically (Fig. 48). Paramere short, broad, not fused (Fig.
49). Endophallus with generally distributed small spines, and two distal, longitudinal patches of larger spines (Fig. 38).
Female. I tentatively accept the female specimen in the BMNFl, identified by Matthews, as a member of this species. The
middle tibia is widened only apically, the micropunctules are more distinct, and the color paler than the lectotype.
Geographic relationships. - This species occurs at Volcan de Chiriqui in Chiriqui Province, Panama.
Matthews also reported C hamatus from this locality.
Cladistic relationships. - C. bihamatus is the sister species of C. spinaculeus. Synapotypic
occurrence of patches of large spines on the endophallus suggest this arrangement.
Material examined. - Type specimens, 1 male, 1 female, only. Lecotype male labelled: SYNTYPE
(circular, ringed with blue); Bihamatus M. (hand-written); Sp. figured; V. de Chiriqui below 4,000 ft.,
Champion; B.C.A., Col. ,11, 1. In this specimen the legs on the right side are detached, on the left side
they are in position (handwritten); Creagrophorus bihamatus M. S , det. Hlisnikovsky 1962;
Creagrophorus bihamatus., M.E. Bacchus det 1978, SYNTYPE; and my lecotype labels. Female
specimen (BMNH), also from Chiriqui, also figured in B.C.A.
Creagrophorus jamaicensis Peck
(Fig. 37,44-46,69)
Creagrophorus jamaicensis Peck, 1972: 56; Peck, 1977a.
Holotype. - Male, Illinois Natural History Survey, original designation (Peck, 1972) (Not
examined).
Type locality. - JAMAICA, St. Andrew Parish, Hermitage Dam.
Diagnostic combination. - Color reddish-brown, luster slightly iridescent; elytral punctules indistinct;
middle femur without hook; male hind femur with two hooks; endophallus with small spines, generally
distributed, lacking patches of large spines (Fig. 37); Jamaican distribution.
Description. — Length about 1.2-1. 4 mm.
Color. Reddish-brown, antennae paler.
Microsculpture. Punctules obscure on head, pronotum, and elytra.
Luster. Shining; somewhat iridescent.
Legs. Male middle femur without hook, hind femur with two hooks; female femora without hooks; tibiae similar in males and
females, middle tibia very wide (see figs, in Peck, 1972), similar to those illustrated for C. spinaculeus.
Male genitalia. Median lobe ventrally curved (Fig. 44), apex not fused, narrowed (Fig. 45). Parameres wide (Fig. 46).
Endophallus with small spines, irregularly distributed (Fig. 37), without patches of large spines.
Female genitalia. Not examined.
Geographic relationships. - This species is the only one which occurs in the Antilles, and is known
only from Jamaica (Peck, 1972; 1977a).
Cladistic relationships. - C. jamaicensis is the sister taxon of the bihamatus-spinaculeus lineage,
based on synapotypic occurrence of two hooks on male hind femur and increased spination of the
endophallus.
Material examined. - I have seen a female paratype (MCZC) and four specimens from Pt. Antonio,
Jamaica (MCZC).
Quaest. Ent. 1979, 15 (4)
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462
Figures 27 - 35. C. spinaculeus: Fig. 27, elytron, dorsal, right; Fig. 28, mandible, right; Fig. 29, maxilla, right; Fig. 30, labium,
ventral; Fig. 31, labrum, dorsal; Fig. 32, meso- and meta-sterna; Fig. 33, female genitalia; Fig. 34, female abdomen, ventral;
Fig. 35, spermatheca, MD (membranous duct), SD (sclerotized duct), DP (distal process), SG (spermathecal gland), BP (basal
process), SC (spermathecal capsule).
Revision and cladistics of Creagrophorus
463
Figures 36 - 40. Creagrophorus spp., endophallus: Fig. 36, C. hamatus\ Fig. 37, C. jamaicensis\ Fig. 38, C. bihamatus\
Fig. 39-40, C. spinaculeus. Fig. 39, ventral; Fig. 40, lateral.
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464
Figures 41 - 52. Creagrophorus spp.: Fig. 41-43, C. spinaculeus. Fig. 41, median lobe, lateral. Fig. 42, same, apex, ventral.
Fig. 43, paramere; Fig. 44-46, C. jamaicensis,. Fig. 44, median lobe, lateral. Fig. 45, same, apex, ventral. Fig. 46, paramere;
Fig. 47-49, C. bihamatus. Fig. 47, median lobe, lateral. Fig. 48, same, apex, ventral. Fig. 49, paramere; Fig. 50-52, C. hamatus.
Fig. 50, median lobe, lateral. Fig. 51, same, apex, ventral. Fig. 52, paramere.
Revision and cladistics of Creagrophorus
465
Larvae of Creagrophorus
No larvae have previously been described for Creagrophorus, or taxa of the Aglyptinus association.
Information about larvae of Leiodinae has, in general, been sparse and fragmentary (Hatch, 1927;
Cornell, 1972). This makes deciding which characters are of value at the genus level problematical, at
best.
Leiodid beetles belong to the “Leptinid Association” of families in the Staphylinoidae (Roving &
Craighead, 1931; Dybas, 1976). Dybas defined this assemblage of families with the following characters
of larvae: (1) denticulate or asperate expanded molar region of mandible; (2) prostheca on inner margin
of mandible; (3) urogomphi one or two segmented and articulated; (4) antennae three-segmented, with
sensory appendage on segment II; and (5) galea fimbriate. Most of these characters, as pointed out by
Dybas, are plesiotypic, but unite several staphylinoid families in a practical sense for purposes of
discussion until phylogenetic lines become evident. Dybas (1976) gives the following combination of
characters for recognition of leiodid larvae: (1) anal membrane without two prominent claws; (2)
epicranial lines present; (3) ocelli present or absent; (4) compound setae often present.
Creagrophorus larvae differ from this typical pattern of characters in the following ways. The mola
has distinct transverse grooves, but not distinct teeth or asperites (Fig. 6). The galea is not fimbriate. No
distinct epicranial lines are apparent in any of my preparations.
It is not presently clear to me which characters will be of use at the generic level, and which at the
species level. Studies of larvae in other genera will eventually make such decisions easier. Meanwhile,
only C. spinaculeus larvae are available, and are described below.
Creagrophorus spinaculeus new species
(Fig. 6,53-64; species described above, based on adults)
Diagnostic combination. - Epicranial lines absent (Fig. 54); galea not fimbriate (Fig. 62); four pairs
of forked setae on each tergum (Fig. 55) of abdomen; urogomphus segment I quadrisetose, one seta very
long, segment II long, crenulate (Fig. 58).
Description. — Late-instar larva (Fig. 53), length about 2.9 mm; width across metathorax about 0.6 mm.
Form. Body subcylindrical in cross-section, widest at metathorax, gradually narrowed posteriorly (Fig. 53).
Color. White to yellowish; mandibles testaceous; more or less distinct brownish spot on each side of midline of abdominal and
thoracic terga.
Setae. Forked or simple (pointed).
Sclerotization. Body lightly sclerotized.
Head capsule (Fig. 54). Without distinct epicranial lines. Clypeus not delimited by suture, except basolateral angles, with single
pair of dorsal setae. Labrum (Fig. 60) separated by fine suture; one dorsal pair large setae medially and two pairs minute setae
laterally on disc; five pairs of setae along anterior margin, medial pair small. Ocelli, two pairs. Antenna (Fig. 61) with three articles: II
long, with large apical sensory appendage and smaller accessory sensory appendage; ill with single stout, apical spicule. Maxilla
(Fig. 62) with three palpomeres: I large, II much shorter than I, III long, thin with thin walled basal process (digitiform organ) nearly
half length of 111; galea not fringed (fimbriate); lacinia with comb of five stout setae, apex blade-like (Fig. 63). Labium with two
palpomeres; 1 large, cylindrical, with apicolateral tuft; II much smaller, cylindrical, with apical tuft; ligula broad, apex rounded.
Mandibles (Fig. 64) similar, except crenulations distal to median dens on right mandible, absent from left; apex pointed; molar region
prominent, with many transverse ridges (Fig. 6); prostheca broad, dentiform.
Thorax. Slightly broader than abdominal segment I, terga not clearly defined. Prothorax with two transverse rows of setae:
anterior row entirely of simple (pointed) setae, posterior row with four medial pairs forked setae, one pair of lateral, simple setae.
Mesothorax and metathorax each with single row of setae consisting of four medial pairs of forked setae, and single pair lateral simple
setae. Mesothoracic spiracle larger than abdominal spiracles, similar in form.
Abdomen. Tergum IX with single pair long, simple, posterolateral setae; all other terga with four pair forked setae, large lateral
pair of simple setae, and pair of smaller, simple, post-spiracular setae (Fig. 55). Sterna with three pairs of simple setae each (Fig. 56).
Spiracles small, annular, present on segments 1-Vlll. Urogomphus (Fig. 58) two-segmented: segment I short, wide, quadrisetose, setae
simple, one very long (about length of segment II); II long, thin, gradually narrowed, basal three-fourths crenulate (“wrinkled”) in
Quaest. Ent. 1979, 15 (4)
466
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Figures 53 - 58. C. spinaculeus, larva: Fig. 53, habitus, dorsal aspect; Fig. 54, head, dorsal aspect; Fig. 55, abdominal
tergum III; Fig. 56, abdominal sternum III; Fig. 57, anal vesicle; Fig. 58, urogomphus.
Revision and cladistics of Creagrophorus
467
appearance with single, long, apical seta. Anal vesicle with two pairs simple setae (Fig. 57).
Legs. Similar in form to hind leg (Fig. 59).
Material examined. - About 40 larvae were collected with the type series from Lycogalopsis solmsii
(Gasteromycetes, Lycoperdales, Lycoperdaceae) fruiting bodies in Panama, on Barro Colorado Island.
About twelve larvae were deposited in the British Museum (BMNH), and remaining larvae retained in
my collection (QDWC).
Comments. Larvae were observed feeding on spores inside puffball fruiting bodies in the field, and
“plugs” of spores were visible in larval digestive tracts in slide preparations. No comparative statements
are yet appropriate regarding relationships to other leiodids.
Figures 59 - 64. C. spinaculeus, larva: Fig. 59, hind leg; Fig. 60, labrum, dorsal aspect; Fig. 61, antenna; Fig. 62, maxilla;
Fig. 63, lacinia apex; Fig. 64, mandible.
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ECOLOGICAL RELATIONSHIPS
My only encounter with Creagrophorus was in the lowland, tropical forest of Barro Colorado Island
(Fig. 65), where both adult and larval Creagrophorus spinaculeus were feeding in puffball fruiting
bodies (Fig. 66). Although there is no way of estimating the degree of host specificity, as much
information as possible about the host should be discussed as a basis for developing explanations for
spatial and temporal distribution of Creagrophorus beetles.
The host puffball, Lycogalopsis solmsii Fisher (Gasteromycetes: Lycoperdales: Lycoperdaceae), is
pantropical and recorded from Java (type locality), Congo, Ceylon, Japan, Puerto Rico, Martinique, and
Panama. Ligneous substrata seem to be preferred, as is generally true of puffballs (Bessey, 1950; Smith,
1951;Dring, 1973).
Lycogalopsis solmsii was collected previously on Barro Colorado Island and in Balboa, both in the
Canal Zone of Panama (Martin, 1939). Abundance of its fruiting bodies was recorded in Balboa during
the rainy summer of 1935, but was not noticed during the drier summer of 1937. Smaller series were
taken from decaying wood on Barro Colorado Island during both of those years. Moisture seems to be an
important parameter for the life cycle of Lycogalopsis.
Figures 65 - 66. Habitat and host of C spinaculeus-. Fig. 65, lowland tropical forest, Barro Colorado Island; Fig. 66, fruiting
body, Lycogalopsis solmsii (Gasteromycetes: Lycoperdales).
Revision and cladistics of Creagrophorus
469
Larvae were found associated with the fruiting bodies in the field, and extracted by hand in the
laboratory. Both larvae and large series of fully-pigmented adults fulfill the criteria established by
Lawrence (1971, 1973) for recognition of valid breeding records for fungus beetles (ciids in particular).
Perhaps we can explain the absence of Creagrophorus from extensive leaf litter samples taken in
Jamaica by Peck (1977a), if Lycogalopsis is fairly resricted to ligneous substrata, primarily decaying
wood. It will be of great interest to collect specifically for both Lycogalopsis and puffballs generally to
find if this is true, and what the range of acceptable hosts are for Creagrophorus.
Mycologists generally believe that the Gasteromycetes are closely related to (and in fact, are
descendants of) the Hymenomycetes (e.g., mushrooms and related fungi). It is not difficult to imagine
an Aglyptinus-Wkt ancestor, already feeding on Hymenomycetes, adapting to puffball hosts.
Voucher specimens of the host fruiting bodies {Lycogalopsis solmsii), identified by
Alexander H. Smith, have been deposited in the National Museum of Natural History (USNM) along
with the type series of C. spinaculeus.
CLADISTICS
Cladistics are used as a method of approximate reconstruction of evolutionary pathways, as in an
earlier paper (Wheeler, 1979). Character polarity determinations were based on out-group comparisons
with Aglyptinus association taxa. Numbers in parentheses refer to characters in Table 2; references to
illustrations are given in the table.
Character polarity (Table 2)
Compact structure of the antennal club (15) is interpreted as synapotypic for Creagrophorus and
Scotocryptini. Similar compaction of clubs occurs in other Leiodini genera (ie., Isoplastus,
Apheloplastus etc.), but I believe it is independently acquired within the Aglyptinus association.
Eventually, intensive out-group comparisons will refine this and similar questions. The broad, flattened,
curved palpomere segment III (1) of the Creagrophorus labium is autapotypic for the genus; Aglyptinus
and Scotocryptini have segment III short, subcylindrical, and more or less truncate apically (the
plesiotypic homologue). The taeniaform seta (2) is unique to Creagrophorus.
Oblique lines on abdominal sternum III (11) are synapotypic for Creagrophorus, Aglyptinus, and
Scotocryptini. In general, larger body size seems typical of ’primitive‘ Creagrophorus (e.g., C. hamatus),
and Aglyptinus and Scotocryptini, suggesting that smaller size (19) is apotypic for some Creagrophorus
species. Black coloration (20) of C. spinaculeus contrasts with the usual reddish-brown color of
Aglyptinus association beetles. Elytral punctation is plesiotypic in relation to the Aglyptinus assoctiation
and Leiodini as a whole, and reduction or absence (21) is presumably apotypic. Iridescence (22) is only
known in one species of Creagrophorus (C. jamaicensis). Nature of this iridescence has not been
determined, but it subjectively looks different than surface gratings which occur in Aglyptinus sp. which
will be discussed elsewhere, and is therefore interpreted as autapotypic. Hind femur hooks (8) are
autapotypic for Creagrophorus, as are middle femur hooks for C. hamatus (18). 3-3-3 tarsi (12) are
apparently confined to Aglyptinus association genera. Compact, subcylindrical tarsi (14) are shared by
Creagrophorus and Scotocryptus. Expansion of the middle tibia is apotypic for Creagrophorus. Greater
degree of expansion (29) is synapotypic in members of the jamaicensis-bihamatus-spinaculeus lineage.
Genital structural information is fragmentary for the out-group; that which is published is not
detailed enough for evaluation here (e.g., Hlisnikovsky, 1964). I interpret available data as follows.
Annulae of most of the distal portion of the flagellum of the endophallus (3) is autapotypic for
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Table 2. Polarity of character states in Creagrophorus Matthews and related genera
Creagrophorus', annulae are absent from or confined to the basal portion of the flagellum in other
Aglyptinus association taxa. Evident, broad spines of the endophallus (4) have so far only been seen in
Creagrophorus, within which a transformational series exists. The trend is toward increased number and
size of spines. C. hamatus is plesiotypic with only a few spines confined to middle of endophallus
(Fig. 36); C. jamaicensis has increased numbers of spines (28); C. bihamatus and C. spinaculeus share
synapotypic patches of spines (24); and spines of the patches are increased in size (25) in C. spinaculeus.
Parameres are partially fused to the median lobe (23) in C. hamatus; the plesiotypic homologue is freely
articulated parameres in related in-group and out-group taxa. Dorsal curvature of the median lobe (26)
is unique to C. spinaculeus within the genus.
Transverse, short coxites (9) and flattened, short, highly sclerotized styli (10) of the female genitalia
Revision and cladistics of Creagrophorus
471
are autapotypic for Creagrophorus. The plesiotypic homologue for both characters involve long, lightly
sclerotized, digitiform shaped structures in Aglyptinus, Scotocryptus, and Leiodinae in general.
Epicranial lines of the larval cranium seem plesiotypic in Staphylinoidea and the “Leptinid
association” of families (Dybas, 1976), and have been seen in larvae identified as Aglyptinus and
Anistoma (tribe Agathidiini) by me. Absence of the lines (6) from Creagrophorus is therefore inferred
to be autapotypic. Similarly, lack of fimbriae from larval galeae (7) seems autapotypic (and was seen in
the same taxa m.entioned above). I suggest that the quadrisetose Article I of the urogomphus (5) may be
autapotypic for Creagrophorus as well.
Based on observations in the field of North and Middle American species of Aglyptinus, I believe
that the general (and plesiotypic) ecological relationship involves feeding on various fleshy
hymenomycete fungi (mushrooms, etc.). Gasteromycete association in Creagrophorus (with
Lycogalopsis) (16) is autapotypic, as is the inquilinism in meliponine bee nests for Scotocryptini (17).
Cladistic relationships
Suprageneric relationships (Fig. 68). - Aglyptinus association genera share synapotypic 3-3-3 tarsi
(12), and oblique lines on abdominal sternum III (11). The sister group has not been identified, but will
probably belong to the Leiodini sensu stricto (Jeannel, 1962). Compact tarsi (14) and antennal club
(15) support CreagrophorusScoiocrypiim as a sister group pair. Their collective sister group is
Aglyptinus, though monophyly has not been shown for that genus. Transitions to puffball feeding (16) in
Creagrophorus and inquilinism (17) with bees in Scotocryptini are not yet understood, and a simple
dichotomy subjectively seems simplistic in explaining the known extant fauna.
Autapotypic support for monophyletic status of Creagrophorus is abundant, including modifications
of labial palpi (1,2), endophallus (3,4), female genitalia (9,10), legs (8,13), larvae (5,6,7) and ecologlical
relationsips (16).
Intrageneric relationships (Fig. 67). - Fused parameres (23) and a hook on the middle femur of the
male (18) are autapotypic for C. hamatus. Support for the sister lineage,
jamaicensis-bihamatus-spinaculeus, includes increased complexity of endophallus spines (28), reduced
elytral punctation (21), smaller body size (19), and greatly expanded middle tibiae (29). C. jamaicensis
is autapotypically iridescent (22). Support for the sister lineage, bihamatus-spinaculeus, includes
patches of large spines on the endophallus (24). Autapotypies for C. spinaculeus are its black coloration
(20), dorsally curved median lobe (26), and large endophallus spines (25).
ZOOGEOGRAPHY
Few specimens and sparse locality information strictly limit discussions of both extant zoogeographic
patterns and historical explanations for them. I concur with Whitehead (1972) and Ball (1975) who
advocate provision of a theoretical basis for additional research, in the form of testable hypotheses, in
every systematic study. I feel this is particularly true during early studies of a taxon, such as this one of
Creagrophorus. General improvement in our understanding of evolutionary patterns will only come
about with concern for forming and testing hypotheses, and dismissal of fears about invalidation of our
ideas, as discussed by Ball (1978). It is in this spirit that I offer some interpretations of the available
zoogeographic data for Creagrophorus and related genera.
Quaest. Ent. 1979, 15 (4)
472
Wheeler
Jamaica O
Mexico 9
Panama 0
o
o
o o
o o
• •
Figures 67 - 68. Ciadograms, numbers refer to Table 2: Fig. 67, intrageneric relationships of Creagrophorus\ Fig. 68,
intergeneric relationships of Aglyptinus association taxa.
Revision and cladistics of Creagrophorus
473
Distribution patterns
For reasons discussed under ‘Speciation patterns’ below, I believe that species of Creagrophorus are
continental in their pattern of species differentiation. I assume that species are widely distributed in
suitable habitats, and that endemic populations are not common.
C. hamatus occurs in Mexico (Puebla), and was reported from Volcan de Chiriqui by Matthews
(1888). I could not locate the latter specimen, but provisionally accept Matthews’ identification as valid.
Remaining species constitute the sister-group of C. hamatus. C. jamaicensis occurs in Jamaica, and the
bihamatus-spinaculeus lineage in Panama. C. bihamatus is known only from the Volcan de Chiriqui
area and C. spinaculeus from Barro Colorado Island, Canal Zone. Collectively, the distribution of
Creagrophorus is Middle America, extending northward into Mexico.
Scotocryptini, sister group of Creagrophorus is primarily South American, with a northern
subtraction pattern (Table 1). Only one species {Parabystus inquilinus, see Hatch, 1929b) occurring
north of South America does not also have populations in South America. I suggest that this one will
also represent a range extension by a South American form, or that its sister species will be South
American. Scotocryptines constrast sharply with the ecological relationships of Creagrophorus, with
their inquiline habits in Meliponinae bee nests.
Speciation patterns
Whitehead (1976) discussed criteria by which two generally distinct patterns of species
differentiation may be recognized: continental and islandic. Briefly, clues for recognizing these patterns
are as follows.
A. Islandic speciation pattern:
1. High-altitude distributions, generally;
2. Disjunct (differentiated) populations including mountaintops, caves, water-locked islands,
etc.;
3. Low-altitude distributions of organisms which are ecologically specialized or
habitat-restricted.
B. Continental speciation pattern:
1. Low-altitude distributions, generally;
2. Continuous (undifferentiated) populations including riparian forms along streams, lowland
forest forms, etc., characterized by wide geographic ranges;
3. Ecological generalists or non-habitat-restricted organisms.
Whitehead stressed that these are only generalizations of patterns, and that every organism must be
evaluated individually. Those found to conform to the islandic pattern are more likely to have been
influenced by cyclic weathering of the Pleistocene than continental forms, and hence often evolve at a
more rapid rate. How should the speciation pattern be interpreted in Creagrophorus!
At first glance, puffball feeding appears to be a rather specialized ecological relationship. Host fungi,
however, are typically cosmopolitan (or in the case of Lycogalopsis, pantropical) within suitable climatic
regions, and occur spottily over wide geographic ranges, presumably occupied by Creagrophorus. Fully
winged adult beetles undoubtedly spend much of their time migrating from one fruiting body to another
in the lowland forest, wherever microclimatic conditions permit successful growth of the host. Thus, we
can assume that they have escaped major pressures from Pleistocene cyclic climatic changes, for reasons
enumerated by Whitehead (1976) for lowland, tropical, terrestrial carabids, and they should be
classified as demonstrating the continental speciation pattern.
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Figure 69. Geographic distribution of species of Creagrophorus, and South American ancestral origin.
Revision and cladistics of Creagrophorus
475
Historical zoogeography
Vicariance biogeography (see Platnick & Nelson, 1978; Rosen, 1978; Croizat et al., 1974) provides a
means of estimating recency of common ancestral ranges from knowledge about cladistic relationships.
Congruence of a number of vicariant sister pairs may indicate a general track, which can potentially be
causally explained through historical geologic events.
Constraints imposed by lack of empirical data about Creagrophorus geographic distribution preclude
detailed vicariant hypotheses (ie., those correlating Creagrophorus vicariant events with those of other
groups, and with known geological events). Albeit, I believe that it would be useful to discuss the general
sequence of vicariant events indicated by present data. Tests and substantiation of these ideas would
make more detailed biogeographic analyses possible.
Historical scenario. - Ancestral Creagrophorus entered Middle America from South America
(Fig. 69), became widespread, and differentiated from the southern ancestral stock which is today
represented by Scotocryptini. The scotocryptines have become specialized for inquilinism in Meliponinae
bee nests. The first vicariant event isolated the hamatus lineage to the north of the Tehuantepec portal.
The second was a division of nuclear Middle American and southern Middle American stocks (the
nuclear stock dispersed to Jamaica). The third vicariant event isolated sister species to the north and
south of the Chiriqui volcanic region {bihamatus to the north, spinaculeus to the south).
Predictions. - Any discussion about zoogeography at this early date in studies of Creagrophorus is
meaningless without some obvious utility to later research. Therefore, I offer the following predictions as
a framework for continued study.
1. Species conform to a continental pattern, enjoying wide geographic ranges.
2. If C. hamatus is actually widespread, no additional species of the hamatus lineage will be found
which have originated in Middle America and/or are adapted to the same hosts.
3. The jamaicensis lineage is represented in nuclear Middle America by either C jamaicensis itself
or its sister species. If C. jamaicensis has mainland populations, then no additional species is predicted.
Similarly, if C. jamaicensis is widespread in the Antilles, no other insular members of the jamaicensis
lineage are expected.
4. C. bihamatus occurs throughout southern Middle America, north of Chiriqui. Absence of this a
species from nuclear Middle America is a corailary of the presence of a member of the jamaicensis
lineage there in prediction 3.
5. C. spinaculeus is the southern vicariant of the Chiriqui barrier. It is either allopatric or parapatric
with C. bihamatus.
6. C. bihamatus and C. spinaculeus actually share a most recent common ancestor (ie., no additional
species of the bihamatus-spinaculeus lineage exist).
7. The sequence of vicariant events hypothesized above require that no additional branching points
exist on the lineage leading to bihamatus-spinaculeus. Specifically, the jamaicensis lineage is the sister
of the bihamatus-spinaculeus lineage, and the hamatus lineage sister to these collectively.
8. Creagrophorus is absent from South America. If any species do exist there, they will not be part of
the Middle American lineage.
Some aspects of the historical scenario and the predictions are bold extrapolations from a limited
amount of empirical data. My intent is to set forth possible explanations for the observed pattern of
distribution, which can subsequently be field-tested and supported or rejected. The eventual outcome
should be the same; approximation of the actual sequence of events leading to present Creagrophorus
patterns. These ideas will minimally serve as a template for arranging later observations and a stimulus
Quaest. Ent. 1979, 15 (4)
476
Wheeler
to continued field work in puffball/leiodid beetle relationships.
CONCLUSIONS
Many conclusions have been reached in this study which are stated or implied within the text. It
would be useful, however, to draw some conclusions of a general nature from the study as a whole. There
remains much to be done in the field before ideas about ecological and zoogeographic relationships of
Creagrophorus beetles become stable. I hope that this study has set the stage for such investigations. I
urge detailed study of the Scotocryptini, and suspect that the tribal status must be abandoned, based on
my cladistic analysis. My discussion of the larvae is a minor contribution to an area of study in the
Leiodinae which is virtually untouched. Ecological data, though limited, should now be sufficient to
allow (and stimulate) the accumulation of specimens and information about hosts and geographic
distributions throughout Middle America.
ACKNOWLEDGEMENTS
Field studies in Panama were supported in part by the Smithsonian Institution’s Environmental
Sciences Program funds awarded to Dr. Terry L. Erwin, National Museum of Natural History. I
sincerely thank Dr. Erwin for his encouragement and the chance to successfully test David Fairchild’s
challenge: “Let the student of life in the temperate zone once (follow) a newly cut trail through a forest
in the dripping tropics and if it does not give him a new outlook upon the world of living organisms, it
will be because he is too sensitive to the humidity and heat or too wet with perspiration to care what he
sees“. (Fairchild, 1924: 99). Alfred Newton and Stewart Peck shared their insights into leiodid
systematics and biology, including ideas basic to this study. M.E. Bacchus, J.M. Campbell, A.F.
Newton, S.B. Peck, and M. Thayer loaned specimens of Creagrophorus and related leiodids; D.S.
Chandler, D.E. Johnston, and L.E. Watrous generously gave me specimens of related genera. All or part
of a draft of this paper was read by Ginter Ekis, Terry Erwin, Candace Martinson, F.C. Thompson,
Charles Triplehorn, Larry Watrous, and Donald Whitehead. The host puffballs were identified through
efforts of Alexander Smith and Roland Seymour. David Stutes and Glen Berkey skillfully prepared
photomicrographs and plate reductions, respectively.
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Universidade degli Studi de Torino 317-357.
Arzone, A. 1971. Nouvi reperti sulla biologia di Leiodes cinnamomea Panzer in Tuber magnatum Pico
(Coleoptera Staphylinoidae). Bollettino dell’Instituto ed Orto Botanico dell’Universita di Torino
17: 121-129.
Ball, G.E. 1975. Pericaline Lebiini: Notes on classification, a synopsis of the New World genera, and a
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Ball G.E. 1978. The species of the Neotropical genus Trichopselaphus Chaudoir (Coleoptera:
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14: 447-489.
Revision and cladistics of Creagrophorus
477
Bessey, E.A. 1950. Morphology and Taxonomy of Fungi. Blakiston, Philadelphia.
Boving, A.G. and F.C. Craighead. 1931. An illustrated synopsis of the principle larval forms of the order
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Cornell, J.F. 1972. Larvae of the families of Coleoptera: a bibliographic survey of recent papers and
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Middlesex.
Croizat, L., G. Nelson, and D.E. Rosen. 1974. Centers of origin and related concepts. Systematic
Zoology 23: 265-287.
Dring, D.M. 1973. Gasteromycetes In Ainsworth, G.C., F.K. Sparrow, and A.S. Sussman (eds.). The
Fungi, an advanced treatise, vol. IVB, A taxonomic review with keys: Basidiomycetes and lower
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Dybas, H.S. 1976. The larval characters of featherwing and limulodid beetles, and their family
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Fairchild, D. 1924. Barro Colorado Island Laboratory. Journal of Heredity 15: 99-112.
Griffiths, G.C.D. 1972. The Phylogenetic Classification of Diptera Cyclorrhapha with Special
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evolution. Systematic Entomology 4: 251-309.
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(Coleoptera: Carabidae: Scaritini). Quaestiones Entomologicae 8: 131-348.
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Cryptorhynchinae), and Quaternary Middle American zoogeography. Quaestiones Entomologicae
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Revision and cladistics of Creagrophorus
419
Index to Names of Taxa
TRIBE AND SUBTRIBES
Agathidiini, 471,477
Bembidiini, 477
Harpalini, 476
Leiodini, 450, 45 1 , 453, 469, 47 1
Pterostichini, 478
Scaritini, 478
Scotocryptini, 447, 448, 450, 452, 453, 469,
471.473.475.476
Tachyina, 477
GENERA AND SUBGENERA
Aglyptinus Cockerell, 448, 449, 450, 452,
453.465.469.470, 471,477
Anistoma, 450, 451,471
Apheloplastus, 469
Creagrophorus Matthews, 448, 449, 450,
451, 452, 453, 458, 465, 468, 469, 470, 471,
473.475.476
Isoplastus, 469
Lathrobium, 451
Lycogalopsis, 468, 469, 473
Parabystus Portevin, 452, 453
Scotocryptodes Portevin, 452 453
Scotocryptus Girard, 452, 453, 469, 47 1
Synaristus Portevin, 449 452, 453
SPECIES AND SUBSPECIES
bihamatus Matthews, Creagrophorus, 456,
458.461.470, 473,475
cinnamomea Panzer, Liodes, 476
germaini Portevin, Scotocryptodes, 453
hamatus Matthews, Creagrophorus, 453, 456,
458.461.469, 470, 471,473,475
inquilinus Matthews, Scotocryptus, 453
jamaicensis Peck, Creagrophorus, 456, 46 1 ,
469, 470, 471,473,475
laevis (LeConte), Aglyptinus, 449
matthewsi Champion, Aglyptinus, 449
meliponae Girard, Scotocryptus, 453
pilosus Portevin, Synaristus, 453
solmsii Fisher, Lycogalopsis, 468
spinaculeus new species,
Creagrophorus, 447, 450, 45 1 , 456, 458,
461. 465. 469, 470, 471, 473, 475
Quaest. Ent. 1979, 15 (4)
481
CORRIGENDA — Quaestiones Entomologicae, Volume 15(3)
Straneo — Classification of South American Pterostichini
page/line
355/ 6 remove “Cephalostrichus Straneo”
355/35 change “Cephalostichus Straneo, 349, 353” to “Cephalostichus Straneo, 345, 349, 353”
Kurtak — Seasonal changes in larval black fly food
page/line
358/16 change “Naumann” to “Neeley”
Quaest. Ent. 1979, 15 (4)
:.i'’^' ■-■■':V ^'' 7.-7
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