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HARVARD UNIVERSITY
LIBRARY
OF THE
Museum of Comparative Zoology
r
Quaestiones
entomologicae
MUS. COMP. ZOOti
LIBRARY
FEB fe I
S A/'
’ . (M j , i-. i v
A periodical record of entomological investigations,
published at the Department of Entomology, Uni-
versity of Alberta, Edmonton, Canada
VOLUME V
1969
ii
CONTENTS
Book Review 1
McFadden - New distributional records for Canadian soldier flies
(Diptera: Stratiomyidae). Part I. Beridinae and Sarginae 5
Tawfik - Effects of population density of Cimex lectularius L 9
Larson - A revision of the genera Philophuga Motschoulsky and
Tecnophilus Chaudoir, with notes on the North American
Callidina (Coleoptera: Carabidae) 15
Book Review 85
Freitag - A r evision of the species of the Genus Evarthrus LeConte
(Coleoptera: Carabidae) 89
Errata 212
Book Review 213
Graham - A comparison of sampling methods for adult mosquito
populations in central Alberta, Canada 217
Chiang - Some pharmacological properties of the nerve cord of the
cockroach, Periplaneta americana (L.) 263
Errata / 307
Graham - Observations on the biology of the adult female mosquitoes
(Diptera: Culicidae) at George Lake, Alberta, Canada 309
Fredeen - Outbreaks of the black fly Simulium arcticum Malloch in Alberta 341
INDEX
iii
Abaris, 90, 94
Abax, 90 133
Abdelnur, O.M., 341 , 347, 370, 371
acetate, 295
acetic acid, 269
acetone, 267
acetyl-beta-methyl choline chloride, 266
acetyl-3-methylcholine, 275
acetylcholine (ACh), 263, 269, 273, 279,
294, 296, 299
acetylcholine chloride, 266, 268
acetylcholinesterase (AChE), 263, 265, 275,
279, 285, 295, 297, 299.
Actina viridis, 5
Adam, J. P. (see Hamon), 333, 337
Adephaga, 94
adrenaline, 297
Aedes, 220, 236, 240, 244, 246, 251, 256,
310,315,322-328,332,333,334
campestris, 3 1 1 , 324
canadensis , 237 , 311,315, 325
cataphylla, 240, 312, 3-15, 325, 331
cinereus, 220, 236, 240, 312, 315, 324, 332
communis, 220, 236, 240, 247, 313, 314,
325,328, 331,333,335
communis nevadensis, 325
diantaeus, 313, 315, 325, 331
dorsalis, 311,315, 324, 325
excrucians, 220, 234, 240, 247, 310, 313,
325,327,331,333,336
fitchii, 220, 234, 236, 240, 310, 314, 325-326,
327, 331,334,336
flavescens, 310, 312, 315, 326, 336
hexodontus, 313, 314, 326, 328, 331
impiger, 312, 324
implicatus, 220, 236, 240, 247, 312, 315,
326,331,332
increpitus, 3 12, 314, 326
intrudens, 240, 247, 313, 326, 328, 331, 335,
336
nigripes, 324
niphadopsis, 312
pionips, 247, 313, 315, 326, 328, 331
pullatus, 313, 315, 327, 331
punctor, 220, 234, 236, 240, 313, 327, 331
riparius, 220, 236, 240, 310, 327, 331
spencerii, 312,315, 327
sticticus, 313, 324, 327, 328, 331
stimulans, 312, 315, 327
(cont.)
trichurus, 312, 315, 327
220, 234, 240, 247, 257, 311,
313,324,331,334,336
Aedimorphus, 315, 322, 324
Albert, A., 275, 295, 299
ali-esterase (AliE), 268
alkaline sulfite solution, 269
Allognosta brevicornis, 5
f us cit arsis, 5
obscuriventris , 5
Alnus tenuifolia, 219
Amara stupida, 63
Ambache, N., 295, 299
Amelanchier alnifolia, 219
amines, biogenic, 265, 290
Anaferonia, 126, 156, 158, 159
distincta, 128, 129
evanescens, 156, 157
fausta, 128, 129
iowana, 128, 129
latebrosus, 159
lixa, 159
pantex, 156, 157
papago, 159, 160
pimalis, 159
pudica, 159, 160
vernicata, 159, 160
analysis of variation (Carabidae), 1 9
Ancylis comptana, 65
Anderson, J.R., W. Olkowski & J.B. Hoy,
255, 257, (see Olkowski, W., 255, 259)
animal bait, 217
Anopheles, 313, 315, 316, 333
earlei, 220, 236, 240, 244, 247, 254,
256, 260, 311,315,321,328, 331
intrudens, 328
maculipennis, 252, 316
occidentalis , 316
quadrimaculatus , 260
anopheline vectors, 25 1
Anoplura, 1,2,3
ant larvae, 63
pupae, 63
anticholinesterase, 164, 167, 283, 288
Aplysia, 296
Arctia caja, 265
arcto-tertiary geoflora, 174
iv
arginine phosphate, 269
Arnason, A.P. (see Fredeen, F.J.H.), 341, 347,
349, 372; (see Rempel, J.G., 341 , 350, 372)
Artemisia , 65
arthropod ecology, 2
population, 2
Arthus’ syndrome, 354
aryl-esterase (ArE), 268
Aster, 219
Astigmata, 2
A triplex, 65
nuttalli, 63
atropine, 265, 295
Auffenberg, W. & W.W. Milstead, 174, 191
Austrogoniodes, 3
gressitti, 3
keleri, 3
Avenzoariidae, 2
Axelrod, D. I., 69, 73
Bach, R.C. (see Huffacker, C.B.), 238, 241, 258
Ball,G.E., 17, 73, 89, 93, 101, 174, 191
Barlow, R.B., 275, 299
Barr, A.R., 217, 235, 257, 309, 321, 336
(see Chapman, H.C., 325, 337)
Barr, A.R., T.A. Smith, M. Boreham & K.E.
White, 254, 257
Barton Brown, L., L.F. Dobson, E.S. Hodgson
&J.K. Kiraly, 297,299
Bar-Zeev, M., 9, 13, 14
Basford, N.L., J.E. Butler, C.A. Leone & F.J.
Rohlf, 94, 191
Bates, H.W., 73, 191
Bates, M., 227, 252, 255, 257
beaver dams, 219
Beddington, A. & R.W. Brimblecombe, 275, 299
Beckel, W.E., 314, 336
Bellamy, R.E. & W.C. Reeves, 227, 238, 255, 257
(see Hayes, R.O., 239, 258)
Belton, P. & M. Galloway, 235, 254, 257
Berck, B. (see Fredeen, F.J.H.), 349, 372
Beridinae, 5
Beris californica, 5, 6
Berry, E.W., 174, 191
Betula papyrifera , 219
Biddlingmayer, W.L., 217, 153, 157 (see Klock,
J.W., 255, 258)
Bigelow, R.S. & C. Reimer, 19, 73
Biram’s anemometer, 222
Birks, R.I. (see Macintosh, F.C.), 273, 303
Birks, R. & F. C. Macintosh, 273, 299
Bisset, G.W., J.F.D. Frazer, M. Rothschild,
& M. Schachter, 265,299
blackfly, 341-371
Blackmore, J.S. (see Rainey, M.B.), 255,
259
Blackwelder, E., 69, 73, 191
Blair, W.F., 174, 186, 191
blatchleyi group, 127, 130-133, 172, 188,
210
Blatchley, W.S., 192
Bodenheimer, F.S., 9, 14
Boistel, J. (see Gahery, Y.), 297, 301
Boreham, M. (see Barr, A.R.), 254, 257
Boullin, J. (see Costa, E.), 290, 301
Bouteloua gracilis, 63
Boura, A.L.A. & A.F. Green, 297, 300
Boving, A.G. & F.C. Craighead, 93, 192
Brady, V.E. & J. Sternburg, 288, 299, 300
Braun, E.L., 174, 184, 192
Brazin, M. (see Hoskin, F.C.G.), 299, 302
Breeland, S.G. & E. Pickard, 220, 235, 237,
25 1 , 1 53, 1 57 (see Smith, G.E., 220, 253,
260
Brimblecombe, R.W. (see Bebbington, A.),
275, 299
Brocus approximatus , 106
Brodie, B.B. & P.A. Shore, 297, 300
Brodie, W.B. (see Costa, E.), 290, 301
(see Shore, P.A., 290, 304)
Broscus, 102, 108
laevipennis, 103
Brown, A., T.H.D. Griffitts, S. Erwin, &
L.Y. Dyrenforth, 354, 371
Brown, A.W.A., 227, 239, 259
Brown, A.W.A., D.S. Sarkaria & R.P.
Thompson, 239, 257
Brown, R.H. (see Mikalonis,S. J.), 264,303
Burdick, D.J. & E.H. Kardos, 252, 257
Burgess, L. & W.O. Haufe, 322, 336,
(see Haufe, W.O., 222, 238, 258)
Burn, J.H., 297, 300
Burn, J.H. & M.J. Rand, 263, 273, 297,
300
Butanol, 268
Butler, J.E. (see Basford, N.L.), 94, 191
Burse 11, E., 217, 251 , 257
Burton, A.N. (see McLintock, J.), 254, 259
Cain, A.J. & G.A. Harrison, 66, 73
V
calcium cyanide, 220
Calleida croceicollis, 44, 46, 60
viridis, 33
Callida, 15, 18,21,23,26,28,32,62,66,67
chloridipennis , 60
cyanea, 38
decora, 22, 29, 31,78, 80
purpurea, 26, 29, 30, 32, 80
viridipennis , 32, 80
Callidina, 15, 20, 24, 28, 43, 64, 66, 68, 72
biology of the subtribe, 62
key to the subtribe, adults, 28
key to the subtribe, larvae, 24
phylogenetic diagram, 67
taxonomy of the subtribe, 22-23
Callidinae, 16
Calliphora erythrocephala , 213
Cameron, A.E., 347, 371
Cameron, M.C., 297, 300
Carabidae, 1 5, 89, 94
Carausius morosus, 213
carbachol (carbamylcholine), 263, 266, 275,
279, 295, 296
carbamate, 266
carbon dioxide baited traps, 227
Carestia, R. R. & L. B. Savage, 238, 241, 257
Car ex, 219
Carlston, C.W., 174, 192
Carpenter, M.J. & W.J. LaCasse, 309, 313, 326,
336
Carpenter, M.J. & L.T. Nielsen, 252, 257
Carpenter, S.J. & L.T. Nielsen, 333, 337
Casey, T.L., 16, 73, 89, 192
catecholamines, 268, 296, 297
Cephalotes, 103, 106, 125
Chadwick, L.E., 268, 279, 300
Chadwick, L.E. & D.L. Hill, 289, 300
Chamberlain, R.W. (see Newhouse, V.F.), 227,
238, 241, 259
Chamberlin, J.C. (see Stage, H.H.), 222, 231, 253,
260
Chamberlin, J.C. & F.R. Lawson, 222, 257
Chang, S.C. & C.W. Kearns, 265, 300
Chang, S.C. (see Sternburg, S.), 266, 283, 298,
304
Chang, V. & M.J. Rand, 297, 300
Chapman, H.C., 335, 337
Chapman, H.C. & A.R. Barr, 325, 337
Chapman, R., 9, 14
Chaudoir, M. de, 23, 73, 192
Chen, G., 275
Chen, G. & R. Portman, 275, 300
Chen, G., R. Portman & A. Wickel, 275,
300
Chevrolat, L.A., 73
Chiang, P.K., 263
chicken baited traps, 222
Chironomidae, 2
Chivers-Wilson, V.S. (see Hutcheon, D.E.),
351,372
choline, 263, 265, 283, 288, 295, 297
choline acetylase (ChA), 263, 273
choline chloride, 266, 269
cholinesterase, 297
choline esters, 264, 265
Christophers, S.R., 337
Cimex lectularius, 9-13
adult stage, 10
fecundity, 10
mortality rate, 1 3
nymphal stadia, 10
population density, 9-13
preoviposition period, 10
Clark, J.C. & F.C. Wray, 254, 257, 324,
337
Clarke, W.B., 174, 192
Clement, A.N., 230, 238, 252, 257, 334,
337
Cnephia saskatchewana, 354
Cobben, R.H., 85
cockroach nerve cord, determination of
AChE activity, 267-268
effect of acetyl choline, 279-283
effect of AChE activity, 289
effect of adrenergic drugs, 290
effect of carbachol (carbamylcholine),
275
effect of choline, 283
effect of choline upon TEpp-treated
n nerve cords, 288
effect of dimethylphenylpiperazinium
(DMPP), 275
effect of eserine, 283
effect of hemicholinium, 269-273
effect of methacholine (acetyl-|3-m ethyl-
choline), 275-279
effect of nicotine, 275
effect of pilocarpine, 279
vi
cockroach nerve cord (cont.)
effect of pyridine-2-aldoxime methiodid
(2-PAM) upon TEPP-treated nerve cords,
285-288
effect of tetraethylpyrophosphate (TEPP),
283
endogenous activity, 269
electrophysiological studies, 266-267
spectrofluorometric determination of
noradrenaline, 268-269, 294
Cohn, T.J., 69, 73
Coleman, A. P., 174, 192
Coleoptera, 15, 89
Colhoun, E.H., 263, 268, 283, 288, 297, 300
Colhoun, E.H. & E.Y. Spencer, 265, 301
Collembola, 1, 2
Coquillettidia, 315, 322
perturbans, 31 1, 313, 315, 322, 332
Corbet, A.S. (see Fisher, R.A.), 235, 258, 316.
Corbet, P.S., 217, 130, 142, 252, 258, 333, 335
Cornus canadensis , 219
stolonifera, 219
Costa, E., D. J. Boullin, W. Hammer, W. Vogel,
& W.B., Brodie, 290, 301
Craig, D.A., 86
Craighead, F.C. (see Boving, A.G.), 93, 192
criteria for species, subspecies & genera
(Carabidae), 17
Crombie, A.C., 9, 14
Cross, H.F. (see Twinn, C.R.), 347, 372
Cryptostigmata, 2
Csiki, E.,73, 89, 192
Culex, 31 1, 313, 315, 322
annulirostris , 243
apicalis, 322
restuans, 31 1 , 322
tarsalis, 238, 252, 256, 260, 31 1, 315, 322
territans, 220, 231, 234, 240, 247, 311,315,
322, 331
tri taeniorhynchus,25S
Culicidae, 309
Culicinae, 314
Culiseta, 220, 236, 31 1, 313, 315-322
alaskaensis, 240, 247, 31 1, 314, 316, 321,
331,336
impatiens, 311, 320
incidens, 31 1 , 320
inornata, 220, 231, 234, 236, 240, 244, 247,
256, 260, 311,314, 320, 331-336
Culiseta morsitans dyari , 311,313,315,
321
sylvestris minnesotae , 311,313,315,
321
Curran, C.H., 5, 7
Curtis, C.L., 321, 322, 337, 347, 371
Curtis, D.R., R.W. Ryall & J.C. Walkins,
295, 301
Cyanogas, G., 320
Cyclotrachelus , 89, 95, 101, 109-1 16,
119, 125, 126, 169, 171, 173, 176,
187, 211
fallaciosus, 125, 126
fucatus, 89
levifaber, 89
macrovulum, 89
parafaber, 89
roticollis, 125, 126
texensis, 89
Cylindronotum, 23, 28, 66, 67
Cymindis, 21 , 32
amoena, 42
viridicollis , 31
viridis, 34, 38
Dahl, E., B. Flack, C. von Mecklanburg,
& H. Myhrberg, 296,301
Dauterman, W.C., A. Talens & K. van
Asperen, 267, 301
Davis, M.B., 174, 192
DDT, 298
De Groat, W.C. & R.L. Voile, 296, 301
Dejean, P.F.M.A., 73, 101, 192
Detinova, T.S., 217, 230, 252, 258
Dettbarn, W. & P. Rosenberg, 269, 301
Dettbarn, W., P. Rosenberg & D. Nach-
mansohn, 298, 301
diazoblue, 267
laurylsulfate solution (DBLS), 268
diisopropyl fluorophosphate (DEP), 299
phosphoric acid, 299
Dillenberg, H. (see McLintock, J.), 254,
259
Dillon, L.S., 69, 74
j3j3-dimethyl acrylcholine, 265
dimethylphenyl piperazinium (DMPP),
263, 266, 275, 295
Diptera, 1,5,255,309,371,372
Dobson, L.F. (see Barton Brown, L.), 297,
299
vii
Dow, R.P., 239, 258
Downey, J.E., 220, 258
Duke, B.D.L., 231, 258
Dunn, E. (see MacLagen, D.S.), 9, 14
Dyar, H.G. (see Howard, L.O.), 321, 337
Dyrenforth, L.Y. (see Brown, A.), 354, 371
Eccles, R.M. & B. Libet, 296, 301
Ehrenpreis, S., 301
Emden, F. I. van, 74
endogenous activity, 263
Engel, L.G. & R. W. Gerard, 269, 301
Ephestia kuhniella, 213
Epilobium angusti folium, 219
Erigonum flavum, 63
Erwin, S. (see Brown, A.), 354, 371
eserine, 263, 264, 267, 283, 288
sulfate, 266
esterase, 295
Euler, V.S. von, 297,301
Eumolops, 127, 146, 152, 160, 163, 165
ampla, 163, 164
decepta, 161, 162
impolita , 161, 162
inflatula, 161 , 162
prominens , 161, 162
sexualis, 127, 161, 162
sulcata, 147, 149
Euproctinus , 16, 23, 24
trivittatus, 80
Euproctus, 16
Evarthrinus, 113, 117, 127, 146, 152, 161
alabamensis , 117
alternans, 153
inflatipennis , 147, 149
lilliputicus , 117, 118
minax, 161, 162
pinorum , 113, 114
retractus, 147
Evarthrops , 113, 117, 127, 147, 152
Evarthrus, biology, 93
centers of concentration, 178-183
distribution pattern, 174-176
effects of the Pleistocene epoch, 183-184
extent of range, 176-178
historical zoogeography, 186-190
key to the species & subspecies, 95-100
material, 90
methods, 90-93
phylogeny, 168-173
Evarthrus , primitive & specialized
character conditions, 170
revision of the species of the genus, 89-
212
sister species, 1 84
species-pairs, 184-186
subgenus, 126
taxonomy, 93-1 68
zoogeography, 174-190
Evarthrus acutus , 104
alabamae, 98, 141-142, 172, 181, 185,
188. 197, 204, 208
alabamensis, 95, 1 15-119, 171, 181, 187,
197, 199, 202, 207
alternans, 98, 146, 153-154, 173, 181,
184. 189. 198. 205. 209
americanus, 131
approximatus , 96, 106-107, 171, 181,
184, 187, 196, 200, 207
blatchleyi, 99, 130, 134, 136, 172, 181,
185, 188, 197, 199, 203, 208
breviformis, 133, 135
brevoorti, 96, 110, 111, 113-115, 171,
181, 187, 196,201,207
constrictus, 99, 117, 1 18, 126, 154, 158-
160. 167. 173. 181. 189. 199. 205. 209
convivus, 99, 133, 134, 137-139, 172, 181,
185. 188. 197, 203, 208
deceptus, 127, 173
engelmanni, 98, 139, 142-143, 172, 181,
185. 197, 204, 208
enormis, 144
faber, 93, 95, 122, 125-126, 172, 176,
181. 185. 188. 197. 199. 202. 207
fatuus, 147, 149
floridensis, 99, 130, 132-133, 136, 181,
185, 188, 197, 203, 208
fucatus, 96, 110, 111-112, 171, 181,
185. 187. 196. 201. 207
furtivus, 98, 127, 146, 152, 173, 181, 185,
189. 198. 205. 209
gigas ,95, 100, 164, 165-166, 167, 173,
181, 185, 198. 206, 210
gravesi, 95, 127, 167-168, 176, 180, 184,
189. 198, 210
gravidus, 97, 100, 160, 161, 163-164, 173,
181, 184, 189, 198, 206, 210
hernandensis , 96, 101-102, 171, 181, 184,
187. 196. 199. 200. 107
viii
Evarthrus heros , 100, 154, 155-157, 173, 181,
198, 199, 205, 210
hypherpiformis,91 , 145-146, 172, 181, 184,
198. 204, 208
incisus, 97,99, 128, 172, 181, 185, 188, 197,
203. 207
iowensis, 97, 100, 129, 135, 147, 154-156,
159, 161, 173, 181, 184, 189, 198, 205
iuvenis, 96, 106, 107-108, 171, 181, 187,
196, 200, 207
laevipennis, 96, 101, 102, 103-105, 171, 181,
187, 195, 200, 207
latebrosus, 156, 157
levifaber, 96, 109, 122-125, 172, 181, 185,
188, 197,202, 207
lodingi, 141, 147, 148, 149
macrovulum, 96, 115-121, 171, 181, 185,
187, 188, 197,202, 207
montanus, 133, 135
morio, 96, 101-104, 171, 181, 184, 187, 196,
200. 207
nonnitens,9S, 139, 143-145, 172, 180, 181,
185. 188. 197. 204. 208
obsoletus, 97, 106, 107, 108-109, 171, 181,
187. 196. 200. 207
or bat us, 137
ovulum, 95, 104, 1 15-1 19, 171, 181, 188,
197. 199. 202. 207
parafaber, 95,117, 122-123, 172, 176, 181,
188. 197. 202. 207
parasodalis, 98, 100, 146, 150-151, 173, 181,
185. 189. 198. 204, 209
roticollis, 109
rotundatus, 113, 114
rubripes , 140
sallei, 100, 164, 173, 181, 185, 198, 206,210
seximpressus , 98, 139-141, 143, 172, 181, 185,
188. 197. 199. 204. 208
sigillatus, 91,99, 126, 133-136, 138, 172, 181,
188, 197, 199, 203,208
sinus, 99, 132, 126-137, 142, 172, 181, 185,
188, 197,203,208
sodalis, 89,94, 146-149, 151, 173, 185, 189,
204
spoliatus, 96, 110, 111-114, 171, 181, 187,
196. 201. 207
substriatus , 97, 99, 100, 129, 155-159, 167,
173, 181, 184, 189, 198, 199, 205, 209
taurus, 102, 103
Evarthrus tenebricus, fossil species, 168
texensis, 96, 115, 121-122, 172, 181,
185, 187, 188, 197, 202, 207
tervus, 89, 160-162, 173, 184, 189
unicolor, 95, 109-111, 114, 169, 171,
181, 187, 196, 199, 201
vagans, 141, 143
vinctus, 96, 115-116, 171, 181, 187,
196, 201,207
whitcombi, 97, 128, 129-130, 172, 181,
188, 197,203,207
Exodontha luteipes, 5, 6
Eyles, D.E. (see Wharton, D.H.), 255, 260
faber group, 109, 122-127, 171, 188,210
Falk, B. (see Owen, C.), 296, 303
Fawcett, D.W., 213
Feldberg, W., 295, 301
Ferestria, 101, 106, 108, 118
acuta, 104, 105
bullata, 104, 105
castigata, 104, 105
nanula, 104, 105
siminola, 104
simiola, 104, 105
Feronia, 108, 1 10, 133, 158, 166
abdominalis, 128, 129
acuminata, 160
americana, 133, 135, 166
brevoorti, 1 1 4
colossus, 146
cons trie ta, 158
corax, 146, 148
heros, 166
incisa, 127
lixa, 127, 129
morio, 102
obsoleta, 108
orbata, 133, 135, 137
ovipennis, 158, 160
ovulum, 1 18
seximpressa, 139
sigillata, 133
sodalis, 146
spoliata, 113, 125
tenebricosa, 125, 126
unicolor, 1 1 0
vagans, 147, 148
vidua, 133, 135
Fisher, R.A., 19, 74
ix
Fisher, R.A., A.S. Corbet & C.B. Williams, 235,
258, 316, 337
Fisher, R.W. (see Smallman, B.N.), 298, 304
Flack, B. (see Dahl, E.,), 296, 301
Flemings, M.P., 255, 258
Flint, R.F., 174, 192
Fortax, 89, 91,95, 101, 108, 169, 171, 187,21 1
iuvensis, 89
fossil material, 168
Franeria dumosa, 64
Frazer, J.F.D. (see Bisset, G.W.), 265, 299
Frazer, W.T. (see Kandel, E.R.), 296, 302
Fredeen, F .J.H., 341 , 347, 349, 350, 355,371
Fredeen, F.J.H., J.G. Rempel & A.P. Arnason,
341,347, 349, 372
Fredeen, F.J.H., A.P. Arnason & B. Berck, 349,
372
Freitag, R., 20, 74, 89
Frontali, N., 265, 297, 301
Fruentov, N.R. (see Magazanik, L.G.), 303
Fukuto, T.R. (see Winton, M.Y.), 264, 306
Gahery, Y. & J. Boistel, 297, 301
Galloway, M. (see Belton, P.), 235, 254, 257
garden tiger moth, 265
Gardiner, J.E., 273, 301
Gater, B.A.R., 255,258
Geber, G.L. & R.L. Voile, 275, 279, 283, 296,
301
Gerard, R.W. (see Engel, L.G.), 269, 301
Germar, E.F., 192
Gershenfeld, H.M. (see Tauc, L.), 296, 305
gigas group, 167-168, 173,210
Ginetsinskii, A.G., 298, 302
Ginsborg, B.L. & S. Guerrero, 275, 302
Ginsburg, S. (see Wilson, I.B.), 285, 306
Gjullin, C.M. (see Stage, H.H.), 324, 338
Gjullin, L.M., W.W. Yates, & H.H. Stage, 324,
337
Glascow, J.P., 231, 253, 258
Glossina, 231 , 253
swynnertoni, 251, 257
Glycia, 29
viridicollis, 3 1
Gnus, 341,347
Gomphiocephalus hodgsoni, 2
Gomori, G., 267, 302
Gomori's technique, 267
Goth, A., 295,302
Gordon, H.T. (see Welsh, J.H.), 275, 306
Goulden, C.H., 19,74
Graham, A., 174, 192
Graham, P„ 214, 156, 158, 309, 337
Grahamely tron crofti, 1
Grauer, F.H. (see Gudgel, E.F.), 354,372
gravesi group, 167-168, 173, 210
Green, A.F. (see Boura, A.L.A.), 297,300
Greenslade, P.J.M., 182, 192
Gregerman, R.I. & G. Wald, 297, 302
Grenier, P. (see Hamon, J.), 333, 337
Gressitt, J.L., 1
Griffitts, T.H.D. (see Brown, A.), 354, 371
Grollman, A., 279, 297, 301
Gudgel, E.F. & F.H. Grauer, 354, 372
Guerrero, S. (see Ginsborg, B.L.), 275, 302
Guilday, J.E. (see Hibbard, C.W.), 174,
193
Gurba, J.B.,355,372
Habu, A., 16,21,23,74
Haddow, A.J., 231, 158
Halacaridae, 2
Halarachnidae, 2
haloalkylamine, 290
Haldeman, S.S., 192
Hamberger, B., K.A. Norberg & F. Sjoqvist,
296, 302
Hamberger, B., K.A. Norberg & U. Ungestedt,
296, 302
Hammer, W. (see Costa, E.), 290, 301
Hammon, McD. (see Reeves, W.C.), 238,
260
Hamon, J. , S. Sales, J.P. Adam & P. Grenier,
333,337
Happold, D.C.B., 256, 258, 314, 316, 320,
322,325,326,328, 337
Harpalinae, 94
Harpalus, 94
Harrell, B.E. (see Martin, P.S.), 186, 194
Harrison, G.A. (see Cain, A.J.), 66, 73
Hartshorn, J.H. (see Schafer, J.P.), 174, 194
Hatch, M.H., 74
Haufe, W.O. (see Burgess, L.), 322, 336
Haufe, W.O. & L. Burgess, 222, 238, 258
Hayes, R.O., R.E. Bellamy, W.C. Reeves,
& M.J. Willis, 239, 258
Hearle, E., 326, 337, 341,372
hemicholinium (HC-3), 263, 265, 266, 269,
273, 295
Heming, B.S., 214
Hess, A. (see Rainey, M.B., 255, 259), 263,302
Heteroptera, 235, 254
evolutionary trends, 85
phylogeny, 85-86
hexamethonium, 295
Hibbard, C.W., D.E. Ray, D.E. Savage, D.W.
Tayler & J.E. Guilday, 174, 193
Hill, D.L. (see Chadwick, L.E.), 289, 300
Hippelates pusio, 258
histamine, 294, 35 1
Hobbiger, F., 285, 288, 302
Hocking, B. (see Klassen, W., 325, 337),
(see Twinn, C.R., 347, 372), 242, 258, 325,
337
Hodgson, E.S. (see Barton Brown, L.), 297, 299
Hokin, M.R., L.E. Hokin & W.D. Shelp, 296,
302
Holmes, R. & E.L. Robins, 285, 288, 302
Holstein, M.H., 328, 337
Horn, G.H., 16,45,74,91, 193
Horsfall, W.R., 324, 325, 337
Hoskin, F.C.G., P. Rosenberg & M. Brazin, 299,
302
Howard, L.O., H.G. Dyar & F. Knab, 321, 337
Howden, H.F., 174, 176, 186, 193
Hoy, J.B. (see Anderson, J.R, & Olkowski, W.),
255,257
Hoyle, G., 264, 302
Hubbell, T.H., 186, 193
Huckett, H.C. (see James, M.T.), 5, 7
Huffacker, C.B., 238, 258
Huffacker, C.B. & R.C. Bach, 238, 241, 158
Hulten, E., 178, 183, 193
human bait, 227
Hutcheon, D.E. & V.S. Chivers-Wilson, 351, 372
hydroxyindoles, 268
hydroxytryptamine (5-HT), 290, 297
h ypheripiformis group, 127, 145-146, 188, 210
Hypherpes, 146
incisus group, 127-120, 172, 188, 210
Infernophilus , 15, 18, 21, 28, 19, 43, 66, 68,71
castaneus, 15, 18, 43, 67, 78, 83
insect saline, 266, 267
iodine, 268, 269
Ixodidae, 2
Iyatomi, K & K. Kanehisa, 264, 302
Jacobowitz, D. & G.B. Koelle, 296, 302
James, M.T., 5, 7
James, M.T. & H.C. Huckett, 5, 7
Jamnback, H.A. (see Stone, A.), 356,372
Javik, M.E., 290, 302
Jeannel, R., 16, 68, 74, 169, 193
Jedlicka, A., 16, 74
Jenkins, D.W. & K.L. Knight, 314, 327,
337-338
Johnson, C.G., 9. 1 4
Johnson, J.G. (see Newhouse, V.F.), 227,
238. 241, 259
Jonkers, A. H. (see Worth, C. B.), 255,
260
Judson, S. (see Richards, H.G.), 174, 194
Juillet, J.A., 231,258
Kandel, E.R. & W.T. Frazier, 296, 302
Kanehisa, K. (see Iyatomi, K.), 264, 302
Kardos, E.H. (see Burdick, D.J.), 252, 257
Kearns, C.W. (see Chang, S.C.), 265,300,
(see Sternburg, J.), 266,283, 298, 304
Kennedy, N.K. (see Roeder, K.), 264, 304
Khan, Z.H. (see Meillon, B. de), 324, 337
Khelevin, N.W., 325, 338
Khromov-Borisov, N.V. & M. J. Michelson,
295, 302
King, P.B., 69, 74, 174, 192
Kiraly, J.K. (see Barton Brown, L.), 297, 299
Klassen, W„ 324, 325, 337
Klassen, W. & B. Hocking, 325, 337
Klock, J.W. & W.L. Biddlingmayer, 255,
258
Knab, F. (see Howard, L.O.), 321 , 337
Knight, K.L. (see Jenkins, D.W.), 324,
327, 337-338; (see Stone, A.), 321 ,
324, 338
Koelle, G.B., 263, 265, 275, 279, 294,297,
302, 303; (see Jacobowitz, D., 296,302),
(see McKinstry, D.N., 275, 303); (see
Voile, R.L, 283,297,305)
Kollros, J. J. (see Tobias, J.M.), 269, 295,
305
Kopine, I.J., 290, 303
Kuntzman, R.G. (see Shore, P.A.), 290,304
Laarman, J.J., 238, 259
LaCasse, W.J. (see Carpenter, M.J.), 309,
313,326, 336
Lacordaire, J.T., 193
Laelapidae, 2
Lampyris noctiluca, 214
Larix laricina, 219
LaRoi, G., 259
xi
Larson, D.J., 1 5
larvicides, chemical, 37 1
Lawson, F.R. (see Chamberlin, J.C.), 222,
257
Leach, G.D.IL, 275, 303
Lebia, 16, 23, 62
Lebiina, 24
Lebiini, 16, 21, 22, 45
key to the larvae of the subtribes, 24
Lecalida, 23, 29, 66, 29, 71
LeConte, J.S., 74, 101, 193, 194
Ledum groenlandicum, 219
Leech, R., 3
Leng, C.W.,74, 101, 194
Leng, C.W. & A.J. Mutchler, 194
Leonard, M.D., 194
Leone, C.A. (see Basford, N.L.), 94, 191
Lepidoptera, 65, 316
larvae, 62
Leptidae, 255
Leptopodoidea,'85
Lesticus, 90
Lewin, V., 63,74
Lewis, S.E., 298, 302
Lewontin, R.C. (see Simpson, G.G.), 231, 260
Libet, B. (see Eccles, R.M.), 296, 301
light traps, 217, 220-222
Lindroth, C.H., 23, 74, 89, 93, 186, 194
Linsley, E.G. (see Mayr, E.), 17, 18, 75
livestock fatalities, 351
suspension of breeding activities, 351-352
declines in the production of milk & beef, 35 2
general losses, 352
Locus ta migratoria, 264
Loding, P.H., 194
Loomis, E.C., 254, 259
Loomis, J., 285, 303
Love, G.J. & W.W. Smith, 222, 23 1 , 238, 259
Lumsden, W.H.R., 255, 259
MacGinitie, H.D., 69, 74
Macintosh, F.C. (see Birks, R., 273, 299), 273,
303
Macintosh, F.C., R.I. Birks & P.B. Sastry, 273, 3(
MacLagen, D.S. & E. Dunn, 9, 14
Madge, R.B., 16,23,74
Magazanik, L.G., N.R. Fruentov, E.R. Roshkova,
R.S. Rybolovlev & M. Mikhelson, 303
Magoon, E.H., 222, 227, 255, 259
Malaise, R.A., 220, 259
Malaise trap, 217, 220
with carbon dioxide, 217, 227, 230
Mallophaga, 1, 2, 3
parasitic on penquins, 2
Mamillaria vivipara, 63
Mannerheim, C. G. von, 75
Mansonia fuscopennata, 242, 243
perturbans , 220, 236, 240
Marshall, J.F., 321,338
Martin, P.S., 69, 75
Martin, P.S. & B.E. Harrell, 186, 194
Maslin, T.P., 169, 194
Matheson, R„ 325, 326, 328, 338
Mattingly, P.F., 252, 259
Maw, M.G., 254, 259
Mayr, E., 17,75
Mayr, E., E.G. Linsley, & R.L. Usinger,
17, 18, 75
McDuffie, W.C. (see Twin, C.R.), 347, 372
McFadden, M.W., 5
McKinstry, D.N. & G.B. Koelle, 275, 303
McLennan, H., 296, 303
McLintock, J., A.N. Burton, II. Dillenberg
& J.G. Rempel, 254, 259
McWade, J.W. (see Steward, C.C.), 310,
322, 338
Mead, J.A.R. (see Shore, P.A.), 290, 304
mealworms, 63
Means, R.G., 322, 338
measurements & ratios (Carabidae), 19
Mecklanburg, C. von (see Dahl, E.), 296,301
Megasteropus , 95, 126, 127, 165
gigas, 126, 165, 166
Meillon, B. de & Z.H. Khan, 324, 337
Mellanby, K„ 9, 14
Mene tries, M., 75
Mesostigmata, 2
Metastigmatia, 2
Metcalf, R.L. (see Winton, M.Y.), 264, 306
methacholine, 263, 275, 279, 295
methylthiocholine, 264
Michelson, M.J. (see Khromov-Borisov, N.V.),
5 295, 302
Microchrysa flavicornis, 5, 7
polita , 5, 6
Mikalonis, S.J. & R.H. Brown, 264, 303
Mikhel’son, M. Ya (see Magazanik, L.G.),303
Milburn, N., E.A. Weiant, & K.D. Roeder,
296, 303
Xll
Millar, J.L. & J.G. Rempel, 351, 372
Milstead, W.W. (see Auffenberg, W.), 174, 191
Mimodromiides, 1 6, 45
Minter, D.M., 255, 259
mites, marine, 2
mesostigmatic, 2
nasal, 2
oribatid, 2
terrestrial trombidiform, 2
Alolops, 89, 90, 94, 127, 146, 156, 158, 169
faber, 109, 125
monoamine oxidase (MAO), 290
Mono group, 101-105, 106, 171, 186, 210
morphological methods (Carabidae), 1 8
mosquitoes, bivdogy of the adult female of
Culicidae, 309- 336
handling & dissection 230
larvae, 13, 219
ovarian development, 242, 244
parous, 2 1 7
sampling methods, 217-261
study area, 217, 219
woodland, 217
Motschoulsky, V. von, 75, 194
Muirhead-Thompson, R.C., 246, 251, 256, 259,
328,338
Mulhern, T.D., 254, 259
Muller, E.H., 174, 194
Mutchler, A.J. (see Leng, C.W.), 194
Myhrberg, H. (see Dahl, E.), 296, 301
Nachmansohn, D. (see Dettbarn, W.), 298, 301
naphthol, 267, 289
naphthylacetate, 267, 289
Narahashi, T. (see Yamasaki, T.), 264, 279, 283,
289, 306
Nelson, R., 252, 259
nematodes, mermithid, 349
neurons, cockroach, 263
Newhouse, V.F., R.W. Chamberlain, J.G. Johnson,
& W.D. Sudia, 227, 238, 241 , 259
Newman, E., 194
Nickerson, M., 290, 297, 303
nicotine, 263, 266, 275, 295
Nielsen, A.T. (see Nielsen, L.T.), 217, 259
Nielsen, L.T. (see M.J. Carpenter), 252, 257
Nielsen, L.T. (see S.J. Carpenter), 333, 334, 337
Nielsen, L.T. & A.T. Nielsen, 217, 259
Nielson, E.T. &D.M. Rees, 326, 338
nitrogen, liquid, 268
nitrogen, mustard, 290
noradrenaline, 263, 268, 290, 294, 297
Norberg, K.A. (see Hamberger, B.), 296,
302
O’Brien, R.D., 264, 303
obsoletus group, 101, 106-109, 171, 187,
210
Ochlerotatus, 313, 322, 324-328
Olin, J. (see Shore, P.A.), 268, 304
Olkowski, W., J.R. Anderson, & J.B. Hoy,
255, 259
Olkowski, W. (see Anderson, J.R.), 255, 257
Olson, A.L., T. H. Hubbell, & H.F. Howden,
194
Omori, N., 9, 14
Oncopeltus fasciatus, 213
Onota, 23, 28, 66, 67
floridana, 80
organophosphates, 264, 265
Oribatidae, 2
Ostlunde, E., 297, 303
ovulum-faber complex, 171
ovulum group, 95, 109, 115-122, 171, 187,
188, 210
Owen, C. & B. Falck, 296, 303
Page, I.H., 290, 303
Panton, W.D.M., 296, 304
Parker, S.L. (see Pearl, R.), 9, 14
Pearl, R. and S.L. Parker, 9, 14
Periplane.ta americana, 213, 163-299
phenoxybenzamine, 263
hydroxide (dibenzyline), 266, 290
Philophuga, biology of, 65-66
key to the species, 30
revision of the genera, 1 5-72
Philophuga amoena, 15, 36
brachinoides, 29, 65, 67, 71 , 79, 80, 83
caerulea, 30, 67,71,78,81
canora, 15, 42
castanea, 18, 43
cobaltina, 15,41
cyanea, 29
horni , 1 5, 36, 41
lauta , 15, 38
obscura, 1 5, 42
puella , 42
purpurea , 31 , 32
uteana , 15,41
viridicollis, 15, 25, 30, 65, 71, 76
xiii
Philophuga viridis, 15, 21, 29, 34, 39, 40, 65,
68, 71, 82
key to the subspecies, 37
Philo tecnus, 44
nigricollis, 44, 60
ruficollis, 60
phylogeny, Carabidae, 66-68
Heteroptera, 85-86
Picea glauca, 2 1 9
Pickard, E. (see Breeland, S.G.), 235, 238, 257;
(see Smith, G.E.), 220, 253, 260; (see Snow,
W.E.), 255, 260
pilocarpine, 263, 266, 279, 296
Pleistocene, 174
Pletscher, A., 290, 304
Plochionus, 16, 21, 28, 30, 66, 68
amandus, 80
pallens , 68
timidus, 25, 76, 78, 80
Populus balsamifera, 219
tremuloides, 219
poplar forest, 219
Portman, R. (see Chen, G.), 275, 300
praying mantis, 279
preservation of larvae (Carabidae), 18
Pringle, J.W.S., 266, 304
Pristonychus complanatus, 63
Procotophyllodidae, 2
Prosimulium gibsoni, 357
Prosser, C.L., 266, 304
Prostigmata, 2
Pterostichini, 89, 93, 94, 169
Nearctic & Palaearctic, 1 69
Pterostichus, 89,94, 102-118, 125, 131, 137, 141,
152, 158, 160, 163, 165
batesellus, 117
carolinensis, 133, 135
chalcites, 94
dejeanellus, 102
lixa ,159
sigillatus, 137
Pucat, A., 314, 322, 325,338
Pumphrey, R.L. & A.F. Rawdon-Smith,265,304
Putnam, P. (see Shannon, R.C.), 9, 13, 14
pyridine-2-aldoxime methiodide (2-PAM), 266,
285, 288, 298
radiant species, 178
Rainey, M.B., G.V. Warren, A.D. Hess & J.S.
Blackmore, 255, 259
Rand, M.J. (see Burn, J.H.), 263, 273,
297, 300; (see Chang, V.), 297, 300
rat baited traps, 222- 227
Rawdon-Smith, A.F. (see Pumphrey, R.J.),
265-304
Ray, D.E. (see Hibbard, C.W.), 174, 193
rearing methods (Carabidae), 19
Rees, D.M. (see Nielson, E.T.), 326,338
Reeves, W.C., (see Bellamy, R.E., 227,
238, 255, 257), (see Hayes, R.O., 239,
258), 238, 259
Reeves, W.C. & McD. Hammon, 238, 260
Reimer, C. (see Bigelow, R.S.), 19, 73
Rempel, J.G. (see Fredeen, F.J.H., 341,
347, 349, 372); (see McLintock, J.,
254, 259); (see Millar, J.L., 351, 372);
309,31 1,324,327,338
Rempel, J.G. & A.P. Arnason, 347, 350,
372
resting mosquitoes, captures in a trailer,
217, 227
Rhinonyssidae, 2
Rhodacaridae, 2
Ribes locus tre, 219
Richards, H.G. & S. Judson, 174, 194
Roberts, R.H., 227, 255, 260
Robertson, F.W. & J. Sang, 9, 14
Robins, E.L. (see Holmes, R.), 285, 288,
302
Roe, A. (see Simpson, G.G.), 231, 260
Roeder, K.D. (see Milburn, V., 296, 303);
263, 275, 279, 283, 295, 304
Roeder, K.D. (see Twarog, B.M.), 264, 279,
283, 297, 305
Roeder, K.D. & N.K. Kennedy, 264, 304
Roeder, K.D. & S. Roeder, 275, 279, 304
Roeder, S. (see Roeder, K.D.), 275, 279,
304
Rohlf, F.J. (see Basford, N.L.), 94, 191
Rohwer, S.A. & G.E. Woolfenden, 174, 194
Rosa acicularis, 219
Rosenberg, P. (see Dettbarn, W.), 269, 298,
301; (see Hoskin, F.C.G.), 299, 302
Roshkova, E.K. (see Magazanik, L.G.), 303
Ross, H.H., 174, 194, 195
rotary sweep net, 217, 222
Rothschild, M. (see Bisset, G.W.), 265, 299
Rubzov, I. A., 347, 372
Rudolfs, W., 238, 260
XIV
Russell, P.F. & D. Santiago, 246, 251, 260
Ryall, R.W. (see Curtis, D.R.), 295, 301
Rybolovlev, R.S. (see Magazanik, L.G.), 303
Saldidae, 85
Sales, S. (see Hamon, J.), 333, 337
Salicornia, 63
Saliternik, Z., 246, 260
Salt c, 219
sand flies, 355
Sang, J. (see Robertson, F.W.), 9, 14
Sarcoptiformes, 2
Santiago, D. (see Russell, P.F.), 246, 251, 260
Sarginae, 5, 6
Sargus bipunctatus, 5, 6
cuprarius, 5, 6
decorus, 5, 6
lucens, 5, 6
viridis, 5, 6
Sarkaria, D.S. (see Brown, A.W.A.), 239, 257
Sastry, P.B. (see Macintosh, F.C.), 273, 303
Savage, D.E. (see Hibbard, C.W.), 174, 193
Savage, L.B. (see Carestia, R.R.), 238, 241, 257
Savit, J. (see Tobias, J.M.), 269, 295, 298, 305
sawfly, larvae of wheat stem, 63
Say, T., 75, 195
Schachter, M. (see Bisset, G.W.), 265, 299
Schafer, J.P. & J.H. Hartshorn, 174, 194
Schaupp, F.G., 195
Schuler, L., 94, 195
sclerophyllous plants, 70
Scott, J., 88
Scudder, S.H., 195
Selander, R.B. & P. Vaurie, 75
Selander, R.K., 174, 195
Sella, stage of, 230
seximpressus group, 127, 139-145, 172,188,210
Shannon, R.C. & P. Putnam, 9, 13, 14
Shannon, R.G., 255, 260
Shelenova, M.F., 333, 338
Shelp, W.D. (see Hopkin, M.R.), 296, 302
Shemanchuk, J.A., 256, 260, 316, 338
Shore, P.A. (see Brodie, B.B.), 297, 300
Shore, P.A. & J. Olin, 268, 304
Shore, P.A., J.A.R. Mead, R.G. Kuntzman,
S. Spector & B.B. Brodie, 290, 304
Shotton, F.W., 186, 195
sigillatus group, 127, 133-139, 172, 188, 210
Simmet, R.P. (see Sommerman, K.M.), 253,260
Simpson, G.G., 94, 195
Simpson, G.G., A. Roe & R.C. Lewontin,
23 1 , 260
Simuliidae, 371 , 372
Simulium arcticum , 341-371
effects on man, 352
aureum, 357
cor bis, 341 , 347
croxtoni, 357
decorum, 357, 359
defoliarti, 341 , 347
furculatum, 357, 359
latipes, 357, 359
luggeri, 357
malyshevi, 341, 347
meridionale , 357, 359
nigricoxum, 341, 347
puge tense, 359
rugglesi, 357
simile, 371
tuberosum, 356, 357, 359
venustum, 354, 357, 359
verecundum, 356, 357, 359
vittatum, 354-359
Sjoquist, F. (see Hamberger, B.), 296, 302
Skiersca, B., 314, 324, 338
Smallman, B.N., 264, 304
Smallman, B.N. & R.W. Fisher, 298, 304
Smith, D.S., 213,263, 304
Smith, D.S. & J.E. Treherne, 264, 297,304
Smith, D.S. (see Treherne, J.E.), 264, 295,
305
Smith, G.E., 251, 260
Smith, G.E., S.G. Breeland & E. Pickard,
220, 253, 260
Smith, T.A. (see Barr, A.R.), 254, 257
Smith, W.W. (see Love, G.J.), 222, 231,
238. 259
Snow, W.E., 251, 260
Snow, W.E., E. Pickard & R.E. Sparkman,
255. 260
sodium chloride, 268
laurylsulfate, 267
hydroxide, 268
sulfite, 268
so Idalis group, 127, 146-155, 173,188,210
Solidago, 219
soldier flies (distribution records in Canada
& Alaska), 5-7
Sommerman, K.M. & R.P. Simmet, 253,260
XV
Southwood, T.R.E., 227, 23 1 , 235, 246, 253,
260
Sparkman, R.E. (see Snow, W.E.), 255, 260
Spector, S. (see Shore, P.A.), 290, 304
spectrofluorometric assay, 263
Spencer, E.Y. (see Colhoun, E.H.), 265, 301
sphagnum, 219
spoliatus group, 95, 109, 110-1 15, 171, 187,
210
spruce, 2 1 9
Stahler, N. (see Terzian, L.A.), 9, 14
Standfast, H.A., 243, 260, 335, 338
Stanley, J., 19, 75
Starke, H. (see Stone, A.), 321, 324, 338
Stebbins, G.L., Jr., 174, 195
Sternburg, J. (see Brady, V.E.), 288, 299, 300
Sternburg, J., S.C. Chang & C.W. Kearns, 266,
283, 298, 304
Steropus, 102, 125
Stevenson screen, 220
Steward, C.C. & J.W. McWade, 310, 322, 338
stimulans group, 3 1 3
Stomis, 90
Stone, A., 321,325,338
Stone, A. & H.A. Jamnback, 356, 372
Stone, A., K.C. Knight & H. Starke, 321, 324,
338
Stratiomyidae, 5
Strickland, E.H., 5,7,341,372
substriatus group, 155-160, 173, 189
Sudia, W.D. (see Newhouse, V.F.), 227, 238, 241,
259
Symphoromyia, 255
synaptic transmission, 263
systematic category, 18
Tabanidae, 255
tabanids, 231
Takeshige, C. & R.L. Voile, 279, 283, 296,
304, 305
Talens, A. (see Dauterman, W.C.), 267, 301
Tauc, L. & H.M. Gershenfeld, 296, 305
Tawfik, M.S., 9, 14
Taylor, D.W. (see Hibbard, C.W.), 174, 193
taxonomic characters (Carabidae), 20
color, 20, 22
external morphology, 20, 22
female ovipositor, 22
male genitalia, 21
Tecnophilus, 15-72
Tecnophilus, biology of, 63-65
key to the species, 45
materials, methods & taxonomic
characters, 16-22
revision of the genera, 15-72
croceicollis, 17, 26, 44,46-59, 65,68,
72,76, 84
key to the subspecies, 60
glabripennis, 60
pilatei, 15, 22, 45, 60, 67, 72, 78, 80
Terzian, L.A. & N. Stahler, 9, 14
tetraethylpyrophosphate (TEPP), 263,266,
268, 288, 298
Theobaldia, 316
thermohygrograph, 220
Thompson, R.P. (see Brown, A.W.A.),
239, 257
ticks, 2
Tobias, J.M., J.J. Kollros & J. Savit, 269,
295, 298, 305
Torre-Bueno, J. R. de la, 75
torvus group, 127, 160-164, 173,210
Townes, H., 220, 260
tranylcypromine, 263, 266, 290
Treherne, J.E., 263, 295, 297, 305; (see
Smith, D.S., 264, 265, 297, 304)
Treherne, J.E. & D.S. Smith, 264, 295,305
Trembley, H.L., 242, 260
tsetse flies, 231, 251, 253
Twarog, B.M. & K.D. Roeder, 264, 279,
283, 297, 305
Twinn, C.R., B. Hocking, W.C. McDuffie
& H.F. Cross, 347, 372
Tydeus tilbrooki, 3
Typha, 219
Udenfriend, S., 269, 305
Unger, H., 294, 297, 305
Unquestedt, U. (see Hamberger, B.), 296,
302
Usinger, R.L., 17, 18 (see Mayre, E., 17,
18,75)
VanAsperen, K., 267, 289, 305 (see
Dauterman, W.C., 267, 301)
Van Dyke, E.C., 195
VanEmden, F.E., 93, 169, 195
Vaurie, P. (see Selander, R.B.), 75
Verheijen, F.J., 254, 260
Viburnum edule, 219
visual attraction trap, 217, 222
xv i
Vockeroth, J.R.. 310, 314, 325, 327, 339
Vogel, W. (see Costa, E.), 290, 301
Voile, R.L., 275, 279, 296, 301; (see De Groat,
W.C., 296, 301 ); (see Geber, G.L., 275, 279,
283, 296, 301 ; (see Takeshige, C., 279, 283,
296, 304)
Voile, R.L. & G.B. Koella, 283, 297, 305
Wada, Y., 9, 13, 314, 320, 324, 339
Wald, G. (see Gregerman, R.I.), 297, 302
Warren, G.V. (see Rainey, M.B.), 255, 259
Warren, M.C.W. (see Wharton, D.H.), 255, 260
Watkins, J.C. (see Curtis, D.R.), 295, 301 ;
(see Milburn, N.), 196, 303
wax moth, 63
Weiant, E.A., 266, 305
Welsh, J.H., 297,306
Welsh, J.H. & H.T. Gordon, 275, 306
Wesenberg-Lund, C., 321, 338
Wharton, D.H., D.E. Eyles & M.C.W. Warren,
255, 260
White, K.E. (see Barr, A.R.), 254, 257
Whitehead, D.R., 174, 195
Wickel, A. (see Chen, G.), 275, 300
Wigglesworth, V.B., 213, 265, 297, 306
Williams, C.B., 235, 260, 314, 316, 338; (see
Fisher, R.A., 235, 258, 316, 337)
Willis, E.R., 238, 260
Willis, M.J. (see Hayes, R.O.), 239, 258
Wilson, I.B., 285, 306
Wilson, I.B. & S. Ginsburg, 285, 306
wing characters, 1 8
Winteringham, F.P.W., 298, 306
Winton, M.Y., R.L. Metcalf & T.R. Fukuto,
264. 306
Woolfenden, G.E. (see Rohwer, S.A.), 174, 194
Worth, C.B. & A.H. Jonkers, 255, 260
Wray, F.C. (see Clark, J.C.), 254, 257, 324, 337
Yamasaki, T. & T. Narahashi, 264, 279, 283,
289. 306
Yates, W.W. (see Gjullin, L.M.), 324, 337;
(see Stage, H.H.), 324, 338
Zhogolev, D.T., 220, 260
zoogeography, Carabidae, 68-72
r-9. - OY)y
Quaestiones
entomologicae
VOLUME V
COMP. ZOOEJ,
LIBRARY
FEB fe; f
V" 1
* i ;v ,
A periodical record of entomological investigations,
published at the Department of Entomology, Uni-
versity of Alberta, Edmonton, Canada.
NUMBER 1
JANUARY 1969
QUAESTIONES ENTOMOLOGICAE
A. periodical record of entomological investigations published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.
Volume 5 Number 1 13 January 1969
CONTENTS
Book Review 1
McFadden - New distributional records for Canadian soldier flies
(Diptera: Stratiomyidae). Part I. Beridinae and Sarginae 5
Tawfik - Effects of population density of Cimex lectularius L 9
Larson - A revision of the genera Philophuga Motschoulsky and
Tecnophilus Chaudoir, with notes on the North American
Callidina (Coleoptera: Carabidae) 15
Book Review
GRESSITT, J. LINSLEY (editor). 1967. Entomology in Antarctica. American Geophysical
Union. Library of Congress Catalogue No. 67-62159. Volume 10, xii + 395 pp., many figs,
and photographs. $17.00 U.S.A.
This book has a large informative introduction by the editor, and then is divided into two
major sections, Systematics and Ecology, for the text.
The Introduction contains five parts as follows: History of Entomological Explorations in
Antarctica; Zoogeographical Summary; The Fossil Record; Dispersal; and Classification.
Entomological exploration began in 1897-1899 with the “Belgica” Expedition under de
Gerlache and is continuing today. The “Belgica” Expedition brought back specimens of
three species of springtails, a wingless midge, specimens of six species of mites, a tick, and
specimens of four species of chewing bird-lice. Up to and including 1967, Gressitt lists 130
species and subspecies (a few are uncertain identifications or records) of terrestrial arthro-
pods in 68 or 69 genera belonging to 29 families in 10 orders, and states that more new
species will be found. Only one marine mite is mentioned by him. The distribution of each
species is given with the list in tabulated form. The Zoogeographic Summary is short but
clear. The Fossil Record is one beetle, Grahamelytron crofti Zeuner, of doubtful family
association, and a few insect wing fragments. Gressitt states that the main dispersal methods
available to insects appear to be winds and birds. The short Classification at the end of the
Introduction is but a key to the orders and some Tamilies and seems a bit out of place. I feel
that it should have been completed to include all the known arthropod fauna of Antarctica.
It is not very useful in its present form.
The Systematics section treats the Acarina, Collembola, Mallophaga, Anoplura, and Dip-
tera.
2
The Acarina are represented by 94 species in the suborders Mesostigmata (mesostigmatic
and nasal mites); Metastigmatia (ticks); Prostigmata (marine and terrestrial trombidiform
mites), Astigmata (sarcoptiformes), and the Cryptostigmata (oribatid mites). Hunter dis-
cusses the mesostigmatic families Rhodacaridae and Laelapidae. He provides neither key nor
diagnoses for all the species. Wilson discusses the remaining mesostigmatic families Rhin-
onyssidae and Halarachnidae (nasal mites) and the metastigmatic Ixodidae (ticks). He pro-
vides diagnoses and drawings for the mites, and keys, diagnoses, and drawings for the ticks.
There are no new species noted. Strandtmann discusses the terrestrial prostigmatic mites. He
provides a key to all the Antarctic species of free-living mites. Brief diagnoses are provided
for the eight previously known species and detailed diagnoses and illustrations are provided
for the 1 4 new species. The known distributions are cited. Newell discusses the marine pros-
tigmatic mites, the Halacaridae. Twenty-eight species are recorded from south of 60° south
latitude. One new species is described and illustrated. Keys are provided for all the genera
and species, and illustrations for the above mentioned species. Newell gives an interesting
account of the family Halacaridae in the polar regions. Atyeo and Peterson discuss the astig-
matic mites, which are represented by Procotophyllodidae and Avenzoariidae. Some illus-
trations are provided. Keys are provided for all genera and species. Wallwork discusses the
cryptostigmatic mites. He lists 18 species and subspecies, three of which are new and pro-
vides a key to the species. He discusses briefly the endemic continental, the South American,
and the circum- Antarctic - Sub-Antarctic elements of the Oribatidae.
The Collembola are represented by 17 species, three of which are new, in 13 genera of
four families in two suborders. Wise provides a key to the species and illustrations of diag-
nostic characters of the new species. He discusses the history of collections of Collembola in
Antarctica, and distribution and origin of this fauna.
The Mallophaga and Anoplura are discussed together, but in two parts. The first, by Clay,
is a short paper on those Mallophaga parasitic on penguins. A key is provided to all the
known species. The second paper, by Clay and Moreby, provides keys and locality lists of
Mallophaga and Anoplura.
Wirth and Gressitt end the Systematic section with the Chironomidae. They discuss the
two known species, one of which is apterous, the other winged. Data on distribution are
given.
The Ecology section provides data on the biology and distribution of selected species.
Janetscheck introduces this section with a broad, but thorough, discussion of Arthropod
ecology of south Victoria land, and a specific discussion on growth and maturity of a
springtail, Gomphiocephalus hodgsoni Carpenter, from the same region. Many graphs, charts
and photographs are included. Gressitt and Shoup follow with a discussion of the ecology of
several free-living mites in the north Victoria land, and Gless with short notes on the bio-
logy of another mite, again from the same region. Wise and Shoup discuss some Collembola
distributions in relation to transects at Cape Hallett, and Tilbrook discusses arthropod ecol-
ogy in the Maritime Antarctic. Strong discusses the ecology and habits of the free-living
arthropods at Palmer Station, Anvers Island, and Gressitt follows with notes on arthropod
populations in the south Shetlands, the Antarctic peninsula, and south Orkneys area.
Murray, Orton, and Cameron end the book with a very short paper on the only Antarctic
3
flea, covering distribution, taxonomic description and biology.
As the editor states in the Preface, “This volume is the first extensive assemblage of stud-
ies on the entomology of Antarctica.” This is true, and it is well done. However, with just a
little more planning, it could have been even better. For instance, the division of the inform-
ation about the Mallophaga and Anoplura into two papers is needless and wasteful. The fig-
ures for both papers are at the end of the second paper, and not with their respective ones.
The division apparently came as a late afterthought, with the result that there is a problem
in the key to the mallophagan genera, and a taxonomic problem. The genus Austrogoniodes
Harrison (1937) is not provided for in the key to the mallophagan genera in the second
paper, and two new species described by Clay in the first paper, A. gressitti Clay 1967, and
A. keleri Clay 1967 are in the second paper as A. gressitti Clay n. sp., and A. keleri Clay n.
sp., despite the fact that they were described in the first paper.
Some of the ecology papers could have been joined together, such as those by Strong and
Gressitt, to make the information more useful and to thus avoid some contradictions. For
instance, Strong writes that the eggs of Tydeus tilbrooki Strandtmann (1967) have not yet
been encountered (page 371 ), yet Gressitt describes them (page 382).
I feel that there should have been some discussion of the single Antarctic flea in the Sys-
tematic section.
It is interesting to note that only one of the acarologists (Newell) mentions the publication
by Dalenius, The Acarology of the Antarctic Regions, in Biogeography and Ecology in
Antarctica (van Mieghem and van Dye, editors), Junk, The Hague, 1965.
As Gressitt gave a good coverage in the Introduction on the history of Antarctic ento-
mology, I feel that there is some unnecessary repetition in several of the systematic papers.
However, for separate papers in standard scientific journals, the introductory history for
each group would have been well placed. I think that some of the ecology titles should have
read “Habitats” rather than “Ecology”.
The book lacks an index, and for the ecological parts this is sorely needed, as there is over-
lap of discussion of many species.
The proof reading for Entomology of Antarctica was well done; most of the illustrations,
figures and graphs are of high quality, though some small graphs are a little “busy” while
others are unnecessarily large and with little information. The printing is clear and of a pleas-
ant reading size. The binding is well done with cloth, and the paper is high gloss kaolin
(which might not be a good feature in humid climates).
I believe that this book is worthwhile in spite of the above mentioned shortcomings and
should get wide circulation. The price, I think, is a little too high.
The value of this book to biologists will increase significantly when it becomes possible to
compare the Antarctic insect fauna with its Arctic counterpart. This will be possible when the
systematic and ecological work done under the program “Studies on Arctic Insects” by the
Canadian government is completed and published.
Robin Leech
'
NEW DISTRIBUTIONAL RECORDS FOR
CANADIAN SOLDIER FLIES (DIPTERA: STRATIOMYIDAE)
PART I. BERIDINAE AND SARGINAE
M. W. McFADDEN Quaestiones entomologicae
Department of Entomology 5 5.7 7959
Washington State University
Pullman, Washington 99163
New distribution records in Canada and Alaska are presented for the following species of
Stratiomyidae: Allognosta fuscitarsis, A. obscuriventris, A. brevicomis, Actina viridis, Beris
califomica, Exodontha luteipes, Sargus bipunctatus, S. cuprarius, S. decorus, S. viridis, S.
lucens, Microchrysa polita, and M. flavicomis.
Except for the contributions of Strickland (1938, 1946), James (1951), and James and
Huckett (1952), information on the distribution of Stratiomyidae in Canada and Alaska has
not been revised since Curran’s synopsis (1927).
Work in progress on a revision of the Stratiomyidae of Alaska and Canada has revealed the
following new records.
SUBFAMILY BERIDINAE
Allognosta fuscitarsis (Say)
Previously recorded from Nebraska to Quebec; the new records include specimens col-
lected from Manitoba and Nova Scotia and indicate that the range of this species covers the
eastern half of Canada.
Allognosta obscuriventris (Loew)
The species, previously known only from Ontario and Quebic, is now known to occur in
Manitoba and Nova Scotia and apparently is sympatric with A. fuscitarsis.
Allognosta brevicomis Johnson
All previous records for the species have been from the northeastern section of the United
States. New records show that its range can be extended in Canada to include Quebec,
Ontario, Alberta, and British Columbia.
Actina viridis (Say)
Previously known to occur in Canada from Newfoundland to Alberta; new records from
Nova Scotia, New Brunswick, and British Columbia confirm that this species extends from
coast to coast.
6
McFadden
Beris califomica James
The species was not previously recorded from Canada or Alaska but is now known to
occur in the southern portion of British Columbia. It has also been recorded from Vancouver
Island.
Exodontha luteipes (Williston)
The species has been recorded from Vermont and New Hampshire in the northeastern
United States and from Idaho in the northwest. It is now known to occur in the provinces of
Alberta and British Columbia.
SUBFAMILY SARGINAE
Sargus bipunctatus (Scopoli)
Previously known only from the states of Oregon and Washington where it was originally
introduced (James 1960), the species has now extended its range to include southeastern
British Columbia.
Sargus cuprarius (Linnaeus)
In Canada, the species is known to occur from Quebec westward to British Columbia.
Specimens collected from Nova Scotia indicate that the range now extends from coast to
coast.
Sargus de corns Say
The species has been reported as having a range extending from Alaska to the province of
Quebec. With new records from Nova Scotia and New Brunswick, the range now extends
from coast to coast.
Sargus viridis Say
Previously known to occur in Alaska, its range extended eastward to the province of
Quebec. New records from New Brunswick indicate that the range actually extends from
coast to coast in Canada.
Sargus lucens Loew
The species has not been previously reported to occur in Canada. All new records are from
the province of Ontario.
Microchrysa polita (Linnaeus)
The species is reported to have a range extending from coast to coast in Canada. The
single new record is from the province of New Brunswick which serves to verify the fact that
the species is established in the Atlantic provinces.
Soldier Flies
7
Microchrysa flavicomis (Meigen)
The species, previously known to have a range extending from the state of Washington to
the province of Newfoundland, is now known to occur in the provinces of Saskatchewan,
Alberta, and British Columbia.
REFERENCES
Curran, C.H. 1927. Synopsis of the Canadian Stratiomyidae (Diptera). Trans. R. Soc. Can.
21 : 191-228.
James, M.T. 1951. The Stratiomyidae of Alberta (Diptera). Proc. ent. Soc. Wash. 53(6) :
342-343.
James, M.T. 1960. The soldier flies or Stratiomyidae of California. Bull. Calif. Insect Survey
6(5) : 79-122.
James, M.T. and H.C. Huckett. 1952. The Diptera collected by I.O. Buss in southwestern
Yukon Territory during the summer of 1950. Can. Ent. 84(9) : 265-269.
Strickland, E.H. 1938. An annotated list of Diptera of Alberta. Can. Res. D. 16 : 175-219.
Strickland, E.H. 1946. An annotated list of Diptera of Alberta. Additions and corrections.
Can. Res. D. 24 : 157-173.
EFFECTS OF POPULATION
DENSITY ON CIMEX LECTULARIUS L.
9
MONIB SA YED TA WFIK Quaestiones entomologicae
Department of Entomology j 9.J4 1969
University of Alberta, Edmonton.
In C. lectularius, population density affects fecundity and the duration of the nymphal
stadia, the preoviposition period, and life, apparently through a contact stimulus. The
effects reach optima at a population density of 4 to 8 insects/ cm2.
Many studies have been made of the environmental factors that influence development
and behavior of C. lectularius. Also, quantitative studies of natural populations have been
conducted by Mellanby (1939) , Omori (1941), and Johnson (1942). Although the effect
of population density on the physiology and ecology of many insects has received much
attention, no detailed investigations have been carried out on C. lectularius. High population
density, or overcrowding, is often accompanied by a shortage of food. The effect of food
quantity free from the effect of population density was studied (Tawfik 1968). The aim
of this study is to separate the effect of density itself from that of food quantity.
Overcrowding has a detrimental effect on insects in various ways; slower growth rate,
increased mortality, smaller adults, and lower fecundity (Pearl and Parker 1922, Chapman
1928, Shannon and Putnam 1934, Bodenheimer 1938 and 1955, MacLagen and Dunn 1936,
Crombie 1942, Robertson and Sang 1944, Terzian and Stahler 1949, Bar-Zeev 1957, Wada
1965). Most investigators have attributed the effect mainly to the mechanical disturbance of
insects by each other. Some have demonstrated that most of these effects are dependent on
the quantity and quality of food.
METHODS
Density ranging between 2 and 128 insects per surface area of 8 cm2 of a piece of folded
filter paper (2x2 cm) in 2 x 7 cm specimen tube was studied. Relating the density to the
surface area of the folded filter paper is more important than relating it to the volume of
the specimen tube because bedbugs are dorso-ventrally flattened, always aggregate on the
surface of the folded filter paper, and thus essentially live in a two dimensional habitat.
Data representing a density equal to 1 per 8 cm2 was taken from Tawfik (1968) when the
insects were fed till engorgement every 2 days. Eggs were taken from the standard culture
and put in 4 x 4 x 1 .5 cm plastic boxes. First instar nymphs were taken as soon as they
hatched and were put in 7 x 2 cm specimen tubes together with 2 x 2 cm folded filter
paper. Eight experiments were conducted in which the numbers of nymphs per tube were
2, 4, 8, 16, 32, 64, and 128 respectively (i.e. 0.25, 0.5, 1, 2, 4, 8, and 16/cm2). In all
the experiments the insects were allowed to feed on the second day after hatching and
twice every week thereafter. For feeding, the insects from each tube were transferred to a
4 x 4 x 1.5 cm plastic box and were allowed to feed on human blood through organdie
which covered a 3 cm diameter hole in the lid of the plastic box. Observations were carried
out daily and the effects of the population density on the duration of the nymphal stadia,
preoviposition period, fecundity, longevity, and mortality rate were studied.
10
Tawfik
With a constant food supply for the insects in all the experiments, the effect of popul-
ation density could be due to either visual stimulus, olfactory stimulus, contact stimulus or
combinations of these stimuli. All the experiments were conducted in the dark and the in-
sects were exposed to the light only for ten minutes for examination to rule out the visual
stimulus. An additional experiment was conducted to test whether the effect of population
density is due to an olfactory or a contact stimulus. In this experiment 40 plastic boxes,
similar to those used for feeding, were used. The lids of each 2 were glued together forming
a combination of 2 chambers separated by the organdie on the inside of the lids. In one of
these chambers a single first instar nymph was put with a piece of folded filter paper. In the
other chamber 64 first instar nymphs were put. The insects were fed twice every week on
human blood and the effect on the duration of the nymphal stadia of the single nymph was
recorded. After the fifth moult two insects of opposite sex were put in one chamber instead
of one insect and the effects on preoviposition period, fecundity, and longevity were
studied. All the experiments were done at 80 F and 75% relative humidity.
On the Duration of the Nymphal Stadia
The effect of population density on the duration of the nymphal stadia is shown in fig.
1 . Population density does not show a clear effect on the duration of the first nymphal sta-
dium as this ranges between 3.6 and 4 days. On the other hand, the duration of the other
four stadia decreased with the increase of population density. Olfactory stimulus does not
seem to be involved in the effect on the duration of the nymphal stadia because there is no
significant difference between the results of the experiment with the double chamber com-
bination and those at density 1 and 2 (fig. 1).
On the Duration of the Preoviposition Period
The results are shown in fig. 2. Increasing the population density appears to decrease the
minimum duration of the preoviposition period but beyond density 32 has no significant
effect. Olfaction has no effect on the duration of the preoviposition period; the value ob-
tained in this experiment does not show a significant difference from that when the popul-
ation density was either 2 or 4.
On Fecundity
The relationship between the number of eggs laid per female and the population density
is shown in fig. 3. Population density of 2 or 4 does not have a significant effect on the
number of eggs laid per female. On the other hand, increasing the population density causes
an increase in the number of eggs laid per female. This increase in the number of eggs
reached a maximum when the population density was 64 and then decreased again. As
shown in fig. 3, the number of eggs laid per female per day in the different population den-
sities ranged between 2.1 and 2.4. Fig. 3 also shows that there is no olfactory stimulus in-
volved in the effect of the population density on fecundity of the females.
On Longevity of the Adult Stage
Fig. 4 shows the effect of population density on the longevity of the adult stage. The
longevity of both the female and the male increases with the increase in the population den-
sity. This increase in longevity reaches a maximum at a population density of 32 and then
decreases with further increase in population density. There is no olfactory stimulus in-
volved in the effect of population density on the longevity of the females or the males (fig.
4).
Duration in days
Population Density
11
Fig. 1 . Effect of population density of C. lectularius on the duration of the nymphal stadia.
Fig. 2. Effect of the population density of C. lectularius on the duration of the preovipos-
ition period.
Fig. 3. Effect of population density of C. lectularius on the number of eggs laid per female
and the number of eggs laid per female per day.
Fig. 4. Effect of the population density of C. lectularius on the longevity of males and fe-
males.
Eggs/?/ day
Population Density
13
On Mortality Rate
Table 1 shows the effect of the population density on the percentage of mortality in
the different instars of C. lectularius. Increasing the population density beyond 8 causes a
decrease in the percentage of insects that reach the adult stage.
TABLE 1. Effect of population density on the percentage mortality in the different instars
of C. lectularius.
Percentage of the total died as
Density
DISCUSSION
From these investigations the effect of population density on C. lectularius was apparent
and it was clear that neither a visual stimulus nor an olfactory stimulus was involved in that
effect. The effect of population density on the duration of the nymphal stadia, preovipos-
ition period, fecundity, and longevity reaches an optimum at a population density of 32 to
64 insects per 2 cm2 folded filter paper in 2 x 7 cm specimen tube. Population density
seems to influence the bedbug through the stimulations of increased mutual contact. This
influence was also suggested for mosquito larvae by Bar-Zeev (1957) and Shannon and
Putnam (1934). However, Wada (1965) claimed that the situation seemed to be more com-
plex and that neurophysiological processes might be involved. Duration of the nymphal
stadia and the preoviposition period in the females decreased with the increase of popul-
ation density. It is difficult from these experiments to specify the manner in which popul-
ation density produces its effect through increased mutual contact. It seems that increasing
the population density may increase the temperature of the microclimate as C. lectularius is
always found in aggregates. This increase in the temperature may cause the decreases in the
duration of the nymphal stadia and the preoviposition period.
Neurophysiological processes may also be involved, but more experiments are required
to prove that. Such experiments should relate population density to endocrine activity,
moulting, and reproductive functions. Although the number of eggs laid per female in-
creased with the increase in population density to a maximum when the density was 64, it
seems that this effect on fecundity resulted from the effect of population density on lon-
gevity of the adult stage rather than the effect of population density on fecundity, since
there was no significant difference in the number of eggs laid per female per day at the dif-
ferent population densities.
14
Tawfik
ACKNOWLEDGMENTS
I am indebted to Professor B. Hocking, Department of Entomology, University of Al-
berta, for his guidance and stimulating discussions during the course of this work. I am
grateful to the late Dr. R.L. Usinger of the University of California, Berkeley, who was my
external examiner. I also wish to acknowledge the financial assistance of the U.S. Army
Grant No. 63-G83 (Hocking trust) which made this study possible.
REFERENCES
Bar-Zeev, M. 1957. The effect of the density of the larvae of a mosquito and its influence
on fecundity. Bull. Res. Coun. Israel (B), 6B: 220-228.
Bodenheimer, F.S. 1938. Problems of animal ecology. Oxford University Press, London.
Bodenheimer, F.S. 1955. Precis d’ecologie animale. Payot, Paris.
Chapman, R.N. 1928. The quantitative analysis of environmental factors. Ecology 9 : 111-
122.
Crombie, A.C. 1942. Effect of crowding upon the oviposition of grain infesting insects. J.
exp. Biol. 19 : 311-340.
Johnson, C.G. 1942. The ecology of the bedbug, Cimex lectularius L., in Britain. J. Hyg.
Camb. 41 : 345-461.
MacLagen, D.S. and E. Dunn. 1936. The experimental analysis of the growth of an insect
population. Proc. roy. Soc. Edinburgh 55: 126-139.
Mellanby, K. 1939. The physiology and activity of the bedbug ( Cimex lectularius L.) in a
natural infestation. Parasitology 31 : 200-211.
Omori, N. 1941. Comparative studies on the ecology and physiology of common and trop-
ical bedbugs, with special references to the reactions to temperature and moisture. J.
med. Ass. Formosa 60(4) : 555-729.
Pearl, R., and S.L. Parker. 1922. Experimental studies on the duration of life. Data on the
influence of density of population on the duration of life in Drosophila. Amer. Nat. 56 :
312-321.
Robertson, F.W. and J.H. Sang. 1944. The ecological determinant of population growth in
Drosophila culture. Proc. R. ent. Soc. Lond. 132 : 258-277.
Shannon, R.C. and P. Putnam. 1934. The biology of Stegomyia under laboratory condi-
tions. (1) The analysis of factors which influence larval development. Proc. ent. Soc.
Wash. 36 : 185-216.
Tawfik, M.S. 1968. Effects of the size and frequency of blood meals on Cimex lectularius
L. Quaest. ent. 4(3) : 92-1 1 1 .
Terzian, L.A. and N. Stahler. 1949. The effects of larval population density on some labor-
atory characteristics of Anopheles quadrimaculatus Say. J. Parasit. 35 : 487-498.
Wada, Y. 1965. Effect of larval density on the development of Aedes aegypti (L.) and
the size of adults. Quaest. ent. 1(4) : 223-249.
A REVISION OF THE GENERA PHILOPHUGA
MOTSCHOULSKY AND TECNOPHILUS
CHAUDOIR WITH NOTES ON THE NORTH AMERICAN
CALLIDINA (COLEOPTERA: CARABIDAE)
DA VID J. LARSON
Department of Biology
Quaestiones entomologicae
5 : 15-841969
University of CalgaryCalgary , Alberta
A taxonomic revision of the species of the North American genera Philophuga Motschoul-
sky and Tecnophilus Chaudoir is presented. Four species and four subspecies of Philophuga
are recognized. The taxa Philophuga homi Chaudoir and Philophuga amoena LeConte are
grouped with P. viridis Dejean to form a polytypic species, and P. viridis klamathea new sub-
species is described. Infernophilus new genus, is erected to contain the species castaneus
Horn, formerly included in Philophuga. The following names are synonymized for the first
time: P. lauta Casey (= P. viridis viridis Dejean), P. canora Casey (= P. viridis amoena
LeConte), P. obscura Casey (= P. viridis amoena LeConte), P. cobaltina Casey (= P. viridis
horni Chaudoir), P. uteana Casey (= P. viridis horni Chaudoir). The name Tecnophilus
pilatei Chaudoir is removed from synonymy to the name of a species. Two subspecies of a
second species of Tecnophilus are recognized; T. c. croceicollis Menetries and T. c. peigani
new subspecies.
Geographical variation among the populations of Philophuga viridis Dejean and Tecno-
philus croceicollis Menetries is analyzed.
The structure of the female stylus is described for the Callidina and the variation pattern
of this sclerite is used to erect a classification of the North American species of this group.
The method of oviposition and the habitats of the adults are described, as well as the lar-
vae of Philophuga viridicolhs and Tecnophilus croceicollis.
My interest in this group of carabids began with the discovery of specimens of Tecno-
philus croceicollis Menetries in southern Alberta. Examination of specimens of T. crocei-
collis from other localities indicated that considerable geographic variation was exhibited by
specimens belonging to this species, and also suggested to me that the genus Tecnophilus
contained more than the one recognized species. I decided to include the genus Philophuga
in this study because of the similarities in both morphological characteristics and distribu-
tion patterns shown by members of the genera Philophuga and Tecnophilus. I undertook a
cursory study of the subtribe Callidina to elucidate the relationship between these two gen-
era, and their relationships to the genus Callida.
16
Larson
The complexity of the tribe Lebiini has led many authors to divide it into subtribes. The
concept of the supra-generic grouping, the Callidina, was first introduced by Chaudoir
(1872). Chaudoir’s Callidides was a heterogeneous assemblage of lebiine genera which in-
cluded some genera not belonging here, and excluded others, such as Philophuga, previously
described by Motschoulsky 1859, which are certainly callidines. When Chaudoir (1877)
later described Tecnophilus, a callidine genus in even the strictest sense, he placed it in his
unspecified grouping Mimodromiides.
Horn’s (1882) synopsis of the tribe Lebiini was the next, and last comprehensive work
on the North American Callidina. Although Horn did not recognize any formal subtribal
groupings, he did indicate that a close relationship existed between Callida, Philophuga and
Plochionus. Without reference to the genitalic structures, Horn described castanea as a mem-
ber of Philophuga. He also suggested that a relationship may exist between Tecnophilus
Chaudoir and Euproctus Sober (= Euproctinus Leng and Mutchler) and that these two gen-
era are rather separate from Callida.
Since Horn’s time, the only work on a large group of North American lebiines has been
Madge’s (1967) revision of Lebia Latreille.
Casey (1913 and 1924) described six species of Philophuga, five of which are here regar-
ded as conspecific with previously described species.
Jeannel (1942 and 1949) treated the lebiine faunas of France and Madagascar respective-
ly. He recognized the formal grouping Callidinae but did not present any truly diagnostic
characters.
V
Recently, two major works by Jedlicka (1963) and Habu (1967) have appeared, dealing
with the Asian Lebiini. Habu presents the best subtribal classification yet proposed, based
largely on the structure of the legs, mandibles and female ovipositor. Habu’s definition of the
Callidina is followed in part in this study.
MATERIALS, METHODS, AND TAXONOMIC CHARACTERS
Materials
I have examined over 1 200 specimens of adults and larvae of the genera Philophuga and
Tecnophilus in the course of this study. Most of these specimens were obtained on loan from
various museums in Canada and the United States. Many specimens of Tecnophilus were col-
lected by myself on a number of field trips to southeastern Alberta, and on an extended trip
through southwestern United States. Larvae were reared in the laboratory from eggs laid by
captive adults.
Following the description and discussion for each species and subspecies, a list of the
localities from which specimens have been examined is presented. The localities are listed
alphabetically, in the following order: country, province or state, county and specific local-
ity. Following this, the collector’s name and the museum in which the specimen is stored are
listed in parentheses.
Abbreviations for museums from which specimens were seen, are as follows: AMNH -
American Museum of Natural History; Car. M. - Carnegie Museum; CAS - California Aca-
demy of Sciences; CNC - Canadian National Collection, Ottawa; CNHM - Chicago Natural
History Museum; CU - Cornell University; DAL - Canada Department of Agriculture Re-
search Station, Lethbridge, Alberta; INHS - Illinois Natural History Survey; IUM - University
Carabidae
17
of Idaho; KUM - University of Kansas; MUB - University of Montana, Bozeman; MCZ
Museum of Comparative Zoology; OUM - Oregon State University; SJSC - San Jose State
College, California; UASM - University of Alberta, Strickland Museum; USNM - United States
National Museum; and WUM - University of Washington, Seattle.
Methods
General
Observation and comparison were the methods used (Ball 1 966). Observations were made
of both morphological and biological features of both preserved and living specimens of the
subtribe Callidina. Using these observations, comparisons were made between different pop-
ulation samples in order to ascertain similarities and differences. Most observed characteris-
tics were compared independently and were weighed subjectively depending upon the cir-
cumstances. That is, in one situation a given character may have been regarded as possessing
no discriminatory value, while in another situation it may have been judged important. How-
ever, in dealing with variation in one species, Tecnophilus croceicollis , several characters
were compared simultaneously and the weights for each of these characters were calculated
according to their individual statistical discriminatory value, regardless of other considera-
tions.
Criteria for Species, Subspecies and Genera
The multi-dimensional definition of the species (Mayr 1963) has been used as the under-
lying basis for the taxonomy in this study. The taxa assigned to the species category are less
arbitrary than taxa assigned to lower or higher categories, hence the species category forms
the basis around which a classification is built. Because it is difficult to make the necessary
tests on a population to prove the principal criterion for recognizing a species, that is, gene-
tical isolation from other such groups, this information must be obtained indirectly.
In this study, evidence interpreted as indicating specific identity, is provided by: forms
which overlap geographically but do not intergrade in their diagnostic characters in the area
of overlap; and allopatric forms in which geographically intermediate specimens do not show
intermediate states in their diagnostic characters. For example, the two species Tecnophilus
croceicollis Menetries and T. pilatei Chaudoir are primarily allopatric, and their members can
be consistently distinguished from one another on the basis of several characters. However,
in the vicinity of Brownsville, Texas, the ranges of these two species overlap, yet the diag-
nostic characters still hold and permit complete segregation of members of each of these
species. I assume that maintenance of these differences is the result of reproductive isolation
between these two groups.
I have recognized subspecies only in cases of concordant non-clinal variation in two or
more characters. I have not recognized populations along, or at the end of dines as being
subspecifically distinct. Rather an attempt has been made to point out such dines and to
describe them when they have been recognized (Mayr, Linsley, and Usinger 1953). In one
18
Larson
species, Tecnophilus croceicollis Menetries, clinal variation occurs in many characteristics;
however, two subspecies of this species have been recognized. The new subspecies, T. c.
peigani, was recognized because a number of characters, many of which varied clinally,
showed a sharp change through a relatively short distance to produce two forms which
were remarkably different in general appearance, and could be readily distinguished from
one another.
A genus has been defined as “a systematic category [= taxon] including one species or a
group of species of presumably common phylogenetic origin, which is separated from other
similar units by a decided gap” (Mayr, Linsley, and Usinger 1953). However, the gap neces-
sary to distinguish between groups of species belonging to different genera cannot be object-
ively defined. Rather, this must be decided by the taxonomist on the basis of his knowledge
and experience in the group. Consistency in deciding the nature and size of the gap (degree
of morphological difference) required to separate related genera is imperative in producing a
uniform and workable classification.
In this study, I have recognized the previously described genera, Callida, Philophuga and
Tecnophilus, and have used the degree of morphological difference between these three
genera as a guideline by which to recognize other genera within this subtribe. The species
Philophuga castanea Horn differs from the other species included in the genus Philophuga
by at least as much as these species differ from the species included in the genera Callida
and Tecnophilus. For this reason, I removed castanea from Philophuga and proposed the
new genus Infemophilus to contain it. Recognition of Infemophilus is consistent with the
maintenance of Callida, Philophuga and Tecnophilus as distinct genera.
Morphological Methods
To examine characters of the wings and of the genitalia, the specimens were first relaxed
by placing them in boiling water for five to ten minutes. After this time, the specimens were
sufficiently pliant to allow the elytra to be lifted so that the hind wings could be examined.
This was adequate to determine the relative size of the wings. However, in cases where it was
necessary to study the pigmentation of the wings, the hind wing was removed by breaking it
off near the base. The wing was unfolded and first studied in water, then flattened out and
glued to a card which was pinned beneath the specimen.
Male and female genitalia were removed from the relaxed specimens by breaking the sup-
porting membranes around the genitalia with a hooked insect pin. The genitalia were re-
moved with a pair of fine forceps. After dissection, the genitalia were placed in cold 10%
KOH and allowed to stand overnight to clear the sclerites. The male genitalia were then ex-
amined with a binocular microscope. It was necessary to mount the female genitalia on slides
and examine them with a compound microscope at higher magnification.
Preservation of Larvae
To prevent distortion and discoloration of specimens, larvae were killed by dropping them
directly into gently boiling water. They were quickly removed and preserved in 70% ethanol.
This method of killing extends the larvae to their maximum length and preserves color and
hence facilitates later study.
Carabidae
19
Rearing Methods
Adult specimens of Tecnophilus were reared in the laboratory to obtain eggs and larvae.
After collection, five or six specimens were placed in each jar, where mating readily oc-
curred. After mating, the females were removed and isolated in small jars which contained
soil a half inch deep and several twigs that permitted the insects to climb. Pieces of meal-
worm were added for food, and water was sprinkled into the jar weekly.
If adequate food and moisture were provided, the females began to oviposit within a week
after mating. The eggs were removed as they appeared, and were placed on a moistened blot-
ter in a petri dish where they were left until hatching occurred. Newly emerged larvae were
isolated in small vials and were fed pieces of various soft-bodied insect larvae. High humid-
ity was maintained by placing a piece of moistened blotting paper in the neck of the vial.
Mortality was high, but a number of larvae were reared to the second and third instars.
Measurements and Ratios
Measurements were made of representative specimens of all species and subspecies dealt
with in this study. Although they did not provide diagnostic characters for any species,
measurements and the ratios derived from them were valuable in the analysis of intra-speci-
fic variation. A micrometer eyepiece in a Wild M5 stereoscopic microscope at a magnifica-
tion of X 50 was used for these measurements.
The total length (TL) of adult specimens was obtained by the addition of three measure-
ments: the length of the head from the anterior margin of the clypeus to the mid-point of
the occipital groove (LH); the length of the pronotum along the mid-line (LP); and the
length of the left elytron from the apex of the scutellum to the apex of the elytron (LE).
Total length (LH + LP + LE) obtained in this way was not affected by contraction or cur-
vature of the specimen. Other measurements taken of adult specimens and their abbrev-
iations are: Width of pronotum (WP) - width across widest point of pronotum; maximum
width of head (WHi) - width of head across eyes; width of head between eyes (WH2) -
minimum width of fro ns between eyes. Measurements taken of larval specimens are: Width
of head (WH) - width across widest part of head capsule; width of pronotum (WP) - width
across widest part of pronotum; total length (TL) - one measurement of length from the
anterior margin of the nasale to the posterior margin of abdominal segment 10. All ratios
used in the text are derived from the above measurements, and are expressed in terms of the
above abbreviations.
Analysis of Variation
Specimens from populations that differ slightly and possibly discordantly in a number of
characters, may be assigned to the correct population by means of the discriminant function.
In the study of such groups, the traditional methods of taxonomy, observation and compar-
ison, do not work well for the mind is incapable of making the numerous summated judg-
ments required.
The discriminant function used in this study has been developed by Fisher (1936) and
outlined by Stanley (McGill University unpublished MS). The method has been used or dis-
cussed by Goulden (1952) and Bigelow and Reimer (1954).
20
Larson
Interpretation of the discriminant function is similar to that used for the hybrid index
(Freitag 1965). Using this method, a specimen which deviates from the other members of its
population in only one or a small percentage of its total characters, is still assigned to its
correct population because of its greater overall similarity to members of that population.
This reduces the possibility of mis-identification due to the over-weighting of one character.
Illustrations
Illustrations are presented to augment descriptions. These were prepared with the aid of
an ocular grid in a stereoscopic microscope.
The male genitalia have been illustrated for many species even though they usually present
few useful characters. Stippling has been used to indicate the folding pattern of the endo-
phallus but as this folding pattern varies with small changes in inversion, it should not be
considered as important in identification.
The slender sub-apical spines on the stylus of the female ovipositor have been included in
illustrations of the styli. Both dorsal and ventral spines have been shown as though they oc-
curred in the same plane, as this is the way they appear when the stylus is cleared in potas-
sium hydroxide and mounted on a microscope slide.
Distribution maps are presented for all species and subspecies. Also maps summarizing
data on intra-specific variation in the species Philophuga viridis and Tecnophilus croceicollis
are included.
Taxonomic Characters
Taxonomic Characters of Adults
Color. - Within the Callidina, color is useful in the identification and classification of
species, and to a limited extent, genera. Non-metallic colors are described by the terms
yellow, testaceous (brownish-yellow), rufous (reddish), brown, piceous and black. The color
black often has a metallic sheen associated with it. Metallic colors are usually blue or green.
To describe variation in certain species, intermediate colors are designated in terms such as
blue-green (green predominant) or greenish-blue.
External morphology. - Punctation and vestiture of the body, while extremely useful for
the recognition of species and subspecies with Philophuga and Tecnophilus, are difficult
characters to interpret and quantify. As the setae composing the dorsal vestiture of the body
are situated in usually distinct punctures, the punctation of the body often gives an indi-
cation of the vestiture. However, in specimens from certain populations, the body is deeply
punctate, but the punctures do not bear setae, or bear very inconspicuous setae. Care must
be taken to avoid confusing these specimens with normally setose specimens that have been
rubbed. The depth and density of the punctures in the elytral striae and on the elytral inter-
vals are important for identification.
To describe the depth of punctation of the body and the depth of the impression of the
elytral striae, the terms lightly, moderately and coarsely have been used. These terms are
strictly comparative, and have been avoided as much as possible in keys. However, they have
been used in descriptions in a comparative sense. Similarily, the terms sparse, moderate and
dense have been used to describe vestiture.
Carabidae
21
The shape of the head shows both intra- and inter-specific variation. Differences in the
convexity of the eyes and the constriction of the neck present characteristic forms for cer-
tain species. For all species, values for the ratio of the width of the head across the eyes to
the minimum width of the head between the eyes has been presented in an attempt to quan-
tify differences in eye convexity. This ratio is not diagnostic of any species but it provides an
index of the head shape.
The post-ocular pinch is an impression just behind the posterior dorsal margin of the eye.
This pinch is evident but small in most of the Callidina. However, in some of the neotropical
species of Callida and in some species of Cymindis this pinch is very strongly developed.
The mouthparts were not found to provide any characters useful for species recognition.
They are of value at the generic level, but even here their importance has been overestimated
in past works.
The shape of the pronotum is of appreciable diagnostic value. In Philophuga, differences
in pronotal shapes between related species are often slight and may be within the range of
normal intra-specific variation. In Tecnophilus the shape of the pronotum is much more
characteristic of a species, and is usually rather constant within a population sample. The
pronotum has been illustrated for most species and subspecies.
The wings present several characters which are considered very important in Philophuga.
Wing reduction occurs in this genus and has been used to distinguish between several sub-
species of viridis. Also, the degree to which the membranous areas of the wings are pig-
mented is useful in the recognition of certain species. The wing venation was not found to
offer any characters of diagnostic value at either the specific or generic levels.
Habu (1967) made extensive use of the structure of the legs in his classification of the
Japanese Lebiini. In the present study, only two characteristics of the legs have been used:
the structure of tarsal article 4; and the structure of the tarsal claws. Two forms of tarsal
article 4 are found among the Nearctic Callidina, the bilobed form and the emarginate form.
These are illustrated in figs. 28 and 29 respectively. The tarsal claws of members of the genus
Tecnophilus are simple, while members of the other North American callidine genera possess
pectinate claws.
Characteristic arrangements of setae on the ventral surface of the abdomen provide useful
characters for the recognition of two genera. The usual arrangement of setae is: visible sterna
2 to 5 each with a pair of medial setae, sternum 6 with one to three pairs of anal setae. How-
ever, members of the genus Plochionus bear, in addition to the above-mentioned setae, a
distinctive arrangement of setae on the lateral margins of sterna 4, 5 and 6 (fig. 26). Mem-
bers of the genus Infernophilus lack the lateral setae, but sternum 6 possesses a pair of dense
brushes of setae (these brushes are much denser in the male than in the female, fig. 27).
Male genitalia. - The structure of the male genitalia is rather consistent within the species
of Philophuga and Tecnophilus. The aedoeagus (intromittent organ, see Torre-Bueno 1962)
is slightly arcuate, with a prominent rounded apex. The basal piece, the portion of the
aedoeagus surrounding the basal orifice, varies slightly, but is of minor taxonomic impor-
tance. The endophallus is usually unarmed and possess only poorly defined spiculate fields.
However, the genitalia of Infernophilus shows a number of peculiarities.
22
Larson
Female ovipositor. - The structure of the apical article of the stylus is of great importance
in determining subtribal affinities. Basically, the stylus is rectangular in shape, lightly sclero-
tized and with a setose apical margin. In Tecnophilus pilatei Chaudoir (fig. 55), and Callida
decora Fabricius (fig. 50), the outer apical corner of the stylus is greatly produced. In Infem-
ophilus (fig. 57) the apex of the stylus is broadly rounded and in Plochionus (figs. 45, 46)
the apex is sharply pointed with a tassel-like arrangement of setae. Small slender spines are
scattered over the surface of the stylus but were found to be of little taxonomic value.
Taxonomic Characters of Larvae
As larvae representing only a few of the species of the Callidina have been seen, the vari-
ation and distribution of the taxonomically important characters within the subtribe are un-
known.
Color. - Color is of considerable value in recognizing the larvae of certain genera. The ter-
minology used to describe color is presented under taxonomic characters of adults.
External morphology . - Most of the characters and terminology used are taken from Van
Emden (1942).
The subtribal character of the presence of a soft membranous pulvillus between the tarsal
claws is difficult to observe in some specimens; hence this structure must be sought for with
care.
The dorsal surface of the maxilla of Plochionus possesses short dense setae (fig. 8). This
surface is glabrous or covered with very short stout sensilla in the larvae of the other cal-
lidine genera.
The shapes of the pronota of Philophuga (fig. 9) and Tecnophilus (fig. 10) have been used
to separate the larvae of these two genera. The number of articles in the cerci of 2nd and 3rd
instar larvae also separates these two genera, but because this character could not be used to
separate the larvae of all instars, it was not used in the key to genera.
TAXONOMY - THE SUBTRIBE CALLIDINA
This sub tribe may be defined as follows:
Adult. - Lebiini with the following characteristics: labrum transverse, with six setae along
anterior margin; mandible widened toward base, usually with scrobe; labial palpus with pen-
ultimate article bisetose, apical article securiform or at least tumid (less in female than in
male) and truncate apically; ligula bi- or plurisetose apically, paraglossae not or but slightly
extending beyond ligula; mentum usually with a tooth; fore tarsus of male with various
number of articles bearing ventral adhesive scales, such scales frequently present on some
articles of middle tarsus; tarsal article 4 bilobed, emarginate or simple; pro thorax not lobed
behind; male genitalia with smaller paramere bilobed apically; female stylus with setose
apex.
Larva. - Head approximately quadrate; epicranial suture wanting, lateral sutures meeting
medially at cervical margin; adnasale prominent, covering base of mandibles; nasale broad
and prominent, entire or variously emarginate apically; antenna shorter than mandible, arti-
Carabidae
23
cle 3 about twice length of article 1, and bearing a short sensory papilla distally; mandible
bisetose externally, with reduced or obsolete retinaculum, penicillus present; maxilla with-
out an inner lobe, palpiger evident; labium with a pair of prominent apical setae; leg with two
equal claws, claws simple or toothed basally; pulvillus present, conspicuous or not;tergites
margined anteriorly; abdominal segment 10 with a pair of protrusible vesicles, each bearing a
semicircular row of crochets; cerci of four or five articles.
This definition of the Callidina is based upon the limited lebiine fauna of the United
States. In this region, the genera Callida Latreille and Dejean, Cylindronotum Putzeys,
Lecalida Casey, Onota Chaudoir, Philophuga Motschoulsky, Plochionus Wiedemann, Tecno-
philus Chaudoir, and Infemophilus new genus may be assigned to the Callidina.
As defined here, this subtribe is more restricted than conceived by either Chaudoir (1872)
or Habu (1967). Habu regarded the shape of the mandibles, the form of the tarsi and the
structure of the female ovipositor as the principal subtribal characters. The structure of the
tarsus does not unite the Nearctic genera of the Callidina. Callida possesses dilated tarsi (fig.
28) which Habu regards as characteristic of this subtribe. However, the closely related genus
Philophuga , has only the stout form of tarsus (fig. 29) and members of the genus Tecno-
philus possess almost slender tarsi. Both stout and dilated tarsi are found in the genus Plo-
chionus.
Habu recognizes three forms of female ovipositor among the Japanese Callidina. These are
as follows: (a) hemisternites transverse, apical segment of stylus long and pubescent apically
(figs. 44-57); (b) hemisternites transverse, apical segment of stylus without terminal setae;
(c) hemisternites not transverse, apical segment of stylus without pubescence but with slen-
der spines near apical angle (fig. 43). In the Nearctic fauna, no genus occurs which possesses
type “b” ovipositor, so I have not been able to determine the relationship between this form
of ovipositor and type “a” ovipositor. Members of Euproctinus Leng and Mutchler possess
type “c” ovipositor (fig. 43). As this genus differs from the North American callidines on the
basis of the female ovipositor and larval characteristics, I exclude it from the Callidina.
Because Euproctinus cannot be assigned to any of the presently recognized subtribes, a new
subtribe may have to be proposed to receive it.
The unique brush-like stylus forms an excellent subtribal character for the Callidina, for
while the structure shows considerable variation among the genera, the basic form remains
recognizable. No other character examined has shown as great a diagnostic value within this
group at the subtribal level.
LARVAE
The larvae of few species of the tribe Lebiini have been described. This may be the result
of at least two factors which make it difficult to obtain the requisite specimens: adults of
many species are rarely collected; and the species of at least one genus, Lebia Latreille, have
a complex developmental cycle (Lindroth 1954, Madge 1967).
24
Larson
Key to the Larvae of the North American Subtribes of Lebiini (modified from van Emden
1942)
1. Retinaculum of mandible vestigial or absent; or if well developed, the cervical groove
and keel are present; antenna with article 1 longer than article 3 2
1*. Retinaculum well developed, projecting by more than a third of width of mandible just
apicad of retinaculum; head without cervical groove or keel; antenna with article 1
shorter than article 3 ; tarsal claws toothed or simple Dromiina
2(1). Tarsus without a pulvillus; tarsal claws with or without a basal tooth, but if tooth
present, head with a cervical groove 3
2\ Tarsus with a soft unpaired but more or less bilobed pulvillus between claws; reti-
naculum vestigial or absent; head without cervical groove; epicranial suture very short
or almost absent 5
3(2). Retinaculum distinct; cervical groove and keel present except if antennal article 1 is
longer than article 3; maxillary stipes rather slender, almost or fully three times as long
as wide Coptoderina
3’. Retinaculum small, vestigial or absent; cervical groove absent; antennal article 1 shorter
than 3; maxillary stipes robust, at most twice as long as wide 4
4(3).Ligula and its two setae well developed; epicranial suture present though short in some
species; cercus of 5-7 articles (but of four articles in at least some first instar specimens)
Cymindina
4’. Ligula and its two setae minute or absent; epicranial suture absent; cercus with at most
four articles Lebiina
5(2). Anterior margin of abdominal tergites with margined border Callidina
5’. Anterior margin of abdominal tergites without margined border
Euproctinus Leng and Mutchler
The Callidina, Cymindina and Lebiina show a number of common features that distin-
guish these groups from the Dromiina and Coptoderina. However, I do not intend to propose
a classification based on characteristics of lebiine larvae. Rather, the known larvae were ex-
amined to determine if their characteristics substantiated the generic separations proposed in
the Callidina on the basis of characters of adults. Where larvae were available, their charac-
ters supported the adult classification.
Key to the known Larvae of the Nearctic Genera of the Subtribe Callidina
1. Anal vesicles of abdominal segment 10 bearing crochets 2
1’. Anal vesicles of abdominal segment 10 without crochets Onota Chaudoir
2(1). Maxilla with dorsal pubescence in addition to the usual fixed setae (fig. 8); nasale nar-
row, without a deep medial incision (fig. 7) Plochionus Weid. p. 25
2’. Maxilla glabrous except for usual fixed setae and short stout sensillae; nasale broad,
with a deep medial emargination (figs. 11-14) 3
3(2).Pronotum quadrate (fig. 9) Philophuga Mots. p. 25
3’. Pronotum conical, widest basally and narrowing anteriorly (fig. 10)
Tecnophilus Chaud. p. 26
Carabidae
25
Plochionus Wiedemann 1 823
Larvae of only P. timidus Haldeman 1843 have been seen. The following description is
based on five specimens from Kirkwood, Missouri (29/VI/ 1892, USNM).
Description. - Values for ratios and measurements are: TL, 2nd instar 5. 9-7. 4 mm; 3rd
instar 8.9-10.4 mm; WH/LH, 2nd instar 1.30-1.36, 3rd instar 1.24-1.28; WP/WH, 2nd instar
1.20-1.24, 3rd instar 1.25.
Color whitish, head and anterior portion of pronotum yellow; nota two and three, legs,
and abdominal tergites testaceous, without metallic lustre; stemites and pleurites testaceous-
yellow.
Head broad, more or less quadrate, but somewhat rounded laterally; nasale convex with a
prominent relatively narrow tri- or quadridentate medial tooth (fig.7); medial suture present
but very short, divided into paired sinuate lateral sutures a short distance anterior to cervical
margin, cervical border narrow.
Antenna shorter than mandible, with four articles; article 1 broad, about one half length
of article 3; article 4 narrow, broadly rounded apically; article 3 with a minute subapical sen-
sillum; subapical setae on articles 3 and 4 only.
Mandible relatively short, slightly and unevenly arcuate, with two external setae; retina-
culum minute, rounded and blunt apically; internal apical half of mandible expanded and
plate-like; penicillus present at base.
Maxilla (fig. 8) hairy above, with internal dorsal row of short stiff setae; ventral surface
glabrous except for usual long setae; internal lobe absent; palpiger well developed, terminal
article of palpus conical and pointed apically; median lobe with two articles, distal article
very small.
Labium with medial pair of setae subapical and dorsal, ventral pair of setae also present;
palpi of two articles, glabrous, apical article elongate, conical.
Prothorax transversely rectangular, broader arid longer than head; lateral margin narrow
and faint.
Legs with tarsal claws toothed at base and equal; pulvilli present but inconcpicuous.
Anterior margins of tergites and nota strongly margined; tergites entire; ventral sclerites
consisting of a ventrite and three pairs of lateral postventrites, all lightly sclerotized and
poorly delimited.
Abdominal segment 1 0 with a pair of protrusible anal tubes, each bearing a semicircular
row of crochets; cerci elongate, slender, with five articles in 2nd and 3rd instar larvae (1st
instar larvae not seen).
Discussion. - The larva of Plochionus stands apart from the known larvae of the other
North American callidine genera on the basis of its setose maxillae, and toothed tarsal claws.
Consequently, the position of this genus among the North American Callidina is unclear, and
probably cannot be settled until the Neotropical genera are better known.
Philophuga Motschoulsky 1859
In this genus, the larvae of P. viridicollis LeConte and P. viridis amoena LeConte are
known. As only first instar larvae of viridis , and second and third instar larvae of viridicollis
were available, I was not able to find characters that would distinguish between the two
26
Larson
species with any degree of certainty. Because of this, a key is not provided, and only the
larva of viridicollis is described below.
Philophuga viridicollis LeConte 1848
A third instar larva, two pupae, an associated adult and a number of exuviae of viridicollis
were seen (Rocky Ford, Colorado, Aug. 17, 1915, Hamilton Coll., USNM). Van Emden
(1942) included this material under Callida, as C. purpurea Say.
Description. - Values for ratios and measurements of 12 larvae and exuviae are: TL, 3rd
instar 9.1 mm; WH/LH, 3rd instar 1.18-1.22 (X = 1.20), 2nd instar 1.15-1.20 (X = 1.18);
WP/WH, 3rd instar 1.18.
Color whitish; head testaceous; body sclerites brown to piceous with a metallic green or
violaceous sheen.
Head (fig. 11) broad, almost quadrate with hind angles broadly rounded; frontal piece
broad; sinuate lateral sutures meeting medially at cervical margin, median suture absent;
nasale very broad, deeply incised medially, lateral portions tridentate; cervical margin nar-
row; ocelli six, strongly pigmented.
Antenna slightly shorter than mandible, of four articles; article 4 narrow, subequal in
length to article 3, article 1 short; article 3 with a small subapical sensillum; subapical setae
on articles 3 and 4 only.
Mandible slender and arcuate, with two lateral setae; retinaculum small and inconsp-
icuous; basal penicillus present.
Maxilla with an internal dorsal row of stout setae plus usual long fixed setae, otherwise
glabrous above; inner lobe absent; palpi with well developed palpiger, terminal article narrow
and pointed.
Labium broad, bisetose apically, also with a pair of subapical ventral setae; palpi of two
articles, glabrous.
Prothorax transversely rectangular (fig. 9), wider than head; lateral margin strong and
complete.
Legs with equal simple claws; pulvilli present.
Tergites entire, with strong anterior borders; lateral margins of meso- and metanota an-
gular; ventral sclerites consisting of a ventrite and three pairs of lateral postventrites.
Cerci with five articles in second and third instars, only subapical setae present; abdominal
segment 1 0 with a pair of eversible anal tubes, each bearing a semicircular row of crochets.
Discussion. - Van Emden (1942), who saw some larval specimens of Callida, keyed Callida
and Philophuga to the same couplet, stressing their similarities especially in the shape of the
nasale, number of articles in the cerci, and the shape of the tarsal claws. Larval characteris-
tics apparently confirm the close relationship suggested for these two genera by adult charac-
teristics.
Tecnophilus Chaudoir 1877
The larvae of both subspecies of Tecnophilus croceicollis Men. are known. They were
obtained by rearing from adults collected at the following localities: junction of the Lost
and Milk Rivers, Alberta (T. c. peigani n. ssp. ); Cuddeback Lake and Alviso, California (71 c.
croceicollis Men.).
Carabidae
27
Description. - Values for ratios and measurements are presented in table 1 . The first instar
larva of T. croceicollis Men. is illustrated in fig. 6.
TABLE 1 . Variation in dimensions and ratios among the larvae of Tecnophilus croceicollis
Menetrie's.
Color whitish; head capsule and in many specimens prosternum and anterior half of pro-
notum yellow; remaining sclerites and appendages piceous, with at least a faint metallic
green cast.
Head rectangular (figs. 12-14); frontal piece broad, medial suture wanting, sinuate lateral
sutures meeting medially just anterior to cervical margin; nasale broad, deeply cleft medially,
lateral pieces shallowly bifid; posterior constriction of head varied; cervical margin narrow;
ocelli six, strongly pigmented.
Antenna shorter than mandible, with four articles; article 1 short, broad, about one half
length of article 3; article 4 narrow, widening apically where broadly rounded; subapical
setae on articles 3 and 4 only; small sensillum on article 3.
Mandible slender, with two lateral setae; retinaculum small, but evident; basal penicillus
present.
Maxilla with internal dorsal row of short, stiff setae and scattered long external setae,
otherwise glabrous; palpiger distinct; palpus with three articles, article 3 short, narrow; inner
lobe absent; outer lobe with two articles, article 2 very short and narrow, basal article with a
single preapical seta.
Labium bisetose apically, also with a pair of ventral setae; palpi with two articles, terminal
28
Larson
article short and narrow, pointed apically.
Prothorax somewhat conical (fig. 10), widest basally and narrowing anteriorly to about
width of base of head; variously margined laterally.
Legs with equal simple claws; pulvilli present but reduced.
Tergites and nota margined anteriorly; ventral sclerites consisting of a ventfite and three
pairs of postventrites.
Cerci of four articles in all instars; abdominal segment 10 with a pair of eversible vesicles,
each bearing a semicircular row of crochets.
Discussion. - I was unable to find characters to distinguish between the larvae of T. c.
croceicollis and T. c. peigani. Table 1 summarizes variation in measured characteristics and
ratios among larvae from three localities.
The larvae of Tecnophilus are similar to those of Philophuga, but they can be distin-
guished from one another by the character presented in the key. Also, larvae of all instars of
Tecnophilus have cerci of only four articles. In Philophuga, on the other hand, the first in-
star larvae have cerci of four articles while later instars have cerci of five articles. These dif-
ferences do not contradict the separation of Tecnophilus and Philophuga at the generic
level.
ADULTS
Characteristics common to all members of the subtribe Callidina, are presented in the
taxonomy section. Below is a key to the adults of the North American genera of the subtribe
Callidina. Following the key, the genera Philophuga, Infernophilus and Tecnophilus are dis-
cussed in detail. I have not had the opportunity to examine the other genera in detail, as
the majority of their members are neotropical. Hence any remarks about the latter group are
tentative.
Key to the Adults of the North American Genera of the Sub tribe Callidina
(modified from Ball 1963)
1. Mandible broadly expanded, without scrobe Onota Chaudoir
1 ’. Mandible with scrobe 2
2(l).Mentum without a tooth; width of pronotum less than maximum width of head ....
Cylindronotum Putzeys
2’. Mentum with a tooth; width of pronotum greater than width of head 3
3(2). Lateral margin of abdominal sternum 4 with one seta, sternum 5 with two setae, and
sternum 6 with one seta (fig. 16); hind femur broad. Plochionus Wiedemann
3’. Lateral margin of abdominal sterna 4, 5 and 6 without such arrangement of long setae;
hind femur slender 4
4(3). Tarsus with fourth article bilobed (fig. 28) 5
4’. Tarsus with fourth article at most emarginate (fig. 29) 6
5(4). Elytra metallic green or blue; pronotum elongate and slender, lateral grooves narrow.
Callida Latreille and Dejean
Carabidae
29
5’. Entire body rufo-piceous, without metallic color; pronotum with lateral grooves broad
Lecalida Casey
6(4). Tarsal claws pectinate 7
6’. Tarsal claws simple Tecnophilus Chaudoir, p. 38
7(6). Color dark with metallic blue or green sheen; abdominal sternum 6 with at most four
pairs of anal setae Philophuga Motschoulsky, p. 24
7’. Color brown, non-metallic; abdominal sternum 6 with at least six pairs of moderate to
long setae in female, distinct anal brushes present on male (fig. 27)
Infernophilus new genus, p. 37
Philophuga Motschoulsky 1859
Philophuga Motschoulsky 1859 - 140.
Philopheuga Bates 1883 - 202.
Glycia LeConte 1851 , not Chaudoir 1842.
Type species - Philophuga cyanea Motschoulsky 1850 (= P. viridis Dejean), here desig-
nated.
The diagnostic features of this genus are presented in the key. Other characteristics, com-
mon to all species of Philophuga , are given in the following description.
Description - Beetles 5.5-10.0 mm in length. Color various, but at least elytra and abdom-
inal sterna dark with a metallic blue or green sheen; antennae black with articles 1 to 3 and
base of 4 pale, at least on ventral surface.
Head with eyes prominent, convex; at least a faint post-ocular pinch present. Genae vari-
ously narrowed behind, neck evident. Labrum slightly emarginate medially; clypeus with a
single seta on each side. Frons with indistinctly defined, rugose frontal furrows; punctate, at
least laterally and posteriorly. Antennal articles 1 to 3 and base of 4 glabrous or very sparse-
ly and finely hairy; remaining articles pubescent. Maxillary palpus with fusiform terminal
article. Labial palpus with penultimate article bisetose; terminal article securiform (narrower
in female). Ligula bisetose. Mentum with a prominent margined tooth.
Pronotum varied in shape; sides usually evidently sinuate behind; disc transversely rugose,
best developed laterally; posterior lateral setae present or absent.
Tarsal articles glabrous or sparsely hairy above, moderately setose beneath; article 4 emar-
ginate; claws pectinate; male with articles 1 to 4 of front and middle tarsi bearing two rows
of scales beneath.
Elytra completely margined basally and apically. Hind wings full or reduced.
Abdominal sterna 3 to 5 with a pair of medial setae; sternum 6 with one to four pair of
anal setae (male with usually one less pair of setae than female); sterna 4 to 6 without long
lateral setae.
Male genitalia on right side in repose, left paramere large; aedoeagus tube-like, with apical
orifice opening slightly to left of midline of aedoeagus; endophallus unarmed.
Female styli of typical callidine form.
Discussion. - Some members of this genus are superficially very similar to certain of the
species included in the genus Callida (for example Philophuga viridis amoena LeConte and
Callida purpurea Say; and Philophuga brachinoides Bates and Callida decora Fab.), and can
often be recognized only on the basis of the character presented in the key. Even in the
30
Larson
structure of tarsal article 4, C. purpurea very closely approaches the condition found in
members of Philophuga. Perhaps when the species of Callida are studied in more detail, it
will be found necessary to include Philophuga in Callida, as a subgenus (a parallel situation
occurs in the two subgenera of Plochionus Weidemann which are separated by differences
in the structure of tarsal article 4). However, in this study the status of Philophuga as a dis-
tinct genus is maintained
Distribution - The species of Philophuga occur in the semi-arid and arid regions of western
North America.
Key to the Species of Philophuga Motschoulsky
1 . Pronotum rufous, contrasting with black head brachinoides Bates, p. 30
1*. Pronotum and head concolorous, black with a metallic blue or green sheen 2
2(1). Hind wings constantly full, membranous areas distinctly pigmented (fig. 23); hind angle
of pronotum with a setiferous puncture 3
2\ Hind wings reduced, without reflexed apex, or if full, membranous areas not or only
lightly pigmented; hind angle of pronotum with or without a setiferous puncture . .
viridis Dejean, p. 34
3(2). Head and pronotum with metallic green lustre, contrasting with blue elytra, or elytra
also green; elytral striae shallowly impressed; intervals flat, finely punctate
viridicollis LeConte, p. 31
3’. Head and pronotum with metallic blue or blue-green lustre, similar in color to elytra
and not contrasting; elytral striae deeply impressed; intervals convex, finely to moder-
ately coarsely punctate caerulea Casey, p. 33
Philophuga brachinoides Bates 1883
Philopheuga rachinoides Bates 1883 - 202. Type locality - Cerro de Plumas, Veracruz
(Selander and Vaurie 1962), Mexico. Blackwelder 1944 - 62 ( Philophuga ).
Description. - Values for ratios and measurements for five specimens are: TL 6. 8-7. 8 mm
(7.2); LE/LP 2.70-2.86 (2.75); WP/LP 1.18-1.28 (1.24); WH1/WH2 1.41-1.48 (1.45).
Color of prothorax, mesothorax, femora and antennal articles 1 to 3 and base of 4 rufous;
remainder of body piceous to black with a metallic blue lustre.
Microsculpture obsolete on frons and disc of pronotum; strong on elytra, isodiametric
medially, slightly stretched laterally.
Eyes prominent; post-ocular pinch faint. Genae strongly constricted behind, forming
relatively narrow neck. Labrum broadly but shallowly emarginate. Clypeus micropunctate.
Frons smooth and shiny medially, deeply but sparsely punctate posteriorly and laterally;
frontal furrows short, deep, confluently punctate.
Prothorax (fig. 33) rounded laterally, sinuate before prominent though obtuse hind
angles; front angles rounded, not or only slightly protruding; hind angle on each side bearing
a setiferous puncture; disc glabrous, transversely rugose. Prosternum with very short fine
setae. Metasternum with scattered moderately long fine setae.
Elytral striae clearly impressed medially on disc, effaced apically and in some specimens
also laterally; discal striae finely punctate, punctures coarser basally and laterally; intervals
Carabidae
31
slightly convex, each bearing an irregular row of fine punctures. Wings full, densely pig-
mented.
Abdominal sternum 2 with a small medial patch of setae; remaining sterna with very short
sparse setae; abdominal sternum 6 with two pairs of anal setae in male, and three pairs in fe-
male.
Aedoeagus elongate, slender. Female stylus as in fig. 54.
Discussion. - This species very closely resembles Callida decora Fab. in color. It is the only
species of Philophuga with pale legs and thorax.
Distribution. - 1 have seen only five specimens of this species from the following locality
(fig. 60).
MEXICO, Oaxaca, Rte. 190, 21.7 miles SE Nochixtlan, 7200’. March 24, 1966 (Ball and Whitehead, UASM).
Philophuga viridicollis LeConte 1848
Cymindis viridicollis LeConte 1848 - 188. Lectotype - (here selected) female, labelled as
follows: green disc, Glycia viridicollis Lee., Type 5821 MCZ , Philophuga viridicollis (Lee.).
Philophuga subcordata Chaudoir 1877 - 246. Holotype not seen. Type locality - “Mexique”.
Philophuga purpurea Chaudoir 1877 - 245, not Say 1823.
Description. - Values for ratios and measurements of 40 specimens are: TL 7. 1-9. 6 mm;
LE/LP 2.77-3.1 3 (2.98); WP/LP 1 .07-1 .29 ( 1 .1 9); WH i /WH2 1 .42-1 .54 (1 .47).
Color of head, pro no turn and ventral parts of thorax and abdomen shiny metallic green
or blue-green; elytra duller, blue or rarely in some specimens green; legs, epipleura and lateral
portions of abdominal sterna piceous to black; antennae with articles 1 to 3 and base of 4
pale, at least on ventral side, outer articles black; palpi dark, with apex of terminal articles
pale.
Microsculpture more or less effaced on head; lightly impressed and transverse on pro-
notum; finely granular on elytra, rarely slightly stretched.
Head with basal constriction gradual, neck evident; eyes prominent, convex. Labrum
slightly emarginate. Clypeus smooth and shiny, irregularly and finely punctate; frons shiny,
often with faint transverse rugae, sparsely but deeply punctate posteriorly and laterally;
frontal furrows broad, poorly delimited, confluently punctate and longitudinally rugose;
genae behind and below eyes with short sparse setae; labial palpus with terminal article
very broadly dilated in male, less so in female.
Pronotum (fig. 31) narrow, broadest in apical third, rounded laterally with a faint sinu-
ation before hind angle; hind angle obtuse and somewhat rounded, bearing a setiferous punc-
ture; front angle rounded, slightly protruding; base clearly margined, at least laterally; pros-
temum and proepisternum sparsely punctate; prosternal punctures bearing very short fine
setae. Metasternum punctate laterally, sparsely setose medially.
Elytra elongate, subparallel; striae fine, often only very faintly impressed apically, mod-
erately punctate, punctures deeper and denser in basal half; intervals flat, with an irregular
row of faint punctulae in each. Wings fully developed, strongly pigmented.
Abdominal sterna with fine setiferous punctures medially; abdominal sternum 6 with two
pairs of anal setae in female and three pairs in male.
Male genitalia (fig. 1 5) with aedoeagus slightly arcuate dorsally; basal piece flattened, bent
32
Larson
somewhat to right. Female stylus as in fig. 51.
Variation. - No marked pattern of variation was noticed for this species. These characters
were observed to vary within a single population sample: shape of the pronotum, microscul-
pture of frons and pronotum, and the depth of the elytral striae. The color of the head and
the pronotum varies from metallic green to blue-green, but remains sufficiently green to con-
trast conspicuously with the blue elytra. This color contrast is lost in the few specimens that
possess greenish elytra. Color, however, is the most reliable character for separating viridi-
collis from the markedly similar caerulea.
Discussion. - 1 have seen two specimens of viridicollis from the following California local-
ities: San Francisco (Hopping, CAS); Needles (CAS). As no specimens of this species have
been seen from Arizona and western New Mexico, these specimens may have been accidently
mislabelled. The two species viridicollis and caerulea appear to be almost completely allo-
patric (see discussion under caerulea ).
Notes on synonomy. - Say’s (1823) original description of Callida purpurea and C. viridi-
pennis (both originally assigned to Cymindis ) stated that article 4 of the tarsus was bilobed
for these two species, thereby confirming their position in Callida. However, Chaudoir
(1877) applied the name purpurea Say to specimens of Philophuga viridicollis LeConte, and
relegated the name viridicollis LeConte to synonymy. Horn (1882), after receiving identified
specimens of Philophuga from Chaudoir, noticed this confusion and corrected it by placing
purpurea Say in the genus Callida , and reinstating viridicollis in Philophuga. In the same
paper, Horn suggested that subcordata Chaudoir was probably a synonym of viridicollis
LeConte. Leng (1920) followed Horn’s suggested synonymy, and although the type of
subcordata Chaudoir has not been seen in this study, this synonymy is followed here.
Specimens of the species viridicollis LeConte and caerulea Casey have been confused with
one another. LeConte’s collection contains specimens of both of these species under the
name viridicollis.
Distribution. - The species viridicollis is found in the southern Great Plains, from Kansas
and eastern Colorado south into eastern New Mexico, Texas (excluding the Edwards Pla-
teau), northeastern Mexico and possibly California - but see above (fig. 58). I have examined
over 260 specimens of this species from the following localities:
United States - ALABAMA: one specimen labelled “Ala.” (WUM). COLORADO: Otero Co., Rocky Ford (Hamil-
ton, USNM). KANSAS: Ellsworth Co., (Martin, KUM); Kiowa Co., (Woodruff, KUM); Reno Co., (Hopping, CAS); Sumner
Co., (USNM). NEW MEXICO: Lincoln Co., Ramon (Ball, UASM); Roosevelt Co., Portales (IUM), Water Canyon (Shaw, CU).
OKLAHOMA: Oklahoma Co., Oklahoma City (Grant, CAS), Stillwater (Whitaker, MCZ). TEXAS: Atascosa Co., Pleasanton
(White, CNHM); Bee Co., Beeville (Tucker, USNM); Bexar Co., San Antonio (CAS, CUM, USNM); Blanco Co., Cypress Mill
(USNM); Brewster Co., Alpine (CAS, USNM), Big Bend Nat’l Park (Becker & Howden, CNQ, Horse Canyon (Becker &
Howden, CNC), Marathon (Malkin, CNHM); Brooks Co., Falfurrias (Beer, Martin, CAS, KUM); Cameron Co., Brownsville
(Glick, USNM), Bruni (Martin, CAS), Childress Co., Childress (Mitchell, USNM); Comal Co., New Braunfels (CAS, MCZ,
USNM); Culberson Co., Van Horn (Barr, IUM); Denton Co., Denton (Bishop, USNM; Duval Co., San Diego (USNM); Frio
Co., Pearsall (Tucker, USNM); Hardeman Co., Quanah (Morrill, USNM); Hemphill Co., Canadian (Mann, USNM); Hidalgo
Co., Edinburg (CUM, USNM); Jeff Davis Co., Davis Mts. (CAS), Ft. Davis (Ball, CAS, UASM); Karnes Co., Kenedy (Marlatt,
USNM); Kerr Co., Kerrville (Becker & Howden, CNC); Kleberg Co., Kingsville (Reed, CUM); LaSalle Co., Cotulla (USNM);
Lubbock Co., (Manis, IUM); Nueces Co., Corpus Christi (CNHM, USNM); Presidio Co., Marfa (Scullen, Wickham, MCZ,
OUM, USNM); Randall Co., Canyon (Stephenson, KUM); Terrell Co., Dryden (Ball, UASM), Sanderson (Martin, Mason,
CAS, CNC); Travis Co., Austin (Darlington, Martin, Pinkus, CAS, MCZ, USNM); Uvalde Co., Sabinal (Pratt, USNM), Uvalde
Carabidae
33
(CAS, CNC, USNM); Val Verde Co., Comstock (Barr, IUM), Del Rio (CAS, CNC, USNM), Devils River (Schwarz, USNM);
Victoria Co., Victoria (USNM); Ward Co., Monahans (Larson, UASM); Webb Co., Laredo (Martin, Werner, CAS, UASM).
Philophuga caerulea Casey 1913
Philophuga caerulea Casey 1913 - 174. Holotype - female labelled as follows: Ariz., Casey
bequest 1925, Type USNM 47668, caerulea Casey.
Calleida viridis Chevrolat 1825 - 155 (not Dejean 1831). Type locality - Las Vigas, Veracruz,
Mexico.
Description. - Values for ratios and measurements for ten specimens from Arizona are: TL
7.46-9.02 mm (8.36 mm); LE/LP 2.73-3.07 (2.94); WP/LP 1.12-1.26 (1.18); WH1/WH2
1.44-1.57 (1.49).
Color of dorsal surface uniformly metallic blue or greenish-blue; head and pronotum more
shining than elytra but of about the same hue; epipleura rufo-piceous; legs piceous to black;
antennal articles 1 to 3 and base of 4 pale, at least on ventral side, outer articles black; palpi
dark with apices of terminal articles paler.
Microsculpture obsolete on frons and disc of pronotum; on elytra isodiametric and finely
granular medially, slightly stretched laterally.
Head as in viridicollis', short sparse setae present on genae behind and below eyes.
Pronotum (fig. 32) varied in shape with seta present near each hind angle. More cons-
tricted basally than in specimens of viridicollis, with a longer more evident lateral sinuation;
posterior lateral impressions slightly broader; lateral reflexion narrower.
Elytral striae clearly impressed, moderately punctate; intervals slightly convex, distinctly
punctate, punctures stronger than in viridicollis. Hind wings fully developed; distinctly pig-
mented (fig. 23).
Male genitalia as in fig. 1 6. Female stylus similar to that of viridicollis.
Variation. - Size was the only character observed to vary geographically. Specimens from
the southeastern portion of the range of caerulea tend to be noticeably smaller than speci-
mens from other populations. The mean length for a sample of five specimens from Las
Vigas, Veracruz, was 6.92 mm (range 6.74-7.22 mm), while the mean of a sample of ten
specimens from Arizona was 8.36 mm (range 7.46-9.02 mm). Too few specimens from inter-
mediate localities were available to determine if this variation was clinal.
Discussion. - Philophuga caerulea strongly resembles viridicollis, and differs little from this
species aside from the characters presented in the key. In the United States, these two spe-
cies are allopatric and are readily separated from one another. However, in Mexico the distri-
butions are too incompletely known to determine if this geographical separation is main-
tained. A specimen from Monterrey, and two specimens from Monclova, Mexico, resemble
viridicollis in color, but caerulea in the convexity of the elytral intervals and the depth of the
striae. Perhaps when more specimens are available from Mexico, it will be necessary to treat
these two species as well marked subspecies.
Distribution. - 1 have examined 92 specimens of this species from the following localities
(fig. 58):
Mexico - AQUASCALIENTES: Aquascalientes (Hendrichs); CHIHUAHUA: Chihuahua (Wickham, MCZ);
COAHUILA: Monclova (Schwarz, USNM); JALISCO: Guadalajara (MCZ, UASM); MEXICO: Lerma (Ball, UASM); Presa
34
Larson
del Angulo (Hendrichs); Toluca (Bowditch, MCZ); Valle de Bravo (Hendrichs); NUEVO LEON: Monterrey; PUEBLA:
Tlachichuca (Ball, UASM); TAMAULIPAS: “Mesa Gonzales” (= Gonzales) (CAS); VERACRUZ: Las Vigas(Hoege, MCZ,
USNM); ZACATECAS: Sombrerete (Evans, UASM).
United States - ARIZONA: Cochise Co., Huachuca Mts. (CAS, USNM); Tombstone (UASM); Gila Co., Pinal Mts.
(USNM); Santa Cruz Co., Nogales (CAS, CNHM); Patagonia (CNHM, MCZ); Sonoita (CAS, KUM); Sta. Rita Mts. (CAS,
MCZ, UASM).
Philophuga viridis Dejean 1831
Cymindis viridis Dejean 1831 - 325. Type locality - California. Holotype not seen. Mots-
choulsky 1859 - 144 ( Philophuga ); Horn 1882 - 144 (. Philophuga ); Leng 1920 - 67.
Hatch 1953 - 157 (not Chevrolat 1835 - 155).
This is the most varied species of Philophuga, containing four well defined subspecies.
Characteristics common to all subspecies of viridis are presented in the following description.
Description. - Values for ratios and measurements are presented separately under each of
the following four subspecies.
Color varied, ranging from dull black with a faint metallic blue sheen to bright metallic
blue or green; palpi, legs and outer antennal articles piceous to black; antennal articles 1 to
3 and base of 4 pale, at least on ventral surface.
Microsculpture varied; effaced on frons and disc of pronotum in many specimens; isodia-
metric or slightly transverse on elytra, faintly impressed to granular.
Dorsal surface of body glabrous or setose ; frons and disc of pronotum punctate.
Pronotum varied in shape; with or without a seta near each hind angle.
Elytra relatively short, oval, with greatest width in apical half. Hind wings varied in devel-
opment, with re flexed apex or reduced to a small scale; when fully developed, membranous
areas only lightly pigmented.
Male genitalia with aedoeagus short, slightly arcuate; basal piece curved (fig. 17).
Female stylus as in figs. 52 and 53.
Geographical variation and subspecies. - The species Philophuga viridis Dejean ranges wide-
ly in the semi-arid and cold desert regions of western North America, from the prairies of
southern Canada west to Washington and Oregon, and south to northern New Mexico, Ari-
zona, and California. Over this range, color, vestiture, wing development, and the presence of
the posterior-lateral pro thoracic setae vary. Data on variation in these characters is sum-
marized in a pie diagram map (fig. 1) and in table 2. Four subspecies are recognized, based
on variation in these characters.
On the eastern side of the Rocky Mountains, and in British Columbia, Washington, and
northern Oregon, populations possessing the following characteristics occur: the posterior-
lateral setae of the prothorax are absent; the wings are fully developed; the dorsal surface of
the body is glabrous; and the color is a dark blue-black. Associated with the absence of the
posterior-lateral setae, the pronotum tends to be less sinuate laterally, with more obtuse
hind angles than in specimens that possess the setae. The name viridis amoena LeConte ap-
plies to specimens with these characteristics.
To the west of the Rocky Mountains, a population centering around the Great Basin
occurs. This subspecies, v. horni Chaudoir, differs from the neighboring v. amoena by pos-
sessing a posterior-lateral seta on each hind angle of the pronotum, and by being bright me-
tallic green or blue in color. The hind angles of the pronotum are more distinct than in v.
Carabidae
35
amoena and the lateral margins are more sinuate posteriorly.
TABLE 2. List of localities from which specimens of Philophuga viridis Dejean were used to
compile pie-diagram map (fig. 1 ).
Map No. of
symbol Locality specimens
A
B
C
D
E
F
G
H
I
J
K
L
M
N
O
P
Q
R
S
T
U
V
W
X
Y
Z
a
b
c
d
e
f
g
Southern British Columbia 4
Southern Alberta 24
Southern Saskatchewan 3
Southern Manitoba 1 5
Valley Co., Montana 1
Lake Co., Montana 1
Lewis & Clark, and Missoula Cos., Montana 1 1
Musselshell Co., Montana 1
Gallatin Co., Montana 6
Bighorn Co., and Yellowstone National Park, Wyoming 6
Carbon and Laramie Cos., Wyoming 4
Delta and Garfield Cos., Colorado 2
Boulder, Jefferson, and Larimer Cos., Colorado 5
Chaffee, El Paso, Park and Teller Cos., Colorado 8
La Plata Co., Colorado 2
Sapello, San Miguel Co., New Mexico 1
Coconino Co., Arizona 7
Washington Co., Utah 1
Soldier Summit, Utah Co., Utah 1
Touele Co., Utah 12
Esmeralda Co., Nevada 2
White Pine Co., Nevada 1
Washoe Co., Nevada 13
Humbolt Co., Nevada 1
Ada, Canyon, Owyhee, and Payette Cos., Idaho 8
Blaine, Cassia, Lincoln, Twin Falls Cos., Idaho 1 3
Bannock and Lake Cos., Idaho 2
Clark and Lemhi Cos., Idaho 2
Adams and Nez Perce Cos., Idaho 4
Grant Co., Washington 3
Seattle, Washington 2
Wasco Co., Oregon 3
Baker Co., Oregon 16
36
Larson
TABLE 2. (cont.).
Map No. of
symbol Locality specimens
h Harney, Lake, and Malheur Cos., Oregon 10
i Klamath Co., Oregon 5
j Jackson Co., Oregon 3
k Yreka, California 1
1 Lassen, and Modoc Cos., California 8
m San Francisco, California 96
TOTAL 308
Although amoena and horni are consistent in their characters over the major portion of
their respective ranges, two areas of intergradation occur; along the Rocky Mountains, and
in the north-western states. In the Rocky Mountains intergradation is most pronounced
around low passes. I have seen two typical specimens of amoena from Pocatello and Salmon,
Idaho. A specimen from Dubois was colored like horni but lacked the posterior-lateral pro-
thoracic setae. All other specimens seen from Idaho were typical horni. Some specimens
from the eastern side of the continental divide in southwestern Montana and northwestern
Wyoming show horni characteristics. One of the six specimens seen from Gallatin Co., Mon-
tana, possessed posterior-lateral pronotal setae. One specimen of the six seen from Yellow-
stone National Park had the coloration of horni.
A similar situation in which the mountains do not form a complete barrier to genetic in-
terchange, occurs in western Colorado and eastern Utah. On the eastern slopes of the Rocky
Mountains in Colorado, some specimens of amoena show a brighter metallic blue or green
coloration, most obvious along the frontal grooves of the head and the lateral margins of the
prothorax. Specimens from localities to the south and west in Colorado show a higher inci-
dence of bright coloration until all specimens seen from Utah, Arizona and New Mexico
show the horni coloration. The posterior-lateral prothoracic setae tend to be absent from
populations of horni colored beetles in eastern Utah, western Colorado and in New Mexico.
Some specimens of horni from as far west as Salt Lake City, Utah, lack these setae even
though the pronotum has the characteristic shape for horni. In this region, the bright color-
ation of horni seems to pass through or around the southern Rocky Mountains and occurs
with limited frequencies in populations on the eastern slopes of the mountains. The amoena
characteristic, the absence of the posteior-lateral pronotal setae, tends to occur in popula-
tions of horni as far west as Touele Co., Utah.
The situation in the northwestern states is similar. The subspecies amoena is found in the
north and western portions of Washington and Oregon. One of the four specimens I have
seen from southern British Columbia is typical horni while the remaining specimens are
amoena. Even more surprising are two specimens of horni labelled Seattle, Washington
(O.B.J.), while all other Washington specimens may be assigned to amoena. On the validity
of O.B. Johnson’s labels, Hatch (1950, p. 21) stated “But if he (O.B. Johnson) believed that
Carabidae
37
a given species occurred in the vicinity of Seattle, he would not hesitate, according to Prof-
essor Kincaid, to put a Seattle label on any specimen that came to hand regardless of its
exact point of origin.” However, a specimen with horni characteristics from southern British
Columbia does not permit me to discount the possibility that horni does occur in the Seattle
region. Hybrid specimens have been seen from Baker and Wasco Counties, Oregon.
The separation of amoena and horni may be primarily ecological. Specimens of amoena
are found on short-grass prairie in southern Alberta, and seem to occur in regions where
grassland is prevalent. The subspecies horni seems to be found in regions of desert scrub
vegetation.
The third subspecies, v. klamathea new subspecies, is known from only a small area in
south-central Oregon and northern California. In coloration and the shape of the pronotum
(the posterior-lateral setae are present), this subspecies resembles horni. However, it is
readily distinguished from the latter by the presence of sparse vestiture on the dorsal sur-
face of the body and by the reduced hind wings which are represented by no more than
small scales. On the basis of these two characters, klamathea is quite distinct from horni.
Nevertheless some introgression does occur. One macropterous specimen was collected along
with four micropterous beetles in a series from Klamath Co., Oregon. All specimens from
the neighboring Lake Co. to the east are macropterous but some specimens show traces of
fine vestiture on the elytral intervals and on the genae below the eyes (such specimens have
been scored as being glabrous in pie-graph map (fig. 1 ). The subspecies klamathea does not
show any signs of intergrading with amoena to the north.
The fourth subspecies, v. viridis Dejean, appears to be isolated in the vicinity of San Fran-
cisco Bay and Sonoma Co., California. Its characteristics are: hind wings reduced to a small
scale; vestiture present on dorsum; color black to dull blue-black, rarely bright blue or green;
posterior-lateral setae of pronotum absent from most specimens (94%); however, the lateral
margin of the thorax is sinuate and the hind angles are distinct though obtuse. The basal bor-
der of the elytra tends to be more sinuate internally in micropterous than in macropterous
specimens. Also, this sinuation is usually stronger in viridis s. str. than in klamathea. These
two subspecies also differ in some characteristics of the elytral striae and microsculpture.
These differences are described below.
The geographically isolated position of v. viridis in conjunction with the differences men-
tioned above, suggest that viridis and klamathea may actually be specifically distinct. How-
ever, some instability in the diagnostic characters of color and the presence of the posterior-
lateral pronotal setae cause me to consider these populations to be conspecific, at least until
better evidence to the contrary is discovered.
Key to the Subspecies of Philophuga viridis Dejean
1 . Hind wing reduced, much shorter and narrower than a single elytron 2
1’. Hind wing full, with reflexed apex 3
2(1). Hind angle of pronotum lacking setiferous puncture in most specimens (fig. 37); elytral
striae finely impressed, outer striae often obsolete and represented by a row of fine
punctures v. viridis Dejean, p. 38
2\ Hind angle of pronotum with a setiferous puncture (fig. 36); elytral striae clearly im-
38
Larson
pressed, outer striae distinct almost to apex v. klamathea new subspecies, p. 40
3(2). Hind angle of pronotum with a setiferous puncture (fig. 35). . v. horni Chaudoir, p. 41
3’. Hind angle of pronotum without a setiferous puncture (fig. 34)
v. amoena LeConte, p. 42
Philophuga viridis viridis Dejean 1 83 1
Cymindis viridis Dejean 1831 - 325. Type locality - California. Type specimen not seen
(not Chevrolat 1835 - 155).
Callida cyanea Motschoulsky 1850 - 36. Type locality - San Francisco, California. Type
specimen not seen.
Philophuga lauta Casey 1913 - 175. Holotype - male, labelled as follows: Cal., Casey be-
quest 1925, Type USNM 47670, lauta Csy. New Synonymy.
Description. - Values for ratios and measurements of fifty-two specimens are: TL 5. 7-7. 5
mm (6.8 mm); LE/LP 2.39-2.80 (2.59); WP/LP 1.10-1.29 (1.21); WH1/WH2 1.41-1.54
(1.46).
Color dark, ranging from dull black to metallic blue or green; palpi and legs dark piceous
to black; abdomen piceous to black.
Micro sculpture varied; isodiametric to slightly transverse on frons, obsolete in many speci-
mens; on disc of pronotum lightly impressed, transverse; isodiametric, faintly impressed to
granular on elytra.
Body setose dorsally; evident setae present on frons behind and below eyes, on disc of
pronotum and on at least odd numbered elytral intervals.
Head broad, genae broadly rounded laterally; postocular pinch faint; frons with setiferous
punctures best developed posteriorly and laterally, also micropunctate.
Pronotum varied (fig. 37); lateral margins rounded, each with a long sinuate posterior con-
striction; sides subparallel or slightly converging behind; front angles rounded, slightly pro-
truding; hind angles prominent but obtuse; most specimens without posterior lateral setae
(setae present in 6% of specimens examined); lateral margin narrow, widening toward hind
angles; disc with sparse setiferous punctures, deeper and denser posteriorly and laterally.
Elytra relatively short, oval, with greatest width in apical half; shoulders broadly rounded,
basal margin with strong anterior convexity; striae lightly impressed, finely punctate, outer
striae often incomplete before apex, stria 8 represented by a row of punctures in many speci-
mens; intervals flat to slightly convex, with an irregular row of small punctures in each. Hind
wings reduced, represented by a small scale.
Abdominal sterna sparsely and finely setose; sternum 6 bearing one pair of anal setae in
the male or two pairs in the female.
Male genitalia (fig. 17) with aedoeagus short, apex short; basal piece strongly curved.
Female stylus as in fig. 52.
Distribution. - Localities from which specimens of this subspecies have been collected are
indicated in fig. 59. I have examined 96 specimens of this subspecies from the following lo-
calities:
CALIFORNIA: Alameda Co., Grizzly Peak (Van Dyke, CAS); San Francisco Co., Lake Merced (Van Dyke, CAS), San
Francisco (BlaisdeU, Van Dyke, CAS, MCZ, USNM); Sonoma Co., Eldridge (CAS, USNM).
Carabidae
39
Fig. 1 . Pie-diagram map illustrating geographic variation in four characteristics: wing develop-
ment; color; vestiture; and presence of posterior-lateral prothoracic setae, of selected popu-
lation samples of Philophuga viridis Dejean. A number external to a quadrant, indicates the
percentage of specimens of that sample that show the black phase of the character. Local-
ities represented by each pie-diagram are listed in Table 2.
40
Larson
Philophuga viridis klamathea new subspecies
Holotype - male, Klamath Falls, Oregon (Van Dyke, CAS).
Allotype - female, Medford, Oregon (USNM).
Paratypes are from the following localities:
CALIFORNIA: “CaL” (MCZ, WUM); Siskiyou Co., Yreka (USNM); OREGON: “Or.” (USNM); Jackson Co., Ashland
(Stephen, OUM), Colestine (Bishop, OUM), Medford (USNM); Klamath Co., (CNHM).
The diagnostic characteristics of this subspecies are presented in the key and in the discus-
sion of variation under Philophuga viridis.
Description. - Values for ratios and measurements of ten specimens are: TL 6.56-8.00 mm
(7.03 mm); WP/LP 1.16-1.29 (1.22); WH1/WH2 1.37-1.50(1.45).
Color bright metallic blue or green. Antennal articles 1 to 3 and base of 4 pale, outer arti-
cles black.
Microsculpture highly effaced on head and pronotum, these areas shiny; irregularly
isodiametric or faintly stretched on elytra, lightly impressed and shiny in many specimens.
Body setose dorsally. Setae evident on head behind and below eyes; on disc of pronotum
most conspicuous along posterior-lateral margins; on disc of elytra forming an irregular row
in at least odd numbered intervals. Ventral portions of body also with short evident setae.
Shape of head similar to v. horni\ eyes convex, neck evidently constricted.
Pronotum as in fig. 36, but in some specimens somewhat narrower with sides more stron-
gly constricted behind; differing from v. viridis by narrowness with greater lateral sinu-
ation; posterior-lateral angles each bearing a setiferous puncture.
Elytra oval, narrowed basally; basal margin strongly sinuate; striae clearly impressed on
disc, outer striae evident almost to apex, strongly punctate; intervals varied, flat to convex,
deeply punctate.
Male genitalia similar in shape to that of v. viridis. Female stylus as in fig. 53.
Variation. - The following characteristics were observed to vary; shape of pronotum; den-
sity of vestiture; microsculpture, and depth of impression of elytral striae. As few specimens
of this subspecies were available, I was unable to determine the pattern of variation in any
of these characters. Variation was most evident among specimens from localities near the
periphery of the range of klamathea, indicating that hybridization with homi may in part at
least be responsible for this variation.
Relationships. - This subspecies is intermediate between viridis s. str. and horni in the diag-
nostic characters presented in fig. 1 . However, on the basis of its peculiar combination of
characteristics, and the apparently isolated geographical position it occupies, I chose to re-
cognize this form as a subspecies.
Etymology. - This name is the latinized form of part of the name of the type locality - Kla-
math Falls, Oregon.
Disposition of type material. -The holotype has been deposited in the California Academy
of Sciences. The allotype is in the United States National Museum. Paratypes have been de-
posited in the Chicago Natural History Museum (4 specimens), Museum of Comparative
Zoology (1), Oregon State University (2), United States National Museum (2) and the Uni-
versity of Washington (3).
Distribution. - Philophuga viridis klamathea is known only from southwestern Oregon and
Carabidae
41
northern California (fig. 59).
Philophuga viridis homi Chaudoir 1877, new combination
Philophuga homi Chaudoir 1877 - 245. Type locality - Nevada.
Philophuga uteana Casey 1924 - 92. Holotype - male, labelled as follows: Stockton, Utah,
IV/7/04, Tom Spalding, Casey bequest 1925, Type USNM 47673, uteana Csy. New Syno-
nymy.
Philophuga cobaltina Casey 1924 - 91. Holotype - male, labelled as follows: Trout Creek
Juab Co., Utah, VII/4/22. Tom Spalding, Casey bequest 1925, Type USNM 47672,
cobaltina Csy. New Synonymy.
Description. - Values for ratios and measurements of ten specimens are: TL, 7.02-8.24 mm
(7.58 mm); LE/LP, 2.77-2.93 (2.83); WP/LP, 1.21-1.27 (1.24); WH1/WH2, 1.38-1.50 (1.45).
Color shiny metallic blue or green. Antennal articles 1 to 3 and base of 4 pale, outer art-
icles black.
Microsculpture absent or highly effaced on head and pronotum; strong on elytra, iso-
diametric medially showing tendency to become strethced and arranged in transverse rows
laterally.
Body glabrous dorsally; even head below eyes without setae.
Head with eyes large, strongly convex; genae constricted posteriorly forming evident
neck.
Pronotum as in fig. 35; lateral margin with long sinuation towards hind angle; hind angles
prominent though somewhat obtuse, each bearing a setiferous puncture.
Elytra with greatest width in apical half but not strongly narrowed basally; basal border
only slightly sinuate; elytral striae clearly impressed, moderately to coarsely punctate; inter-
vals slightly to strongly convex, finely to coarsely punctate. Hind wings fully developed.
Male with three, female with two pair of anal setae on abdominal sternum 6.
Male genitalia similar to that of v. viridis. Female stylus as in v. klamathea.
Distribution. - This subspecies occurs throughout the Great Basin province, and extends a
little outside of this region especially in the north (fig. 59). I have seen more than 90 speci-
mens from the following localities:
United States - ARIZONA: Coconino Co., Flagstaff (USNM), Pinal Mts. (CU), William (Wickhanj USNM).
CALIFORNIA: Lassen Co., Bridgeport (Wickham, USNM), Hallelujah Junction (Westcott, UASM), Susanville (Martin,
CAS), Modoc Co., Davis Creek (Fox, CAS), Hackmore (Van Dyke, CAS). COLORADO: La Plata Co., Pagosa Springs
(Bowditch, MCZ); IDAHO: Ada Co., Nampa (Barr, IUM), Regina (Barr, IUM); Adams Co., Martin (Houk, WUM), Mesa
HUM); Bear Lake Co., Bear Lake (AMNH); Blaine Co., Carey (Hewitt, IUM), Crystal Ice Cave (Westcott, IUM), Magic
Reservoir (Barr, IUM); Canyon Co., Caldwell (Barr, IUM), Parma (WUM); Cassia Co., Elba-Basin Pass (Stecker, SJSC),
Malta (Henry, IUM), Rupert (Shull, IUM); Clark Co., Dubois (Penrose, UASM); Lincoln Co., Richfield (Barr, IUM); Nez
Perce Co., Lewiston (Shull, IUM, USNM); Owyhee Co., Bruneau (Fillmore, IUM), Jordan Valley (Henry, IUM), Reynolds
(Hewitt, IUM); Payette Co., Payette (Shull, IUM); Twin Falls Co., Hollister (Fox, IUSNM), Twin Falls (Homan, IUM).
NEVADA: Esmeralda Co., Lida (O’Brien, UASM); Humbolt Co., Winnemucca (Barr, IUM); Washoe Co., Reno (Wickham,
CAS, MCZ, USNM), Steamboat Springs (Van Dyke, CAS), Verdi (Blaisdell, CAS); White Pine Co., Ely (Van Dyke, CAS).
OREGON: Baker Co., Baker (Van Dyke, CAS, WUM), Durkee (WUM), Pleasant Valley (Fender, WUM), Sparta (Van
Dyke, CAS), Wallowa Mts. (Van Dyke, CAS), Harney Co., Frenchglen (Malkin, WUM), Tencent Lake (Malkin, CNHM
WUM); Lake Co., Hart Mtn. Antelope Refuge (Nelson, OUM); Malheur Co., Harper (OUM), Rome (CAS, OUM). NEW
MEXICO: San Miguel Co., SapeUo (Ball, UASM); UTAH: Touele Co., Stanisbury I. (USNM), Stockton (Spalding, CAS, MCZ);
American Fork (H & S, USNM); Washington Co., St George (AMNH). WASHINGTON: King Co., Seattle (O.B.J., WUM).
42
Larson
Philophuga viridis amoena LeConte 1848, new combination
Cymindis amoena LeConte 1848 - 188. Lectotype - (here selected) female, labelled as fol-
lows: green disc, amoena LeC., Type 5822 MCZ ,P. amoena (LeC.).
Philophuga canora Casey 1913 - 174. Holotype - female, labelled as follows: Tex., Casey
bequest 1925, Type USNM 47669, canora Csy. New Synonymy.
Philophuga puella Casey 1913 - 176. Holotype - male, labelled as follows: Boulder Co.,
Colo., Casey bequest 1925, Type USNM 47671 , puella Csy. New Synonymy.
Philophuga obscura Casey 1924 - 91. Holotype - female, labelled as follows: N.Y., Casey
bequest 1925, Type USNM 47-674, obscura Casy. New Synonymy.
Description. - Values for ratios and measurements of ten specimens are: TL 7.20-8.32 mm
(7.52 mm); LE/LP, 2.68-2.90 (2.80); WP/LP, 1.18-1.24 (1.21); WH1/WH2, 1.28-1.46(1.40).
Color black (elytra piceous in some specimens) with a dull metallic blue sheen on disc of
pro no turn and elytra; very rarely with faint blue-green sheen, usually restricted to frontal
furrows and lateral margins of pronotum. Antennae with articles 1 to 3 and base of 4 pale, at
least ventrally.
Body glabrous dorsally; glabrous even laterally behind and below eye.
Head broad; eyes flattened, somewhat reduced; neck broad. Antennae with outer articles
relatively short and stout.
Pronotum (fig. 34) convex; sides slightly rounded and only shallowly sinuate posteriorly;
hind angles broadly rounded and lacking setiferous puncture on each side; front angles roun-
ded, not or only slightly protruding; lateral margin narrowly and uniformly reflexed.
Elytra relatively convex, elongate oval, with greatest width in apical third; shoulders
broad, basal border slightly sinuate interiorly; striae finely impressed, finely to moderately
punctate; intervals faintly convex basally, flat apically, sparsely and finely punctate.
Abdominal sternum 6 with one pair of anal setae in male, and two pairs in female.
Male genitalia as in v. viridis.
Female stylus as in the other subspecies of viridis.
Variation. - Hybridization between v. amoena and v. horni has been discussed above. Some
variation has been observed in the eastern portion of the range of amoena. Here, hybrid-
ization with horni would not occur.
Only a single female specimen has been seen from Kansas. It differs from the more typical
western and northern specimens of amoena by a greenish-blue cast dorsally, and by
a strongly rounded pronotum with a strong lateral reflexion, narrow anteriorly and broad-
ening behind. The single female specimens seen from Nebraska has a normally propor-
tioned pronotum, but the elytral striae are rather coarsely punctate, and the elytral intervals
are convex and coarse punctate.
Distribution. - This species occurs primarily on the Great Plains east of the Rocky Moun-
tains, and along the northern periphery of the Great Basin (fig. 59).
Canada - ALBERTA: Lethbridge (Larson, UASM); Lost River nr. Onefour (Larson, UASM); Medicine Hat (Carr,
Pepper, CAS, CNC, CU, UASM); Milk River (Pepper, CNC); Ralston (Ball, UASM); Taber (White, DAL). BRITISH COLUM-
BIA: Mts. between Hope and Okanagan (Bowditch, MCZ); “Van.” (= Vancouver?) (MCZ). MANITOBA: Aweme (Criddle,
White, AMNH, CAS, CNC, CU, DAL, USNM); Brandon (AMNH, USNM); SASKATCHEWAN: Cypress Hills (Ball & Lin-
droth, UASM); Roche Percee (Criddle, CNC); Saskatoon (McMillan, CNC).
United States - COLORADO: Boulder Co., Nederland (Stainer, CNC); Chaffee Co., Buena Vista (Wickham,
USNM); Delta Co., Paonia (Van Dyke, CAS); El Paso Co., Colorado Springs (Soltau, Wickham, MCZ, USNM); Garfield Co.,
Carabidae
43
Glenwood Springs (AMNH); Jefferson Co., Denver (Soltau, USNM); Mesa Co., Grand Junction (Titus, USNM); Park Co.,
Trout Creek Pass (Beamer, KUM), Wilkerson Pass (White, KUM); Teller Co., Florissant (Bowditch, MCZ). IDAHO: Bannock
Co., Pocatello (Bruner, USNM); Lemhi Co., Salmon (Wakeland, IUM); KANSAS: “Ks.” (MCZ); MONTANA: Gallatin Co.,
(MUB); Lake Co., (MUB); Lewis and Clark Co., Helena (many collectors, CAS, MCZ, USNM); Missoula Co., Missoula
(MUB); Musselshell Co., (MUB); Valley Co., Hinsdale (MUB); NEBRASKA: “Neb.” (MCZ); OREGON: Wasco Co., Maupin
(Van Dyke, CAS), Warm Springs Indian Reserve (WUM); WASHINGTON: Grant Co., Dry Falls (Burnes, WUM), Grand
Coulee (MCZ), Soap Lake (Hatch, WUM); WYOMING: Bighorn Co., Bighorn Mts. (Edwards, SJSC); Carbon Co., Rawlins
(AMNH); Laramie Co., Cheyenne (Soltau, USNM); Yellowstone National Park (Hatch, Van Dyke, CAS, WUM).
Infernophilus new genus
Type species. - Philophuga castanea Horn 1882, here designated.
The species /. castaneus Horn was described originally as a member of the genus Philo-
phuga, but it differs in several important details from the other species included in that
genus. Further, this species cannot be included in any other known callidine genus. For these
reasons, the genus Infernophilus is established to include castaneus.
The most striking external features of this genus are indicated in the preceding key. Ano-
ther remarkable feature is the orientation of the male genitalia. In repose, the aedoeagus is on
its left instead of on its right side; the latter is the usual position among the members of the
Lebiini. Associated with this, the parameres are reversed. They are of the typical form for
the Callidina, but the large paramere is on the right rather than on the left side of the aedoea*
gus. Also, the apical orifice opens dorso-laterally to the right. I know of no other North Am-
erican callidine, or even lebiine, that shows a similar condition. As no intermediate stages are
known between the normal and the reversed position of the genitalia, it seems plausible that
the initial 1 80° rotation arose in a single step. Other modifications such as the reversal of the
parameres and the shift of the apical orifice may have been secondary.
Etymology. - The name is derived from the Latin noun inf emus - m., hell; and the Greek
philia - fondness. The name refers to the hot desert region in which this insect is found.
Infernophilus castaneus Horn 1882
Philophuga castanea Horn 1882 - 144. Type locality - Kern Co., California. Leng 1920 -
67. Csiki 1932- 1462.
Description. - Values for ratios and measurements for fourteen specimens are: TL 7.9-9. 1
mm (8.6 mm); LE/LP 2.62-2.84 (2.74); WP/LP 1.07-1.25 (1.17); WH1/WH2 1.46-1.56
(1.52).
Color brown; elytra and abdomen darker brown to piceous.
Microsculpture on frons isodiametric, but effaced on most specimens; obsolete on disc of
pronotum; isodiametric on elytra, but partly effaced and shiny. Body sparsely setose above.
Head with eyes prominent, convex. Genae broad behind eyes, with faint postocular pinch;
narrowing posteriorly to form an evident neck. Labrum slightly emarginate. Clypeus with a
single seta on each side; sparsely punctulate medially. Frons with poorly defined rugose
frontal furrows; sparsely punctate medially with deeper setiferous punctures posteriorly
and laterally. Antennae with articles 1 to 3 and base of 4 sparsely hairy; remaining articles
pubescent. Maxillary palpus with terminal article cylindrical. Labial palpus with penultimate
article bisetose; terminal article securiform (narrower in female). Ligula bisetose. Mentum
44
Larson
with a prominent margined tooth.
Pronotum (fig. 30) varied in shape; lateral sinuation long, lateral margins subparallel to-
ward base; hind angles obtuse, with setiferous puncture present; base slightly sinuate later-
ally; frontal angles broadly rounded, slightly protruding; posterior lateral impressions indis-
tinctly limited and continuous with broad lateral grooves; lateral margins uniformly re flexed;
disc transversely rugose laterally.
Tarsal articles sparsely hairy above, densely setose beneath; article 4 emarginate; basal ar-
ticle of middle and hind tarsi with faint median dorsal groove; claws pectinate; male with
articles 1 to 3 of front tarsus and articles 1 and 2 of middle tarsus bearing two rows of scales
beneath.
Elytra completely bordered both apically and basally; striae clearly impressed and well
defined to apex, lightly punctate; intervals broadly convex, bearing small irregular setiferous
punctures; odd-numbered intervals and apex of even numbered intervals with an irregular
row of larger setiferous punctures. Hind wings fully developed.
Abdominal sternum 6 of male with a paired brush of setae along hind margin (fig. 27); in
female brush reduced and in some specimens it is represented by as few as six pairs of mod-
erate to long setae.
Male genitalia (figs. 21-22) with reversed parameres; endophallus with a field of short
spines.
Stylus of ovipositor (fig. 57) short, broad; broadly rounded apically, sparsely setose.
Distribution. - 1 have examined fifteen specimens of this species from the following loc-
alities (fig. 61).
CALIFORNIA: Kem Co. (AMNH); Mono Co., Bridgeport (O’Brien, UASM, Coleville (O’Brien, CAS, DAO, UASM); San
Diego Co., (USNM). NEVADA: (AMNH, USNM).
Biology.- Several specimens in a series taken at Coleville, California (July 6-10, 1966) were
slightly teneral.
Tecnophilus Chaudoir 1877
Tecnophilus Chaudoir 1877 - 240. Type species - Calleida croceicollis Menetries 1843, here
designated.
Philotecnus LeConte 1851 - 175, not Mannerheim 1837 - 42. Type species - Philotecnus
nigricollis LeConte 1851 (= T. croceicollis Menetrie's), here designated.
This is a small genus closely related to Philophuga. Aside from the characters presented
in the key to Callidina, the adults can be recognized on the basis of the cordate pronotum
and the usually dense vestiture of the body.
Characteristics common to all species of Tecnophilus are given in the following des-
cription.
Description. - Beetles 5.7 to 8.0 mm in length. Color various, but elytra always metallic
blue or greea Basal three articles of antenna similarly colored to outer articles.
Body variously punctate and setose dorsally; ventral portions of body moderately to
densely setose.
Eyes prominent, of various convexity; hairy or glabrous. Labrum truncate apically with
slight medial elevation. Frontal furrows shallow, broad, deeply punctate. Frons punctate.
Carabidae
45
Antennal articles 1 to 3 and base of 4 hairy; outer articles pubescent. Maxillary palpus with
fusiform terminal article. Labial palpus with terminal article securiform ; penultimate article
bisetose. Ligula bisetose. Mentum with a prominent margined tooth.
Pronotum cordate, with strong posterior-lateral sinuations; lateral setae present; hind
angles lacking setiferous punctures.
Tarsal articles slender; article 4 at most emarginate; hairy dorsally; tarsal claws simple (in
some specimens may be minutely serrulate); articles 1 to 3 of front tarsi, and 2 and 3 of mid-
dle tarsi of male bearing two rows of scales beneath.
Elytra parallel sided or oval; completely margined both basally and apically; striae evident
and punctate; intervals variously punctate. Hind wings fully developed.
Abdominal sterna 3, 4 and 5 with a pair of medial setae; sternum 6 with at most three
pairs of anal setae; sterna 4 to 6 without long lateral setae.
Male genitalia on right side in repose. Left paramere large; right paramere smaller, bi-
lobed apically. Aedoeagus simple, tube-like, with apical orifice opening somewhat laterally to
the left; endophallus unarmed.
Female stylus with densely setose apex.
Discussion. - Chaudoir (1877) separated Tecnophilus from his Callidides on the basis of the
structure of the ligula and paraglossae, and placed the genus in his Mimodromiides. However,
Horn (1881) carefully studied the ligula and paraglossae of the Lebiini and concluded that
they were of little value in defining groups of higher rank than the genus, and even here he
suggested that they be used with caution. Tecnophilus is certainly a callidine in the sense
used here.
Many of the peculiar features of Tecnophilus , such as the simple tarsal claws, slender tarsi,
cordate pronotum and setose body, may be regarded as adaptations to a terrestrial mode of
life.
Distribution. - The members of this genus are found in alkaline or saline situations in west-
ern North America.
Key to the Species of Tecnophilus Chaudoir
1 . Apex of femur infuscated and contrasting with the otherwise rufous legs
pilatei Chaudoir, p. 45
1’. Femur concolorous, rufous to black; or if femur rufous and infuscated apically, tarsi
also infuscated croceicollis Mene trie's, p. 46
Tecnophilus pilatei Chaudoir 1877
Tecnophilus pilatei Chaudoir 1877 - 239. Type locality - Texas. Horn 1882 - 137. Leng
1920 -67. Csiki 1932- 1462.
Specimens of this species can be readily recognized by possession of the infuscated apices
of otherwise rufous femora. Also, the elytral striae are much more coarsely punctate than in
croceicollis.
Description. - Values for ratios and measurements of ten specimens are: LE 4.44-4.76 mm
(4.56 mm); WP/LP 1.06-1.15 (1.10);WHi/WH2 1.61-1.70(1.65).
Color of head and thorax rufous; abdomen piceous medially, paler laterally; elytra metal-
46
Larson
lie blue or green, often with rufmistic background; antennae and palpi pale; legs pale except
for conspicuous infuscated apical spot on femur.
Microsculpture obsolete on head; lightly impressed, transversely stretched on disc of pro-
notum; isodiametric but shallowly impressed on elytra.
Body densely pubescent, both dorsally and ventrally.
Head narrow, with temporal region strongly constricted; eyes very convex and protruding,
glabrous.
Pronotum (fig. 38) narrow; sides slightly rounded laterally, relatively shallowly sinuate be-
hind; front angles rounded, not protruding; hind angles obtuse, broadly rounded; lateral re-
flexion narrow.
Tarsal article 4 deeply emarginate, almost bilobed but not bearing dense setiferous pads
beneath.
Elytra elongate, parallel sided; striae moderately impressed medially on disc, becoming
obsolete laterally; striae with very coarse often confluent punctures, deepest in basal half
but also evident apically; intervals flat; basal margin strongly sinuate internally; apex ob-
liquely truncate and slightly sinuate.
Abdominal sternum 6 of male with two pairs of anal setae, with three pairs in female.
Male genitalia as in fig. 18; aedoeagus slightly arcuate, apex elongate.
Female stylus (fig. 55) with external apical angle greatly produced.
Relationships. - Specimens of the species pilatei superficially resemble Texas specimens of
the species croceicollis. However, the strikingly different styli of the female ovipositor and
the elongated apex of the male genitalia, associated with the external characteristics men-
tioned above, consistently separate specimens of these two species. The species pilatei is
isolated from the croceicollis complex.
Distribution. - I have seen specimens of this species only from localities along the Gulf
Coast of Texas (fig. 62). I examined 140 specimens from the following localities:
TEXAS: Aransas Co., Goose Island State Park (Larson, UASM), Brazori Co., Freeport (Evans, UASM); Cameron Co.,
Brownsville (UASM, USNM), Port Isabel (Ball, UASM), Cedar Lane (Shaw, KUM); Nueces Co., Corpus Christi (Hubbard &
Schwarz, USNM).
Tecnophilus croceicollis Menetries 1 843
Calleida croceicollis Menetries 1843 - 54. Type locality - California.
The most diagnostic characteristic of this highly variable species is presented in the pre-
ceding key to the species of the genus Tecnophilus.
Description. - Values for ratios and measurements of selected population samples are pres-
ented in tables 6 to 10.
Color highly varied; head and pronotum black to rufous with head always same color or
darker than disc of pronotum; appendages black to rufous, femur uniformly colored, or if
rufous and infuscated apically, tarsus and clypeus also infuscated; elytra black to piceous
with metallic green, blue or purple sheen.
Vestiture varied, but even in least setose specimens setae present on head behind and be-
low eyes, along lateral margins of pronotum and on at least odd numbered elytral intervals;
dorsal vestiture of most specimens quite conspicuous.
Carabidae
47
Microsculpture obsolete on frons; on pronotum consisting of partially effaced transverse
meshes; on elytra isodiametric but shallowly impressed and shiny on many specimens.
Head with neck variously constricted; eyes of various size and convexity (figs. 24 and 25).
Pronotum varied; more rounded laterally and more constricted behind than in pilatei
(figs. 39-42).
Elytra with basal border shallowly sinuate; apex truncate or slightly sinuate; striae finely
to moderately punctate, never as coarsely punctate as in pilatei.
Aedoeagus of male genitalia (figs. 19 and 20) short, arcuate, with short apex.
Female stylus (fig. 56) with apex truncate, not produced.
Geographical variation and subspecies. - The complex geographical variation displayed by
this species makes it very difficult to organize the available population samples into well de-
fined subspecies. Clinal variation occurs in several characteristics, and the general pattern of
variation is discordant. Geographically terminal populations are comprised of individuals
easily distinguished by distinctive combinations of characters, but specimens from inter-
mediate areas are often difficult to associate with the terminal groups. However, as is shown
below, it is possible to distinguish two groups of subspecific rank.
The pattern of variation of each character is described below, and this section is con-
cluded with a summary of this information.
Color. - Over much of the range of croceicollis, the basic color pattern is as follows: elytra
metallic blue or green; head, pronotum, and legs rufous; abdominal sterna piceous. However,
considerable variation in color of each of these body parts occurs.
Data on variation in the color of the elytra among selected population samples of the
species croceicollis are presented in table 3. Briefly, the color of the elytra varies in the fol-
lowing way: populations from the vicinity of San Francisco Bay, and from the Central
Valley of California, possess dark blue or purple elytra; populations from southern Cali-
fornia, Arizona, Nevada, and southern Utah generally have blue elytra, with a very small per-
centage of the specimens possessing greenish-blue or green elytra. The incidence of green
elytra increases southwardly along the Gulf of California, and eastwardly through New Mex-
ico, until green or blue-green is the predominant color in Texas populations. All specimens
seen from Wyoming and Alberta show dark blue elytra. West of the Rocky Mountains in
Idaho and northern Utah, the color of the elytra is highly varied.
Data on variation in the color of the head and pronoturrt are presented in tables 4 and 5.
In specimens that show infuscation of the head and pronotum, the head is always the same
color or darker than the disc of the pronotum. All specimens that have been seen from Mex-
ico, and the United States from southeastern California east to Texas, and north to central
Colorado, Utah, and Nevada, possess rufous heads and pronota. Specimens from the eastern
side of the Rocky Mountains from northern Colorado to southern Alberta, possess black
heads and pronota. I have not seen any specimens from Colorado that were intermediate be-
tween the black color of northern specimens, and the rufous color of specimens from south-
ern localities. This suggests that no hybridization occurs between specimens belonging to
these two different color classes in this region. However, west of the Rocky Mountains in
Idaho and northern Utah, specimens of croceicollis are highly varied in color, with the color
ranging from entirely black to rufous with only a light infuscation of the tarsi and the cly-
48
Larson
peus. This variability in color probably results from hybridization between the black mem-
bers of the northern population, and rufous colored specimens which occupy the Great
Basin. Similar variation in color occurs among populations in the Central Valley of Califor-
nia. Many specimens occurring around San Francisco Bay are black in color. Specimens
from further inland tend to be paler, and specimens from the southern end of the San
Joaquin Valley are almost as pale as specimens from the Mojave Desert. This evidence sug-
gests that there is gene flow from the Mojave Desert, into the San Joaquin Valley.
TABLE 3. Geographical variation in color of elytra of Tecnophilus croceicollis Menetries.
TABLE 4. Geographical variation in color of head of Tecnophilus croceicollis Menetries.
Carabidae
49
TABLE 5. Geographical variation in color of pronotum of Tecnophilus croceicollis Menetries.
Vestiture. - Specimens from southeastern Alberta, and from southeastern California and
the neighboring portion of Arizona, possess very short sparse setae. Over the remainder of
the range of croceicollis, specimens are moderately setose, or in southern New Mexico and
along the Rio Grande in Texas, specimens are very densely setose.
Measurements and ratios. - Five measurements were taken of specimens of Tecnophilus
croceicollis. Various combinations of these measurements were made up to produce four
ratios to map out the general shape of the insects and to quantify observed differences in
habitus between members belonging to different populations of this species. Values for
measurements and ratios of four population samples of the species croceicollis are pre-
sented in tables 6 to 10. Data are given for only four population samples, as these four
are the only homogeneous samples available of sufficient size to yield statistically signi-
ficant results. The analysis of variation which follows is based on these four samples.
Below, the pattern of variation shown by each of these characters is discussed separately.
Length of elytra ( table 6). - This measurement is taken as giving an index of the size of the
beetle. The largest specimens seen were from coastal localities (Brownsville, Texas, X= 4.38
mm; Newark, California, X = 4.21 mm). Specimens from intermediate inland localities were
smaller (Cuddeback Lake, California, X = 4.03) and the smallest specimens seen were from
northern inland localities (Lost River Ranch, Alberta, X = 3.73 mm).
Width of pronotum /length of pronotum (table 7). -The population sample with the highest
mean value for this ratio is from Newark, California (X= 1 .22). The sample from Cuddeback
Lake, California has the lowest mean value for this ratio (X = 1.14). Population samples
from Brownsville, Tocas and Lost River Ranch, Alberta possess intermediate mean values,
and do not differ significantly between themselves.
50
Larson
TABLE 6. Geographical variation in length of elytra (mm) among selected population sam-
ples of Tecnophilus croceicollis Menetries.
Length of elytra/ length of prono turn (table 8). - The pattern of variation shown by this
character is similar to the pattern of variation shown by the length of the elytra. The sam-
ples with the largest mean values are from coastal localities while samples from inland and
northern localities possess smaller mean values for this ratio.
Width of prono turn /maximum width of head ( table 9). - The pattern of variation in this char-
acter is similar to the pattern of variation shown by the character width of pronotum/length
of pronotum. The sample from Newark, California possesses the greatest mean value for this
ratio (X = 1.18) while the sample from Cuddeback Lake, California has the smallest mean
value (X = 1.10). The means for the samples from southeastern Alberta and from Browns-
ville, Texas are identical in value to one another (X= 1.14) and are intermediate between the
mean values for the two California population samples.
Maximum width of head /minimum width offrons between eyes ( table 10). - The sample
with the largest mean value for this ratio is from Brownsville, Texas (X = 1 .63). This is fol-
lowed in descending order by the samples from Cuddeback Lake, California (X = 1.54).
Newark, California (X = 1.45), and southeastern Alberta (X = 1 .41 ). The value for this ratio
becomes smaller for samples from east to west and from south to north.
The above five characters show several different patterns of variation. Because it is dif-
ficult to justify the weighting of any one of these characters above the other four in any
given comparison of population samples, all characters were used simultaneously to develop
discriminant functions to compare population samples with one another. For each compar-
ison of a pair of population samples, a weight was calculated for each character used. The
calculation was such that the value of the weight depended upon the discriminatory value of
Carabidae
51
the character in that comparison (see Stanley MS for details on calculation). That is, if a
character can separate members of two populations consistently, that character has a higher
weighting coefficient than a character that gives inconsistent separations.
For the sake of brevity in the following discussion the population samples used in the
comparisons are designated alphabetically, as follows:
A = Brownsville, Texas
B = Cuddeback Lake, California
C = Newark, California
D = Lost River Ranch, Alberta
Not all possible combinations of population comparisons were made. The Brownsville,
Texas population sample was not compared with the population sample from Newark, Cali-
fornia because both of these populations were independently compared with the geograph-
ically intermediate population sample from Cuddeback Lake, California. Thus, the A versus
C comparison was made indirectly. Similarly, an indirect comparison between Newark, Cali-
fornia (C) and Lost River Ranch, Alberta (D) was made. The populations of croceicollis in
the central valley of California appear to be effectively isolated to the north and east by the
Klamath Mountains and Sierra Nevadas respectively. This means that any genetical connec-
tion between these two populations would have to be indirect, through southern California.
For this reason, the population sample from Cuddeback Lai® was also taken as being inter-
mediate between these two populations.
TABLE 8. Geographical variation in the ratio of length of elytra/length of pronotum among
selected population samples of Tecnophilus croceicollis Menetries.
52
Larson
TABLE 10. Geographical variation in the ratio of maximum width of head/minimum width
of head between eyes among selected population samples of Tecnophilus croceicollis
Me ne tries.
Comparisons between pairs of populations were made in the following way. A Zj value
(the sum of the mean character values multiplied by their individual weighting coefficients)
was calculated for each of the two population samples compared. An axis was plotted on a
map joining the localities from which these population samples were obtained. Using the
weighting coefficient calculated for the original comparison, Zj values were calculated for
samples from geographically intermediate localities that lay along the axis and these Zj val-
ues were plotted on a map. Where the Zj value obtained from geographically intermediate
population samples was equal to or approached the mid-point value, a transverse line was
drawn across the axis. This transverse line theoretically divided members of the two popu-
lations and their associated specimens from one another. If the position in which this trans-
verse line occurred was concordant with a change in the state of one or more other char-
acters such as color, the populations separated by this line were considered to be subspecifi-
cally distinct.
A summary of the population comparisons that have been made is presented below. Table
1 1 gives the weighting coefficients and the calculated Zj values of the population samples
used for each set of comparisons.
TABLE 1 1. Summary of calculated weighting coefficients (W*) and Zj values obtained for
each comparison of a pair of population samples of Tecnophilus croceicollis Menetries.
. . % overlap
Mid-
Comparison point Ob- Calcu-
Wj to W5 are weighting coefficients for characters X\ to X5 respectively, where character
Xj = WP/LP; X2 = LE mm; X3 = WHj /WH2; X4 = SE/LP; and X5 = WP/WHj .
Population samples have been designated alphabetically as follows: A = Brownsville, Texas;
B = Cuddeback Lake, California; C = Newark, California; and D = Lost River Ranch, Alberta.
Carabidae
53
Brownsville, Texas versus Cuddeback Lake, California (Comparison A-B). - Calculated Zj
values for population samples from Brownsville, Texas, Cuddeback Lake, California and for
specimens from intermediate localities are presented in table 12, and are plotted on a map
in fig. 2. The line transverse to the A-B axis separates populations with Zj values above the
mid-point value. This line is not concordant with that of any other character observed to
vary between the two populations.
TABLE 12. Zj values calculated for selected population samples of Tecnophilus croceicollis
Menetries from the discriminant function developed for the Brownsville, Texas (A) vs
Cuddeback Lake, California (B) populations comparison (mid-point Zj value is 1.667).
54
Larson
Fig. 2. Plotted mean Zj values of selected population samples of Tecnophilus croceicollis
Menetries used in the comparison of populations from Brownsville, Texas (A) to Cuddeback
Lai®, California (B). The mid-point Zj value equals 1.667. Association of letters and locality
names are made in Table 1 3.
Brownsville, Texas versus Lost River Ranch, Alberta (Comparison A-D). - Calculated Zj
values for the two population samples compared and for specimens from intermediate local-
ities are presented in table 13 and are plotted on a map in fig. 3. The transverse line represen-
ting the position in which mid-point values are expected to occur extends across northern
Colorado. This line is concordant with a change in color and a change in eye shape.
Cuddeback Lake, California versus Newark, California (Comparison B-C). - Calculated Zj
values are presented in table 14 and fig. 4, in the same way as above. The discriminant func-
tion gives poor separation of specimens from the two populations used in the original com-
parison. It provides even poorer separation of specimens from intermediate localities. Speci-
mens with intermediate Zj values are found in the northern portions of the San Joaquin Val-
ley and this change is not concordant with a change in the state of any other character. Like
color, Zj values vary clinally.
Carabidae
55
Fig. 3. Plotted mean Z\ values of selected population samples of Tecnophilus croceicollis
Menetries used in the comparison of populations from Brownsville, Texas (A) to Lost River
Ranch, Alberta (D). The mid-point Z\ value equals 8.596. Association of letters and locality
names are made in Table 14.
56
Larson
TABLE 13. Zj values calculated for selected population samples of Tecnophilus croceicollis
Menetries from the discriminant function developed for the Brownsville, Texas (A) vs Lost
River Ranch, Alberta (D) populations comparison (mid-point Zj value is 8.596).
Cuddeback Lake, California versus Lost River Ranch, Alberta (Comparison B-D). -Calcul-
ated Z j values are presented in table 1 5 and fig. 5 in the manner outlined above. In this case,
the transverse line representing the approximate location of specimens possessing mid-point
Zj values, is in northern Utah. Z j values for specimens collected from localities in close prox-
imity to this line, tend to intergrade into each other. Color and vestiture also change through
this region.
Summary. - Because of lack of concordance in the variation patterns among the characters
studied, the population samples A, B and C are considered to be consubspecific. However,
several characteristics differentiating population samples A and D change in the same area.
Also, several characteristics differentiating population samples B and D change in the same
area. I conclude therefore, that population sample D and specimens associated with it are
sub specifically distinct from samples A, B and C and their associated specimens.
Carabidae
57
Fig. 4. Plotted mean Z\ values of selected population samples of Tecnophilus croceicollis
Menetries used in the comparison of populations from Cuddeback Lake, California (B) to
Newark, California (C). The mid-point Zj value equals 0.621 . Association of letters and local-
ity names are made in Table 15.
Fig. 5. Plotted mean Zi values of selected population samples of Tecnophilus croceicollis
Menetries used in the comparison of populations from Cuddeback Lake, California (B) to
Lost River Ranch, Alberta (D). The mid-point Zj value equals 6.410. Association of letters
and locality names are made in Table 1 5.
TABLE 14. Zj values calculated for selected population samples of Tecnophilus croceicollis
Menetries from the discriminant function developed for the Cuddeback Lake, California (B)
vs Newark, California (C) populations comparison (mid-point Zj value is 0.621) (fig. 4).
Carabidae
59
TABLE 15. Zj values calculated for selected population samples of Tecnophilus croceicollis
Menetries from the discriminant function developed for the Cuddeback Lake, California (B)
vs. Lost River Ranch, Alberta (D) populations comparison (mid-point Zj value is 6.410).
60
Larson
Key to the Subspecies of Tecnophilus croceicollis Menetries
1 . Specimens from California c. croceicollis Menetries
1*. Specimens from localities other than California 2
2(1). Color of head, pronotum and legs rufous; eyes large and convex (fig. 25)
c. croceicollis Men., p.60
2’. Color not as above; if head, pronotum and legs basically rufous in color, at least tarsi,
clypeus and frons between eyes somewhat infuscated: eyes small and flat (fig. 24) ....
c. peigani new subspecies, p. 61
OR
2a(l).
2a’.
3(2).
3’.
4(2).
Specimens from west of Rocky Mountains 3
Specimens from east of Rocky Mountains 4
(-3.28DWP/LP +(-0.223)LE + (2.023)WHi/WH2 + (1.396)LE/LP + (3.768)WP/WH
>6.410 c. croceicollis Men.
‘WW* < 6.410 peigani n. ssp.
(0)WP/LP + (-0.053)LE + (3.643)WHi/WH2 + (1.165)LE/LP + (0)WP/WH > 8.596 .
4’.
‘’+‘’+‘W’ < 8.596.
c. croceicollis Men.
. c. peigani n. ssp.
Tecnophilus croceicollis croceicollis Mene trie's 1 843
Calleida croceicollis Menetries 1843 - 54. Type locality - California.
Callida chloridipennis Motschoulsky 1850 - 39. (from Horn 1882)
Philotecnus nigricollis LeConte 1851 - 176. Type locality - San Jose, California.
Philo tecnus ruficollis LeConte 1851 - 176. Type locality - San Diego, California.
Tecnophilus glabripennis Chaudoir 1877 - 242. Type locality - Nevada.
Specimens of this subspecies may best be recognized on the basis of the combination of
characters presented in the key to subspecies of croceicollis .
Description. - Values for ratios and measurements of selected population samples of this
subspecies are presented in tables 6 to 10.
Color varied in specimens from the Central Valley of California; head, pronotum and legs
basically rufous with at least some infuscation and in some specimens these parts are black;
elytra dark blue or purple. Specimens from remainder of range with head, pronotum and legs
uniformly rufous; elytra blue to green.
Microsculpture lightly impressed or obsolete on head and disc of pronotum; varied on
elytra from granular to lightly impressed and shiny.
Head with eyes large and convex (fig. 25). Antenna relatively long and slender.
Pronotum varied in shape (figs. 39 to 41); not so cordate as in peigani.
Elytra varied in shape, short and oval in some specimens to more elongate and parallel
sided; striae evidently impressed and distinctly punctate; intervals flat to convex, variably
punctate.
Male genitalia as in fig. 19. Similar to that of T. pilatei but with a shorter apex.
Female stylus (fig. 56) truncate apically, not produced.
Discussion. - The high degree of variation shown by this subspecies has led past authors to
describe a number of species, or to regard croceicollis as a single highly variable species.
Carabidae 61
Notes on synonomy. - The name pilatei Chaudoir is not a synonym for croceicollis
Menetries as is shown above.
Mexico - DURANGO: Durango (Emburg, CAS); SINALOA: El Camaron (Ball, UASM).
United States - ARIZONA: Cochise Co., San Bernardino Ranch (Snow, UASM); Gila Co., Globe (USNM); Navajo
Co., Winslow (USNM); Pima Co., Tucson (MCZ); Yuma Co., Palomas (CU), Wellton (CU). CALIFORNIA: Alameda Co.,
Bay Farm Island (Barr, Dahl, CAS, IUM, MCZ), Newark (Tyson, UASM); Butte Co., Oroville (Kelfer, CAS); Contra Costa
Co., Antioch (Rose, CAS), Brentwood (Van Dyke, CAS), Vine Hill (Blaisdell, CAS); Fresno Co., Mendota (MCZ); Imperial
Co., El Centro (Hanson, Van Dyke, CAS, CU, WUM); Palo Verde (Barr, IUM); Inyo Co., Lone Pine (Van Dyke, CAS) Pana-
mint Valley (USNM), (Nunenmacher, CNHM); Kern Co., Koehn Lake (Erwin, UASM); Kings Co., (CNHM); Lassen Co.,
(Nunenmacher, CNHM); Merced Co., Los Banos (O’Brien, Van Dyke, CAS, UASM); Orange Co., Seal Beach (Gillogly, CAS);
Riverside Co., Coachella (Van Dyke, CAS); San Bernardino Co., Barstow (Hayward, MCZ), Cuddeback Lake (Larson &
Sharp, UASM); Needles (Kusche, CAS); Santa Clara Co., Alviso (Erwin, Larson & Sharp, UASM); San Diego Co., San Diego
(CAS); San Joaquin Co., Weston (Van Dyke, CAS); San Mateo Co., San Mateo (Nunenmacher, Van Dyke, CAS, CNHM);
Solano Co., Benica (Car. M); Sonoma Co., Glen Ellen (Kusche, CAS, CNQ; Stanislaus Co., Patterson (Ross, CAS); Yolo
Co., Davis (Erwin, Hatch, Car. M., UASM, WUM); Yuba Co., Marysville (Van Dyke, CAS). COLORADO: Bent Co., La Junta
(Hayward, MCZ); El Paso Co., Colorado Springs (Soltau, USNM); Freemont Co., Florence (Soltau, USNM); Pueblo Co.,
Pueblo (Soltau, USNM). NEVADA: Churchill Co., Humbolt Lake (Wickham, USNM); Clark Co., Las Vegas (Barr, Johnston,
CNC, IUM); Pershing Co., Lovelock (Baker, SJSC). NEW MEXICO: Hidalgo Co., Lordsburg (Howden, CNC); McKinley Co.,
Ft. Wingate (Dow, CAS); Otero Co., Alamogordo (Wickham, USNM); Taos Co., San Juan Valley (Bowditch, MCZ).
TEXAS: Cameron Co., Brownsville (many collectors, CNC, INHS, MCZ, UASM, USNM); El Paso Co., Fabens (Howden,
CNC); Hidalgo Co., Mission (Gurney, USNM); Reeves Co., Pecos (Johnston, CNC), Pecos River? (Odenbach, USNM).
UTAH: Washington Co., St George (AMNH, CAS, MCZ, USNM).
Tecnophilus croceicollis peigani new subspecies
Holotype - male, Milk River near junction with Lost River, Lost River Ranch, Alberta (12
V/1965, Getty & Larson, CNC).
Allotype - female, same locality (CNC).
Paratypes. - All other specimens from Alberta have been labelled as paratypes (see below for
list of localities).
The diagnostic characteristics of this subspecies are presented in the above key, and in the
discussion of geographical variation in T. croceicollis.
Description. - Values for ratios and measurements of this subspecies are presented in tables
6 to 10.
Color black, with elytra and in some specimens disc of pronotum with dull metallic blue
or blue-green lustre; appendages black or piceous (see following discussion for variation in
color).
Microsculpture lightly impressed or obsolete on head and disc of pronotum ; isodiametric
and coarse on elytra.
Vestiture very short and sparse on specimens from Alberta. Specimens from more south-
erly localities moderately setose.
Head broad; eyes small and little convex (fig. 24). Antenna shorter and stouter than in
c. croceicollis.
Pronotum strongly cordate (fig. 42), lateral margins strongly constricted behind, lateral
reflexion narrow.
Elytra short and oval, with greatest width in apical half; striae deeply impressed and evid-
ently punctate; intervals convex, each bearing an irregular row of coarse punctures.
62
Larson
Male genitalia (fig. 20) and female stylus similar to c. croceicollis.
Variation. - Vestiture and color vary over the range of this subspecies. Specimens from the
eastern side of the Rocky Mountains are black or at least dark piceous in the color of the
head and pronotum. Specimens with this coloration have also been seen from eastern Idaho
and extreme northern Utah. However, specimens from central and western Idaho and the
Salt Lake region of Utah are much paler in color. In specimens from these regions, the
ground color of the head, pronotum and legs is usually rufous, with at least some infuscation
occurring on the legs, clypeus and the frons between the eyes. This tendency towards pale
coloration in specimens of peigani from Idaho and Utah is probably a result of hybridization
with c. croceicollis.
Specimens of peigani from Alberta appear to be almost glabrous dorsally because of the
greatly reduced length of the setae. Over the remainder of the range, the dorsal pubescence
is longer and quite distinct.
Etymology. - The subspecific name is the latinized form of the word Peigan, the name of a
tribe of Indians of the Blackfoot Confederation, which inhabited the prairies of southern
Alberta. Confusion exists as to the correct spelling of this name, but the Edmonton office of
the Canadian Government Department of Indian Affairs accepts “Peigan” as the correct
spelling.
Disposition of type material. - The holotype and allotype have been deposited in the
Canadian National Collection, Ottawa. Paratypes have been deposited in California Academy
of Sciences, Canadian National Collection, Museum of Comparative Zoology, University of
Alberta, Strickland Museum, and the United States National Museum.
Distribution. - Localities from which specimens of peigani have been collected are plotted
in fig. 63. I have seen 73 specimens of this subspecies from the following localities.
Canada - ALBERTA: Forty Mile Coulee, 5 miles north Etzikom (Getty, Larson, Whitehead, UASM); Milk River near
junction with Lost River, Lost River Ranch (Ball, Erwin, Freitag, Getty, Larson, CAS, CNC, MCZ, UASM, USNM); Picture
Butte (Getty & Larson, UASM).
United States - COLORADO: “CoL” (USNM). IDAHO: Butte Co., Howe (Barr, IUM); Cassia Co., Malta (Henry,
IUM); Strevell (Barr, IUM); Owyhee Co., Grandview (Fumiss, IUM). MONTANA: Beaverhead Co., Dillon (Jellison, MUB).
UTAH: Cache Co., Logan (Henderson, CAS); Davis Co., (Stafford, USNM); Salt Lake Co., Salt Lake City (J.B., Henderson,
Huelleman, Klages, Car. M., IUM, WUM). WYOMING: Carbon Co., Medicine Bow (AMNH), Saratoga (Bryant, CAS);
Sublette Co., Pinedale (Alexander, MCZ); Sweetwater Co., Creston (Bryant, CAS), Green River (Bowditch, MCZ).
BIOLOGY
Although little is known of the biology of beetles of the subtribe Callidina, members of
many of the species and genera appear to be at least partly arboreal. Habu (1960, 1967)
summarized what is known of the biology of members of the genus Callida. Generally, these
insects are arboreal predators, feeding mostly on lepidopterous larvae. These insects do not
show any of the parasitic adaptations possessed by the members of the genus Lebia.
What I have observed of the biology of the genera Tecnophilus and Philophuga is presen-
ted below.
Carabidae
63
Biology of Tecnophilus Chaudoir
The biology of these insects is poorly known. Adults of this genus tend to be more ter-
restrial than are adults of any of the other North American callidine genera. This is reflected
in the slender legs and tarsi, setose body and the cordate prothorax of Tecnophilus ; char-
acters which are often found in terrestrial carabids, but seldom found in arboreal forms.
Adults are found on saline or alkaline soils in eroding areas. Table 1 6 presents some of the
physical characteristics of the top six inches of the soils on which these insects have been
collected. The high soil salinity and the arid conditions in regions in which members of the
genus Tecnophilus occur, suggests that desiccation may be an important factor with which
these insects must contend. During relaxation of specimens prior to dissection, I noticed that
the dried tissues were quite impermeable to water. Immersion in near-boiling water for per-
iods of fifteen minutes to half an hour was not sufficient to soften the abdominal memb-
ranes enough to permit the genitalia to be withdrawn easily. This is a longer time than is re-
quired to relax most species of carabids. This impermeability may be an adaptation to resist
desiccation under arid conditions.
In southeastern Alberta, specimens of Tecnophilus croceicollis peigani are usually found
along the bases of south facing coulees, where considerable erosion has occurred and alluvial
fans have been deposited. These fans are composed of a clay-sand soil which bakes and
cracks extensively when dry. The sparse vegetation consists principally of Bouteloua gracilis
(Lag.) H.B.K., Atriplex nuttalli Wats., Opuntia spp., Mamillaria vivipara Haw., and Erigonum
flavum Nutt. A more complete description of the region is presented by Lewin (1963).
On sunny days in May and early June, specimens of c. peigani were found running over
the surface of the ground, or climbing on very low vegetation. Infrequently specimens have
been taken under cover or from cracks in the ground. At this time, the insects are often
found in copulation, and mating occurs readily among captured specimens. Fertile females
collected at this time and brought into the laboratory, usually laid eggs within a week or
two. Larvae emerging from these eggs were reared as far as the third instar on the following
foods: larvae of several species of curculionids, wheat stem sawfly larvae, wax moths, meal
worms, and ant larvae and pupae. Mortality was high on all of these foods.
This subspecies probably hibernates in the adult stage, for one specimen from Green
River, Wyoming (20-27/VII/l 877), was teneral, indicating that emergence from the pupal
stage occurs during the middle of the summer. All specimens are fully winged but flight has
not been observed.
The habitats in which specimens of c. croceicollis were found, vary over the wide range of
this subspecies. The most consistent features of the habitats are saline eroding soils and
sparse vegetation.
In the vicinity of San Francisco Bay, beetles have been collected on salt flats, usually in
areas dominated by plants of the genus Salicomia. During the winter months, these insects
are found near or at the surface of the soil, under the Salicomia mats or other loose cover
(Erwin, pers. comm.). However, on July 5, 1966 (Alviso, California, Larson & Sharp, col-
lectors) seven specimens were collected at depths of four to eight inches in the sandy soil
beneath Salicomia plants. The beetles were on the upper surface of the clay hardpan, and in
company with specimens of Amara stupida LeConte and Pristonychus complanatus Dejean.
64
Larson
The two teneral specimens that I have seen from the central valley of California (Marysville,
Yuba Co., VI/1908; and Mendota, Fresno Co., V-VIII) suggest that the larval stage occurs
during the spring and summer in this region.
Specimens of c. croceicollis have been collected from the margins of several alkaline
playas in the Mojave Desert of southeastern California. A long series of beetles collected at
Cuddeback Lake (June 30, 1966, Larson & Sharp) was found among the buried portions of
the crowns of bur sage ( Franeria dumosa Grey) at depths of three to ten inches. Specimens
collected at the neighboring Koehn Lake in the spring (April 10, 1965, Erwin) were found at
the surface of the soil, under loose cover. I have seen teneral specimens from Lone Pine,
California (20/V/1937). The larval stage probably occurs in the early spring as this is the
season of greatest rainfall in the Mojave Desert.
All long series of c. croceicollis that I have seen from Texas, have been collected at light. I
do not know the habitat of this insect in Texas, but it appears to occur only in inland local-
ities and not along the Gulf Coast. On the other hand, T. pilatei is found primarily along the
coast and only for a short distance inland. A mixed series of both of these species was col-
lected at light in Brownsville, Texas (May 1967).
Outside of Texas, I have seen only one specimen of croceicollis (s.e. California) labelled as
being collected at light.
Most specimens of the species pilatei have been collected at light. However, some at least
have been collected by sweeping vegetation along sea beaches (Port Isabel, Ball).
In order to obtain the larvae of Tecnophilus, I maintained adults of c. croceicollis and c.
peigani alive. The method of oviposition was observed, and was found to proceed as follows
for both subspecies: when ready to lay an egg the female collected a small ball of soil parti-
icles on the apex of her abdomen. These particles were first loosened with the mandibles
then picked up singly with the apex of the abdomen to which they adhered, probably by
means of an adhesive substance produced by the accessory glands. When sufficient material
was collected, the female climbed some object such as a twig. When she had climbed to a
height of several inches, the beetle turned and faced downward. The abdomen was pressed
against the twig, and a drop of fluid was released. The abdomen was then moved away from
the twig, drawing the drop of fluid out until it hardened into a silk-like strand. This strand
was formed partly by the movement of the abdomen, and partly by means of an apposable
motion of the styli. When the thread reached a length of about 1 to 6 mm, the female in-
jected an egg into the ball of soil particles that she still carried on the apex of her abdomen.
The egg and soil covering were then released, and were left dangling in the air, suspended
from the twig by means of the silken thread. The egg remained suspended in this way until
the larva hatched.
This method of oviposition may be related to the peculiar form of stylus that is found in
members of the Callidina. Philophuga viridis amoena, which possesses a similar form of ovi-
positor, lays its eggs in the same way. This form of egg laying could certainly be a valuable
adaptation for desert species, for it would protect the eggs to some extent from most pre-
dators, flash floods, and soil erosion. On the other hand, it would make the eggs more sus-
ceptible to desiccation. As the rest of the members of the Callidina possess this form of ovi-
Carabidae
65
positor, and are more or less arboreal, this form of egg laying was probably an adaptation to
an arboreal existence, and has secondarily proved useful to desert dwelling species.
TABLE 16. Some physical characteristics of soils on which specimens of Tecnophilus
croceicollis Mene tries have been collected.
Biology of Philophuga Motschoulsky
As I have collected few specimens of the genus Philophuga , I know little of the habits of
these insects. In general, members of this genus occur in arid and semi-arid regions of west-
ern North America. They are not usually found in saline situations as are members of the
genus Tecnophilus , and they appear to be much more arboreal than the latter, for specimens
of Philophuga are often found on vegetation.
In southern Alberta, specimens of P. viridis amoena are found in the same general region
as are specimens of T. croceicollis peigani. However, rather than occurring along the bases
of coulees in regions of baked soil and sparse vegetation, they are usually found on short
grass prairie.
Many of the specimens of P. viridis homi that I have examined, are associated with labels
indicating that they have been collected on vegetation, usually Artemisia or A triplex. I know
nothing of the biology of the other subspecies of viridis except that most specimens of v.
viridis that I have examined were collected during the month of December.
The labels associated with many adult specimens of P. viridicollis indicate that the beetles
were collected on vegetation. Four specimens of this species collected at Monahans, Texas
(Larson & Sharp, June 12,1966) were found on a short legume, in leaf rolls made by a lepi-
dopterous larva. Larvae of viridicollis have been collected at Rocky Ford, Colorado (Hamil-
ton, USNM) feeding on the larvae of Ancylis comptana W. & R. (Lepidoptera : Olethreut-
idae). Teneral adults of this species have been collected from April 28 to July 5.
Specimens of P. brachinoides have been collected in bromeliads (Ball & Whitehead).
I have not seen any records of specimens of the genus Philophuga collected at light.
66
Larson
In this genus, oviposition has been observed only for the subspecies viridis amoena. It pro-
ceeds in the same way as outlined under Tecnophilus croceicollis.
PHYLOGENY
The following discussion is an explanation of the rationale used to construct the phylo-
genetic diagram presented in table 17. In order to construct this model, two assumptions had
to be made: (a) it is possible to discover those characteristics that are widespread in the
group due to common ancestry, and to distinguish these from similar characteristics that
have resulted from convergence (Cain & Harrison 1960); and (b) the rate of evolution has
been uniform throughout the group, so that the degree of difference between two taxa is
directly proportional to the length of time for which the two ancestral stocks have been iso-
lated. These assumptions are necessitated by the complete lack of fossil evidence.
Based on these considerations, the following hypothetical model of the evolution of the
North American Callidina is presented. The ancestral stock was probably arboreal, and pos-
sessed the following characteristics: color of elytra metallic blue or green, color of remainder
of body rufous to black; dorsal surface of body glabrous; labial palpus with terminal article
securiform, penultimate article bisetose; ligula bi- or pluri-setose;prothorax broad, with wide
lateral reflexion; legs stout, with tarsal article 4 bilobed, claws pectinate, ventral surface of
tarsal articles 1 to 4 of front and middle tarsi of male bearing two rows of scales beneath;
elytra completely bordered basally and apically, striae evident and punctate, intervals flat or
slightly convex; hind wings fully developed and lightly pigmented; abdominal sterna 4, 5, 6
lacking lateral setae, sternum 6 with a small number of anal setae (probably three pairs or
less); male genitalia on their right side in repose, with left paramere large; endophallus armed
or unarmed; female stylus rectangular, lightly sclerotized, with a setose apical margin.
An early differentiation of the basic callidine stock may have given rise to the ancestor of
the Plochionus-Onota-Cylindronotum lineage. This group differs in many details from the re-
maining members of the North American Callidina, and has been included in the Callidina
largely on the basis of the structure of the female ovipositor. The structure of the ovipositor
as well as other external features of this group may resemble the condition found in the
hypothetical callidine ancestor simply through convergence. If this is the case, the Callidina
represents a grade, resulting from parallel adaptation to a common way of life, an arboreal
habitat. As mentioned above, the affinities of this group are obscure and probably are among
the poorly understood neotropical fauna. For this reason, this group will not be considered
further.
A later differentiation of the ancestral stock of the Callidina probably gave rise to the gen-
era Callida and Lecalida with very little additional modification. The genera Philophuga,
Infemophilus and Tecnophilus are closely related, and were probably derived from a com-
mon stock. This stock differentiated from the main Callida lineage by acquiring slender legs
(tarsal article 4 not bilobed, but only shallowly emarginate) and a setose body.
The most distinctive of these three genera is the genus Infemophilus . For this reason, the
ancestor to the genus Infemophilus may have diverged from the Philophuga-Tecnophilus
Carabidae
67
TABLE 17. Hypothetical phylogeny of the endemic North American genera of the subtribe
Callidina and their included species.
Hypothetical ancestor to the Callidina
castaneus
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Larson
stock early in its history, in order to permit the Infemophilus stock sufficient time to ac-
quire the characteristics listed below. This genus exhibits a peculiar form of male genitalia
which has undergone 1 80° rotation. The mechanism which brought this reversal about is un-
known, but it seems reasonable to postulate that the initial change involved a 1 80° shift in
the orientation of the aedoeagus. The reversal of the parameres may have accompanied this
initial reversal of the aedoeagus or they may have rotated at a later date. In this genus, other
peculiar characters include the anal brushes, the non-metallic elytra, and the short, rounded
female styli.
The stocks giving rise to Philophuga and Tecnophilus separated sometime after the separ-
ation of the Infemophilus stock. The stock giving rise to the genus Philophuga remained es-
sentially unmodified after this point, but the stock ancestral to Tecnophilus acquired simple
tarsal claws, a more cordate pronotum, and a more densely setose body.
The stock giving rise to the genus Philophuga probably possessed the following characters,
as these are the most widespread characteristics in species of this genus: color of entire body
except for basal antennal articles, black with a metallic blue or green lustre dorsally; body
sparsely setose; pronotum sub-cordate, bearing posterior-lateral seta on each hind angle;
elytra elongate and more or less parallel sided, with greatest width in apical half; hind wings
fully developed, lightly pigmented.
An early derivative of this stock gave rise to the ancestor to the viridis complex. This
stock was characterized by shorter, more oval shaped elytra and by a more cordate pro-
notum. The stock giving rise to viridis has undergone considerable recent diversification to
give rise to the four extant subspecies. These four subspecies may have developed simultan-
eously from four isolated populations, as no subspecies is strikingly distinct.
The other ancestral stock of the genus Philophuga gave rise to the extant species viridi-
collis, caerulea and brachinoides . The ancestor to these species possessed deeply pigmented
wings and elongate elytra. This stock gave rise first to the uniquely colored species brach-
inoides which also differs from the other two species in this group by having a more cor-
date pronotum, and also possesses several patches of setae on the ventral surface of the
body. The two extant species caerulea and viridicollis are the result of relatively recent di-
versification in the ancestral stock.
The ancestral stock of the genus Tecnophilus may have divided early in its history to give
rise to the species pilatei and croceicollis. Recent differentiation has produced two sub-
species and a great deal of geographical variation in the species croceicollis .
ZOOGEOGRAPHY
The subtribe Callidina is represented in all major zoogeographical regions of the world.
However, the greatest diversity in both numbers of genera and species occurs in tropical and
subtropical areas. Only two genera contain members in both the Old and New Worlds. The
majority of the species included in the genus Plochionus are restricted to the New World, but
the species Plochionus pallens Fabricus has been widely distributed by commerce, and is now
almost cosmopolitan. Only the genus Callida is represented naturally in Asia, the Americas
and perhaps Africa (but see Jeannel 1949).
Carabidae
69
At present, the range of the genus Callida is disjunct. The majority of the American
species occur in the tropics, but some species in eastern North America extend north into
the southern portion of the cold temperate region. No members of the genus Callida occur
in western North America, hence the New World species of Callida are separated from the
oriental members of this genus by a wide geographical gap of apparently unsuitable habitat.
In western North America, the genera Lecalida, Philophuga, Infernophilus and Tecno-
philus, replace the genus Callida.
Many of the morphological and biological adaptations shown by members of the subtribe
Callidina, are adaptations for an arboreal existence. It then seems reasonable to assume that
present and past distributions of these insects have paralleled the distributions of floral types
that have provided suitable habitats.
While no fossil record is available for this group of carabids, adequate fossils have been
discovered to enable us to extrapolate the movements of Tertiary floras in North America.
Thus, it is necessary to review the events influencing floral dispersal in western North Amer-
ica during the Tertiary and Pleistocene in order to reconstruct the history of the callidine
genera which are endemic to this region.
The following review is based largely on a paper by Axelrod (1959). Other papers from
which this brief review is drawn are: Axelrod 1948, 1950, 1957, 1958; Blackwelder 1948,
1954; Cohn 1965; Dillon 1956; King 1958; MacGinitie 1958, and Martin 1958.
During the early Cenozoic, three geofloras were found in western North America. The
southern portion of the continent was occupied by the broad-leaved evergreen Neotropical-
Tertiary Geoflora, while the northern and central portions were covered by the temperate
Arcto-Tertiary Geoflora of mixed deciduous hardwoods and conifers. Between them, in
central southwestern North America, small areas of semi-arid vegetation of the Madro-Ter-
tiary Geoflora were making their initial appearances.
The Eocene environment was one of low relief and low altitude, and the climate was
warm and humid. These conditions permitted widespread distribution and migration of
plants, and during this time, the Neotropical-Tertiary Geoflora moved northward along the
Pacific coast into Alaska, and to or near the Canadian border interiorly. This Geoflora was
dominated by broad-leaved evergreens of tropical families and genera which now find their
closest counterparts in subtropical forests such as those presently found from southern Mex-
ico to Panama, and in southern Asia, where annual rainfall is high and the climate is uni-
formly warm. The Coast Ranges and the eastern Cordilleras did not exist at this time, and the
desert flora that now occupies the intervening region was not then in existence.
The cooler, drier climate following the Eocene gradually restricted the Neotropical-Ter-
tiary Geoflora southward, and coastward where it persisted in warm coastal valleys of Cali-
fornia and Oregon until into the Miocene, with a few relicts surviving into Pleistocene times.
No plants in the United States today represent direct descendants from this flora. However,
some genera and species were secondarily derived from it. These derivatives represent sub-
tropic to warm temperate groups that became adapted to the expanding dry subtropical and
warm temperate areas of southwestern North America.
During the early Tertiary, the Arcto-Tertiary Geoflora had a holarctic distribution at high
and middle latitudes, and probably was continuous across the Bering Land Bridge that was
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Larson
present during early Eocene times. In western North America, three principal elements com-
prised this Geoflora: the western American element consisting of species related to present
day dominants of the coastal and mountain forests; the eastern North American element
consisting of deciduous hardwoods related to those of present eastern North America; and
the East Asian element consisting of mixed deciduous forest species which are no longer
native to North America, or have a discontinuous distribution in eastern North America and
eastern Asia.
Species of these three elements were mixed in a forest of a rather generalized floristic
composition. However, the Geoflora was not homogeneous throughout its area. It was com-
posed of several forest types, depending upon geographical occurrence and climatic condi-
tions. In response to gradual development of increasingly more emergent continents follow-
ing early Tertiary, and to the accompanying trend toward lower temperature, this Geoflora
gradually migrated southward from near the Canadian border where it occurred in the early
Eocene, to central Nevada and northern California by the middle Oligocene. During the
Miocene and early Pliocene, the increasing aridity further restricted this flora to more humid
coastal and upland sites. Species belonging to the eastern North American and eastern
Asian elements were rapidly reduced in western North America during the late Tertiary due
to the reduction in summer rains brought about by uplifts in the coast ranges and Sierra
Nevadas. Only a few of these forms lingered on into the late Pliocene in the mild coastal
strip from central California north, becoming extinct here in the early part of the Pleisto-
cene.
The species now dominating the western coniferous forests represent the surviving west-
ern American element of the old Arcto-Tertiary Geoflora. In the west, its species became
adapted to a climate which was typified by winter rain and summer drought. Its major com-
munities were differentiated chiefly in the latter Pliocene when important topographical
changes such as the rise of the Sierra Nevada and the coast ranges produced more diverse
climates.
The Madro-Tertiary Geoflora which originated in southwestern North America, was com-
prised of small-leaved, drought-deciduous, sclerophyllous plants. These plants were derived
primarily from the Neotropical-Tertiary Geoflora and to a lesser extent from the Arcto-
Tertiary Geoflora in response to the expanding dry climate. Fossil plants, apparently an-
cesteral to Madro-Tertiary species have been discovered in late Cretaceous and Paleocene
floras of southwestern North America. This Geoflora had come into existence by the middle
Eocene, and spread over the southwestern part of the continent during succeeding epochs
as dry climates expanded. Part of the desert flora as well has been derived from this Geo-
flora.
In the middle Pliocene, the rapid expansion of dry climates almost completely eliminated
woodland and chaparral from the lowlands of the present desert regions. In the later Plio-
cene and Pleistocene, the rapidly rising Cascades, Sierra Nevada and more easterly mountain
ranges brought a drier climate to their lee, and a desert flora came into existence. Its species
were derived from those represented in the Geofloras that dominated the area earlier in the
Pliocene. The colder Great Basin received increments from the surviving Arcto-Tertiary
Geoflora; the Sonoran and Mojave Deserts from the Madro-Tertiary Geoflora.
Considering the present distribution of the genus Callida, it seems that the coniferous
Carabidae
71
forests, semi-arid regions and colder climates of northern and western North America pro-
vide unsuitable habitats for the species of this genus, and effectively isolate the few Asian
species of Callida from the diverse American fauna. This distribution suggests that a faunal
connection between Asia and North America occurred in the past.
The Bering Land Bridge which existed during Eocene times was probably forested by
elements of the Arcto-Tertiary Geoflora, which at this time had a holarctic distribution at
high latitudes. This would provide a pathway through which the ancestors of the Oriental
species of Callida could disperse from the New World. At this time, the ancestral stock of
Callida was probably trans-continental in North America. Following the Eocene, cooler cli-
mates restricted the Arcto-Tertiary Geo flora southward, away from the Bering region, per-
manently isolating the American and Asian callidine faunas. The latest at which this separ-
ation could have occurred was the Oligocene or the early Miocene. During the Miocene and
Pliocene epochs, increasing aridity in western North America gradually eliminated much of
the deciduous element of the Arcto-Tertiary Geoflora from this region, and with this sector
of the flora, the ancestral species of Callida were restricted to the eastern and southern por-
tions of the continent.
With the expansion of dry climates and the Madro-Tertiary Geoflora during the late Ter-
tiary, a stock of the ancestral Callida developed adaptations for a more terrestrial existence,
and occupied the semi-arid and arid regions of the west.
The descendants of this stock, Philophuga, Tecnophilus, and In fernophilus , more closely
resemble eastern North American species of Callida than they do Mexican species. Hence,
they were probably derived from the Arcto-Tertiary fauna, rather than from a Neotropical-
Tertiary ancestor. On the other hand, the genus Lecalida resembles many of the Mexican
species of Callida , and may be a northern extension of this group into the North American
deserts. Thus, it is probably derived from a Neotropical-Tertiary ancestor.
The time and place of origin of the genus Infemophilus are obscure, for I know nothing
of the habits of this genus and hence cannot trace the development of the necessary habitat.
After acquiring the basic terrestrial adaptation of slender legs, the ancestral stock of
Philophuga has persisted with little modification. The derivative stock giving rise to
Tecnophilus has acquired a number of specializations. This stock was derived from the stock
ancestral to Philophuga probably during the Pliocene, as increased aridity resulted in the
production of suitable habitats in the form of large salt flats and alkaline playas.
During the early Pliocene, the environment of western North America was one of low re-
lief, and extensive plains covered much of this region. Mountains were not effective barriers
at this time. During this period, the ancestral stock of Philophuga may have ranged widely
throughout this area. Crustal unrest and the renewed mountain building processes of the
later Pliocene probably isolated populations that evolved into the ancestors of brachinoides,
viridis and caerulea-viridicollis.
The subspecies of viridis and the species caerulea and viridicollis do not appear to be com-
pletely separated from one another at present by geographical barriers such as mountains.
The isolations that led to the evolution of these groups probably developed during glacial
periods of the Pleistocene when mountain glaciers separated populations in the northern
deserts and increased rainfall resulted in expanding forests which produced ecological bar-
72
Larson
riers in the southern deserts.
At this time, the same factors isolated populations of Tecnophilus croceicollis , and pro-
duced the geographical variation now shown by this species. I do not know what factors led
to the isolation of croceicollis and pilatei. Perhaps pilatei was isolated on the Gulf Coast of
Texas, and the presence of croceicollis in Texas is the result of a recent invasion from the
west.
Floral movements have profoundly influenced the distribution and evolution of the Cal-
lidina. Thus, the endemic callidine fauna now found in western North America is the result
of climatic, geological and resulting floral changes which have occurred in this region in Ter-
teary times.
ACKNOWLEDGEMENTS
It is a pleasure to acknowledge the contributions of the following people and to extend my
thanks to them.
G.E. Ball, my supervisor, was most helpful with his kind assistance and guidance during
the course of this study. The National Research Council of Canada provided financial sup-
port through Grant No. NRC-A1399, held by G.E. Ball. Much of the Mexican material re-
ported above was collected in the course of an expedition financed by the National Science
Foundation through Grant GB 3312, also held by Ball.
B. Hocking and D.A. Boag, members of my committee, read and criticized the manuscript.
My wife, Margaret Larson, offered many valuable suggestions and much encouragement.
My colleagues, J. Barron, T.L. Erwin, R. Freitag and D.R. Whitehad contributed many
ideas in the discussions I had with them. Richard Freitag examined Casey’s and LeConte’s
type material for me.
Specimens of Tecnophilus , both preserved and living, were provided by T.L. Erwin, D.
Kavanaugh and W. Tyson. Mr. W. Sharp accompanied me on a collecting trip through western
United States and assisted me in the collection of many specimens. My friend, Mr. R. Getty,
collected many specimens of Tecnophilus croceicollis in southern Alberta.
G.A. Hobbs, N.D. Holmes, Ruby I. Larson, C.E. Lilly, and R.W. Salt and many others of
the Canada Department of Agriculture Research Station, Lethbridge, Alberta, did much to
stimulate my interest in entomology and to provide continued encouragement. A.L. Lagler
of that institution wrote the computer program for the analysis of variance used in this
study. E.T. Gushul assisted in the preparation of the illustrations.
I thank the following individuals for lending me material in their care: R.T. Allen, Illinois
Natural History Survey;N. Anderson, University of Montana; W.F. Barr, University of Idaho;
E.C. Becker, Canada Department of Agriculture, Ottawa; P.J. Darlington, Jr., Museum of
Comparative Zoology; H. Dybas, Chicago Natural History Museum; M.H. Hatch, University
of Washington; H.B. Leech, California Academy of Sciences; L.L. Pechuman, Cornell Univer-
sity; P. Spangler, United States National Museum; P. Vaurie, American Museum of Natural
History; and G.E. Wallace, Carnegie Museum.
Carabidae
73
REFERENCES
Axelrod, D.I. 1948. Climate and evolution in western North America during middle Pleocene
time. Evolution 2 : 127-144.
Axelrod, D.I. 1950. Evolution of desert vegetation in western North America. Publ. Cameg.
Instn. 590 : 215-306.
Axelrod, D.I. 1957. Late Tertiary floras and the Sierra Nevadan uplift. Bull. geol. Soc. Amer.
68 : 19-46.
Axelrod, D.I. 1958. Evolution of the Madro-Tertiary Geoflora. Bot. Rev. 24 : 433-509.
Axelrod, D.I. 1959. Geological history. In Munz and Peck, A California flora. University of
California Press, Los Angeles, Calif. 1681 pp.
Ball, G.E. 1963. Carabidae, Fascicle 4. In Arnett, R.H. The beetles of the United States (A
manual for identification). The Catholic University of America Press, Washington, D.C.
55-210, 58 figs.
Ball, G.E. 1966. A revision of the North American species of the subgenus Cryobius Chaud-
oir ( Pterostichus , Carabidae, Coleoptera). Opusc. ent., Suppl. 28, 166 pp.
Bates, H.W. 1883. Biologia Centrali-Americana, Insecta, Coleoptera, Carabidae, Cicindelidae
suppl., l(pt. 1) : 1 53-256, plates vi-xii.
Bigelow, R.S., and C. Reimer. 1954. An application of the linear discriminant function to in-
sect taxonomy. Can. Ent. 86(2) : 69-73.
Blackwelder, E. 1948. The Great Basin. 1. The geological background. Bull. Univ. Utah geol.
Ser. 38(20), 16 pp.
Blackwelder, E. 1954. Pleistocene lakes and drainage in the Mojave region of southern Cali-
fornia. Bull. Calif. Min. Bur. 170 : 35-40.
Blackwelder, R.E. 1944. Checklist of the Coleopterous insects of Mexico, Central America,
the West Indies, and South America. Part 1. U.S. Natl. Mus. Bull. 185, 188 pp.
Cain, A.J. and G.A. Harrison. 1960. Phyletic weighting. Proc. zool. Soc. Lond. 135 : 1-31.
Casey, T.L. 1913. Studies in the Cicindelidae and Carabidae of America. In Casey, T.L. Mem-
oirs on the Coleoptera. IV. New Era Printing Company, Lancaster, Pa., 192 pp.
Casey, T.L. 1924. Additions to the known Coleoptera of North America. In Casey, T.L.,
Memoirs on the Coleoptera XI. New Era Printing Company, Lancaster, Pa., 347 pp.
Chaudoir, M. de. 1842. Description de quelques genres nouveaux de la famille des carabi-
ques. Bull. Soc. Imp. Nat. Moscou. 15 : 832-857.
Chaudoir, M. de. 1872. Monographic des callidides. Ann. Soc. ent. Belg. 15 : 97-204.
Chaudoir, M. de. 1877. Genres nouveaux et especes inedites de la famille des carabiques.
Bull. Soc. Imp. Nat. Moscou 52 : 188-268.
Chevrolat, L.A.A. 1835. Coleopteres du Mexique, fasc 5. Strasbourg, 50 pp.
Cohn, T.J. 1965. The arid-land katydids of the North American genus Neobarrettia (Orth-
optera: Tettigoniidae): Their systematics and a reconstruction of their history. Misc. Publ.
Mus. Zool. Univ. Mich. 126, 179 pp.
Csiki, E. 1932. Harpalinae VII. In Junk, W., and S. Schenkling (editors). Coleopterorum
Catalogus‘11. Berlin and s’Gravenhage. 1279-1598.
Dejean, P.F.M.A. 1831. Species general des coleopteres de la collection de M. le compte
74
Larson
Dejean. Vol. V. 883 pp. Paris.
Dillon, L.S. 1956. Wisconsin climate and life zones in North America. Science 123 : 167-176.
Emden, F.I. van. 1942. A key to the genera of larval Carabidae. Trans. R. ent. Soc. Lond. 92:
1-99, 100 figs.
Fisher, R.A. 1 936. The use of multiple measurements in taxonomic problems. Ann. Eug.
London 7 : 179-188.
Freitag, R. 1965. A revision of the North American species of the Cicindela maritima group
with a study of hybridization between Cicindela duodecimguttata and oregona. Quaest.
ent. 1(3) : 87-170.
Goulden, C.H. 1952. Methods of statistical analysis. J. Wiley & Sons, Inc., New York. 467
pp.
Habu, A. 1960. Callida species of Japan (Coleoptera, Carabidae). Bull. Nat. Inst. Agr. Sci.(C)
12 : 155-169.
Habu, A. 1967. Fauna Japonica: Carabidae Truncatipennes group (Insecta: Coleoptera).
Tokyo Electrical Engineering College Press Tokyo. 338 pp., 27 plates.
Hatch, M.H. 1950. An account of the life of Orson Bennett Johnson. In Studies honoring
Trevor Kincaid, 11-28.
Hatch, M.H. 1953. The beetles of the Pacific northwest. Univ. Wash. Publ. Biol. 16 : 340
pp.
Horn, G.H. 1881. On the genera of Carabidae, with special reference to the fauna of Boreal
America. Trans. Amer. ent. Soc. 9 : 91-196, 10 plates.
Horn, G.H. 1882. Synopsis of the tribe Lebiini. Trans. Amer. ent. Soc. 10: 126-183.
Jeannel, R. 1942. Coleopteres Carabiques, 2me partie. Faune de France 40: 573-1 173, figs.
214-368. Paris.
Jeannel, R. 1949. Cole'opt'eres carabiques de la Region malgache (troisieme partie). Faune de
l’Empire francais, vol. XI, 767-1146, figs. 365-548.
v
Jedlicka, A. 1963. Monographic der Truncatipennen aus Ostasien. Lebiinae-Odacanthinae-
Brachyninae (Coleoptera, Carabidae). Ent. Abh. Ber. Mus. Tierk. Dresden 28 : 269-579.
King, P.B. 1958. Evolution of modem surface features of western North America, p. 3-60.
In C.L. Hubbs, (ed.), Zoogeography. Amer. Assoc. Adv. Sci. Symp., Washington, D.C.
LeConte, J.L. 1848. A descriptive catalogue of the geodephagous Coleoptera inhabiting the
United States east of the Rocky Mountains. Ann. Lyc. Nat. Hist. New York 4: 173-474.
LeConte, J.L. 1851. Descriptions of new species of Coleoptera from California. Ann. Lyc.
Nat. Hist. New York 5 : 125-184.
Leng, C.W. 1920. Catalogue of the Coleoptera of America, north of Mexico. Mount Vernon,
N.Y., 470 pp.
Lewin, V. 1963. The herpetofauna of southeastern Alberta. Canad. Fid. Nat. 77(4) : 203-
214.
Lindroth, C.H. 1954. Die larve von Lebia chlorocephala Hoffm. (Col. Carabidae). Opusc.
ent. 20 : 10-34.
MacGinitie, H.D. 1958. Climate since the late Cretaceous. In C.L. Hubbs (ed.), Zoogeo-
graphy. Amer. Assoc. Adv. Sci. Symp., Washington, D.C., 61-79.
Madge, R.B. 1967. A revision of the genus Lebia Latreille in America north of Mexico (Col-
Carabidae
75
eoptera, Carabidae). Quaest. ent. 3(3) : 139-242.
Mannerheim, C.G. von. 1837. Memoire sur quelques genres et especes de carabiques. Bull.
Soc. Imp. Nat. Moscou. 10(2) : 3-49.
Martin, P.S. 1958. Pleistocene ecology and biogeography of North America. In C.L. Hubbs
(ed.), Zoogeography. Amer. Assoc. Adv. Sci. Symp., Washington, D.C., 375-420.
Mayr, E. 1 963. Animal species and evolution. The Belknap Press of Harvard University Press,
Cambridge, Mass. 797 pp.
Mayr, E., E.G. Linsley, and R.L. Usinger. 1953. Methods and principles of systematic zool-
ogy. McGraw-Hill Book Co., Inc., New York. 336 pp.
Menetries, M. 1843. Sur un envoi d-insectes de la cote N.O. d’Amerique. Bull. Acad. Imp.
Sci. St. Petersbourg Phys.-Math. 2 : 49-64.
Motschoulsky, V. von. 1850. Die Kafer Russlands, 9 pp. Moscou.
Motschoulsky, V. von. 1859. Coleopteres nouveaux de la Califomie. Bull. Soc. Imp. Nat.
Moscou 32(2) : 122-185.
Say, T. 1823. Descriptions of coleopterous insects collected in the late expedition to the
Rocky Mountains, performed by order of Mr. Calhoun, Secretary of War, under the com-
mand of Major Long. J. Acad. nat. Sci. Philad. 3(1) : 139-216.
Selander, R.B., and P. Vaurie. 1962. A gazetteer to accompany the “Insecta” volumes of the
“Biologia Centrali-Americana”. Amer. Mus. Novit. no. 2099. 70 pp.
Stanley, J. The discriminant function. A mathematical method for the taxonomic separation
of species. MS.
Torre-Bueno, J.R. de la. 1962. A glossary of entomology. Brooklyn Entomological Society,
Brooklyn, N.Y. 336 pp.
76
Larson
Figs. 6-10. 6. Larva of Tecnophilus croceicollis croceicollis Menetries, first instar (Newark,
California). 7. Nasale and mandibles of Plochionus timidus Haldeman. 8. Maxilla of
Plochionus timidus Haldeman. 9. Pronotum of larva of Philophuga viridicollis LeGbnte, third
instar, dorsal aspect. 10. Same of Tecnophilus croceicollis Menetri6s.
Carabidae
77
Figs. 11-14. Head capsule, third instar larva. 1 1 . Phophuga viridicollis LeConte. 1 2. Tecno-
philiis croceicollis croceicollis "Menetries. 13. Tecnophilus croceicollis peigani new subspecies,
dorsal aspect. 14. Same, ventral aspect.
78
Larson
Figs. 1 5-29. 15. Male genitalia of Philophuga viridicollis LeConte, lateral aspect. 1 6. Same of
Philophuga caerulea Casey. 17. Same of Philophuga viridis viridis Dejean. 18. Same of Tecno-
philus pilatei Chaudoir. 19. Same of Tecnophilus croceicollis croceicollis Menetries. 20. Same
of Tecnophilus croceicollis peigani new subspecies. 21. Same of Infemophilus castaneus
Horn, dorsal aspect, parameres included. 23. Left wing of Philophuga caerulea Casey. 24.
Head and pronotum of Tecnophilus croceicollis peigani new subspecies, dorsal aspect. 25.
Same of Tecnophilus croceicollis croceicollis Me'netries (Brownsville, Texas). 26.Abdomen of
Plochionus timidus Haldeman, ventral aspect. 27. Same of Infemophilus castaneus Horn.
28. Front tarsus of Callida decora Fabricius. 29. Same of Philophuga viridicollis LeConte.
Carabidae
79
Figs. 30-42. Pronotum, left half, dorsal aspect. 30 . Infernophilus castaneus Horn (Coleville,
California). 31. Philophuga viridicollis LeConte (Monahans, Texas). 32. Philophuga caerulea
Casey (Nogales, Arizona). 33. Philophuga brachinoides Bates (Nochixtlan, Oaxaca). 34.
Philophuga viridis amoena LeConte (Medicine Hat, Alberta). 35. Philophuga viridis homi
Chaudoir (Stockton, Utah). 36. Philophuga viridis klamathea new subspecies (holotype,
Klamath Fall, Oregon). 37. Philophuga viridis viridis Dejean (San Francisco, California). 38.
Tecnophilus pilatei Chaudoir (Brownsville, Texas). 39-41 . Tecnophilus croceicollis croceicol-
lis Menetries: 39. Brownsville, Texas; 40. Cuddeback Lake, California; 41 . Alviso, California.
42. Tecnophilus croceicollis peigani new subspecies (paratype, Lost River Ranch, Alberta).
80
Larson
Figs. 43-57. Right stylus of female ovipositor, dorsal aspect. 43. Euproctinus trivittatus
LeConte. 44. Onota floridana Horn. 45. Plochionus amandus Newman. 46. Plochionus
timidus Haldeman. 47. Callida punctata LeConte. 48. Callida viridipennes Say. 49. Callida
purpurea Say. 50. Callida decora Fabricius. 51. Philophuga viridicollis LeConte. 52. Philo-
phuga viridis viridis Dejean. 53 JPhilophuga viridis klamathea new subspecies. 54. Philophuga
brachinoides Bates. 55. Tecnophilus pilatei Chaudoir. 56. Tecnophilus croceicollis croceicol-
lis Menetries. 57 . Infemophilus castaneus Horn.
Carabidae
81
Fig. 58. Distribution of Philophuga viridicollis IeConte (★), and Philophuga caerulea
Casey ( • ).
82
Larson
Fig. 59. Distribution of the subspecies of Philophuga viridis Dejean: ■ , v. viridis Dejean
▲ , v. klamathea new subspecies;^v. horni Chaudoir, v. amoena LeConte.
Carabidae
83
Fig. 60. Distribution of Philophuga brachinoides Bates. Fig. 61 . Distribution of Infernophilus
castaneus Horn. A represents a state locality only. Fig. 62. Distribution of Tecnophilus pilatei
Chaudoir.
84
Larson
Fig. 63. Distribution of the subspecies of Tecnophilus croceicollis Menetries: • croceicollis
croceicollis Menetries; A croceicollis peigani new subspecies.
Quaest
lones
entomologicae
iVTUS.
COfVfp
FEB
'JNh
fc /
A periodical record of entomological investigations,
published at the Department of Entomology, Uni-
versity of Alberta, Edmonton, Canada.
VOLUME V
NUMBER 2
APRIL 1969
QUAESTIONES ENTOMOLOGICAE
A periodical record of entomological investigations published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.
Volume 5 Number 2 April, 1969
CONTENTS
Book Review 85
Freitag — A Revision of the Species of the Genus Evarthrus LeConte
(Coleoptera:Carabidae) 89
Errata 212
Book Review
COBBEN, R. H. 1968. Evolutionary trends in Heteroptera. Part I. Eggs, architecture of the
shell, gross embryology and eclosion. 475 pp, 316 Figs. Centre for Agricultural Publishing
and Documentation, Wageningen, Holland. $16.50 U. S.
This is the first of a series of three volumes by Cobben on the phylogeny of the supra-
generic taxa of the Heteroptera, and there will be many biologists awaiting the next two
volumes.
Cobben approaches the phylogeny of the Heteroptera on a wide front and this can best
be summed up by a quotation from the page preceding the preface, “In its best practice,
taxonomy is a wonderfully promising area of synthesis for all biological knowledge.”
In this volume Cobben has examined the oviposition stance of the female, the detailed
structure of the egg and the embryogenesis of as many species of Heteroptera as there was
material available.
As might be expected in such a detailed study as this, many new characters of taxonomic
value came to light. These, along with previously used characters have all been utilized in
synthesizing a phylogenetic scheme which forms the basis for the next two volumes.
Cobben’s work on the chorion of the eggs is in such detail that little of previous descrip-
tions can be homologized to the structures he shows. Of particular interest are the “airo-
static inner layer” of the Geocorisae chorion and the “porous inner layer” of that of the
eggs of Saldidae. These structures along with other porous structures act as a plastron.
The importance that Cobben attaches to the Leptopodoidea is obvious from the arrange-
ment of the contents of the volume. The first chapter is devoted entirely to the eggs of this
family. The other families are dealt with in the second chapter. For each family the
literature, material used, egg shape, oviposition behaviour, chorion, embryogenesis and eclo-
sion are discussed in turn, enabling detailed information to be extracted rapidly. Chapter
III is devoted to evaluating the egg characters. Chapter IV is a preliminary discussion of the
86
phylogeny of the suborder with a short review of the investigations that will be reported in
the other volumes. There is an excellent summary. The book will no doubt become a
standard reference text on Heteroptera.
Although the book has been well produced it is surprising in a work of this nature that
the abbreviations of morphological terms are not given in the text when the structure is
referred to in a figure. Another disturbing feature is that the scales on the figures have not
been given any absolute value, although they are probably in millimeters. Proof reading is
good apart from some minor spelling mistakes and a series of transpositions in the caption
of Figure 273. The line figures throughout the volume are of high standard and are explicit,
as are the transmission and scanning electronmicrographs.
If the next two volumes of this series maintain the high quality of the first, then Cobben
will have made a significant contribution to the understanding of Heteropteran biology and
phylogeny. His approach is one that future taxonomists might well emulate.
D. A. Craig
Department of Entomology
University of Alberta
-
Evarthrus sodalis sodalis LeConte
Lexington, Kentucky
Photograph by J. Scott
A REVISION OF THE SPECIES OF THE GENUS EVARTHRUS LECONTE
(COLEOPTERA :CARABIDAE)
RICHARD FREITAG
Department of Biology
Quaestiones entomologicae
5 : 89-212 1969
Lakehead University
Port Arthur, Ontario
Within the genus Evarthrus, three subgenera, 43 species, and five subspecies are recog-
nized. The genus Evarthrus is described, and evidence is presented which removes Evarthrus
as a unit from the Pterostichus complex to a position near the genus Molops in the tribe
Pterostichini. A key to the species and subspecies is given. Each subgenus, species group, and
species is described and synonymies are listed. The distribution of each species is presented
by locality records and distribution maps. Structures which are used in identification are
illustrated.
The subgenus Fortax comprises six species of which one, iuvenis, is described as new. One
genus group name and six species names are reduced to synonymy.
The subgenus Cyclotrachelus includes 12 species, of which five, fucatus, macrovulum,
texensis, parafaber, and levifaber are described as new. Six species names are listed as
synonyms.
The subgenus Evarthrus includes 25 species of which seven are new. The species sodalis
LeConte and torvus LeConte are polytypic. Five genus group names and 26 species names
are relegated to synonymy.
A phytogeny is postulated for the subgenera, species groups and species. The geographical
distribution of the genus is discussed.
The endemic flightless Pterostichini of eastern North America are arrayed in a series of
supraspecific taxa each of which is more or less easily defined; however, the relationships of
these groups are at best uncertain. The most diverse of these groups is the genus Evarthrus,
a complex of species included by some (Csiki, 1930; Lindroth, 1966) in the genus
Pterostichus, and by others (Casey, 1918) treated as a group of related genera. (Ball, 1967
drew attention to this problem). I decided to attempt to solve these problems of relation-
ships and classification, but learned in the initial stages of the study that I would first have
to undertake a revision of the species. The results of this investigation are presented in this
paper.
The revision is based on a study of adult specimens. The species were defined on the basis
of evaluation of morphological and geographical evidence. Names were applied on the basis
of study of the relevant type material. Sixteen species names were found to be junior
synonyms, and 13 previously undescribed species were discovered and named.
In addition to the formal taxonomic treatment of this group, I have presented my views
on the phylogeny and geographical distribution of the extant species.
90
Freitag
MATERIALS AND METHODS
Materials
The material examined consisted of 7,600 adult specimens, which included the type
specimens of Casey and LeConte. I have also briefly examined external structures and male
genitalia of species of an additional 35 Nearctic and Palearctic subgenera of Pterostichus
Bonelli, and of Abaris Dejean, Pseudabarys Chaudoir, Oxycrepis Reiche, Cratocerus Dejean,
Catapiesis Brulle, Abax Bonelli, Molops Bonelli, Percus Bonelli, Lesticus Dejean, Piesmus
LeConte, Stomis Clairville and My as Dejean.
Names of individuals and institutions from which material was borrowed are abbreviated
in the text as follows: AMNH — American Museum of Natural History; ANSP — Academy of
Natural Sciences; AU - Auburn University; BM - British Museum (Natural History); CAS -
California Academy of Sciences; CM — Carnegie Museum; CNC — Canadian National
Collection; CNHM — Chicago Natural History Museum; CU — Cornell University; DL —
David Larson; DRW - D. R. Whitehead; FDPI - Florida Division of Plant Industry; GEB —
G. E. Ball; INHS — Illinois Natural History Survey; ISU — Iowa State University; KLE —
Kansas State University; MCZ — Museum of Comparative Zoology; MHNP — Museum
d’Histoire Naturelle, Paris; MSU - Montana State University; NCSU - North Carolina State
University; RCG - R. C. Graves; RF — R. Freitag; RTB - R. T. Bell; RU — Rutgers
University; TAM - Texas A & M University; TCB - T. C. Barr; TE - j Erwin: TH - T.
Hlavac; UA — University of Arkansas; UASM — University of Alberta Strickland Museum;
UK — University of Kansas; UL — University of Louisville; UMMZ — University of Michigan
Museum of Zoology; UP — Purdue University; USNM — United States National Museum; UW
— University of Wisconsin; VMK — V. M. Kirk.
Methods
General methods
By making comparisons among their characteristics, specimens were sorted into demes,
subspecies, species, and species groups according to degree of similarity and difference. The
characteristics used were arbitrarily weighed, and the same characteristics were given
different weights in different situations. The relationships revealed by these comparisons
were interpreted and the evolution of the species and species groups was then inferred.
Characters of adults
Some structures which are used in the identification of the species of Evarthrus are
discussed below to facilitate their use in the text.
The lines of microsculpture of the integument of the dorsal surface are almost effaced,
disoriented, and do not form meshes in specimens of some species. In specimens of other
species the lines are close together and sinuate, but they do not form meshes. More
frequently meshes are formed and are amorphic or isodiametric. The interspaces are flat or
raised and bead-like. The lustre of the integument correlated with the microsculpture is as
follows: shiny in the absence of meshes; dull with isodiametric meshes and flat interspaces;
matte or velvet with isodiametric meshes and bead-like interspaces; iridescent with dense
sinuate parallel lines. Females are always duller than males of the same species.
Revision of Evarthrus
91
The frontal grooves of the head are usually of a particular shape, and are useful in
recognizing specimens of a number of species. The grooves are straight, or crescent-shaped
with the convexity directed medially or laterally (figs. 70-7 1 ).
The number of setae on the penultimate article of the labial palpus is useful in delimiting
species. Two dorsal ‘‘primary’’ setae are always present near the halfway point of the article.
Three “secondary” setae are also present. One is apical, and arises from the ventral side of
the article. Another is near the apical end of the article. It is dorsolateral and directed dorso-
laterally. In a complementary position is the third seta. It is also near the apical end of the
article on the dorsomedial side and is directed dorsomedially. The truly apical seta occurs
more frequently than the other “secondary” setae but it is not always present. Other setae
occasionally occur here and there near the “primary” setae (figs. 66-69). Horn (1881) noted
that the number of setae on the labial palpus are not constant in Evarthrus and suggested
that two groups may be recognized: a group with bisetose labial palpi and a group with
plurisetose labial palpi.
The mandibles of the type species, sigillatus Say, are illustrated (fig. 72).
Details of form and structure of the pronotum are useful in recognizing subgenera and
species. The general outline of the pronotum ranges in form from rectangular to cordiform
(figs. 1-62). Another useful structure is the shape of the basal lateral fovea, which is puncti-
form (figs. 1-7), monostriate or bistriate (figs. 8-20, 21-62). The position of the basal lateral
seta is on (figs. 8-20), or beside the lateral bead (figs. 1-7 and 21-62). The lateral bead in
specimens of a few species is broad posteriorly, but in most species it is narrow posteriorly
as in fig. 25. The prostemal process which projects posteriorly between the front coxae has
a longitudinal medial groove. This groove is deep or shallow. The apex of the prosternal
process is or is not marginate.
At least four setae are present on the anterior face of the middle femur and always in the
same positions. A proximal pair of setae are located near the ventral side of the femur, and a
distal pair near the dorsal side. In specimens of some species additional setae usually occur
near either pair. The total number of setae ranges from four to eleven in the genus and
seems to be a good character for grouping species (figs. 74-76). Setae are absent from the
lateroventral margin of the claw bearing tarsal articles in specimens of four species of the
subgenus For tax.
I have used the term “last abdominal sternum” in the text. This is morphological sternum
VII, which is the apparent sternum VI in beetles (sternum 1 has disappeared).
The male genitalia are very important structures in defining species and species groups. In
fact, it would be exceedingly difficult to recognize and classify the species of Evarthrus
without reference to these structures. Two parameres and a median lobe which contains an
internal sac constitute the external male genitalia. The median lobe is a tube with a central
bend. The portion of the lobe posterior to the bend is referred to here as the apical half, and
that anterior to the bend the basal half. The lobe is always bent dorsoventrally with the
convexity directed dorsally. In specimens of some species the apical half of the median lobe
is bent laterally. The posterior extremity of the median lobe is flattened and heavily
sclerotized, and is called here the apical blade. The posterior edge of the apical blade is the
apex. Within the median lobe is a membranous sac known as the internal sac, which is
92
Freitag
everted during copulation. In studying this organ the following technique was used: the
beetle was relaxed in boiling water; then by inserting a pair of fine forceps into the end of
the abdomen the genitalia were grasped and pulled out; these structures were cleared in a
hot 10% solution of potassium hydroxide for about 10 minutes and then washed in water;
the internal sac was everted by gently pulling the sac through its open end with a pair of
fine forceps. In specimens of some species the internal sac bears serrulate fields and an
apical sclerite (the sclerite is apical when the sac is everted). The shape of the apical sclerite
is commonly used for grouping species as well as separating closely related species. The
shape of the whole everted internal sac is probably of taxonomic value but I have not used it
here. Joined to the left and right sides of the basal portion of the median lobe near the
anterior end are the parameres. The left paramere is broad and somewhat disc shaped in all
of the known species of the genus. The right paramere varies in form among the species of
the genus but does not vary interspecifically.
The sclerites of the female ovipositor exhibit slight variation. With the exception of the
stylus they do not provide taxonomically useful features. Fig. 73 is a drawing of the
ovipositor and bursa of sigillatus. The bursa copula trix is short, and the anterior end is a flat,
lightly sclerotized plate, which has a marked anteriorly-directed central mound. A small
dark sclerite rests on the tip of the mound and is joined to the base of the spermathecal
duct. The common oviduct enters the bursa beside the sclerite. A long accessory gland is
joined to the spermathecal duct. The spermatheca is a simple sausage-shaped sac. The
pygidial reservoir is rather large and it has a short thick duct which appears to open
externally near the posterior end of the gonangulum. The pygidial gland duct is short and
narrow. The stylus is typical of the genus. Slight variation in the form of the stylus occurs
throughout the genus and is referred to in the text.
Measurements
The range of body size for each species was determined. A calibrated eyepiece in a Wild
M5 stereoscopic binocular microscope was used. The body length is indicated by the sum of
three measurements: length of head — distance from the base of the mandible to the hind
margin of the eye; length of pronotum — distance between the anterior margin of the
pronotum to the margin behind the basal angle; length of elytra — distance from the apex of.
the scutellum to the apical tip of an elytron. The widths of the head, pronotum, and
abdomen are defined as follows: head — maximum distance behind the eyes dorsally;
pronotum — maximum transverse distance; abdomen — maximum transverse distance across
both elytra.
Illustrations and maps
The drawings were made with the aid of a Wild drawing tube, on the M5 stereo micro-
scope.
Distribution maps are given for all species. Most maps comprise the distributions of
species of a single species group.
Revision of Evarthrus
93
Criteria for species and subspecies
Two forms with overlapping ranges are regarded as distinct species if they do not inter-
grade in at least one morphological character. If a clinal series of intermediate populations is
intercalated between two morphologically distinct populations that are widely allopatric the
entire complex is treated as a single variable species, but subspecific names are not assigned.
Subspecies are recognized only in cases of steep clinal variation in at least one characteristic.
BIOLOGY
Little is known about the biology of this genus. Probably the members are omnivorous, as
are most Carabidae. I have found spores of fungi in the gut of E. faber Germar, and ant
remains in the gut of E. sodalis colossus LeConte.
All of the species are flightless: not only are the hind wings of all individuals atrophied,
but the metathorax is reduced, and the elytra are fused along the suture. It is not surprizing,
therefore, that geographical variation is marked, that most of the species have restricted
ranges, and that closely related species are often allopatric — facts which indicate restricted
powers of dispersal.
Members of the genus inhabit deciduous forests or open country. Those species which
occur in open places are northern and western in distribution. Conversely, the ranges of the
more numerous forest species are generally southern and east of the Mississippi River.
TAXONOMY
The Genus Evarthrus LeConte
Characteristics
Adults. — small to large Pterostichini (see Ball, 1966 for characterization of tribe); color
of body black, legs usually black sometimes red; penultimate article of labial palpus pluri-
setose (usually) or bisetose; pronotum rectangular to cordate, basal lateral fovea of prono-
tum bistriate, monostriate, or a single puncture, always distinctly impressed; basal lateral
seta of pronotum on lateral bead or beside it; elytron with seventh interval usually raised at
base, 1-5 punctures on medial side of third interval; hind wings absent; metepisternum short,
with lateral margin equal in length to anterior margin; article five of tarsus usually with a
row of setae on each ventrolateral margin; venter impunctate, usually slightly rugose;
females with two setae on last sternum of abdomen; eversion of internal sac of median lobe
of male genitalia usually to right, less often dorsoapical, and rarely to left. Larva —
Pterostichini; antenna with five articles; urogomphi short, terete, curved toward each other
(Van Emden, 1942, and Boving and Craighead, 1930).
Type species. - Evarthrus sigillatus Say, 1823a (designated by Lindroth, 1966:473).
94
Freitag
The species of Pterostichus Bonelli resemble species of Evarthrus, but in the former group
the lateral areas of the ventral surface of the body are usually punctate, the eversion of the
internal sac is to the left or dorsal, the females have usually four to eight setae on the last
abdominal sternum, exceptionally two in some individuals and the larvae have four antennal
articles and long multinodose urogomphi.
Some species of Pseudabarys and Abaris vaguely resemble species of Evarthrus in having a
plurisetose penultimate labial palpus and a single puncture in the third interval of an
elytron. Their general habitus is different, however. Species of Abaris have pectinate claws,
and the internal sac seems to be telescopic rather than of the eversion type.
Members of the genus Evarthrus are like those of Molops. They have the following
characteristics in common: Adult — similar body shape particularly the pronotum; ventral
side of body not punctate; elytron usually with seventh interval raised at base; and setae
usually present on each lateroventral side of the last tarsal article. Larvae — antenna with
five articles. Specimens of Molops differ by having a ninth elytral interval which is lateral to
the umbilicate series, setae on the dorsal side of the last tarsal article, and four setae on the
last abdominal sternum of the females. The larvae of Molops and Evarthrus differ in
characteristics of the urogomphi.
Schuler (1962, 1963a, 1963b) has studied the taxonomic importance of the spermatheca
of female carabids. He points out that the spermatheca of Molops is a simple sac while that
of Pterostichus is not. The spermatheca of Evarthrus is also a simple sausage-shaped sac, like
that of Molops. This similarity may not be in itself important, but it adds to the characters
that Evarthrus and Molops share.
Basford et alii (1968) used immunological techniques to investigate the classification of
the Adephaga. Among other species of carabids, they studied Evarthrus sodalis LeConte and
Pterostichus chalcites. They found the samples of these two species to be markedly different
from one another, and this in itself could be accepted as additional evidence to support the
ranking of Evarthrus as generically distinct from Pterostichus. However, this evidence is of
doubtful value because the other results obtained are at such variance with the generally
accepted classification. Indeed, these authors write that “(6) the distinct position of the
genus Harpalus and the failure of the immunological results to cluster other Harpalinae with
them suggested that, within the Carabidae, large amounts of random molecular variation
exist (1968:405).”
I believe that the treatment of Evarthrus, Pterostichus, and Molops as separate genera in
the tribe Pterostichini is justified. Simpson (1961) points out that criteria derived from
relative divergence apply to the ranking of taxa. and he suggests several criteria of which one
is as follows: in a group of related taxa it is desirable that differences between most similar
taxa should be approximately equal. In addition to this the general feeling among taxono-
mists is that taxa of the same rank should have the same amount of diversity.
In treating Evarthrus, Pterostichus , and Molops as separate genera both of the above
criteria are followed. The differences among the three genera are approximately the same in
numbers of weighted characteristics, which are widespread in each genus. Each of the taxa
contain many species, although Pterostichus , as regarded here, is the most diverse. The genus
Evarthrus is a polythetic group, but nevertheless such groups are acceptable in taxonomic
practice.
Revision of Evarthrus
95
Subgenera and Species Groups
On the basis of similarities and differences of external structures and male genitalia the
species are grouped into three subgenera. The species of each subgenus are arranged in
species groups. The subgenus Fortax Motschulsky includes six species which constitute two
species groups. The subgenus Cyclotrachelus Chaudoir contains 12 species which are
arranged in three species groups. Twenty-five species are included in the subgenus Evarthrus
and are placed in ten species groups. The names of most of the species groups are based on
the name of the first described species contained in each. Two species groups have been
given the names of the most well known species included in each: the spoliatus group and
the ovulum group. Th egigas group is so named because it includes E. gigas Casey, which was
designated as the type species of Megasteropus Casey.
Key to the species and subspecies of the Genus Evarthrus LeConte
1 Plica of elytron present 2
Plica of elytron absent E. gravesi new species, p. 167
2( 1) Basal setae of pronotum in lateral bead (figs. 8-20); basal foveae of pronotum
monostriate 3
Basal setae of pronotum beside lateral bead (figs. 1-7, 21-*61); basal foveae of
pronotum punctiform OR bistriate 15
Gula with anterior end flanked by raised knobs (fig. 63); body longer than 17.7 mm
E. unicolor Say, p. 1 10
Knobs absent; body shorter than 17.7 mm 4
Prosternal process with longitudinal groove deep and sharply defined 5
Prosternal process with longitudinal groove shallow and not sharply defined ... 9
Penultimate article of labial palpus with two medial and two apical setae; pronotum
circular (fig. 20); front tarsi of males with ventral rows of cup-like scales
E. faber Germar, p. 125
Penultimate article of labial palpus with two medial setae only; pronotum cordiform
OR sides not produced (figs. 15-16, 18-19); males with typical scales on front tarsi
6
6( 5) Pronotum with basal angles sharp and produced (figs. 15-16); microsculpture open
and not dense 7
— Pronotum with basal angles broadly rounded and not produced (figs. 18-19),
micro sculpture open but dense 8
7( 6) Frontal grooves crescent -shaped, widely separated, and oblique (fig. 71); range,
Florida and Georgia E. ovulum Chaudoir, p. 1 18
Frontal grooves straight, closer together, and more parallel (fig. 70); range, Mobile,
Alabama area E. alabamensis Casey, p. 117
8( 6) Basal foveae of pronotum with almost effaced long and shallow anterior extensions
that together form a lyre-shaped figure; pronotum oval shaped because of gradual
constriction of anterior half (fig. 18); range, Mobile, Alabama area
E. parafaber new species, p. 122
3( 2)
4( 3)
5( 4)
*In a few specimens one seta on one side in bead.
96
Freitag
Basal foveae of pronotum without long anterior extensions; pronotum cordiform
(fig. 1 9); range, Georgia, South Carolina, and North Carolina
E. levifaber new species, p. 123
9( 4) Pronotum cordiform (figs. 14, 17) 10
Pronotum more oval (figs. 9-13) 12
10( 9) Range, Georgia, Mississippi, and Tennessee; pronotum with basal sinuations elongate
(Fig. 1 4); males with obsolete punctures in elytral striae
E. vinctus LeConte, p. 1 1 5
Range, southern Alabama, Mississippi and Texas; pronotum with shorter basal
sinuations; (fig. 17) males with large punctures in elytral striae 11
11(10) Range, northeastern Texas; male with apex of median lobe broader (fig. 95 e-g)
E. texensis new species, p. 121
Range, coastal Alabama and Mississippi; male with apex of median lobe narrower
(fig. 95 a-c) E. macrovulum new species, p. 1 19
12( 9) Range, east of the Appalachian Mountains 13
Range, south and west of the Appalachian Mountains 14
13(12) Range, eastern South Carolina north to Maryland; male with apex of median lobe
evenly rounded E. spoliatus Newman, p. 1 13
Range, western South Carolina southward; male with apex of median lobe truncate
E. brevoorti LeConte, p. 1 14
14(12) Range, northern Georgia, northern Alabama, Tennessee, Kentucky, Ohio, West
Virginia, western Pennsylvania; apex of median lobe of male evenly rounded;
pronotum of male glossy, microsculpture varying from open and sparse to obsolete
E. fucatus new species, p. 1 1 1
Range, northern Georgia, northern Alabama, south to Florida, southern Alabama
and southern Mississippi; apex of median lobe of male truncate; pronotum of male
semi-glossy, microsculpture open but dense E. brevoorti LeConte, p. 114
15( 2) Basal foveae of pronotum punctiform (figs. 1-7) 16
Basal foveae of pronotum bistriate (figs. 21-61 ) 21
16(15) Apex of prosternal process marginate E. hernandensis Van Dyke, p. 101
Prosternal process not marginate 17
17(16) Pronotum with incomplete marginal groove between lateral setae (fig. 2)
E. morio Dejean, p. 102
Pronotum with complete marginal groove between lateral setae 18
18(17) Pronotum with basal setae near basal angles (figs. 3-4)
E. laevipennis LeConte, p. 103
Pronotum with basal setae in front of basal angles (figs. 5-7) 19
19(18) Pronotum with anterior transverse impression complete (fig. 5)
E. approximates LeConte p. 106
Transverse impression incomplete (figs. 6-7) 20
20(19) East of the Appalachian Mountains, in North Carolina, and Virginia
E. iuvenis new species, p. 107
Revision of Evarthrus
97
West and South of the Appalachian Mountains, in Indiana, Illinois, Ohio, Michigan,
Tennessee, Mississippi, Alabama, Georgia E. obsoletus Say, p. 108
21(15) Elytron with 3-5 setae in third interval 22
— Elytron with one seta in third interval, occasionally one or two setae on one elytron
and two setae on the other 26
22(21) Pronotum quadrate with smooth lateral margins (fig. 37); range, east of the
Mississippi River E. hypherpiformis new species, p. 145
— Pronotum more cordate OR quadrate with lateral crenulations (figs. 54, 57-58);
range, west of the Mississippi River 23
23(22) Elytra with striae almost impunctate E. substriatus LeConte, p. 155
— Striae distinct and deeply punctate 24
24(23) Pronotum 6-8 mm wide, quadrate, with lateral crenulations, particularly in basal
sinuation (fig. 58) E. gravidus Haldeman, p. 163
Pronotum less than 6 mm wide, more cordiform without lateral crenulations (figs.
53,57) 25
25(24) Elytra dull; range, Oklahoma, Texas E. torvus deceptus Casey, p. 1 60
— Elytra glossy; range, Iowa, Minnesota, South Dakota
E. iowensis new species, p. 154
26(21) Pronotum with anterior transverse impression obsolete medially 27
Pronotum with anterior transverse impression complete and clearly impressed OR
complete with short interruptions 36
27(26) Middle femur with four setae on anterior face, occasionally four setae on one femur
and five on other 28
— Five or more setae on anterior face of both middle femora 29
28(27) Median lobe of male strongly arcuate and apical blade short with edges only slightly
bent (fig. 100); body length 1 1.4 — 15.4 mm; legs always black; pronotum (fig. 22);
in Arkansas elytral intervals with micropunctures indistinct; range, Arkansas,
Oklahoma E. whitcombi new species, p. 129
— Median lobe of male moderately arcuate and apical blade long with edges strongly
bent (fig. 99); total length 9.02 — 12.3 mm; legs black OR ferrugineous; pronotum
(fig. 21); in Arkansas elytral intervals with distinct micropunctures; range, Arkansas,
Iowa, Kansas, Missouri, Nebraska, Oklahoma, Pennsylvania, South Dakota
E. incisus LeConte, p. 1 27
29(27) Elytron with striae almost effaced; first three anterior umbilicate punctures with
slight mounds between them E. substriatus LeConte, p. 1 56
Elytron with striae distinct, higher ridges present between first three umbilicate
punctures 30
30(29) Body length 11.2 — 13.9 mm; pronotum (fig. 53); range, Iowa, Minnesota, South
Dakota E. iowensis new species, p. 154
— Body longer than 13.9 mm 31
31(30) Pronotum with longer constriction before basal angles which are about 90° or less
(figs. 45—47, 52); range, mainly west of the Mississippi River 32
98
Freitag
— Pronotum with basal angles shorter and greater than 90° (figs. 38-44, 48-51); range,
mainly east of the Mississippi River AND eastern Iowa and Arkansas 33
32(31) Pronotum with basal angles laterally prominent; basal foveae more V-shaped than
U-shaped, relatively short and inner edge anteriorly not markedly deflected laterally
(figs. 45-47); range, mainly west and southwest of the Missouri River AND western
Iowa E. sodalis colossus LeConte, p. 146
Pronotum with basal angles less produced laterally (fig. 52); basal foveae more
U-shaped than V-shaped, relatively longer and anterior end of inner edge deflected
laterally ; range, Illinois, Iowa, Missouri, South Dakota, Wisconsin
E. alternans Casey, p. 1 53
33(31) Range, Arkansas; pronotum with basal angle obtuse (fig. 49)
E. parasodalis new species, p. 1 50
Range, north and east of Arkansas; specimens near Arkansas have pronotum with
more distinct sinuation and basal angles more acute (figs. 38-44, 50-51 ) 34
34(33) Elytra of male with microsculpture stretched transversely; pronotum (fig. 48); range
Alabama, Tennessee E. sodalis lodingi Van Dyke, p. 146
Elytra of males with microsculpture isodiametric 35
35(34) Pronotum with basal angles round, and more obtuse in southern Pennsylvania (figs.
38-44); range, New York west to Iowa, and Minnesota south to northern Mississippi
E. sodalis sodalis LeConte, p. 146
Pronotum with basal angles sharp in south Pennsylvania and more obtuse in Virginia
(figs. 50-51); range, southern Pennsylvania, Virginia, Maryland, southern New Jersey
E. furtivus LeConte, p. 152
36(26) Apex of prosternal process with apical setae 37
Prostemal process without setae 40
37(36) Pronotum with sides slightly sinuate near base, basal angles slightly obtuse and
prominent (figs. 35-36) 38
Pronotal sinuation obsolete, basal angles very obtuse, broadly rounded, not prom-
inent (figs. 33-34) 39
38(37) Pronotum quadrate, margin slightly expanded near base (fig. 36); elytra dull,
particularly in females; elytron of female with stria 8 and marginal groove widely
separated; range, southern Arkansas, northern Louisiana, western Mississippi and
northeastern Texas E. nonnitens LeConte, p. 144
Pronotum with sides more acutely sinuate near base, margin more broadly expanded
near base (fig. 35); elytra slightly glossy; elytron of female with stria 8 and marginal
groove approximate; range, southeastern Texas
E. engelmanni LeConte, p. 142
39(37) Pronotum at widest point 4-5 mm; body length 10.3 — 15.9 mm; legs black or red;
pronotum more rectangular than circular (fig. 33); males almost always with flat
elytral intervals E. seximpressus LeConte, p. 139
Pronotum at widest point 5.5 — 6.5 mm; body length 14.6 — 18.7 mm; legs black
only; pronotum more circular than rectangular (fig. 34); males almost always with
convex elytral intervals E. alabamae Van Dyke, p. 141
Revision of Evarthrus
99
40(36) Middle femur with four setae on anterior face AND pronotum typically cordiform,
strongly constricted posteriorly (fig. 21 ) E. incisus LeConte, p. 1 27
Middle femur with more than four setae on anterior face OR pronotum not as in
fig. 21 41
41(40) Range, east of the Mississippi River 42
Range, west of the Mississippi River 47
42(41) Pronotum moderately sinuate near base (fig. 44); range, Tishomingo County, Missis-
sippi E. sodalis sodalis LeConte, p. 146
Pronotum more quadrate, less sinuate near base (figs. 23-32) 43
43(42) Range, Florida east of the Suwannee River and coastal Georgia 44
Range, other than above 45
44(43) Pronotum with the width of deplanate area between lateral ridge and disc nearly
even throughout (fig. 23); body length 14.8 — 17.6 mm: elytron with two setae,
in seventh stria near plica E. blatchleyi Casey, p. 1 3 1
Pronotum with deplanate area broad near base (fig. 24); body length 13 — 15 mm;
elytron with one seta, rarely two, in seventh stria near plica
E. floridensis new species, p. 1 32
45(43) Range, mainly east of the Appalachian Mountains and southeastern Alabama,
Florida west of the Suwannee River, eastern Tennessee*, Pennsylvania west to
Pittsburg; Pennsylvania specimens with laterally arcuate and glossy elytra; pronotum
(figs. 25-28); male genitalia (fig. 103) E. sigillatus Say, p. 133
— Range, west of the Appalachian Mountains, Pennsylvania specimens with parallel
and dull elytra; pronotum (figs. 29—32) 46
46(45) Pronotum bell-shaped (fig. 29); range, coastal Alabama and Mississippi
E. sinus new species, p. 1 36
— Pronotum rectangular (figs. 30-32); range, north of E. sinus
E. convivus LeConte, p. 137
47(41) Body length 9.5 - 14.5 mm 48
— Body longer than 14.5 mm 52
48(47) Elytra with striae almost effaced; range, Mexico, Texas, New Mexico
E. substriatus LeConte, p. 156
— Elytra with distinct impressed striae 49
49(48) Umbilicate series with first three anterior punctures small and separated from one
another by low raised areas; pronotum strongly constricted at base (fig. 55) .... 50
Umbilicate series with first three anterior punctures of normal size separate from
one another by normal ridges; pronotum less strongly constricted at base (figs.
53, 57) 51
50(49) Plica large; last abdominal segment with prominent dorsal knob that fits onto plica,
especially distinct in females (fig. 77); elytra markedly sinuate posteriorly (fig. 78)
E. substriatus LeConte, p. 156
Plica small; knob obsolete (fig. 79); elytra not markedly sinuate (fig. 80)
E. constrictus Say, p. 158
*The geographic ranges of convivus and sigillatus are approximate in eastern Tennessee. For
certain identification of specimens occurring in this region, examine the male genitalia.
100
Freitag
51(49) Elytra dull; range, Oklahoma, Texas E. torvus deceptus Casey, p. 160
Elytra glossy; range, Iowa, Minnesota, South Dakota
E. iowensis new species, p. 1 54
52(47) Pronotum slightly or moderately constricted near base, sides not prominent (figs.
49,56-58) 53
Pronotum more strongly constricted near base, sides convex (figs. 45—47, 54, 59, 61)
56
53(52) Pronotum with posterior angles not prominent (fig. 49); range, Arkansas
E. parasodalis new species, p. 1 50
Pronotum with posterior angles more prominent 54
54(53) Pronotum quadrate, lateral margin crenulate particularly in basal sinuation, basal
foveae not complete (fig. 58) E. gravidus Haldeman, p. 163
Pronotum less quadrate and more constricted near base, lateral margin smooth or
with indistinct crenulations, basal foveae complete (figs. 56—57) 55
55(54) Elytra dull; pronotum smooth (fig. 57) E. torvus deceptus Casey, p. 160
Elytra glossy; pronotum rugose (fig. 56) E. torvus torvus LeConte, p. 160
56(52) Elytra with striae very shallow, almost effaced, impunctate, sometimes represented
by a series of extremely shallow dashes rather than continuous lines, intervals always
flat; pronotum (figs. 54, 60) 57
Elytra with striae deeper, punctate, and sometimes represented by a row of punc-
tures or distinctly impressed dashes; intervals flat or convex; pronotum (figs. 45—47,
59,61) 58
57(56) Very large species; body length 19.4 — 23.8 mm; range, Texas
E. gigas Casey, p. 1 65
Smaller; body length 9.5 — 14.5 mm; range Mexico, Texas, New Mexico
E. substriatus LeConte, p. 156
58(56) Elytron with scutellar stria long and always separated from stria 2; first complete
stria (stria 2) begins at basal seta (fig. 65); elytra of females with intervals completely
flat; stria 7 with four to five setae near apex; pronotum (fig. 61)
E. heros Say, p. 166
Elytron with scutellar stria always joined to stria 2 and base of stria 2 indicated near
basal seta or absent (fig. 64), elytra of females with raised intervals and striae more
impressed, stria 7 with two to three, rarely four, setae near apex; pronotum (figs.
45-47,59) 59
59(58) Elytra of males with transversely stretched microsculpture, pronotum with base of
basal foveae straight (fig. 59) E. sallei LeConte, p. 165
Elytra of males with isodiametric microsculpture, pronotum with the base of the
basal fovea curved (figs. 45—47) E. sodalis colossus LeConte, p. 146
Revision of Evarthrus
101
The Subgenus For tax Motschulsky
Fortax Motschulsky, 1865:246. — Ball, 1960:129. TYPE SPECIES — Evarthrus morio
Dejean, 1 828 (here designated).
Ferestria Leng, 1915:576. TYPE SPECIES — Evarthrus laevipennis LeConte. 1848 (desig-
nated by Leng, 1915:576).
Characteristics. — The following combination of characteristics is diagnostic for the
subgenus Fortax: species of small size (body length 7.1 — 12.8 mm); penultimate article of
labial palpus bisetose (usually) to quadrisetose; pronotum with sides strongly constricted
posteriorly, posterior lateral foveae each completely punctiform or punctiform posteriorly
with short anterior extension, posterior lateral setae situated beside lateral bead (figs. 1—7);
middle femur with four setae on anterior face (fig. 74); last tarsal article with or without
setae on latero ventral margins; eversion of internal sac of median lobe of male genitalia
dorsoapically or ventrally on left side of median lobe.
The absence of setae on the ventral side of the last tarsal article and the left ventral
eversion of the internal sac are characteristics found in the subgenus Fortax but not in the
subgenera Cyclotrachelus and Evarthrus.
The two species groups in Fortax are the morio group and the obsoletus group.
The morio Group
Characteristics. — Pronotum with basal lateral foveae punctiform posteriorly, briefly and
shallowly extended anteriorly; basal seta situated near basal angle; claw-bearing article of
tarsus without setae on lateral ventral margins.
This group includes the species hernandensis Van Dyke, morio Dejean and laevipennis
LeConte. The members of this group are found on the Gulf Coastal Plain and on the
Piedmont in the southeastern United States.
Evarthrus hernandensis Van Dyke, 1943
Figures 1,66, 74, 81, 125
Evarthrus (Ferestria) hernandensis Van Dyke, 1943:26. HOLOTYPE, male, labelled as
follows: “Brooksville Fla 1-20. 30/40; Van Dyke Collection; HOLOTYPE No. 5308
Evarthrus hernandensis Van Dyke”. CAS. ALLOTYPE, labelled as follows: “Brooksville
Fla 1—20. 30/40; Van Dyke Collection; Allotype No. 5309 Evarthrus hernandensis Van
Dyke.” CAS. TYPE LOCALITY, near Brooksville, Hernando County, Florida. Black-
welder and Blackwelder, 1948:3 ( Ferestria ).
Recognition. — The following combination of characteristics is diagnostic for hernan-
densis: prosternal process with marginate apex; elytra with strongly convex intervals and
deep striae; eversion of internal sac left and ventral around median lobe; stylus of female
ovipositor elongate and narrow. The species morio is similar to hernandensis but is distin-
guished by the absence of a raised margin at the apex of the prosternal process and
an incomplete groove along the lateral margin of the pronotum.
Description. — Body length 8.1 — 9.3 mm. Form small, short and robust.
Microsculpture of head between eyes and intervals of elytra isodiametric meshes, or
highly sinuous, entwined lines. Disc of pronotum with microsculpture usually effaced, or of
sinuous lines.
102
Freitag
Head glossy; frontal grooves short, shallowly impressed, and not sharply defined, parallel
or slightly oblique. Penultimate article of labial palpus with two medial setae (fig. 66).
Pronotum glossy; form circular in outline as in fig. 1 ; disc convex laterally but flattened in
center; sides produced, constricted slightly anteriorly and strongly posteriorly, not sinuate
near posterior margin; posterior angles obsolete, very broadly obtuse; anterior transverse
impression incomplete, impressed laterally only; basal lateral foveae deep and punctiform
posteriorly, short and shallow anteriorly. Prosternal process with marginate apex, and
medially with short, distinctly impressed longitudinal groove.
Elytra glossy, slightly sinuate apically; intervals strongly convex; striae deep anteriorly,
obsoletely or indistinctly punctate posteriorly ; stria 7 with apical end distinctly impressed,
obsolete anteriorly.
Male genitalia (fig. 81) with median lobe strongly arcuate, angle approximately right;
apical blade spatulate and slightly deflected dorsally. Right paramere fairly short, slightly
tapered apically, not extending to apical half of median lobe. Eversion of internal sac to left,
around left and ventral sides of median lobe; apical sclerite absent, dark serrulate fields
present apically on finger-like projections. The genitalia of two males were studied in detail.
Stylus of female ovipositor elongate and narrow.
Geographical distribution (fig. 125). — This species is found in western peninsular Florida.
I have seen six specimens from the following localities.
United States - FLORIDA: Citrus County: (CAS). Hernando County: Brooksville (CAS). Hillsborough County: Tampa
(ANSP, MCZ, USNM). Marion County: Juniper Springs (FDPI).
Evarthrus mono Dejean, 1828
Figures 2, 82, 125
Feronia (Steropus) morio Dejean 1828:302. TYPE, Labelled as follows: “morio M. in
America borealis”, MHNP. TYPE LOCALITY, Alma, Georgia (here selected). LeConte,
1848:355 ( Broscus ). - LeConte, 1852:231 (Evarthrus). - LeConte, 1863a:8. -
Motschulsky, 1865:264 ( Fortax ). LeConte, 1873:319 ( Evarthrus ). — Schaupp, 1880:49.
— Casey, 1918:364 ( Ferestria ). — Leng 1920:57. — Csiki, 1930:674 ( Pterostichus ).
Pterostichus (Pterostichus) (Sect. Fortax) dejeanellus Csiki, 1930:674.
Evarthrus (Ferestria) taurus Van Dyke, 1943:25. HOLOTYPE, labelled as follows: “Punta
Gorda Fla. 2.5-12.40; Van Dyke Collection”. CAS. ALLOTYPE, labelled the same as
holotype. CAS. TYPE LOCALITY - near Punta Gorda, Fla. NEW SYNONYMY. - Black-
welder, 1 948 : 3 ( Ferestria ).
Recognition. — the following characteristics are diagnostic for morio : pronotum with
incomplete lateral grooves, absent between the lateral and basal setae, and complete anterior
impression; prosternal process with apex unmargined; eversion of internal sac of male
genitalia to the left and around left and ventral sides of median lobe. The species laevipennis
is similar to morio but has crescent-shaped frontal grooves on the head, pronotum with
complete lateral grooves, and the male genitalia are different (fig. 82 cf. fig. 83).
Description. — Body length 7.7 — 10.2 mm. Form robust.
Micro sculpture of head between eyes, disc of pronotum and elytral intervals with sinuous
lines often entwined forming amorphic or isodiametric meshes and partially effaced.
Revision of Evarthrus
103
Head glossy; frontal grooves short, shallowly and broadly impressed, not sharply defined,
slightly oblique. Penultimate article of labial palpus with two to four setae.
Pronotum glossy; form subcordiform in outline as in fig. 2; disc moderately convex; sides
produced, constricted slightly anteriorly and strongly posteriorly, obsolete ly sinuate in front
of posterior angles; posterior angles obsolete, very broadly obtuse; anterior transverse im-
pression complete; basal lateral foveae deep posteriorly, short and shallow anteriorly.
Prosternal process with unmargined apex; longitudinal groove short and distinctly impressed.
First articles of middle and hind tarsi with lateral grooves.
Elytra glossy, slightly sinuate apically; intervals completely flat or slightly raised and
convex; striae 1 to 5 obsolete or distinctly impressed ; striae 6 and 7 obsolete, obsoletely or
indistinctly punctate.
Male genitalia (fig. 82) with median lobe strongly arcuate, angle approximately right;
apical blade spatula te. Right paramere narrow apically and extending to apical half of
median lobe. Eversion of internal sac to left and when everted, curled closely around left
and ventral sides of median lobe; apical sclerite absent; serrulate field present apically. The
genitalia of four males were studied in detail.
Stylus of female ovipositor average size for the morio group.
Variation . — The striae and intervals of the elytra vary. A few individuals have distinctly
impressed and punctate striae and slightly raised and convex intervals. Generally, however,
the striae are obsoletely impressed and the intervals are flat.
Notes on synonymy. — Van Dyke proposed the name taurus for the species. Fie wrote
that the presence of the marginal groove of the pronotum was characteristic of morio
Dejean. The groove is, in fact, absent in morio but it is present in the similar species
laevipennis LeConte.
Collecting notes. — H. J. Weems, Jr. has collected this species in oak leaf litter.
Geographical distribution (fig. 125). — This species ranges from southwestern Florida to
southern Georgia. I have seen 1 1 5 specimens collected in the following localities.
United States - FLORIDA: Alachua County: Archer (FDPI); Gainesville (CNC, FDPI, UMMZ); High Springs (UMMZ);
Micanopy (UMMZ); Newnan’s Lake (FDPI, UMMZ); University Farm (UMMZ); Warren’s Cave (UMMZ). Baker County: Glen
St. Mary (FDPI); Macclenny (FDPI). Charlotte County: Punta Gorda (CAS). Citrus County: (CAS). Collier County: Naples
(CAS). Dixie County: Cross City (UMMZ). Duval County: Jacksonville (AMNH). Hernando County: Brooksville (CAS).
Hillsborough County: Tampa (ANSP, MCZ, USNM). Jackson County: Florida Caverns State Park (FDPI). Manatee County:
Bradenton (GEB); Manatee (UMMZ). Orange County: Winter Park (CU). Palm Beach County: Boynton (CAS); Lake Worth
(AMNH), Putnam County: Camp Rosa, Bostwich (FDPI); Crescent City (USNM); Florahome (UMMZ); Welaka (CU).
Suwanee County: Wellborn (UMMZ). Volusia County: Enterprise (ANSP, RU, USNM). County not determined: North
Smyrna (CAS). GEORGIA: Bacon County: Alma (UMMZ). Bryan County: Lanier (UMMZ).
Evarthrus laevipennis LeConte, 1848
Figures 3-4, 83, 125
Broscus (Cephalotes) laevipennis LeConte, 1848:354. LECTOTYPE (here selected) a female,
labelled as follows: “orange disc; Type 5627; E. laevipennis Lee.” MCZ. TYPE LOCAL-
ITY, Georgia. - LeConte, 1852:231 {Evarthrus). - LeConte, 1863a:8. - LeConte, 1873:
319. - Schaupp, 1880:49. - Leng, 1915:577 ( Ferestria ). - Casey, 1920:193. - Leng,
1920:57. - Csiki, 1930:674 ( Pterostichus ). - Loding, 1945:16 ( Ferestria ).
104
Freitag
Evarthrus acutus LeConte, 1852:231. LECTOTYPE (here selected) a female, labelled as
follows: “orange disc: Type 5626; E. acutus Lee.” MCZ. TYPE LOCALITY, Louisiana.
NEW SYNONYMY. - LeConte, 1863a:8 ( Evarthrus ). - LeConte, 1873:319. - Schaupp,
1880:49. - Leng, 1915:577 ( Ferestria ). - Leng, 1920:57. - Csiki, 1930:674 ( Pterosti -
chus).
Evarthrus ovulum ; Horn, 1 875 : 1 26 (not Chaudoir).
Ferestria nanula C asey, 1918:364. HOLOTYPE, female, labelled as follows: “Mobile Ala;
CASEY bequest 1925; TYPE USNM 47111; nanula Csy.” USNM PARATYPE, female,
labelled as follows: “Mobile Ala; CASEY bequest 1925; nanula — 2 PARATYPE USNM
471 11.” USNM. NEW SYNONYMY. - Casey, 1920:192 {Ferestria). - Leng, 1920:57. -
Csiki, 1930:67 4 (Pterostichus). —Loding, 1945:16 {Ferestria).
Ferestria simiola Casey, 1920:192. HOLOTYPE, female, labelled as follows: “Mobile Ala;
CASEY bequest 1925; TYPE USNM 47112; simiola Csy.” USNM NEW SYNONYMY. -
Leng and Mutchler, 1927:10 {Ferestria). — Csiki, 1930:674 {Pterostichus).
Ferestria seminola Loding, 1945:16 (misspelling for simiola Casey).
Ferestria castigata Casey, 1920:192. HOLOTYPE, male, labelled as follows: “Mobile Ala;
H. P. Loding; male; CASEY bequest; TYPE USNM 47110; castigata Csy.” USNM. PARA-
TYPE, female, labelled as follows: “Mobile Ala.; CASEY bequest 1925; castigata — 2
PARATYPE USNM 47110. NEW SYNONYMY. - Leng and Mutchler, 1927:10
{Ferestria). — Csiki, 1930:674 {Pterostichus). — Loding, 1945:16 {Ferestria).
Ferestria bullata Casey, 1920:193. HOLOTYPE, female, labelled as follows: “Mobile Ala.
H. P. Loding, CASEY bequest 1925; TYPE USNM 47113; bullata Csy.” USNM NEW
SYNONYMY. - Leng and Mutchler, 1927:10 {Ferestria). — Csiki, 1930:674
{Pterostichus). Loding, 1945:16 {Ferestria).
Evarthrus (Ferestria) morio-, Van Dyke, 1 943:26 (not Dejean).
Ferestria acuta-, Loding, 1945:16 (not LeConte).
Recognition. — Specimens of laevipennis are distinguished by the following combination
of characteristics: head with sharply defined, crescent-shaped frontal grooves, oblique, and
widely separated. Pronotum with complete lateral grooves between the lateral and basal
setae; prosternal process shallow and broadly impressed or obsolete; internal sac everts
apicodorsally and to the left. Specimens of morio and laevipennis can be confused. However,
they are distinguished by a number of differences in structures that are described in the
recognition section of morio.
Description. — Body length 7.1 — 9.0 mm. Form relatively slender for the morio group.
Microsculpture of head between eyes, disc of pronotum, and intervals of elytra, com-
prised of generally effaced sinuous lines.
Head glossy; frontal grooves sharply defined, crescent -shaped with convexity directed
laterally, oblique and widely separated. Penultimate article of labial palpus with two medial
setae.
Pronotum glossy; form subcordiform or cordiform in outline as in figs. 3 and 4; disc
moderately convex; sides produced, constricted slightly anteriorly and strongly posteriorly,
obsoletely sinuate in front of posterior angles when posterior angles obsolete (fig. 3) ,
Revision of Evarthrus
105
distinctly sinuate when angles distinct (fig. 4); posterior angles obsolete and broadly
rounded or produced and acute; anterior transverse impression complete or absent medially;
basal lateral foveae deep and short. Prosternal process with shallow and broadly excavated
or obsolete longitudinal groove. First articles of middle and hind tarsi with lateral grooves.
Elytra glossy, obsoletely sinuate apically; intervals completely flat or slightly raised and
slightly convex; striae obsolete and impunctate or distinctly impressed and punctate, 6 and
7 always obsolete and impunctate.
Male genitalia (fig. 83) with median lobe strongly arcuate, angle slightly obtuse; apical
blade slightly tapered and evenly rounded at apex. Right paramere narrow apically and
extending to apical half of median lobe. Eversion of internal sac dorsoapically and when
everted, dorsoapically and to left; apical sclerite absent; dark serrulate field apically. The
genitalia of four males were studied.
Stylus of female ovipositor short, tapered apically and slightly sinuate preapically.
Geographical variation. — Individuals from southern localities are characterized by
obsolete basal angles of the pronotum, distinctly impressed and complete anterior transverse
impression of the pronotum (fig. 3), and elytra with completely flat intervals and obsolete
striae. In central areas of the species range populations are composed of some individuals
with the above characters, and some with more distinct basal angles of the pronotum, an
incomplete anterior transverse impression, and more or less raised intervals and impressed
punctate striae of the elytra. Specimens of northern localities have produced, sharp angles
of the pronotum, an incomplete anterior transverse impression (fig. 4), and elytra with
somewhat raised intervals and distinctly impressed and punctate striae. Because of the
apparent clinal nature of the changes in these structures, I believe northern and southern
populations, although distinct, do not merit subspecific status.
Notes on synonymy. — LeConte was not aware of the geographic variation in laevipennis.
In 1848 he proposed the name laevipennis for the northern form, and in 1852, he recog-
nized the southern form as a separate species to which he gave the name acutus.
Casey provided the names nanula, simiola, castigata, and bullata, the types of which are
of the average form of laevipennis found in Mobile, Alabama.
Collecting notes. — D. Larson and I collected specimens of laevipennis near Grey,
Georgia, in deciduous forest in leaf litter.
Geographical distribution (fig. 125). — This species inhabits the Gulf Coastal Plain and
southern Piedmont. I have seen 343 specimens from the following localities.
United States - ALABAMA: Baldwin County: (UASM). Barbour County: Eufaula (USNM), Clark County: Salt
Mountain, six miles south of Jackson (UMMZ). Colbert County: Tuscumbia Mountains, southwest of Tuscumbia (UMMZ).
Elmore County: Wetumpka (USNM). Houston County: Chatahoochee State Park (GEB). Lee County: Auburn (CAS, KSU,
VMK), Madison County: Monte Sano State Park (CAS), Mobile County: Alabama Port (GEB); Citronelle (CAS); Grand Bay
(AMNH); Mobile (AMNH, CAS, CNC, CU, MCZ, UASM, USNM); Mount Vernon (CU). Randolph County: Wadley (USNM).
Tallapoosa County: Alexander City (KSU). County not determined: Dog River (UK); FLORIDA: Jefferson County:
Monticello (UMMZ). Liberty County: Camp Totreya (CU, UMMZ); Torreya State Park (FDPI). GEORGIA: Cobb County:
Austell (CAS). Hall County: White Sulfur Springs (UMMZ). Jones County: seven miles south of Gray (RF). Rabun County:
(USNM); Clayton (USNM); Pinnacle Park (USNM). MISSISSIPPI: George County: Lucedale (CU). Greene County: Leaf
(CU). Jackson County: Ocean Springs (CU). Lamar County: Lumberton (CU). Perry County: New Augusta (CU); Richton
(CU). Stone County: Wiggins (CU). SOUTH CAROLINA: Greenville County: Greenville (USNM); 17 miles west of
Spartenburg (DL). Oconee County: CCC Camp f2 (CAS); Clemson (GEB); Clemson College (CAS, USNM). Pickens
County: Nine Times (RCG).
106
Freitag
The obsoletus Group
Characteristics. — Penultimate article of labial palpus with two medial setae. Pronotum
cordiform in outline; basal lateral foveae completely punctiform with no anterior exten-
sions; basal seta situated in front of the basal angle. Prosternal process with obsolete medial
longitudinal groove. Claw-bearing tarsal article usually with setae on lateroventral margins.
Right paramere of male genitalia elongate.
This group is generally distributed north of the morio group in regions of the Piedmont
flanking the Appalachian Mountains (fig. 126).
Evarthrus approximatus LeConte, 1848
Figures 5, 84, 126
Broscus (Cephalotes) approximatus LeConte, 1848:354. LECTOTYPE (here selected) a
female, labelled as follows: “pink disc; Type 5628; E. approximatus Lee.” MCZ. TYPE
LOCALITY, Pennsylvania. — LeConte, 1852:231 C Evarthrus ). — LeConte, 1863a:8. —
LeConte, 1873:319. — Schaupp, 1880:49. — Leng, 1920:57 ( Ferestria ). - Csiki, 1930:
674 ( Pterostichus ). - Brimley, 1938:120 C Ferestria ).
Recognition. — This species is characterized by the combination of a complete anterior
transverse impression of the pronotum, male genitalia, and geographical range restricted to
areas east of the Appalachian Mountains. Specimens of approximatus are distinguished from
those of obsoletus by the less arcuate median lobe of the male genitalia. These very similar
species are allopatric in relation to one another.
The species iuvenis resembles approximatus but is distinguished by an incomplete anterior
transverse impression of the pronotum.
Description. — Body length 8.4 — 10.9 mm. Form average for obsoletus group.
Microsculpture of head between eyes and disc of pronotum completely effaced. Micro-
sculpture at intervals of elytra effaced or consisting of indistinct isodiametric meshes. Micro-
punctures present on head and pronotum.
Head glossy; frontal grooves sharply defined, crescent-shaped with convexity directed
laterally, oblique and widely separated.
Pronotum glossy; cordiform in outline as in fig. 5, disc moderately convex; sides
produced, constricted slightly anteriorly and strongly posteriorly, obsoletely sinuate in front
of posterior angles; posterior angles obsolete; anterior transverse impression complete.
Lateroventral margin of last article of tarsus with setae.
Elytra glossy; obsoletely sinuate apically; intervals slightly raised and slightly convex;
striae clearly impressed and indistinctly punctate.
Male genitalia (fig. 84) with median lobe strongly arcuate, angle approximately right,
apical half deflected to right; apical blade evenly rounded at apex and slightly deflected
dorsally. Right paramere tapered apically, long, extending to apical half of median lobe.
Eversion of internal sac apical and to left; apical sclerite absent, serrulate fields present
apically. The genitalia of three males were studied in detail.
Stylus of female ovipositor tapered apically and sinuate preapically.
Geographical distribution (fig. 126). — This species is found in North Carolina, Virginia
and Washington, D. C. According to LeConte, it also occurs in Pennsylvania, but I have not
been able to verify this record. I have seen 41 specimens collected in the following localities.
Revision of Evarthrus
107
United States - DISTRICT OF COLUMBIA: Washington (ANSP, CAS, MCZ, UK, USNM). NORTH CAROLINA:
Guilford County: High Point (USNM). Arlington County: Rosslyn (UASM, USNM). Fairfax County: (USNM); Blackpond
(USNM): Hemdow (USNM). Giles County: Mountain Lake (UMMZ). Henrico County: Richmond (AMNH).
Evarthrus iuvenis new species
Figures 6, 85, 1 26
Recognition . — The internal sac of the median lobe of iuvenis everts to the left and curls
ventrally on the left side of the median lobe. This feature alone sets iuvenis apart from the
similar obsoletus. Also obsoletus inhabits areas west of the Appalachian Mountains, while
iuvenis occurs east of that mountain range. Another diagnostic characteristic of iuvenis is
the shape of the median lobe of the male (fig. 85).
Description. — HOLOTYPE, male, labelled as follows: “24 miles north of Roanoke,
Virginia Blue Ridge Parkway 21 October 1962 leg. D. R. Whitehead; HOLOTYPE Evarthrus
iuvenis R. Freitag (red label).” MCZ.
Body length 10.3 mm; width 4.1 mm. Form robust. Microsculpture effaced on head
between eyes and disc of pronotum. Isodiametric meshes on intervals of elytra.
Head glossy; length 1.2 mm; width 2.5 mm; frontal grooves sharply defined, crescent-
shaped with convexity directed laterally, oblique toward one another, and widely separated.
Pronotum glossy; length 2.8 mm, width 3.4 mm; form cordiform in outline as in fig. 6;
disc moderately convex; sides produced, constricted slightly anteriorly, strongly posteriorly,
obsoletely sinuate in front of posterior angles; posterior angles obsolete; anterior transverse
impression absent medially. First articles of middle and hind tarsi with lateral grooves;
lateroventral margins of claw-bearing article of tarsus with setae.
Elytra glossy; length 6.2 mm, width 4.1 mm; slightly sinuate apically; intervals slightly
convex; striae clearly impressed and indistinctly punctate.
Male genitalia (fig. 85) with median lobe slightly arcuate; apical blade narrow and evenly
rounded at apex, deflected dorsally and to right; right paramere with markedly tapered
apical half, extending to apical half of median lobe; eversion of internal sac to left and in
everted position curled ventrally around median lobe on left side; apical sclerite absent,
serrulate fields present apically.
ALLOTYPE, female labelled as follows: “Raleigh, N. C., April 14’49 H. F. Howden;
under board; near 1043 (yellow label) loan from CNC; ALLOTYPE Evarthrus iuvenis. R.
Freitag”. CNC.
Body length 11.1 mm; width 4.8 mm. Form same as in holotype.
Microsculpture of head between eyes and disc of pronotum consists of partially entwined
lines. Intervals of elytra with isodiametric microsculpture.
Head slightly glossy; length 1.4 mm, width 3 mm.
Pronotum, form same as in holotype; length 3 mm; width 3.8 mm.
Elytra slightly glossy; intervals flat; striae distinctly impressed and clearly punctate;
length 6.7 mm; width 4.8 mm.
Stylus of ovipositor short and tapered apically, slightly sinuate preapically.
Variation among paratypes (six males, seven females. North Carolina, Virginia). — Total
length 9.8 - 12.8 mm. The basal angles of the pronotum are slightly produced and sharp in
108
Freitag
six specimens. The last tarsal article may or may not have setae on the ventral side. The
apical blade of three males is half the width of that of the holotype, while that of the other
three is approximately the same width as that of the holotype. In other respects the
paratypes resemble the holotype and allotype.
Derivation of specific name. — The word iuvenis is a Latin noun, meaning warrior. I have
given the name to this species because its members seem warrior-like, robust and large in
size for the subgenus Fortax.
Disposition of type material. - The holotype is in the MCZ. The allotype was returned to
the CNC. The paratypes are deposited in the collection of CAS, DRW, RCG, RTB, UASM,
UMMZ, and USNM.
Collecting notes. — This species is found in leaf litter in forested places.
Geographical distribution (fig. 126). — Evarthrus iuvenis is known from western Virginia
and North Carolina. I have seen eight specimens from the following localities.
United States - NORTH CAROLINA: Stokes County: Hanging Rock State Park (RTB). Wake County: Raleigh (CNC).
VIRGINIA: Bland County: Summit of Walker Mountain (UMMZ). Campbell County: (USNM). Floyd County: (UASM);
Buffalo Mountain, five miles southeast of Willis (DRW). Rocky Knob Recreation Area, Blue Ridge National Parkway
(RCG). Giles County: Cascades (TCB); Mountain Lake (RTB). Nelson County: (USNM). Roanoke County: Blue Ridge
Parkway, 24 miles north of Roanoke (MCZ).
Evarthrus obsoletus Say, 1823
Figures 7, 86, 126
Feronia obsoleta Say, 1 823a :57. Type lost. Type Locality — Indiana (here selected). — Say,
1834:424 (Feronia). — LeConte, 1848:354 (Broscus). — LeConte, 1852:231 (Evarthrus).
— LeConte, 1863a:8. - LeConte, 1873:319. - Schaupp, 1880:49. - Blatchley, 1910:91
(Pterostichus). - Casey, 1918:364 (Ferestria). — Casey, 1920:193. - Leng 1920:57. -
Csiki, 1930:674 (Pterostichus). — Loding, 1945:16 (Ferestria).
Recognition. — The shape of the median lobe of the male is markedly arcuate, and differs
strongly from that of approximate and iuvenis (fig. 86 cf. figs. 84 and 85). The species
obsoletus alone in this group occurs to the west and south of the Appalachian Mountains.
Description. — Body length 7.9 — 9.9 mm. Form average for obsoletus group.
Microsculpture of head between eyes of obsolete sinuous impressions or completely
effaced. Microsculpture of disc of pronotum and intervals of elytra completely effaced.
Head glossy; frontal grooves sharply defined, slightly curved, with convexity directed
laterally, oblique toward one another and widely separated.
Pronotum glossy, cordiform in outline, as in fig. 7; disc moderately convex; sides
produced, constricted slightly anteriorly and strongly posteriorly; posterior angles obsolete;
anterior transverse impression incomplete, impressed laterally only. First articles of middle
and hind tarsi with lateral grooves; last article of tarsus with setae on lateroventral margin.
Elytra glossy; obsoletely sinuate apically; medial intervals highly convex, lateral intervals
flatter; striae deeply impressed, indistinctly punctate.
Male genitalia (Fig. 86) with median lobe strongly arcuate, angle slightly obtuse; apical
blade deflected to right, left edge deflected dorsally, evenly rounded at apex. Right
paramere tapered apically, long, extended apically well beyond halfway point of median
lobe. Eversion of internal sac apical and to left; apical sclerite absent; serrulate fields present
apically. The genitalia of three males were examined.
Revision of Evarthrus
109
Stylus of female ovipositor pointed at apex, tapered apically.
Notes on synonymy. — Say (1834) wrote that this species occurred in Indiana. For this
reason I have selected Indiana as Type area. This species was identified by the original
description.
Collecting notes. — This species is found in deciduous forests in damp leaf litter.
Geographical distribution (fig. 126). — This species ranges from southern Alabama north
to Michigan west of the Appalachian Mountains. I have seen a total of 131 specimens.
United States - ALABAMA: Bibb County: The Sinks (UMMZ). Cherokee County: Leesburg (UMMZ). Colbert County:
(USNM); Barton (CAS). Fayette County: Berry (GEB). Jackson County: Paint Rock (UMMZ). Madison County: Monte
Sano State Park (CAS, CNHM, UASM). Mobile County: Mobile (CAS). Monroe County: Claiborne (UMMZ). Randolph
County: Wadley (USNM). St. Clair County: Blount Mountains (GEB). Talladega County: Talladega (UMMZ). Tuscaloosa
County: Hurricane Creek, near Peterson (GEB). Hurricane Creek, seven miles north of Tuscaloosa (RF); Lock 14 (CAS);
Tuscaloosa (GEB); University (UMMZ). County not determined: National Forest (CAS, USNM); Tumblin Gap (USNM).
GEORGIA: Cherokee County: Galt’s Lodge (TCB). Fulton County: Silver Lake (USNM). Morgan County: four miles north
of Madison (DL, RF); Madison (UMMZ). ILLINOIS: Cook County: Palos Park (UMMZ). Vermilion County: Camp Robert
Drake, near Fairmount (RTB). Washington County: Dubois (UMMZ). INDIANA: Crawford County: (CNHM). Fulton
County: (CAS). Monroe County: Bloomington (UMMZ). KENTUCKY: Fleming County: Blue Briar Springs (GEB).
MICHIGAN: Washtenaw County: Cady’s Woods, Ann Arbor (UMMZ). MISSISSIPPI: Lauderdale County: Meridian
(UMMZ). Pontotoc County: Pontotoc (UMMZ). NORTH CAROLINA: Cherokee County: Murphy (CAS). OHIO: (CAS).
TENNESSEE: Blount County: Chilhowee Mountains (CNQ; Great Smoky Mountains National Park (CNC). Cumberland
County: Grassy Cove (CAS). Hamilton County: Chattanooga (UMMZ). Lauderdale County: South Fulton (UMMZ). Morgan
County: Burrville (UMMZ); Deer Lodge (USNM); Environs (CNHM). Obion County: Obion (UMMZ). Sevier County:
Elkmont (USNM). County not determined: Cades Cove, Blounto (MCZ); Cedar Glade Area (USNM); Cove Mountain Trail
(TCB).
The Subgenus Cyclo track elus Chaudoir
Cyclotrachelus Chaudoir, 1838:27. Casey, 1918:348. — Van Dyke 1943:27. - Ball 1960:
129. TYPE SPECIES - Molops faber Germar, 1824 (here designated).
Evarthrus roticollis Casey (designated type species by Casey, 1918:348).
Characteristics. — The subgenus Cyclotrachelus is distinguished from the other subgenera
of Evarthrus by the following combination of characteristics: pronotum with sides strongly
constricted posteriorly, posterior lateral foveae monostriate, posterior lateral setae situated
on bead (figs. 8—10), anterior transverse impression incomplete and impressed laterally
only; middle femur with four setae on anterior face except for that of the species unicolor
which has up to seven; eversion of internal sac of median lobe of male genitalia dorsoapical,
internal sac not curled ventrally in everted position; right paramere of male genitalia without
“elbow” at bend, and only slightly tapered apically. The styli of female ovipositor short and
broad, in a few species sinuate preapically and tapered apically. With the exception of the
right paramere all the above characteristics are present in each species of this subgenus. The
species levifaber has an elbow in the right paramere.
The three species groups in Cyclotrachelus are: the spoliatus group, the ovulum group,
and the faber group.
Notes on synonymy *• — Casey’s designation of E. roticollis Casey as type species, was in-
correct because this name was not included with the original description of Cyclotrachelus.
110
Freitag
The spoliatus Group
Characteristics. — Pronotum almost circular in dorsal aspect and basal angles broadly
obtuse; longitudinal groove of the prostemal process usually shallow, but if deep not sharply
defined; internal sac of the median lobe of the male genitalia without apical sclerite, with
serrulate fields apically.
This group includes the following species: unicolor , fucatus, spoliatus, and brevoorti.
Members of these species inhabit the Coastal Plain and Piedmont regions of southeastern
United States.
Evarthrus unicolor Say, 1823
Figures 8, 63, 67, 87, 127
Feronia unicolor Say, 1823a:40. Type lost. TYPE LOCALITY, Georgia (here selected). -
LeConte, 1848:352 {Feronia). - LeConte, 1852:230 C Evarthrus ). - LeConte, 1863a:8. —
LeConte, 1873:319. — Schaupp, 1880:49. — Casey, 1918:349 ( Cyclotrachelus ). - Leng,
1920:56. - Csiki, 1930:672 ( Pterostichus ). - Loding, 1945:15 ( Cyclotrachelus ).
Recognition. — The large body size, four to six setae on the penultimate article of the
labial palpus (fig. 67), raised knobs flanking the anterior end of the gula, and dark apical
serrate field in the internal sac of the median lobe of the male genitalia, combined, dis-
tinguish specimens of unicolor from those of the similar species fucatus, spoliatus and
brevoorti. The species fucatus is further distinguished by frontal grooves of the head that
are oblique to one another and a subcordiform glossy pronotum. All specimens of brevoorti
are smaller than those of unicolor. The species spoliatus and unicolor are allopatric, and can
also be distinguished by structural features of the male genitalia (fig. 87 cf. fig. 89).
Description. — Body length 17.7 — 22.0 mm. Form parallel and elongate.
Microsculpture of head between eyes, pronotal disc, and elytral intervals with lines
distinctly impressed, highly sinuous and entwined, often forming isodiametric or amorphic
meshes.
Head dull or slightly glossy; frontal grooves deeply impressed, straight, and parallel to one
another. Penultimate article of labial palpus with four to six setae. Anterior end of gula
flanked by raised knobs (fig. 63).
Pronotum dull or slightly glossy; form as in fig. 8; disc moderately convex; sides slightly
constricted anteriorly, strongly constricted posteriorly, and moderately sinuate in front of
posterior angles; posterior angles right or slightly obtuse and produced; anterior transverse
impression incomplete; basal foveae of average length and moderately impressed. Longi-
tudinal groove in prosternal process shallow or deep. Anterior face of middle femur with
four to seven setae.
Elytra dull or slightly glossy, sinuate apically; intervals usually flat, occasionally slightly
raised; striae shallowly impressed, small punctures confined to anterior two-thirds, impunc-
tate posteriorly.
Male genitalia (fig. 87) with median lobe strongly arcuate, angle approximately right;
apical blade produced, of average width to relatively broad, apex evenly rounded: right
paramere typical Cyclotrachelus form, not reaching apical half of median lobe; eversion of
internal sac apicodorsal and to the right; internal sac with a dark serrate field apically,
apical sclerite absent. The male genitalia of three specimens were examined in detail.
Revision of Evarthrus
111
Stylus of female ovipositor short, broad, and evenly rounded at tip.
Notes on synonymy. — This species was identified by the original description, and by an
examination of the LeConte unicolor specimen in the LeConte Collection. I have selected
Georgia as type locality because many unicolor specimens which I have seen are from
Georgia.
Geographical distribution (fig. 127). — This species inhabits the Gulf Coastal Plain and
southern Piedmont. I have seen 18 specimens from the following localities.
United States - ALABAMA: Cherokee County: Leesburg (UMMZ). Lee County: Auburn (AL, UMMZ). FLORIDA:
Jackson County: (FDPI). GEORGIA: Dodge County: Chester (CAS, CU). Dooley County: Umadilla (UMMZ), Morgan
County: Madison (UMMZ). Upson County: (MCZ, USNM).
Evarthrus fucatus new species
Figures 9, 88, 127
Recognition. — The following three characters combined are diagnostic for the species
fucatus : subcordiform pronotum; highly glossy dorsum, and form of male genitalia. Al-
though there are striking similarities among fucatus, spoliatus, and brevoorti they differ in
the following respects.
The species fucatus and spoliatus are allopatric. Specimens of fucatus have a subcordi-
form pronotum, and sometimes three setae on the penultimate article of the labial palpus,
while spoliatus specimens have a more circular pronotum and always two setae on the
penultimate article of the labial palpus. In addition the frontal grooves of the head are
oblique in fucatus and parallel in spoliatus.
There is some overlap in the distributions of fucatus and brevoorti. Specimens of fucatus
normally have a more cordiform pronotum. However the most distinguishing character is
the male genitalia. The apex of the median lobe of fucatus is evenly rounded, but is truncate
in brevoorti.
Description. - HOLOTYPE, male, labelled as follows: “Cherokee Co., Ala. Leesburg VII
- 25 - 1929 54. T. H. Hubbell; loan from UMMZ; HOLOTYPE Evarthrus fucatus R. Freitag
(red label).” UMMZ.
Body length 14.1 mm; width 5.7 mm. Form typical of this group, with robust pronotum.
Microsculpture of head between eyes, disc of pronotum, and elytral intervals, with highly
sinuous dense and closely entwined lines, often forming amorphic meshes.
Head glossy; length 1.7 mm, width 3.4 mm; frontal grooves straight, deep and sharply
defined, oblique and widely separated; penultimate article of labial palpus with three setae,
two medial and one apical.
Pronotum with disc glossy; length 3.9 mm, width 4.6 mm; subcordiform (fig. 9); disc
moderately convex; sides slightly constricted anteriorly and markedly constricted posteri-
orly, obsoletely sinuate in front of posterior angles; posterior angles not produced and
broadly obtuse; anterior transverse impression absent medially; median longitudinal im-
pression slightly deeper at either end; basal foveae deepest at bend, of average length.
Longitudinal groove of prostemal process broad, indistinct and very shallowly impressed.
Anterior face of middle femur with four setae.
112
Freitag
Elytra glossy; length 8.5 mm. width 5.7 mm; obsoletely sinuate apically; intervals
moderately raised but slightly flattened; striae deep with indistinct punctures in apical half,
punctures obsolete in apical third.
Male genitalia (fig. 88) with median lobe strongly arcuate, angle slightly acute; apical
blade moderately produced, slightly tapered apically, and evenly rounded; right paramere
typical Cyclotrachelus form and of average length reaching apical half of median lobe,
eversion of internal sac apicodorsal and slightly to right; internal sac with apical serrulate
field; apical sclerite absent.
ALLOTYPE, female, labelled as follows: “Monte Sano State Park, ALABAMA 7-VI-1960
B. Benesh; CNHM 1965 Bernard Benesh General Coleop. Coll.; ALLOTYPE Evarthrus
fucatus R. Freitag”. CNHM.
Body length 14.3 mm, width 8.9 mm. Form same as in holotype.
Microsculpture of head between eyes and disc of pronotum same as in holotype. Elytra
with microsculpture mainly composed of amorphic or isodiametric raised meshes that
appear beady.
Head glossy; length 1.7 mm, width 3.2 mm.
Pronotum, shape, same as in holotype; length 3.7 mm, width 4.7 mm.
Elytra not highly glossy; intervals slightly convex, almost flat; striae deep and obsoletely
punctate anteriorly, impunctate posteriorly; length 8.9 mm, width 5.7 mm.
Stylus of ovipositor obsoletely sinuate preapically and tapered apically.
Derivation of specific name. — The name fucatus is a Latin adjective which means deceit-
ful and has been given this species because of the remarkable similarity of its members to
those of spoliatus.
Variation among paratypes (19 males, 13 females, Georgia, Alabama, Tennessee, West
Virginia, Pennsylvania, Ohio, CM, CNHM, UASM, UMMZ, USNM). - Total length 12.0 -
14.6 mm. The penultimate article of the labial palpus bears two or three setae. A minimal
amount of variation is evident in coloration and structural features among the paratypes
which resemble closely the holotype and allotype. The male genitalia of five specimens were
examined in detail.
Disposition of type material. — The holotype and allotype are in the collections of the
UMMZ and CNHM, respectively. The paratypes are in the collections of the following
institutions: CM, CNHM, UASM, UMMZ, and USNM.
Collecting notes. — Specimens of fucatus have been collected in deciduous forest in leaf
litter.
Geographical distribution (fig. 127). — This species inhabits the Piedmont on the western
and southern sides of the Appalachian Mountains. I have seen 41 specimens collected in the
following localities.
United States - ALABAMA: Cherokee County: Leesburg (UMMZ). Madison County: Huntsville (UMMZ); Monte Sano
State Park (CNHM, UASM). GEORGIA: Floyd County: Armuchee (UMMZ). KENTUCKY: Edmonson County: near Hist.
Ent. (TCB); Mammoth Cave (TCB). OHIO: Hamilton County: Cincinnati (UMMZ). PENNSYLVANIA: Allegheny County:
Pittsburg (CM). Westmoreland County: Jeanette (CM). TENNESSEE: Cumberland County: Grassy Cove (UMMZ). Hamil-
ton County: Signal Mountain (UMMZ). Montgomery County: Clarksville (USNM). Moigan County: Bunville (CNHM).
WEST VIRGINIA: Marion County: Fairmount (CM).
Revision of Evarthrus
113
Evarthrus spoliatus Newman, 1 838
Figures 10, 89, 127
Feronia spoliata Newman, 1838:386. TYPE, male, labelled as follows: “Type H. T.; Ent.
Club. 44—12; J. Ingall, Canada.” BM. TYPE LOCALITY, Southern Pines, N. C. (here
selected).
Evarthrus rotundatus LeConte, 1852:230. LECTOTYPE (here selected) a female, labelled as
follows: “Va; rotundatus 2”. MCZ. NEW SYNONYMY. — LeConte, 1863a:8 ( Evarthrus ).
- LeConte, 1873:319. - Schaupp, 1880:49. - Casey, 1918:349 ( Cyclotrachelus ). -
Leng, 1920:56. - Csiki, 1930:671 ( Pterostichus ). — Brimley, 1938:119 ( Cyclotrachelus ).
— Loding, 1945: 14.
Evarthrinus (Evarthrops) pinorum Casey, 1920:198. HOLOTYPE, male, labelled as follows:
“Southern Pines, A. H. Manee. NC.; CASEY bequest 1925; TYPE USNM 47135; pinorum
Csy”. USNM. TYPE LOCALITY, Southern Pines, N. Carolina. NEW SYNONYMY. -
Leng and Mutchler, 1927:10 {Evarthrinus). — Csiki, 1930:673 {Pterostichus). — Brimley,
1 938 : 1 1 9 {Evarthrinus).
Recognition. — The combination of geographical distribution, generally parallel frontal
grooves on the head, and form of the median lobe of the male genitalia is characteristic of
spoliatus. The differences among spoliatus, fucatus and unicolor have been discussed in
connection with the recognition of the last two species. The remaining species in this group,
brevoorti, can also be mistaken for spoliatus. The frontal grooves of the head are parallel in
spoliatus and oblique in brevoorti, and the apex of the median lobe is rounded in spoliatus,
truncate in brevoorti.
Description. — Body length 12.8 — 15.8 mm. Form elongate.
Head between eyes, disc of pronotum, and elytral intervals, with lines of microsculpture
dense, highly sinuous, entwined, forming amorphic raised meshes.
Head moderately or slightly glossy. Frontal grooves broadly but deeply impressed,
normally with a slight bend the convexity of which is directed medially. Penultimate article
of labial palpus with two setae.
Pronotum with slightly glossy disc; form as in fig. 10; disc moderately convex; sides
moderately constricted anteriorly, more strongly so posteriorly, and distinctly sinuate in
front of posterior angles; posterior angles slightly to broadly obtuse, and slightly produced;
basal foveae moderately impressed. Longitudinal groove in prostemal process shallow.
Middle femora each with four setae on anterior face; occasionally four setae on one middle
femur and five or six on opposite one.
Elytra dull or slightly glossy; slightly sinuate apically; intervals slightly raised and flat;
striae shallow or moderately impressed with small punctures anteriorly, impunctate posteri-
orly.
Male genitalia (fig. 89) with arcuate median lobe, apical half deflected to right; apical
blade slightly tapered apically and apex evenly rounded; right paramere short and rather
stout; internal sac with serrulate field apically, apical sclerite absent. Elongate, left lateral
sclerotized flap of median lobe near opening of invaginated internal sac extending onto basal
half of sac when everted; eversion of internal sac apicodorsal and to right. The genitalia of
four males were studied in detail.
114
Freitag
Stylus of female ovipositor tapered apically and slightly sinuate behind apex.
Notes on synonymy. — I have selected Southern Pines as the type locality because
spoliatus specimens have been collected there. Also, it is centrally located in the species
range. The name Canada on the label of the type specimen indicates the country of the
collector, J. Ingall. The type specimens of rotundatus LeConte and pinorum Casey are
average specimens of spoliatus.
Collecting notes. — V. M. Kirk collected specimens of spoliatus in litter on the ground in
deciduous forest.
Geographical distribution (fig. 127). — This species is found on the Piedmont and Coastal
Plain west of the Appalachian Mountains from District of Columbia south to South
Carolina. I have seen 61 specimens from the following localities.
United States - DISTRICT OF COLUMBIA: Rock Creek (USNM, CNHM). NORTH CAROLINA: Duplin County;
Faison (CNC). Durham County: Durham (USNM). Franklin County: Louisbuig (CNC) Moore County: Pinehurst (MCZ):
Southern Pines (USNM). New Hanover County: Wilmington (USNM). Orange County: Chapel Hill (CAS, CU). Union
County: (GEB). Wake County: Raleigh (CNC). SOUTH CAROLINA: Bamberg County: Bamberg (VMK). Darlington
County: Darlington (UMMZ). Florence County: Florence (GEB, VMK); Scranton (UMMZ); Three miles east of Florence
(GEB). Sumter County: Sumter (GEB). VIRGINIA: County not determined: Virginia (CAS).
Evarthrus brevoorti LeConte, 1848
Figures 11-13,90-91, 127
Feronia (Pterostichus) brevoorti LeConte, 1848:352. LECTOTYPE (here selected) a male,
labelled as follows: “orange disc: Type 5625; E. spohatus (Newm). Brevoorti Lee.” MCZ.
TYPE LOCALITY, Alabama. — LeConte, 1852:230 (Evarthrus). — LeConte, 1863a:8. —
LeConte, 1873:319. — Schaupp, 1880:49. — Leng, 1920:57. - Csiki, 1930:671
(Pterostichus).
Evarthrus spoliatus ; LeConte, 1873:319 (not Newman). — Schaupp, 1880:49. — Csiki,
1930:671.
Recognition. — The most distinctive feature of this species is the truncate apex of the
median lobe of the male. Additional characteristics of this species have been mentioned
above in connection with recognition of unicolor , spoliatus , and fucatus.
Description. - Body length 10.4 — 16.3 mm. Form typical of the spoliatus group.
Head between eyes, disc of pronotum, and elytral intervals, with lines of microsculpture
highly sinuous, entwined and usually forming amorphic meshes.
Head slightly or markedly glossy. Frontal grooves sharply defined, straight, slightly
oblique. Penultimate article of labial palpus with two to four setae.
Pronotum with disc slightly or markedly glossy; form as in figs. 1 1-1 3; disc moderately
convex; sides produced medially, constricted posteriorly slightly or moderately sinuate in
front of posterior angles; posterior angles slightly produced, broadly obtuse; basal foveae
moderately impressed. Longitudinal groove in prostemal process shallow and poorly defined
or obsolete. Front face of middle femur with four setae.
Elytra slightly glossy and usually iridescent; slightly sinuate apically; intervals raised and
weakly convex; striae deep, distinctly punctate anteriorly, obsoletely punctate or impunc-
tate posteriorly.
Revision of Evarthrus
115
Male genitalia (figs. 90—91) with median lobe arcuate, angle slightly obtuse, and apical
half slightly deflected to right, apex truncate and often with a short, right, lateral spine;
right paramere broad and reaching apical half of median lobe; internal sac with dark
serrulate fields basally and very light, highly folded, serrulate fields apically; apical sclerite
absent. Eversion of internal sac apicodorsal. The genitalia of nine males were studied in
detail.
Stylus of female ovipositor sinuate preapically and tapered apically.
Geographical variation. — Individuals from coastal populations are generally somewhat
larger in body size and slightly duller than those of inland areas. In some males the median
lobe has a short spine jutting out of the right side of the apical blade (fig. 91). The prono-
tum varies slightly in form, but there is no geographical pattern to the variation.
Collecting notes. — Specimens of brevoorti are found in forested areas. H. V. Weems, Jr.
collected a specimen in leaf mold on a bank of a stream in Florida. They are also found in
rotting logs.
Geographical distribution (fig. 127). — This species is found on the Gulf Coastal Plain and
southern Piedmont. I have seen 92 specimens from the following localities.
United States - ALABAMA: Clarke County: Salt Mountain, six miles south Jackson (UMMZ). DeKalb County:
Mentone (GEB). Lee County: Auburn (AU). Mobile County: Alabama Port (GEB); Calvert (CAS, NCSU); Mobile (CAS,
CU, MCZ, UMMZ, USNM). Perry County: Felix (UMMZ). FLORIDA: Liberty County: Camp Torreya (UMMZ); Torreya
State Park (FDPI). GEORGIA: Cobb County: Austell (CAS). Floyd County: 2 mi. s. Armuchee (UMMZ). Fulton County:
Atlanta (CAS). MISSISSIPPI: Choctaw County: Little Mountain Camp Ground (RCG). George County: Lucedale (CU).
Jackson County: Ocean Springs (CU); Pascagoula (CU). Oktibbeha County: State College (CAS). Wayne County: Waynes-
boro (UMMZ). County not determined: Oneca Springs (CU). SOUTH CAROLINA: Oconee County: Clemson (CAS, GEB);
Clemson College (AMNH, USNM). Pickens County: Kedwee River (RCG). Saluda County: Saluda (UMMZ).
The ovulum Group
Characteristics. — Small to medium size beetles; pronotum cordate with basal angles pro-
duced, relatively sharp and slightly obtuse; apical sclerite present in internal sac of median
lobe.
This group includes the following species: macr ovulum, texensis, ovulum, alabamensis
and vinctus.
The group occurs on the Coastal Plain, except for the species vinctus, which lives in the
higher altitudes of northern Georgia and western North Carolina, and in the Great Smoky
Mountains.
Evarthrus vinctus LeConte, 1852
Figures 14, 92, 128
Evarthrus vinctus LeConte, 1852:232. LECTOTYPE (here selected) a female, labelled as
follows: “orange disc; Type 5623; E. vinctus Lee.” MCZ. TYPE LOCALITY, Nakutshi
Valley, Habersham Co., Georgia. — LeConte, 1863a:8 (Evarthrus). — LeConte, 1873:319.
- Schaupp, 1880:49. - Casey, 1918:350 ( Cyclotrachelus ). - Leng, 1920:57. - Csiki,
1930:672 (Pterostichus).
Recognition. - The following characters of this species combined are diagnostic: sharply
defined and oblique frontal grooves of the head ; elongate and deeply impressed basal foveae
116
Freitag
of the pronotum; very shallow longitudinal groove in the prostemal process; very convex
and iridescent intervals of the elytra in the males; obsoletely punctate or impunctate elytral
striae; and male genitalia (fig. 92).
Specimens of vinctus and those of its congeners in the ovulum group are similar in appear-
ance. The impunctate or obsoletely punctate elytral striae are characteristics of vinctus and
distinguish specimens of this species from those of alabamensis, macrovulum, texensis and
ovulum.
Description. — Body length 8.5 — 1 1 .1 mm. Form typical of this group.
Microsculpture on head between eyes with slightly sinuous, entwined lines, and raised
amorphic meshes. Disc of pronotum with microsculpture same as that on head but occasion-
ally partially effaced. Micro sculpture of elytral intervals same as that on head only more
stretched transversely and slightly effaced.
Head slightly or markedly glossy. Frontal grooves deep, sharply defined and oblique.
Penultimate article of labial palpus with two or three medial setae.
Pronotum slightly or markedly glossy; form cordiform as in fig. 14; disc slightly convex;
sides slightly constricted anteriorly and strongly constricted posteriorly, very broadly sinu-
ate in front of posterior angles; posterior angles prominent, slightly obtuse; anterior trans-
verse impression incomplete, very rarely complete, impressed laterally only; basal foveae
elongate, sharply impressed throughout, deep posteriorly, crescent-shaped with convexity
directed medially. Prostemal groove distinct or obsolete, but always very shallow. Four
setae on front face of middle femur.
Elytra highly glossy and slightly iridescent in males, slightly glossy and slightly iridescent
in a few females; intervals moderately to strongly convex in males, slightly convex in
females; striae deeply impressed and obsoletely punctate or impunctate.
Male genitalia (fig. 92); angle of median lobe slightly obtuse, apical half deflected to right,
apical blade moderately tapered apically and evenly rounded; right paramere of average
length extending to apical half of median lobe, shape typical of Cyclotrachelus , eversion
of internal sac to right and slightly dorsal of median lobe, and a basal bulbous serrulate field
directed to left; internal sac with serrulate field basally and apically, preapical sclerite lightly
sclerotized, large and hemispherical. The genitalia of two males were studied in detail.
Stylus of female ovipositor short with preapical, lateral sinuation, and tapered apically.
Collecting notes. — Members of this species inhabit leaf litter of forests (information
obtained from locality label).
Geographical distribution (fig. 128). — This species is found in the high Piedmont of
northern Georgia, western South Carolina and eastern Tennessee. I have seen 29 specimens
from the following localities.
United States - GEORGIA: Rabun County: Black Rock Mountain (CU); Clayton (AMNH, CAS, CU, MCZ, UMMZ,
USNM). Rabun - Towns Co. line, Appalachian Trail, 1-2 mi. s. Dicks Ck. Gap, 3000-4000’ (TCB). Towns - White Co. line,
Rt. 75-17, Appalachian Trail, Unicoi Gap, 2950’ (TCB). NORTH CAROLINA: eight miles northeast of Highlands (RTB).
SOUTH CAROLINA: Oconee County: Walhalla (CAS). County not determined: Walnut Creek Gap, Cowee Mountains
(RTB). TENNESSEE: Clingman’s Dome, Great Smoky Mountains National Park (DRW).
Revision of Evarthrus
1 17
Evarthrus alabamensis Casey, 1920
Figures 15,70,93, 128
Evarthrus constrictus Bates, 1882:80 (not Say, 1823b). TYPE, female, labelled as follows:
“Type H. T.; Mexico Salle Coll.; B. C. A. Col. I. 1. Evarthrus constrictus Bates; Evarthrus
constrictus Bates.” BM. NEW SYNONYMY. - Horn, 1886:9. - Blackwelder and Black-
welder, 1948:2.
Evarthrinus (Evarthrops) alabamensis Casey, 1920:198. HOLOTYPE, male, labelled as
follows: “Allen Ala los; CASEY bequest 1925; TYPE USNM 47136; alabamensis Csy.”
USNM. TYPE LOCALITY, Allen, Alabama. — Leng and Mutchler 1927:10 (Evarthrinus).
— Csiki, 1930:673 (Pterostichus). Loding, 1945:16 (Evarthrinus).
Evarthrinus (Evarthrops) lilliputicus Casey, 1920:199. HOLOTYPE, male, labelled as fol-
lows: “Mobile Ala. VII— 17 H. P. Loding; CASEY bequest 1925; TYPE USNM 47137; lil-
liputicus Csy.” USNM. PARATYPE, female, labelled as follows: “Mobile, Ala. II — 5 — 1 5.
H. P. Loding; CASEY bequest 1925; lilliputicus - 2 PARATYPE USNM 47137.” USNM.
NEW SYNONYMY. - Leng and Mutchler 1927:10 (Evarthrinus). - Csiki, 1930:
673 (Pterostichus). - Loding, 1945:16 (Evarthrinus).
Pterostichus batesellus Csiki, 1930:671. — Blackwelder and Blackwelder, 1948:1 (Evar-
thrus). New name for constrictus Bates, not Say, 1823b.
Recognition. — Specimens of this species are most easily recognized by their glossy
pronota and dull elytra. Other diagnostic characters are: distinctly impressed straight frontal
grooves (fig. 70); shape of the pronotum (fig. 15); and form of male genitalia (fig. 93).
This species, macrovulum, texensis and ovulum are separable by a number of characters
that are given in the recognition sections of the last three species. However individuals of the
species parafaber may also be mistaken for those of alabamensis. These can be distinguished
by several characters. Specimens of alabamensis have: a cordiform pronotum with produced
basal angles and glossy disc with partially effaced microsculpture; and moderately impressed
elytral striae with distinct punctures. In contrast specimens of parafaber have: an oval
pronotum with more parallel sides, recessed basal angles, and a semi-glossy disc with dense,
closed, slightly transversely stretched meshes comprising the microsculpture; and deeply
impressed elytral striae with coarse and broad indistinct punctures. In addition the male
genitalia are diagnostic (fig. 96).
Description. — Body length 8.8 — 12.6 mm. Form of body typical of ovulum group.
Head between eyes and disc of pronotum with lines of microsculpture sinuous, entwined
and forming open meshes. Microsculpture of elytral intervals with isodiametric, raised and
beady meshes in females, flatter in males.
Head glossy; frontal grooves (fig. 70) straight, sharply defined, slightly oblique and
moderately separated. Penultimate article of labial palpus with two medial setae.
Pronotum (fig. 15) with disc glossy; sides strongly constricted posteriorly and very
broadly sinuate in front of hind angles; posterior angles small, prominent and slightly
obtuse; anterior transverse impression only impressed laterally, absent medially; basal foveae
moderately impressed. Prosternal process with deep and sharply defined longitudinal groove.
Anterior face of middle femur with four setae.
118
Freitag
Elytra dull in females, slightly glossy in males; intervals moderately convex in males,
distinctly flatter in females; striae moderately impressed in males shallow in females; punc-
tures of striae coarse in males, small and distinct in females, obsolete posteriorly in both
sexes. First articles of middle and hind tarsi with lateral grooves.
Median lobe (fig. 93) of male genitalia strongly arcuate, angle almost right; apical blade
short, broad, almost truncate; right paramere of average length, just short of reaching apical
half of median lobe; eversion of internal sac dorsoapical and slightly to right ; apical sclerite
of internal sac with two horns — one fairly tapered and one other broad and blunt and more
like a serrulate field than a sclerite. The genitalia of three males were studied in detail.
Notes on synonymy. — The type specimen of lilliputicus Casey is an average male of
alabamensis. The name constrictus cannot be used because it is a junior homonym of
const rictus Say.
Geographical distribution (fig. 128). - This species is known only from Mobile County,
Alabama. I have seen 87 specimens from the following localities.
United States - ALABAMA: Mobile County: Citronelle (CAS); Grand Bay (USNM); Mobile (CAS, CNC, CU, MCZ,
NCSU, USNM); Spring Hill (CAS, USNM).
Evarthrus ovulum Chaudoir, 1 868
Figures 16, 71,94, 128
Feronia (Evarthrus) ovulum Chaudoir, 1868:52. TYPE, female, labelled as follows:
“Steropus picipes, Sturm, Georgetown”. MNHP. - LeConte, 1873:319 ( Evarthrus ). -
Schaupp, 1880:49. - Leng, 1920:57 ( Ferestria ). - Csiki, 1930:674 ( Pterostichus ).
Recognition. — The diagnostic characters are a combination of crescent-shaped frontal
grooves; sharp basal angles of the pronotum; deep short groove of the prosternal process;
form of the median lobe and sclerite of the internal sac of the male genitalia; and small body
size. Specimens of ovulum can be confused with specimens of three other small species of
Cyclotrachelus . This species is most similar to macrovulum and texensis, but in specimens of
the last two species the prosternal process is shallowly grooved. The three are also dis-
tinguished by differences in the shape of the median lobe and apical sclerite of the internal
sac (fig. 94 cf. fig. 95a-d and 95e-h). In addition these species are allopatric.
The smaller body size, glossy elytra, crescent-shaped frontal grooves and male genitalia
distinguish ovulum from alabamensis. Further, the two species are allopatric.
Description. — Body length 8.5 — 11.0 mm. Form narrow but rather typical of the
ovulum group.
Head between eyes with micro sculpture composed of highly sinuous, entwined or sparse
lines. Micro sculpture of disc of pronotum same as on head but generally effaced. Elytral
intervals with isodiametric meshes, partially effaced in males.
Head between eyes, glossy; frontal grooves (fig. 71) sharply defined, oblique, crescent-
shaped with bend produced laterally, widely separated. Penultimate article of labial palpus
with two to four setae.
Pronotum (fig. 16) glossy; sides strongly constricted posteriorly and moderately sinuate
in front of hind angles; posterior angles small, prominent and slightly obtuse; anterior trans-
verse impression incomplete, impressed laterally only; basal foveae moderately impressed.
Revision of Evarthrus
1 19
Prostemal process with deep and sharply defined longitudinal groove deepest near apex.
Anterior face of middle femur with four setae.
Elytra glossy in males and slightly duller in females; intervals not markedly convex;
striae of average depth, coarsely punctate anteriorly and impunctate posteriorly ; umbilicate
series markedly impressed.
Male genitalia (fig. 94) with median lobe strongly arcuate, angle slightly obtuse; apical
blade short and broadly rounded at apex; right paramere of average length reaching halfway
point of median lobe; eversion of internal sac dorsoapical and slightly to right; apical sclerite
of internal sac U-shaped with horn-like projections narrow, sturdy, and slightly curved. The
genitalia of three males were studied in detail.
Stylus of female ovipositor evenly rounded apically without preapical sinuations.
Notes on synonymy. — Chaudoir thought the type locality of ovulum was Georgetown,
South Carolina. Georgetown, Georgia is closer to the range of this species and it is more
likely the correct type locality.
Collecting notes. — This species has been collected in pine forests of Florida and Georgia.
Specimens have also been found under bark and caught in malt bait traps
Geographical distribution (fig. 128). — Evarthrus ovulum inhabits Florida and southern
Georgia. I have seen 29 specimens collected in the following localities.
United States - FLORIDA: Alachua County: Gainesville (FDPI). Baker County: Glen St. Mary (FDPI); Macclenny
(FDPI). Gadsden County: Quincy (FDPI). Leon County: Tallahassee (CNHM, CNC, USNM). GEORGIA: Thomas County:
Thomasville (ANSP). Toombs County: Lyons (UMMZ).
Evarthrus macrovulum new species
Figures 17, 95a— d, 128
Recognition. — The following combination of characters is diagnostic for specimens of
macrovulum : crescent-shaped frontal grooves on the head; sharp basal angles of the prono-
tum; very shallow longitudinal groove in prostemal process; glossy pronotum and elytra,
and very short right paramere of the male genitalia (fig. 95). This species, texensis, ala-
bamensis, and ovulum are remarkably similar.
The differences between macrovulum and ovulum are given in the diagnosis of the latter
species.
The frontal grooves of the head of macrovulum are distinctly crescent-shaped and oblique
but they are straight and more parallel in alabamensis. A glossy pronotum and equally
glossy elytra is characteristic of macrovulum and contrasts with the combined glossy prono-
tum and dull elytra of alabamensis . In addition macrovulum has a shallow groove in the
prostemal process while it is deep in alabamensis . Males of the species can be separated by
characteristics of the genitalia (fig. 93 cf. fig. 95a— d).
The species texensis is the only representative of Cyclotrachelus occurring to the west of
the Mississippi River. Structurally, specimens of texensis closely resemble specimens of
macrovulum, but they differ in details of the male genitalia (figs. 95a-c; cf. figs. 95e-g).
Description. - HOLOTYPE, male, labelled as follows: “Mobile, Ala XI— 1 1-39; Van Dyke
Collection; HOLOTYPE Evarthrus macrovulum R. Freitag (red label).” CAS.
Body length 10.8 mm, width 4.1 mm. Form average for group.
120
Freitag
Head between eyes and disc of pronotum with microsculpture composed of isolated
sinuous lines or effaced. Microsculpture of elytral intervals sinuous, closely entwined lines
often forming amorphic meshes, and partially effaced.
Head glossy; length 1.8 mm, width 2.3 mm; frontal grooves sharply defined, oblique,
crescent-shaped with lateral bend, and widely separated. Penultimate article of labial palpus
with two medial setae.
Pronotum glossy on disc; length 2.8 mm, width 3.4 mm; form as in fig. 17; greatest
width slightly anterior to transverse mid-line; disc moderately convex; sides strongly con-
stricted posteriorly, and moderately sinuate in front of posterior angles; posterior angles
small, prominent and slightly obtuse; anterior transverse impression incomplete, impressed
laterally only; median longitudinal impression shallow throughout; basal foveae deepest
posteriorly. Prostemal process with shallow longitudinal groove. Middle femur with four
setae on anterior face.
Elytra glossy; length 6.2 mm, width 4.1 mm; intervals rather flat; striae shallow, distinctly
but not coarsely punctate anteriorly, and impunctate posteriorly; umbilicate series deeply
impressed. First articles of middle and hind tarsi with lateral grooves.
Male genitalia as in fig. 95a— d with median lobe strongly arcuate, apical blade short and
evenly rounded at apex; right paramere short, not reaching apical half of median lobe;
eversion of internal sac dorsoapical and slightly to right; apical sclerite of internal sac with
two horns, one blunt and one sharp and twisted.
ALLOTYPE, female, labelled as follows: “Mobile Ala XII-1-39; Van Dyke Collection;
ALLOTYPE Evarthrus macrovulum R. Freitag (green label).” CAS.
Body length 1 1 .4 mm, width 4.4 mm. Form as in holotype.
Microsculpture of head between eyes and pronotal disc same as in holotype. Elytral
intervals with microsculpture less effaced than that of holotype, formed by close and
distinct closed amorphic meshes.
Head length 1.8 mm, width 2.5 mm.
Pronotum shape as in holotype; length 2.9 mm, width 3.6 mm.
Elytra slightly duller than holotype; length 6.8 mm, width 4.4 mm; sides more parallel
posteriorly than those of holotype. Stylus of ovipositor tapered slightly toward apex.
Derivation of specific name. — This species name suggests that specimens of macrovulum
are large and like ovulum in appearance. Specimens of macrovulum are not necessarily
longer but appear more robust than those of ovulum.
Variation among paratypes (91 males, 89 females, Mobile, Alabama. CAS). — Slight and
inconsequential. Total length, 8.5 — 1 1.8 mm. The genitalia of three males were examined
in detail.
Disposition of type material. — The holotype and allotype are in the collection of the
CAS. One paratype is in the UASM collection, and the others are in the collection of CAS,
and RCG.
Geographical distribution (fig. 128). — This species is known from southern Alabama and
southern Louisiana. I have seen 1 82 specimens from the following localities.
United States - ALABAMA: Baldwin County: Fairhope (CAS). Mobile County: Mobile (CAS). LOUISIANA: Saint
Tammany County: Slidell (RCG). County not determined: Hart (CAS).
Revision of Evarthrus
121
Evarthrus texensis new species
Figures 17a, 95e— h, 128
The following combination of characters is diagnostic for specimens of texensis ; crescent-
shaped frontal grooves on the head; very shallow longitudinal groove in prosternal process;
broadly rounded apex of the median lobe of the male, length of right paramere, and shape
of the apical sclerite of the internal sac.
Recognition. — The differences between the very similar species macrovulum and texensis
are described in the recognition section of the former species.
Description. - HOLOTYPE, male, labelled as follows: “U. S. A., TEXAS, Tyler Co. 12
mi. W. Kirbyville, Rte. 1013 XII— 6— 68 G. E. Ball; HOLOTYPE Evarthrus texensis R.
Freitag (red label).” MCZ.
Body length 7.5 mm, width 3.3 mm. Form average for group.
Microsculpture of head between eyes and disc of pronotum almost effaced; microsculp-
ture of elytra composed of isodiametric meshes.
Head glossy; length 1.0 mm, width 1.6 mm; frontal grooves sharply defined, oblique,
crescent-shaped, and widely separated. Penultimate article of labial palpus with two medial
setae.
Pronotum glossy on disc; length 1.8 mm, width 2.6 mm; form as in fig. 17a; greatest
width anterior to transverse mid-line; disc moderately convex, sides strongly constricted
posteriorly, and moderately sinuate in front of posterior angles; posterior angles small,
prominent and slightly obtuse; anterior transverse impression incomplete, impressed laterally
only; median longitudinal impression moderately impressed; basal foveae deepest posteri-
orly; prosternal process with shallow longitudinal groove. Middle femur with four setae on
anterior face. First article of middle and hind tarsi with faint lateral grooves.
Elytra slightly duller than head and prothorax; length 4.7 mm, width 3.3 mm; intervals
slightly convex; striae moderately impressed, distinctly punctate anteriorly, impunctate
posteriorly; umbilicate series deeply impressed.
Male genitalia as in fig. 95e— g with median lobe strongly arcuate, apical blade short and
broadly rounded at apex; right paramere almost reaching apical half of median lobe; eversion
of internal sac dorsoapical and to right; apical sclerite of internal sac with two horns, one
very broad and blunt and one sharp and twisted (fig. 95h).
ALLOTYPE, female, labelled as follows: “USA., TEXAS, Tyler Co. 12 mi. W. Kirbyville
Rte. 1013 XII— 6— 1968 G. E. Ball; ALLOTYPE Evarthrus texensis R. Freitag (green label).”
MCZ.
Body length 8.4 mm, width 3.6 mm. Form as in holotype.
Microsculpture of head between eyes, pronotal disc, and intervals of elytra same as in
holotype.
Head length 1.0 mm, width 1.8 mm.
Pronotum shape as in holotype; length 2.4 mm, width 2.9 mm.
Elytra length 5.0 mm, width 3.6 mm. Stylus of ovipositor tapered slightly toward apex.
Derivation of specific name. — This species has been named texensis since it is known
only from Texas.
Variation among paratypes (four males, one female, Orange Co., Jasper Co., Tyler Co.,
Texas). — Total length, 7.0 — 9.2 mm. The genitalia of four males were examined.
122
Freitag
Disposition of type material. — The holotype and allotype are in the MCZ collection and
the paratypes are in the collections of CAS, UASM, and USNM.
Geographical distribution (fig. 128). - This species is known from eastern Texas. I have
seen seven specimens from the following localities.
United States - TEXAS: Orange County nr. Lakeview. Jasper County: Rte. 63, 11 mi. N. Jasper. Tyler County: Rte.
1013, 12 mi. W. Kirbyville.
The faber Group
Characteristics. — Small to medium size beetles; pronotum circular to subcordate, with
basal angles recessed and broadly obtuse; longitudinal groove of prostemal process long,
deep and sharply defined; dark apical sclerite present in internal sac of median lobe of male
genitalia.
The species included in this group are faber , levifaber and parafaber. All of these are
found only on the Coastal Plain of southeastern United States.
Evarthrus parafaber new species
Figures 18, 96, 129
Recognition. — This species is distinguished from its relatives by the following combina-
tion of characters: frontal grooves of head fairly straight, slightly oblique in relation to one
another; pronotum with sides not produced, basal angles almost obsolete; long deep, and
sharply defined longitudinal, medial groove in prostemal process; and form of male genitalia.
This species can be confused with most members of the ovulum group but is distinguished
by the obsolete posterior angles of the pronotum. There is no overlap in the geographical
ranges of the closely related species parafaber, faber and levifaber. Specimens of faber are
distinguished from those of parafaber by their larger size, four setae on the penultimate
article of the labial palpus, shape of the pronotum, and details of the male genitalia. The
smaller species levifaber also resembles parafaber in habitus, but it has a pronotum with
markedly produced sides that contrast with the more parallel sides of that of parafaber.
Furthermore there are striking differences in the structures of the male genitalia of these
two species (fig. 96 cf. fig. 97).
Description. - HOLOTYPE, male, labelled as follows: “Mobile, Ala XI -4-39; Van Dyke
Collection; HOLOTYPE Evarthrus parafaber R. Freitag (red label).” CAS.
Body length 9.2 mm, width 3.8 mm. Form average for faber group.
Microsculpture of head between eyes with lines dense, highly sinuous or closed and
forming bead-like meshes. Disc of pronotum with impressions of microsculpture highly
sinuous or closed meshes slightly stretched transversely. Microsculpture of elytral intervals
composed of isodiametric meshes.
Head semi-glossy; length 1.1 mm, width 2.1 mm; frontal grooves sharply defined, slightly
curved but not crescent-shaped, widely separated. Penultimate article of labial palpus with
two medial setae.
Pronotum with disc semi-glossy; length 2.7 mm, width 3.1 mm; form as in fig. 18;
greatest width slightly anterior to transverse midline; disc moderately convex; sides slightly
prominent laterally, moderately constricted anteriorly and strongly constricted posteriorly,
Revision of Evarthrus
123
slightly sinuate in front of posterior angles; posterior angles almost obsolete, widely obtuse
and recessed; anterior transverse impression incomplete, impressed laterally only; median
longitudinal impression distinctly deeper at both ends; basal foveae deep posteriorly, shallow
and elongate anteriorly. Prosternal process with long, deep, sharply defined medial, longi-
tudinal groove. Middle femur with four setae on anterior face.
Elytra semi-glossy; length 5.4 mm, width 3.8 mm; markedly sinuate apically; intervals
slightly convex; striae moderately impressed and coarsely punctate in anterior half, obso-
letely punctate apically.
Male genitalia as in fig. 96 with angle of median lobe almost right, apical blade short and
slightly produced medially and deflected ventrally; right paramere not reaching apical half
of median lobe; eversion of internal sac dorsoapical and slightly to right; apical sclerite of
internal sac with two horns, one blunt, one sharp and twisted.
ALLOTYPE, female, labelled as follows: “Mobile Ala XI— 4— 39; Van Dyke Collection;
ALLOTYPE Evarthrus parafaber R. Freitag (green label).” CAS.
Body length 9.5 mm, width 4.1 mm. Form as in holotype.
Microsculpture of head between eyes with distinct closed meshes. Microsculpture of disc
of pronotum and intervals of elytra same as in holotype.
Head, length 1 mm, width 2.1 mm.
Pronotum shape as in holotype; length 2.7 mm, width 3.7 mm.
Elytra shape, intervals and striae same as in holotype; length 5.7 mm, width 4.1 mm.
Stylus of ovipositor short and broad, not sinuate apically.
Derivation of specific name. — This species is closely related to faber, which is what the
name parafaber suggests.
Variation among paratypes (29 males, 26 females, Mobile, Ala. CAS). — Total length, 9.8
— 12.8 mm. Variation in color is moderate in the elytra ranging from light rufopiceous to
deep piceous. Other parts of the body vary slightly from those of the type specimens. The
genitalia of two males were carefully examined.
Disposition of type material. — The holotype, allotype, and 53 paratypes are in the collec-
tion of the CAS. Two paratypes are in the UASM collection.
Geographical distribution (fig. 129). — This species is known only from the type locality.
I have seen 57 specimens.
United States - ALABAMA: Mobile County: Mobile (CAS, UASM).
Evarthrus levifaber new species
Figures 19, 97, 129
Recognition. — Specimens of levifaber are characterized by a combination of the follow-
ing features: penultimate article of labial palpus bisetose; straight frontal grooves; slightly
cordiform pronotum with produced sides; right paramere of male genitalia with distinct
elbow; and apical sclerite of internal sac crescent-shaped. Characters that distinguish levi-
faber and parafaber are presented in the recognition section of the latter species. The species
levifaber and faber are allopatric. The four setae on the penultimate article of the labial
palpus, inwardly curved frontal grooves of the head, and male genitalia of faber distinguish
it from levifaber. Furthermore the pronotum of faber is circular but in levifaber this sclerite
is more cordiform.
124
Freitag
Description. - HOLOTYPE, female, labelled as follows: “Camden S. C.; Roland Hayward
Coll.; HOLOTYPE Evarthrus levifaber R. Freitag (red label).” MCZ.
Body length 10.1 mm, width 4.2 mm. Form robust. Head with microsculpture composed
of highly sinuous entwined lines occasionally forming amorphic meshes. Disc of pronotum
and elytral intervals with microsculpture same as head except amorphic meshes raised.
Head glossy; length 1.2 mm, width 2.5 mm; frontal grooves sharply defined and straight,
slightly oblique toward one another. Penultimate article of labial palpus with two medial
setae.
Pronotum with semi glossy disc; length 2.7 mm, width 3.5 mm; form as in Fig. 19; disc
moderately convex; sides broadly rounded and prominent, slightly constricted anteriorly
and strongly constricted posteriorly, obsoletely sinuate in front of posterior angles; posterior
angles not prominent and broadly rounded; anterior transverse impression incomplete,
impressed laterally only; median longitudinal impression slightly deeper at posterior end;
basal fovea deep posteriorly, moderately deep and short anteriorly. Prostemal process with
long, deep, sharply defined longitudinal groove. Middle femur with four setae on anterior
face.
Elytra semi-glossy; length 6.2 mm, width 4.2 mm; margin at shoulder broad; apex dis-
tinctly sinuate; intervals slightly convex; striae moderately impressed and distinctly punctate
anteriorly, impunctate posteriorly.
Stylus of ovipositor short and broad, not sinuate apically.
ALLOTYPE, male, labelled as follows: “Ga.; Horn Coll H536; ALLOTYPE Evarthrus
levifaber R. Freitag (green label)”. ANSP.
Body length 9.1 mm, width 3.7 mm. Form same as in holotype.
Microsculpture of head between eyes and disc of pronotum same as in holotype. Micro-
sculpture of elytra with sinuous lines that often form longitudinally stretched meshes.
Body mainly rufopiceous, antennae, palpi, legs and epipleura light rufopiceous.
Head glossy; length 1.1 mm, width 2.2 mm.
Pronotum shape same as in holotype; length 2.5 mm, width 3.0 mm.
Elytra glossy, appearance velvet; intervals more convex and striae more impressed than
in holotype; length 5.5 mm, width 3.7 mm.
Genitalia (fig. 97) with strongly arcuate median lobe, particularly apical half, apical blade
elongate, narrow, and evenly rounded at apex; right paramere with produced elbow, tapered
apically and extended to apical half of median lobe; eversion of internal sac apicodorsal and
to right; apical sclerite of internal sac dark and C-shaped.
Derivation of specific name. — Specimens of this species appear to be lighter in weight
than those of the closely related species faber, which is implied in the name levifaber.
Variation among paratypes (three males, one female, Georgia, South Carolina, North
Carolina, ANSP, MCZ and UASM). - Total length, 11.1 - 13 mm. Except for one teneral
male the coloration of the paratypes is approximately the same as that of the holotype and
allotype, and similar in all other respects. The genitalia of one male was examined in detail.
Disposition of type material. — The holotype and allotype are in the collection of the
MCZ and ANSP respectively. The paratypes are in the collection of MCZ, ANSP, and UASM.
Geographical Distribution (fig. 129). — I have seen six specimens from the following
localities.
United States - GEORGIA: (ANSP, UASM). NORTH CAROLINA: (MCZ). SOUTH CAROLINA: Kershaw County:
Camden (MCZ).
Revision of Evarthrus
125
Evarthrus faber Germar, 1 824
Figures 20, 68, 98, 129
Molops faber Germar, 1824:23. Type not seen. TYPE LOCALITY, “America septentrionali
(Kentucky),” [this locality is probably incorrect]. — LeConte, 1848:353 ( Steropus ). -
LeConte, 1852:230 {Evarthrus). - LeConte, 1863a:8. - LeConte, 1873:319. - Schaupp,
1880:49. - Casey, 1918:349 {Cyclotrachelus). - Leng, 1920:56. - Csiki, 1930:671
( Pterostichus ).
Feronia tenebricosa Dejean, 1828:301. Type seen by C. H. Lindroth (1955). TYPE LOCAL-
ITY, “l’Amerique septentrionale.” MHNP. - Chaudoir, 1838:30 {Cephalotes). - Le-
Conte, 1848:353 {Steropus). - LeConte, 1868a:8 {Evarthrus). - LeConte, 1873:319. -
Casey, 1918:349 {Cyclotrachelus). — Leng, 1920:56. - Csiki, 1930:671 {Pterostichus).
Feronia spoliatus\ LeConte, 1848:353 (not Newman). - LeConte, 1852:230. — LeConte,
1863a:8.
Cyclotrachelus roticollis Casey, 1918:349. HOLOTYPE, male, labelled as follows: “Fla;
CASEY bequest 1925; TYPE USNM 47108; Cyclotrachelus roticollis Csy.” USNM TYPE
LOCALITY, Dunedin, Florida. PARATYPES, two males, labelled as follows: “Dunedin;
Fla. W. S. B. coll. 3-23 1913 and 17-7 2-1 5 ; roticollis - 2 and - 3 PARATYPE USNM
47108.” NEW SYNONYMY. - Casey, 1924:78 {Cyclotrachelus). - Leng, 1920:56. -
Csiki, 1930:671 {Pterostichus).
Cyclotrachelus fallaciosus Casey, 1924:77. HOLOTYPE, male, labelled as follows: “Dune-
din, Fla. W. S. B. coll. 4-5-1915; TYPE USNM 47109; fallaciosus Csy,” USNM NEW
SYNONYMY. - Leng and Mutchler, 1927:10 {Cyclotrachelus). - Csiki, 1930:671
{Pterostichus).
Recognition. — The following combination of characters is diagnostic of the species faber:
penultimate article of labial palpus with four setae, frontal grooves on head crescent-shaped
with the convexity directed medially; sides of pronotum strongly arcuate; long, deep,
sharply defined longitudinal groove in prostemal process; cup-like scales on the ventral
side of the front tarsi of the males; and details of the male genitalia.
The differences among the closely related species faber , levifaber, and parafaber are
described in the recognition sections of the last two species.
Description. — Body length 8.5 — 11.1 mm. Form robust and typical of the faber group.
Head between eyes, disc of pronotum, and intervals of elytra with lines of microsculpture
distinctly impressed, very dense, and sinuous, forming raised amorphic meshes.
Head glossy, dull or slightly glossy; frontal grooves moderately impressed, crescent-
shaped with convexity directed medially, and moderately separated. Penultimate article of
labial palpus with two medial and two apical setae (fig. 68).
Pronotum dull or slightly glossy; form of sides circular in outline, as in fig. 20; disc
markedly convex; sides moderately constricted anteriorly and strongly constricted posteri-
orly, slightly sinuate in front of posterior angles; posterior angles broadly obtuse; anterior
transverse impression incomplete, impressed laterally only; basal foveae deep posteriorly,
and often anterior end very shallowly extended onto anterior half of. disc. Prostemal process
with long, deep, sharply defined longitudinal groove. Middle femur with four or five setae
on anterior face. Males with even rows of cup-like scales on ventral side of front tarsi.
126
Freitag
Elytra dull or slightly glossy; margin near shoulder slightly narrow; sinuate apically;
intervals slightly convex; striae deeply impressed and distinctly punctate anteriorly and on
disc, impunctate posteriorly.
Male genitalia (fig. 98) with moderately arcuate median lobe, apical portion more acute,
apical blade resembling a two-edged sword with produced medial apex; right paramere short,
not extended to apical half of median lobe, shape typical of Cyclotrachelus with recessed
elbow and not strongly tapered apically; eversion of the internal sac apicodorsal and to right;
internal sac with light basal serrulate field and darker apical serrulate field, apical sclerite
dark and C-shaped. The male genitalia of three specimens were examined in detail.
Stylus of female ovipositor short, broadly rounded apically.
Variation. — The number of setae on the front face of the middle femur varies from four
to five, but there is no geographical pattern to the variation.
Notes on synonymy. — In the MNHP collection there are six specimens of the species
determined as faber Germar, the first of which bears the label tenebricosa m. which was
written by Dejean. The specimen was probably given to him by Joseph E. LeConte with
whom Dejean traded specimens.
The type specimens of roticollis Casey and fallaciosus Casey are average specimens of
faber.
Collecting notes. — Specimens of this species have been found in leaf litter (label data).
The gut of one specimen contained a mixture of sand and fungus zygotes, which were
identified by Dr. L. L. Kennedy.
Geographical distribution (fig. 129). — This species inhabits Florida and southern Georgia.
The New York and Ohio records are certainly incorrect. I have seen 132 specimens collected
in the following localities.
United States - FLORIDA: Alachua County: Archer (FDPI); Gainesville (UMMZ): High Springs (UMMZ): R.-24-E
T-10-S (UMMZ). Baker County: Glen St. Mary (FDPI). Brevard County: Melbourne (USNM). Calhoun County: near
Clarksville (CNC). Charlotte County: Punta Gorda (CAS, CNHM). DeSoto County: Arcadia (GEB, UMMZ); Fort Ogden
(CNC). Dixie County: Cross City (UMMZ); Shamrock (CAS). Duval County: Jacksonville (USNM). Gadsden County:
Quincy (FDPI). Hendry County: LaBelle (CU). Hernando County: Brooksville (CAS, UMMZ). Lee County: Fort Myers
(CNC). Leon County: (CU); Tallahassee (CNC, UMMZ). Liberty County: Camp Torreya (UMMZ). Manatee County:
Bradenton (CAS). Marion County: (ANSP); Ocala (CNC); Ocala National Forest (UMMZ). Monroe County: Big Pine Key
(UMMZ). Okaloosa County: Delaco (UMMZ). Orange County: Winter Park (MCZ). Osceola County: Deer Park (MCZ);
Kissimmee (AMNH). Pasco County: Elfers (CNC). Pinellas County: Dunedin (AMNH, CAS): St. Petersburg (AMNH). Polk
County: Lakeland (USNM). Sarasota County: Sarasota (USNM). Seminole County: Sanford (MCZ). Walton County:
DeFuniak Springs (UMMZ). County not determined: Fringers (USNM); Iuka Island (USNM); North Smyrna (CAS); 15
miles south of Wadky (CNC). GEORGIA: Camden County: Kingsland (UMMZ); St. Mary’s (MCZ). Decatur County:
Faceville (UMMZ). NEW YORK: Westchester County: Peekskill (CU). OHIO: (CMNH).
The Subgenus Evarthrus LeConte
Evarthrus LeConte, 1852:225, TYPE SPECIES — Evarthrus sigillatus Say, 1823a (designated
by Casey, 1918:322).
Anaferonia Casey, 1918:341. TYPE SPECIES — Evarthrus constrictus Say, 1823b (desig-
nated by Casey, 1918:321).
Megasteropus Casey, 1918:350. TYPE SPECIES - Megasteropus gigas Casey, 1918 (desig-
nated by Casey, 1918:322).
Revision of Emrthrus
127
Eumolops Casey, 1918:351. TYPE SPECIES - Eumolops sexualis Casey, 1918 (designated
by Casey, 1918:322).
Evarthrinus Casey, 1918:357. TYPE SPECIES — Evarthrus deceptus Casey, 1918 (here
designated).
Evarthrops Casey, 1920:194. TYPE SPECIES — Evarthrus furtivus LeConte, 1852 (here
designated).
Characteristics. - Penultimate article of labial palpus with three (rarely) or five to seven
setae; pro no turn with sides parallel or constricted posteriorly, posterior lateral foveae
bistriate, posterior lateral setae usually beside bead (figs. 21—61), but in gravesi on bead;
middle femur with 4—1 1 setae on anterior face; last tarsal article with setae on ventral side;
eversion of internal sac of median lobe of male genitalia right.
Notes on synonymy. — Casey established the above genera and subgenus Evarthrops on
characters which are common throughout the subgenus Evarthrus. The description of
Anaferonia provided by Casey can be applied to most species groups in the subgenus
Evarthrus. He established Megasteropus on features such as size of head and impunctate
striae of elytra. I do not accept these as generic or subgeneric characters. He separated
Eumolops from Evarthrus mainly because of differences in the form of the last article of the
maxillary palpus, a characteristic which varies intraspecifically throughout Evarthrus. Casey
believed that species with three punctures on the third interval of the elytron constituted a
separate genus which he named Evarthrinus. This characteristic is present in a number of
unrelated species in the subgenus Evarthrus.
The following species groups compose the subgenus Evarthrus: the incisus group, the
blatchleyi group, the sigillatus group, the seximpressus group, the hypheripiformis group,
the sodalis group, the substriatus group, the torvus group, and the gigas group.
The incisus Group
Characteristics. — Penultimate article of labial palpus with five setae. Pronotum cordiform
in outline; anterior transverse impression usually absent medially, complete in a few speci-
mens. Prosternal process with longitudinal groove shallow. Middle femur with four setae on
anterior face. Median lobe of male genitalia with hump medially on ventral surface; pig-
mented apical sclerite in internal sac; right paramere very short.
The species incisus and whitcombi are included in this group, which is represented on the
Great Plains from Arkansas and Oklahoma north to South Dakota.
Evarthrus incisus LeConte, 1 848
Figures 21, 99, 130
Feronia (Molops) incisa LeConte, 1848:345. LECTOTYPE (here selected) a male, labelled
as follows: “green disc; Type 5620; E. incisus Lee.” MCZ. TYPE LOCALITY, Missouri
Territory. - LeConte, 1852:232 {Evarthrus). - LeConte, 1863a:8. — LeConte, 1873:
319. — Schaupp, 1880:49. — Casey, 1918:348 ( Anaferonia ). — Leng, 1920:56. - Csiki,
1930:671 ( Pterostichus ).
Feronia (Molops) lixa LeConte, 1848:346. LECTOTYPE (here selected) a female, labelled
as follows: “green disc; Type 5622; E. lixa LeC; abdominalis 3”. MCZ. TYPE LOCAL-
ITY, near Long’s Peak. — LeConte, 1863a:8 (Evarthrus). — LeConte, 1873:319. —
Schaupp, 1880:49.
128
Freitag
Feronia (Molops) abdominalis LeConte, 1848:347. LECTOTYPE (here selected) a male,
labelled as follows: “green disc; Type 5621 ; E. abdominalis Lee.” MCZ. TYPE LOCAL-
ITY, near Long’s Peak. — LeConte, 1852:232 ( Evarthrus ). — LeConte, 1863a:8. -
LeConte, 1873:319. - Schaupp, 1880:49. - Casey, 1918:347 ( Anaferonia ). — Leng,
1920:56. - Csiki, 1930:671 (. Pterostichus ).
Anaferonia distincta Casey, 1918:342. HOLOTYPE, male, labelled as follows: “la; CASEY
bequest 1925; TYPE USNM 47103; distincta Csy.” USNM. TYPE LOCALITY, Iowa.
NEW SYNONYMY. - Leng, 1920:56 {Anaferonia). - Csiki, 1930:671 (Pterostichus ).
Anaferonia iowana Casey, 1918:347. HOLOTYPE, male, labelled as follows: “la; CASEY
bequest; TYPE USNM 47107; iowana Csy.” USNM. TYPE LOCALITY, Iowa. NEW
SYNONYMY. - Leng, 1920:56 (Anaferonia). - Csiki, 1930:671 ( Pterostichus ).
Anaferonia fausta Casey, 1918:348. HOLOTYPE, male, labelled as follows: “Penn; CASEY
bequest 1925; TYPE USNM 47104; fausta Csy.” USNM; PARATYPE, male, labelled as
follows: “Penn; CASEY bequest 1925; fausta -2; PARATYPE USNM 47104. USNM.
TYPE LOCALITY, Pennsylvania. NEW SYNONYMY. - Leng, 1920:56 (Anaferonia). -
Csiki, 1930:671 (Pterostichus).
Recognition. — Specimens of this species are easily confused with specimens of whit-
combi, but they are distinguished by their smaller size and by differences in male genitalia
(fig. 99 cf. fig. 100). In Arkansas the micropunctures in the elytral intervals are distinct in
specimens of incisus but indistinct in specimens of whitcombi.
Description. - Body length 9.0 - 12.3 mm. Form robust anteriorly, average for incisus
group.
Microsculpture of head between eyes and disc of pronotum effaced. Intervals of elytra
with isodiametric meshes forming microsculpture, occasionally almost effaced; integument
of dorsum glossy.
Head with frontal grooves distinctly but not deeply impressed, straight or slightly curved,
usually oblique but occasionally parallel toward one another, not widely separated.
Pronotum form as in fig. 21 ; disc moderately convex; sides slightly constricted anteriorly,
strongly constricted posteriorly, short and slightly sinuate in front of posterior angles;
posterior angles small, produced, slightly obtuse; anterior transverse impression usually
incomplete, in a few specimens complete with medial portion obsolete or interrupted; basal
lateral foveae with sides usually continuous near base, separated in a few specimens.
Elytra obsoletely sinuate apically; intervals of average convexity or flattened, striae
moderately impressed, indistinctly or obsoletely punctate in anterior half, obsoletely punc-
tate apically.
Male genitalia (fig. 99) with median lobe slightly arcuate, angle broadly obtuse, low
median ventral hump present; apical blade elongate with apical lateral edges strongly de-
flected dorsally, apex evenly rounded; right paramere short, broadly rounded at apex, not
extended to apical half of median lobe; internal sac with serrulate field apically, apical
sclerite dark elongate and slightly curved basally. The genitalia of five males were studied in
detail.
Stylus of female ovipositor narrow, gradually tapered apically.
Revision of Evarthrus
129
Geographical variation. — Individuals of incisus have red or black legs. Red legs are
common in Nebraska and appear occasionally throughout the rest of the species range.
Notes on synonymy. — The lectotypes of abdominalis and lixa LeConte are average speci-
mens of incisus. The type specimen of distincta Casey has a basal fovea of the pronotum
with the sides continuous posteriorly, and is average for this species in most other structures.
The type specimens of fausta and iowana Casey are normal incisus specimens.
Collecting notes. - D. L. Larson and I collected specimens of incisus in leaf litter of
deciduous forest near Morrilton, Arkansas. Some of the specimens were taken at the soil
surface beneath moist leaf litter and some were in the litter itself.
Geographical distribution (fig. 130). — This species inhabits the central states from Kansas
and Arkansas north to Illinois and South Dakota. The Pennsylvania record is probably
incorrect. I have seen 222 specimens collected in the following localities.
United States - ARKANSAS: Carroll County: Eureka Springs (INHS). Conway County: six miles south of Morrilton
(DL, RF). Johnson County: ten miles east of Ozark (DL, RF). Madison County: 45 miles east of Fayetteville (RF).
Marion County: (USNM); Buffalo River State Park (CU). Pope County: (UA). Washington County: Cove Creek (CU, DL,
RF); Devil’s Den State Park (RTB); Fayetteville (UA). ILLINOIS: Knox County: Galesburg (INHS). Piatt County: Robert
Allerton Park, Monticello (RTB). IOWA: Johnson County: Iowa City (MCZ, UASM, USNM). KANSAS: Dickinson County:
(CNHM). Wabaunsee County: McFarland (USNM). MISSOURI: Carter County: Van Buren (UMMZ). Crawford County:
Onandaga Cave (UMMZ). NEBRASKA: Douglas County: Omaha (CAS). Fillmore County: (USNM). Red Willow County:
McCook (USNM). OKLAHOMA: Comanche County. Wichita National Forest (CAS, UMMZ). Oklahoma County: (CAS).
PENNSYLVANIA: Allegheny County: (CAS, CNC, USNM). SOUTH DAKOTA: Hutchinson County: Menno (VMK).
Yankton County: Yankton (VMK).
Evarthrus whitcombi new species
Figures 22, 100, 130
Recognition. — Several characteristics, previously described in connection with the recog-
nition of incisus, distinguish specimens of whitcombi from those of incisus. Both incisus and
whitcombi can be mistaken for specimens of the somewhat similar species substriatus or
iowensis. Individuals of substriatus are distinguished by a large elytral plica and distinct
dorsolateral knob on the last abdominal segement that fits onto the plica (fig. 77). Speci-
mens of iowensis are characterized by having five or six setae on the anterior face of the
middle femur, which contrasts with the four setae on the same structure of incisus and
whitcombi.
Description. — HOLOTYPE, male, labelled as follows: “Hot Springs, Ark. X— 1—39; Van
Dyke Collection ; HOLOTYPE Evarthrus whitcombi R. Freitag (red label).” CAS.
Body length 13.4 mm, width 5.7 mm. Larger and more robust than specimens of incisus
LeC.
Head between eyes and disc of pronotum with microsculpture composed of highly
sinuous lines, entwined, but rarely forming meshes. Microsculpture of elytral intervals with
isodiametric meshes. Integument of dorsum slightly glossy.
Head length 1.5 mm, width 3.3 mm; frontal grooves distinctly but broadly impressed,
straight, parallel to one another.
Pronotum length 3.8 mm, width 4.6 mm; shape cordiform in outline as in fig. 22; disc
moderately convex, somewhat flattened medially; sides constricted slightly anteriorly,
strongly posteriorly, briefly sinuate in front of posterior angle; posterior angles small,
130
Freitag
produced, slightly obtuse; anterior transverse impression absent medially; basal lateral foveae
with sides not continuous near base. First articles of middle and hind tarsi with lateral
grooves.
Elytra 8.1 mm in length, width 5.7 mm; sides parallel, slightly sinuate apically ; intervals
of low convexity almost flat; striae moderately impressed anteriorly, punctate anteriorly,
indistinctly or obsoletely impressed posteriorly.
Male genitalia (fig. 100) with median lobe moderately arcuate, and with marked median
ventral hump; apical blade short and deflected to right, edges of apex not deflected dorsally;
right paramere short, apical half evenly tapered to apex; internal sac serrulate apically,
apical sclerite very dark with broad tooth apically and hook basally.
ALLOTYPE, female, labelled as follows: “Hot Springs Ark. X— 1— 39; Van Dyke Collec-
tion; ALLOTYPE Evarthrus whitcombi R. Freitag (green label).” CAS.
Body length 14.0 mm, width 5.7 mm. Form same as in holotype.
Head between eyes and disc of pronotum with highly sinuous dense, entwined lines
comprising microsculpture. Intervals of elytra with isodiametric meshes forming micro-
sculpture.
Head dull; length 1.7 mm, width 3.5 mm.
Pronotum dull, form same as in holotype; length 3.7 mm, width 4.8 mm.
Elytra dull; intervals somewhat flattened ; striae moderately impressed, distinctly punctate
anteriorly, indistinctly or obsoletely punctate posteriorly; length 8.7 mm, width 5.7 mm.
Stylus of ovipositor narrow, gradually tapered apically.
Derivation of specific name. — This species is named in honour of Dr. W. H. Whitcomb,
formerly Professor of Entomology, University of Arkansas, who has made important con-
tributions in the field of terrestrial arthropod biology.
Variation among paratypes (five males, eight females, Ark., Okla.). - Body length 11.4 —
15.4 mm. The genitalia of two males were examined.
Disposition of type material. — The holotype and allotype are in the collection of the
CAS. The paratypes are in the collection of AMNH, CAS, CNHM, INHS.
Geographical distribution (fig. 130). — This species inhabits eastern Oklahoma and south-
ern Arkansas. I have seen 1 5 specimens from the following localities.
United States - ARKANSAS: Garland County: Hot Springs (CAS, INHS, UASM). Logan County: Mount Magazine
(CNHM). County not determined: Southwest (AMNH). OKLAHOMA: LeFlore County: Page (UMMZ). McCurtain County:
Beavers Bend State Park (UMMZ).
The blatchleyi Group
Characteristics. — Penultimate article of labial palpus with five setae (fig. 69), pronotum
quadrate with obtuse basal angles; prosternal process with deep, medial, longitudinal groove;
middle femur with four setae on anterior face; male genitalia with median lobe slightly
arcuate; right paramere short and broad ; internal sac very lightly sclerotized or with serrulate
field apically.
The group is composed of the species blatchleyi and floridensis whose collective range
includes Florida, and southern and eastern Georgia.
Revision of Evarthrus
131
Evarthrus hlatchleyi Casey, 1918
Figures 23,69,101,131
Evarthrus blatchleyi Casey, 1918:360. HOLOTYPE, male, labelled as follows: “Dunedin
Fla. W. S. B. coll. 3-22-18; CASEY bequest 1925; TYPE USNM 471 22; blatchleyi Csy.”
USNM, PARATYPES, two females, labelled as follows: “Dunedin Fla. W. S. B. coll. 3 —
18-16 and 3-14-16; CASEY BEQUEST 1925; blatchleyi -2 and -3 PARATYPE
USNM 47122.” USNM. - Leng, 1920:57 (Evarthrus). - Csiki, 1930:673 (Pterostichus).
Evarthrus americanus ; LeConte, 1852:228 (not Dejean). — LeConte, 1863a:8. — LeConte,
1873:318. - Leng, 1915:577. - Leng, 1920:57.
Recognition. — The following combination of characteristics is diagnostic of this species:
clearly impressed basal foveae of the pronotum, width of lateral bead of pronotum even
throughout length, elongate apical blade of median lobe of male genitalia, and relatively
large body size.
Specimens of blatchleyi are normally larger than specimens of the similar species floriden-
sis. They are further distinguished by differences in the male genitalia (fig. 101 cf. fig. 102).
Description. — Body length 14.8 — 17.6 mm. Form broad with parallel sides.
Head between eyes and disc of pronotum with microsculpture composed of extremely
tiny, densely distributed, amorphic meshes. Microsculpture of elytral intervals forming
raised, bead-like isodiametric meshes. Micropunctures present on head between eyes. Integ-
ument of dorsum slightly glossy.
Head with frontal grooves deep, sharply defined, straight or slightly curved with con-
vexity directed medially, parallel to one another.
Pronotum somewhat quadrate in outline as in fig. 23; disc slightly convex anteriorly, flat-
ter posteriorly, sides constricted moderately anteriorly and slightly posteriorly, sinuation in
front of basal angles obsolete or absent; posterior angles not produced but not broadly
rounded, slightly obtuse; anterior transverse impression complete and distinctly impressed;
basal lateral foveae with sides not continuous near base, inner side with extension from base
toward middle longitudinal line; width of lateral bead even throughout.
Elytra slightly sinuate apically; intervals flat or slightly raised, striae moderately or shal-
lowly impressed, punctate anteriorly, impunctate posteriorly.
Male genitalia (fig. 101) with median lobe slightly arcuate, angle broadly obtuse; apical
blade elongate and narrow, deflected dorsally and to left, apex evenly rounded ; right para-
mere short and broad, not extended to apical half of median lobe; internal sac serrulate
apically, apical sclerite light amorphic. The genitalia of six males were examined.
Stylus of female ovipositor slightly tapered apically, broadly rounded at apex.
Collecting notes. — This species is found in open disturbed places. G. E. Ball collected
specimens with a pitfall trap in an orange grove near Oneco, Florida.
Geographical distribution (fig. 131). — This species ranges from southwestern Florida to
southeastern Georgia. I have seen 88 specimens from the following localities.
United States - FLORIDA: Alachua County: Gainesville (FDPI), (UMMZ); Newnan’s Lake (UMMZ); Route 18 east
(FDPI). DeSoto County: Arcadia (UMMZ). Duval County: Jacksonville (AMNH, CAS, CNHM, MCZ). Highlands County:
Hammock State Park (GEB). Hillsborough County: Tampa (ANSP). Lee County: Fort Myers (UP). Manatee County:
Oneco (GEB). Marion County: Ocala National Forest (UMMZ). Orange County: Orlando (GEB). Osceola County:
Kissimmee (AMNH). Pasco County: Elfers (CNC). Pinellas County: Dunedin (UP). Putnam County: Welaka (UMMZ).
Suwannee County: Wellborn (UMMZ). GEORGIA: Bryan County: Lanier (UMMZ). Camden County: Kingsland (UMMZ).
Charlton County: Billy’s Island, Okefenokee Swamp (CU). Ware County: Waycross (UMMZ).
132
Freitag
Evarthrus floridensis new species
Figures 24, 102, 131
Recognition. — The combination of the flattened area between the basal fovea and margin
of the pronotum, and unique shape of the apical blade of the median lobe of the male (fig.
102) sets this species apart from the closely similar species blatchleyi. Specimens of another
species, sinus, also resemble specimens of floridensis but these two groups are allopatric and
possess different male genitalia (fig. 102 cf. fig. 104) among other distinguishing features.
Description. — HOLOTYPE, male, labelled as follows: “Winter Park. 2-15—28 Fla.; John
George Gehring Collection; HOLOTYPE Evarthrus floridensis R. Freitag (red label); loan
from MCZ.” MCZ.
Body length 13.1 mm, width 5.3 mm, sides parallel, not robust.
Head between eyes and disc of pronotum with microsculpture formed of highly sinuous,
densely distributed lines, occasionally forming amorphic meshes. Elytral intervals with
amorphic or isodiametric meshes composing microsculpture. Integument of dorsum slightly
glossy.
Head length 1.7 mm, width 3.6 mm; frontal grooves deep and sharply defined, slightly
curved with convexity directed medially.
Pronotum length 3.7 mm, width 4.6 mm; form quadrate in outline as in fig. 24; disc quite
convex; sides not prominent, slightly constricted anteriorly and posteriorly, not sinuate in
front of posterior angles; posterior angles not prominent, slightly obtuse and broadly
rounded; anterior transverse impression complete and clearly impressed ; basal lateral foveae
not continuous posteriorly, inner groove with extension from base toward median longi-
tudinal impression; lateral bead wider near basal foveae, and area between bead and foveae
flat.
Elytra length 7.7 mm, width 5.3 mm, sides parallel, slightly sinuate apically; intervals
almost flat; striae distinctly but not deeply impressed, indistinctly punctate anteriorly,
impunctate posteriorly.
Male genitalia (fig. 102) with median lobe slightly arcuate, angle broadly obtuse; apical
blade with ridge on ventral side, apex deflected dorsally; right paramere short and broad, not
extended to apical half of median lobe; internal sac serrulate near apex, apical sclerite not
present.
ALLOTYPE, female, labelled as follows: “Winter Park 2.15.28 Fla,; John George Gehring
Collection; ALLOTYPE Evarthrus floridensis R. Freitag (green label); loan from MCZ.”
MCZ.
Body length 14.7 mm, width 6.1 mm. Form same as in holotype.
Microsculpture of head between eyes, disc of pronotum and intervals of elytra same as in
holotype; integument of dorsum slightly glossy.
Head length 1.9 mm, width 3.6 mm. Pronotum form same as in holotype; length 4.1 mm,
width 5.1 mm. Elytra length 8.7 mm, width 6.1 mm. Stylus of ovipositor gradually tapered
apically.
Derivation of species name. — The name floridensis was given to this species because its
members are known from Florida only.
Variation among paratypes (18 males, 13 females, Fla.). — Total length 13.0 — 15.0 mm.
The genitalia of five males were examined and no variation was observed.
Revision of Evarthrus
133
Disposition of type material. — The holotype and allotype are in the collections of the
MCZ. Two paratypes are in the UASM collection and the others are in the collections of the
CU and MCZ.
Geographical distribution (fig. 131). — This species is endemic to Florida. I have seen 46
specimens from the following localities.
United States - FLORIDA: Orange County: Winter Park (CU, MCZ, UASM). Osceola County: Deer Park (MCZ);
Kissimmee (AMNH). Seminole County: Sanford (MCZ). Volusia County: Enterprise (CAS). Counties not determined:
Haw Creek (USNM); North Smyrna (CAS).
The sigillatus Group
Characteristics. — Pronotum quadrate with obtuse and broadly rounded basal angles; male
genitalia with median lobe moderately or strongly arcuate, right paramere tapered apically
and slightly to markedly elongate.
The sigillatus group is composed of the species sigillatus, sinus and convivus. This group
occupies the eastern side of the Mississippi River Valley, Piedmont and Coastal Plain areas.
Evarthrus sigillatus Say, 1823
Figures 25-28, 72-73, 103, 131
Feronia sigillata Say, 1823a:42. Type lost. TYPE LOCALITY, Mr. R. Haines farm, German-
town (Pa.). - LeConte, 1848:350 (Feronia). - LeConte, 1863a:8 (Evarthrus). - LeConte,
1873:318. - Schaupp, 1880:49. - Leng, 1920:57. - Leonard, 1926:222. - Brimley,
1938:119.
Feronia (Omaseus) vidua Dejean, 1828:278. Type seen by C. H. Lindroth (1955). MNHP.
TYPE LOCALITY, l’Amerique Septentrionale. - LeConte, 1848:350 (Feronia). - Le-
Conte, 1852:228 (Evarthrus). - LeConte, 1863a:49. — Leng, 1920:57. - Csiki, 1930:
675 (Pterostichus).
Feronia (Abax) americana Dejean, 1828:392. TYPE, male, labelled as follows: “americanus
m.” MHNP. TYPE LOCALITY, “l’Amerique Septentrionale.” NEW SYNONYMY. -
Schaupp, 1880:49 (Evarthrus). — Casey, 1918:361. — Csiki, 1930:673 (Pterostichus).
Feronia orbata Newman, 1835:386. TYPE, female, labelled as follows: “Type H. T.; Ent.
Club. 44-12; J. Ingall Canada; Feronia Latreille orbata Newman Ent. Mag. V.386.” BM.
NEW SYNONYMY. - Motschulsky, 1865:261 (Evarthrus). - Leng, 1920:57. - Csiki,
1930:673 (Pterostichus).
Evarthrus breviformis Casey, 1918:360. HOLOTYPE, female, labelled as follows: “Southern
Pines; A. H. Manee. NC; CASEY bequest 1925; TYPE USNM 47120; breviformis Csy.”
USNM. TYPE LOCALITY, Southern Pines, N. Carolina. NEW SYNONYMY. - Leng,
1920:57 (Evarthrus). — Csiki, 1930:673 (Pterostichus). — Brimley, 1938:1 19 (Evarthrus).
Evarthrus montanus Van Dyke, 1926:116. HOLOTYPE, male, labelled as follows: “Black
Mts. NC VII. 1902; collector E. C. Van Dyke; Van Dyke Collection.” ALLOTYPE,
labelled the same except for “Black Mts. NC VI. 1902.” CAS. TYPE LOCALITY, in the
valley at the base of the Black Mountains, North Carolina. NEW SYNONYMY. - Csiki,
1930:673 (Pterostichus). — Leng and Mutchler, 1933: 13 (Evarthrus).
Pterostichus (Pterostichus) (Sect. Evarthrus) carolinensis Csiki, 1930:673. NEW SYNON-
YMY. - Leng and Mutchler, 1933:13 (Evarthrus). - Brimley, 1938:1 19.
134
Freitag
Recognition. — The following combination of characters separates specimens of sigillatus
from specimens of all similar species of Evarthrus : pronotum quadrate, sides not strongly
constricted posteriorly; basal angles slightly or broadly obtuse, not prominent, evenly
rounded; male genitalia, with left side of apex of median lobe sharply deflected dorsally, in-
ternal sac with characteristic apical sclerite; range mainly east of the Appalachian Mountains.
Specimens of sigillatus in western areas of the range can be confused with convivus indivi-
duals. It is usually necessary to compare the male genitalia for a certain identification. The
right paramere is long and tapered in convivus but short and broader in sigillatus (fig. 103 cf.
fig. 105).
Specimens of blatchleyi resemble those of sigillatus in North and South Carolina. These
can be distinguished as follows: basal fovea of the pronotum of blatchleyi simply and clearly
impressed, but it is more complex in sigillatus (fig. 23 cf. figs. 25-28); apical blade of median
lobe of male in blatchleyi is elongate, narrow, and evenly deflected dorsally and to the left,
but it is short, broader, and left side of apex sharply deflected dorsally in sigillatus (fig. 101
cf. fig. 103).
Description. - Body length 13.4 - 18.3 mm. Form narrow with sides of elytra somewhat
convex or broad with parallel sides of elytra.
Microsculpture of head between eyes and disc of pronotum with highly sinuous, entwined
lines, often forming amorphic meshes, usually partially effaced. Intervals of elytra with
micro sculpture formed by amorphic or isodiametric meshes. Integument of dorsum mark-
edly glossy, elytra dull in some specimens.
Head with frontal grooves fairly deep and sharply defined, usually short with middle
bend, convexity directed medially, or straight and slightly oblique to one another. Penul-
timate article of labial palpus with four or five setae.
Pronotum shape somewhat variable but essentially quadrate as in figs. 25-28; disc of aver-
age convexity; sides not strongly produced, usually fairly parallel, slightly constricted anteri-
orly and posteriorly, sinuation in front of basal angle slight or absent; posterior angles not
produced, obtuse and broadly rounded; anterior transverse impression complete and dis-
tinctly impressed, basal lateral foveae with sides continuous or not posteriorly, inner side
with interrupted extension from base toward middle longitudinal line; lateral bead slightly
broader posteriorly. Prostemal process with deep, sharply defined longitudinal groove.
Middle femur with four to six setae on anterior face.
Elytra slightly sinuate apically; intervals slightly raised or flat; striae distinctly and moder-
ately impressed, punctate anteriorly, obsoletely punctate or impunctate posteriorly.
Male genitalia (fig. 103) with median lobe moderately arcuate, angle distinctly obtuse;
apical blade elongate, left comer of apex deflected dorsally ; right paramere s^ ort not ex-
tending to apical half of median lobe, fairly broad, with slight tapering apically; internal sac
with serrulate field apically, apical sclerite light, amorphic plate with darker basal tooth. The
genitalia of 22 males were examined in detail.
Stylus of female ovipositor with relatively parallel sides and broadly rounded apex.
Geographical variation. — This is one of the most variable species of Evarthrus. The
variable features which I have noted are: the shape of the pronotum, shape of the elytra,
glossiness of the elytra and form of the male genitalia. In northern areas of the species range,
Revision of Evarthrus
135
in Pennsylvania for example, the pronotum is fairly rectangular in outline (fig. 25), elytra
are produced laterally and slightly glossy. At higher altitudes in western North Carolina the
pronotum is more elongate and the sides are more sinuate in front of the posterior angles
(fig. 26). The shape and glossiness of the elytra are the same as those of specimens in
Pennsylvania. On the Piedmont and Coastal Plain regions of North and South Carolina,
Georgia, Florida panhandle and eastern Alabama, the pronotum is relatively broader, with-
out sinuate sides in front of the posterior angles (figs. 27—28), and the sides of the elytra
are more parallel and surface of the elytra are duller than those of specimens further north
in Pennsylvania or at higher elevations. These three morphologically distinct populations are
linked by populations with intermediate structures that intergrade clinally.
Notes on synonymy . — The species sigillatus was identified by the original description.
The type specimen of vidua Dejean resembles sigillatus specimens from northern limits of
this species range. The type specimen of americana Dejean is a sigillatus specimen of the
form that occurs in central and eastern North and South Carolina. The type specimen of
orbata Newman is a sigillatus specimen of the kind that composes populations in Pennsyl-
vania and Virginia. The type of breviformis Casey is a sigillatus specimen of the sort found
in eastern and southern North Carolina. The type specimen of montanus is a sigillatus speci-
men of the average form which inhabits western North Carolina. Csiki lumped Evarthrus
and Pterostichus which brought into one genus the names montanus Motschulsky and
montanus Van Dyke. The new name carolinensis Csiki was created to replace montanus Van
Dyke.
Collecting notes. — This species is found in leaf litter of deciduous forests as well as under
cover in open places such as pastures.
Geographical distribution (fig. 131). — Evarthrus sigillatus ranges from the Florida pan-
handle to southern New York primarily east of the Appalachian Mountains. I have seen 432
specimens collected in the following localities.
United States - ALABAMA: Lee County: Auburn (AU, CAS, MCZ). Randolph County: Wadley (USNM). Tallapoosa
County: Alexander City (AU). DISTRICT OF COLUMBIA: Pincy Bridge (CNHM); Washington (USNM). FLORIDA: Jack-
son County: Grand Ridge (FDPI). Leon County: Tallahassee (FDPI, UMMZ). Liberty County: Camp Torreya (UMMZ);
Rock Bluff (UMMZ). GEORGIA: Camden County: Kingsland (UMMZ). Liberty County: Riceboro (UMMZ). Morgan
County: four miles north of Madison (DL); Madison (UMMZ). Rabun County: Clayton (AMNH, UMMZ, USNM). County
not determined: Wilson Gap (CU). MARYLAND: Ann Arundel County: Odenton (CU); Baltimore (CAS). Harford
County: Edgewood (CU). Montgomery County: (USNM). County not determined: Yellow Springs (RTB). MASSACHU-
SETTS: Middlesex County: Woburn (USNM). NEW JERSEY: Bergen County: Hillsdale (MCZ, USNM); Palisades (USNM);
Ramsey (AMNH). Essex County: Newark (AMNH); South Orange (USNM). Hudson County: Arlington (USNM). Morris
County: Boonton (USNM); Chester (AMNH); Lake Hopatcong (MCZ). Passaic County: Oak Ridge (USNM). Somerset
County: Bound Brook (USNM). Sussex County: Hopatcong (AMNH); Sparta (DRW). Counties not determined: Dundsel
(MCZ); Durh. P. (USNM); Fulerton (CU); Lahaway (USNM); Springdale Park (USNM). NEW YORK: Nassau County: Sea
Cliff (MCZ). Rockland County: Bear Mountain (CAS, UASM). NORTH CAROLINA: Buncombe County: (GEB); Ashe-
ville (MCZ); Black Mountains (AMNH, CAS, CNHM, MCZ, USNM). Burke County: Linn Falls (USNM). Catawba County:
Hickory (CNC). Haywood County: Crestmont (UMMZ); Lake Junaluska (FDPI); Mount Sterling (UMMZ). Henderson
County: 5 mi. north of Bat Cave (AMNH); Hendersonville (USNM); Mills River (CNC). Jackson County: Dillsboro
(AMNH). Madison County: Hot Springs (USNM). McDowell County: Marion (MCZ). Mecklenburg County: Charlotte
(MCZ). Moore County: Southern Pines (CAS, KSU, MCZ, RTB, UW). Orange County: Chapel Hill (CU). Polk County:
Tryon (MCZ). Randolph County: Julian (MCZ). Robeson County: Lumberton (UMMZ). Wake County: Raleigh (CNC,
NCSU, USNM). Wilkes County: Wilkesboro (CU, USNM). Counties not determined: Beaver Creek (NCSU); Black Camp
Gap (TCB); Graybeard Mountain (AMNH); Morrison Mountain (USNM); Mount Pisgah (USNM); Peano Rendezvous (GEB)
136
Freitag
Round Knob (USNM); Stony Creek (RTB). PENNSYLVANIA: Bucks County: (RU) Cumberland County: Enola (MCZ).
Fayette County: Uniontown (CAS). Montgomery County: Whitemarsh (USNM). Northampton County: Easton (CAS,
CNHM, TE, UASM); Wind Gap (CNHM). Philadelphia County: Frankford (USNM); Germantown (ANSP); Mount Airy
(CAS, RU); Philadelphia (MCZ). Counties not determined: Abbotsford (MCZ); Angord (CAS); Femwald (CNHM; Ingle-
nook (CAS); Lehigh Gap (USNM); Rockville (MCZ); Water Gap (AMNH); Wissahickon Creek (RU). SOUTH CAROLINA:
Beaufort County: Hardeeville (UMMZ). Berkeley County: Goose Creek (UMMZ). Colleton County: Round O (UMMZ).
Darlington County: Hartsville (UMMZ). Florence County: Florence (GEB); Scranton (UMMZ). Greenville County: Green-
ville (UMMZ). Greenwood County: Greenwood (UMMZ). Kershaw County: Camden (MCZ, UMMZ). Oconee County:
CCC Camp F-2 (CAS); Clemson College (USNM). Richland County: Columbia (UMMZ). Saluda County: Saluda (UMMZ).
County not determined: Meredith (CAS). TENNESSEE: Blount County: Chilhowee Mountain (CNC). Carter County:
Roan Mountain (UMMZ). Cocke County: French Broad River (MCZ). Knox County: Knoxville (CNC). McMinn County:
1.5 mi. north of Athens (UMMZ). Morgan County: (CNHM); Deer Lodge, Environs (CNHM). Sevier County: Gatlinburg
(UMMZ, USNM). Counties not determined: Crabtree (CU); Great Smoky Mountains National Park (CNC); Unaka Moun-
tains (ANSP); Unaka Springs (RTB). VIRGINIA: Arlington County: Rosslyn (MCZ). Bedford County: Blue Ridge
National Parkway (RCG). Fairfax County: (ANSP, USNM); Dead Run (USNM). Nansemond County: Cypress Chapel
(UMMZ). Counties not determined: Black Pond (USNM); Diamond Springs (USNM); Great Falls (USNM); Stony Man
Mountain (MCZ).
Evarthrus sinus new species
Figures 29, 104, 131
Recognition. — The following combination of characteristics is diagnostic for this species:
pronotum with sides more constricted anteriorly than posteriorly, not sinuate in front of
posterior angles; lateral bead rather broad posteriorly; male genitalia with strongly arcuate
median lobe and narrow parameres; coastal or near coastal distribution. The species which
are most similar in external structural characteristics to sinus are blatchleyi and floridensis.
The geographical ranges of sinus and the last two species are different. However specimens
of these species can also be easily separated by differences in their male genitalia (figs. 101,
102, 104).
Description. - HOLOTYPE, male, labelled as follows: “Alabama Port, Mobile Co. Ala.
June 6, 1950 Ball-Wilson; HOLOTYPE Evarthrus sinus R. Freitag (red label).” MCZ.
Body length 13.7 mm, width 5.7 mm. Form approximately parallel at sides.
Head between eyes, disc of pronotum, and intervals of elytra with microsculpture con-
sisting of sinuous, entwined, dense lines often forming amorphic meshes.
Head glossy; length 1.7 mm, width 3.3 mm; frontal grooves distinctly and sharply im-
pressed; slightly curved away from one another posteriorly. Penultimate article of labial pal-
pus with five setae, three medial and two apical.
Pronotum moderately glossy; length 3.9 mm, width 4.8 mm; shape somewhat cordiform
in outline as in fig. 29; disc of average convexity, sides more constricted anteriorly than
posteriorly, not sinuate in front of posterior angles; posterior angles not produced, obtuse
and broadly rounded; anterior transverse impression complete and deeply impressed; basal
lateral foveae with sides not continuous posteriorly, inner side with extension from base
toward middle longitudinal line; lateral bead distinctly broader posteriorly. Prostemal pro-
cess with deep, sharply defined longitudinal groove. Middle femur with four setae on
anterior face.
Elytra slightly glossy, somewhat velvety in appearance; length 8.1 mm, width 5.7 mm;
slightly sinuate apically; intervals almost flat; striae moderately impressed, distinctly punc-
tate anteriorly, obsoletely punctate posteriorly.
Revision of Evarthrus
137
Male genitalia (fig. 104) with median lobe strongly arcuate, angle approximately right;
apical blade fairly short, round at apex, and deflected to right; right paramere of average
length just extended to apical half of median lobe, distinctly tapered apically, apex narrow;
internal sac with serrulate field apically, apical sclerite light amorphic plate with serrulate
basal portion.
ALLOTYPE, female, labelled as follows: “Alabama Port, Mobile Co. Ala. June, 6, 1950
Ball-Wilson; ALLOTYPE Evarthrus sinus R. Freitag (green label).” MCZ.
Body length 13.9 mm, width 5.8 mm. Form same as in holotype.
Micro sculpture on head, pronotum and elytra same as in holotype. Head glossy; length
1.7 mm, width 3.3 mm. Pronotum glossy; form same as in holotype; length 3.7 mm, width
4.7 mm. Elytra slightly glossy; length 8.4 mm, width 5.8 mm. Stylus of ovipositor with
somewhat parallel sides, apex evenly rounded.
Derivation of specific name. — This species is given the name sinus, a latin noun meaning
gulf, because its members live in the vicinity of the Gulf Coast.
Variation among paratypes (five males, nine females, Mississippi, Alabama). — Total
length 13.1 — 15.9 mm. The variation in the features which I examined is no greater than
that between the, holotype and allotype. The genitalia of five males were examined.
Disposition of type material. — The holotype and allotype are in the collections of the
MCZ. The paratypes are in the following collections: CAS, CU, GEB, MCZ, UASM, UMMZ,
USNM.
Collecting notes. — This species has been collected in pine-oak coastal forest by G. E. Ball.
Geographical distribution (fig. 131). - This species is represented on the Coastal Plain of
Alabama and Mississippi. I have seen 1 9 specimens from the following localities.
United States - ALABAMA: Mobile County: Alabama Port (GEB); Mobile (CAS, MCZ, UASM, USNM). MISSISSIPPI:
George County: Lucedale (CU). Harrison County: Gulfport (UMMZ). Perry County: Richton (CU).
Evarthrus convivus LeConte, 1852
Figures 30-32, 105, 131
Evarthrus convivus LeConte, 1852:229. LECTOTYPE (here selected) a male, labelled as fol-
lows: “orange disc; Type 5654, E. orbatus (Newm) conviva LeC.” MCZ. TYPE LOCALI-
TY, Alabama. — LeConte, 1863a:8 (Evarthrus). - LeConte, 1873:318. - Schaupp, 1880:
49. - Leng, 1920:57. - Csiki, 1930:673 ( Pterostichus ). - Loding, 1945:16 (Evarthrus).
Feronia (Pterostichus) orbata \ LeConte, 1848:348 (not Newman).
Evarthrus orbatus ; LeConte, 1852:229 (not Newman). - LeConte, 1863a:8. - LeConte,
1873:318. - Schaupp, 1880:49. - Blatchley, 1910:101. - Lbding, 1945:16.
Evarthrus sigillatus; LeConte, 1852:228 (not Say). — Blatchley, 1910:101. - Casey, 1918:
359. -Loding, 1945:16.
Evarthrus sigillatus parallelus Casey, 1918:359. HOLOTYPE, male, labelled as follows:
“Ind; CASEY bequest 1925; TYPE USNM 47119, parallelus Csy.” USNM. TYPE LO-
CALITY, Indiana. NEW SYNONYMY. - Leng, 1920:57 (Evarthrus). - Csiki, 1930:674
(Pterostichus). - Loding, 1945:16 (Evarthrus).
Pterostichus (Pterostichus) (Sect. Evarthrus) sigillatus-, Csiki, 1930:674 (not Say).
138
Freitag
Recognition. — A combination of flat and dull elytral intervals, male genitalia with an
elongate and narrow right paramere and C-shaped sclerite of the internal sac, and geographic
distribution west and southwest of the Appalachian Mountains, is characteristic of convivus.
The species sigillatus and convivus are for the most part allopatric but their ranges overlap
in western Pennsylvania and eastern Tennessee. In Pennsylvania convivus specimens have de-
cidedly more parallel sides and are duller than those of sigillatus. The distinction between
these two species is more obscure in eastern Tennessee and indeed it is usually necessary to
examine the male genitalia for a certain identification.
The closely related species sinus and convivus are distinguished by the basal lateral foveae
of the pronotum (fig. 29 cf. figs. 30-32) male genitalia (fig. 105), and geographical range.
Description. — Body length 1 1.6 — 19.2 mm. Form typical of sigillatus group.
Head between eyes with highly sinuous lines and amorphic meshes or meshes alone
composing microsculpture. Disc of elytra with highly sinuous lines, usually very densely
distributed, forming microsculpture. Microsculpture of intervals of elytra with highly sinu-
ous lines and raised bead-like meshes.
Head moderately glossy; frontal grooves fairly deep and sharply defined, with middle
bend, convexity directed medially. Penultimate article of labial palpus with five setae.
Pronotum moderately glossy; shape quadrate as in figs. 30 — 32; disc of average convexity
or slightly flatter; sides not strongly produced, slightly constricted anteriorly and posteri-
orly, obsoletely sinuate in front of posterior angles; posterior angles not produced, obtuse
and broadly rounded; anterior transverse impression complete and distinctly impressed;
basal lateral fovea with sides continuous or not posteriorly, inner side with interrupted
extension from base toward median longitudinal impression; lateral bead slightly broader
posteriorly. Prostemal process with deeply or shallowly impressed longitudinal groove.
Middle femur with four to five setae on anterior face.
Elytra dull, matte, slightly sinuate apically; intervals flat or almost flat; striae not deeply
impressed, distinctly punctate anteriorly, indistinctly or obsoletely punctate posteriorly.
Male genitalia (fig. 105) with median lobe strongly arcuate, angle slightly obtuse; apical
blade short, evenly rounded at apex, curved to right; right paramere elongate, apical half
narrow, extended to apical half of median lobe; internal sac with serrulate field apically,
apical sclerite distinctly C-shaped. The genitalia of 26 males were examined.
Stylus of female ovipositor slightly tapered apically.
Geographical variation. — Body size is notably variable. Tiny specimens are uncommon
and appear sporadically throughout the range. In contrast giant forms are rather common in
northeastern Alabama and in all directions from that region the body size decreases clinally.
In other characteristics variation is minimal and specimens in Mississippi are grossly similar
to those at the other end of the range in Pennsylvania or Illinois.
Notes on synonymy. - The type specimen of parallelus Casey is a slightly smaller than
average convivus specimen.
Collecting notes. — D. J. Larson and I have taken some specimens in leaf litter in mixed
pine and deciduous forests in Mississippi.
Geographical distribution (fig. 131). — This species ranges from the Mississippi River east
to the Appalachian Mountains and from southern Alabama north to Illinois and western
Pennsylvania. Some populations occur beside the Mississippi River in Louisiana. I have seen
491 specimens from the following localities.
Revision of Evarthrus
139
United States - ALABAMA: Bibb County: The Sinks (UMMZ). Blount County: Blount Springs (CM). Cherokee
County: Leesburg (UMMZ). Clarke County: six miles south of Jackson (UMMZ). Conecuh County: Brooklyn (TCB).
DeKalb County: Desoto State Park (CAS). Franklin County: Russellville (GEB). Jackson County: Point Rock (UMMZ);
Sand Mountain, Bryant (UMMZ). Madison County: Huntsville (UMMZ); Monte Sano (CNC, UMMZ). Marengo County:
south of Demopolis (UMMZ). Mobile County: (CAS); Chickasaw (CU); Mobile (ANSP, CAS, MCZ); Mount Vernon (CU);
Spring Hill (CAS). St. Clair County: Blount Mountains (GEB). Talladega County: Talladega (UMMZ). Tuscaloosa County:
Lock 14 (CAS); Peterson (GEB); Talladega State Forest (GEB); Tuscaloosa (GEB, UMMZ); Windham Springs (GEB).
ILLINOIS: Alexander County: Cache River (RTB); Olive Branch (CAS, CNHM). Cass County: Virginia (CU). Champaign
County: Urbana (CNHM, MCZ, RTB). Clark County: Martinsville (UMMZ). Coles County: Fox Ridge State Park (RTB).
Vermilion County: Kickapoo State Park (RTB). Washington County: Dubois (INHS). County not determined: Bottoms
(INHS). INDIANA: Crawford County: (UP). Clark County: (CAS, UP). Elkhart County: Goshen (UMMZ). Floyd County:
(UP, MCZ). Gibson County: Oakland City (UMMZ). Greene County: (UP). Hendricks County: Stilesville (CAS). Jefferson
County: Clifty Falls State Park (GEB). Knox County: (CAS). LaGrange County: Lagrange (UMMZ). Marion County
(UASM). Monroe County: Bloomington (UMMZ). Montgomery County: (UP). Putnam County: (UP). St. Joseph County:
Mishawaka (UMMZ). Starke County: Bass Lake (CNHM). Tippecanoe County: LaFayette (UMMZ). County not deter-
mined: Turkey Run (INHS). KENTUCKY: Barren County: Cave City (USNM). Edmonson County: Mammoth Cave
National Park (TCB). Hardin County: Fort Knox (GEB). Jefferson County: Anchorage (UL); Prospect (UL). Jessamine
County: (GEB). Wayne County: Wolf Creek Lake (UL). County not determined: Sleepy Hollow (UL). LOUISIANA:
East Baton Rouge County: Baton Rouge (UMMZ). St. Tammany County: Covington (USNM). MISSISSIPPI: Adams
County: Natchez (CAS, USNM). Calhoun County: Vardaman (UMMZ). Clairbome County: Edwardsville (RCG). Forrest
County: Hattiesburg (AMNH). George County: Lucedale (CU). Granada County: Dubard Station (UMMZ). Greene
County: Leakesville (CU). Lauderdale County: five miles south of Toomsuba (DL, RF); Meridian (UMMZ). Perry County:
Richton (CU). Simpson County: (UMMZ). Tishomingo County: Iuka (UMMZ). Counties not determined: North Augusta
(CU); six miles east of Iuka (FDPI). OHIO: Adams County: (UMMZ). Allen County: Lima (UMMZ). Brown County:
Russelsville (CU). Cuyahoga County: Cleveland (MCZ, UMMZ). Darke County: Beamville (RU, UMMZ). Hamilton County:
Cincinnati (UMMZ). Licking County: West Alexandria (RTB). Preble County: Eaton (UMMZ). PENNSYLVANIA: Alle-
gheny County: (CM, CU); Fair Oaks (CM); Millvale (CM); Pittsburgh (CAS, CM); Wall (CM, UK). Fayette County: Dunbar
(CM); Union Town (CAS, MCZ, UMMZ, USNM). Westmoreland County: Jeanette (CM). County not determined: Alle-
gheny (ANSP, MCZ, UK, USNM). TENNESSEE: Knox County: 25 miles south of Knoxville (AMNH). Lake County:
Gray’s Landing (RTB). Lauderdale County: South Fulton (UMMZ). Morgan County: Burrville (CNHM, CU). Obion
County: Obion (UMMZ). WEST VIRGINIA: Monongalia County: Greer (GEB). Taylor County: Grafton (USNM). Webster
Springs (MCZ).
The seximpressus Group
Characteristics. — Penultimate article of labial palpus with five setae, two medial and
three apical. Pronotum more or less quadrate in outline; anterior transverse impression com-
plete and distinct throughout. Prosternal process with setae on apex. Middle femur with
five to seven setae on anterior face (fig. 75). Prosternal process with longitudinal groove
shallow, but sharply defined. Male genitalia with median lobe moderately arcuate; internal
sac with apical sclerite light with darker oval basal portion; right paramere markedly tapered
apically.
This group includes the species seximpressus, alabamae, engelmanni, and nonnitens. They
occur on the Gulf Coast in Texas, Louisiana, Mississippi, Alabama, the Ozark Plateau, and
Mississippi Valley north to Wisconsin.
Evarthrus seximpressus LeConte, 1848
Figures 33, 75, 106, 132
Feronia (Pterostichus) seximpressa LeConte, 1848:350. LECTOTYPE (here selected) a
male, labelled as follows: “dark green disc; Type 5653; E. seximpressus LeConte.” MCZ.
140
Freitag
TYPE LOCALITY, Rocky Mountains near Long’s Peak. - LeConte, 1852:228 ( Evar -
thrus). - LeConte, 1863a:8. - LeConte, 1873:318. - Schaupp, 1880:49. - Blatchley,
1910:100. - Casey, 1918:361. - Leng, 1920:57. - Csiki, 1930:673 ( Pterostichus ). -
Loding, 1 945 : 1 6 {Evar thrus).
Evarthrus rubripes Casey, 1918:359. HOLOTYPE, male, labelled as follows: “Mo.; CASEY
bequest 1925; TYPE USNM 47121; rubripes Csy.” USNM. TYPE LOCALITY, Saint
Louis, Missouri. PARATYPE, female, labelled as follows: “la; CASEY bequest 1925;
rubripes - 2; PARATYPE USNM 47121.” USNM. TYPE LOCALITY, Keokuk, Iowa.
NEW SYNONYMY. - Leng, 1920:57 {Evarthrus). - Csiki, 1930:673 {Pterostichus).
Recognition. — The species seximpressus is distinguished from other species of th esexim-
pressus group by the following combination of characteristics: body size relatively small;
pronotum (fig. 33) more quadrate than circular in outline, posterior angles obsolete, lateral
bead broad near base; male genitalia with apical blade of median lobe relatively narrow and
evenly rounded at apex (fig. 106).
Description. — Body length 10.3 — 15.9 mm. Form relatively less robust than other
species of the seximpressus group.
Microsculpture on head between eyes with markedly sinuous lines and amorphic meshes.
Disc of pronotum with microsculpture formed of highly sinuous lines. Microsculpture of
elytral intervals with amorphic or isodiametric meshes.
Head slightly or moderately glossy; frontal grooves fairly deep and distinct, with slight
middle bend, posterior halves directed laterally. Penultimate article of labial palpus with
five setae.
Pronotum moderately or slightly glossy; shape quadrate, as in fig. 33; disc of average con-
vexity; sides produced, slightly constricted anteriorly and posteriorly, not sinuate in front
of posterior angles; posterior angles not produced, obtuse and broadly rounded; anterior
transverse impression complete and distinctly impressed; basal fovea with sides continuous
or not posteriorly; lateral bead much broader posteriorly. Prosternal process with longi-
tudinal groove distinctly but not deeply impressed. Legs red or black; middle femur with
five or six setae on anterior face (fig. 75).
Elytra usually dull, slightly glossy in a few specimens; slightly sinuate apically; intervals
flat or almost flat; striae not deeply impressed, distinctly punctate anteriorly, impunctate or
obsoletely punctate posteriorly.
Male genitalia (fig. 106) with median lobe moderately arcuate, angle clearly obtuse;
apical blade evenly rounded at apex, very slightly deflected to right; right paramere dis-
tinctly tapered apically, not extended to apical half of median lobe; internal sac with
serrulate field apically, apical sclerite light amorphic plate. The genitalia of four specimens
were examined in detail.
Stylus of female ovipositor slightly tapered apically.
Geographical variation. — Leg color varies from red to black. Specimens with red legs are
most abundant in Wisconsin and northern Arkansas but they are uncommon throughout the
rest of the range of this species.
Notes on synonymy. — The type specimen of rubripes Casey is an average, red-legged
seximpressus specimen.
Revision of Evarthrus
141
Collecting notes. — Specimens of E. seximpressus have been collected under cover in
pastures.
Geographical distribution (fig. 132). - This species inhabits eastern areas of the Great
Plains, and the Mississippi Valley, from Oklahoma or possibly Texas in the south, north to
Michigan, Indiana and possibly western Pennsylvania. I have seen 45 1 specimens from the
following localities.
United States - ARKANSAS: Benton County: Rogers (KSU). Bradley County: (UA);Pine Oak Woods (UA). Conway
County: (UA). Garland County: Hot Springs (CAS). Hempstead County: Hope (CAS, MCZ, UMMZ). Lawrence County:
(CAS). Searcy County: Leslie (CAS). Sebastian County: Greenwood (INHS). Washington County: (INHS, UA); Cove
Creek, 27 miles from Fayetteville (DL); Cove Creek Valley (UA); Devil’s Den State Park (RTB). Yell County. (UA).
ILLINOIS: Adams County: Camp Point (INHS). Champaign County: ONHS); Urbana (INHS). DeKalb County: Hinckley
(UP). Hardin County: Shane Landing (RTB). McHenry County: Harvard (RCG). Macoupin County: Chesterfield (RTB).
Peoria County: Hanna City (INHS). Richland County: Wabash Valley (CM, MCZ). Rock Island County: Rock Island
(UMMZ). County not determined: Grand Detour (CNHM). INDIANA: Greene County: (UP). Knox County: (UP). Marion
County: (UP). Putnam County: (UP). IOWA: Davis County: (UMMZ). Henry County: Mount Pleasant (MCZ, MSU).
Johnson County: Iowa City (CNC, MCZ, USNM); Solon (USNM). Polk County: Des Moines (ISU). Pottawattomie County:
Council Bluffs (CAS, USNM). Story County: Ames (ISU). KANSAS: Clay County: (ANSP, CAS, UMMZ, USNM).
Doniphan County: Wathena (USNM). Douglas County: (MCZ, UK, USNM); Lawrence (ANSP, CAS, CNC, UMMZ, UW).
Pottawatomie County: Onaga (KSU). Riley County: Manhatten (KSU); Popenoe (KSU, USNM). Shawnee County: Topeka
(KSU, USNM). Woodson County: two miles east of Yates Centre (DL). Wyandotte County: Argentine (AMNH, RU).
MICHIGAN: Lenawee County: Adrian (MCZ). MINNESOTA: Houston County: Caledonia (KSU). MISSOURI: Buchanan
County: St Joseph (MCZ, USNM). Camden County: Camdenton (UMMZ). Carter County: Van Buren (UMMZ). Greene
County: Willard (ANSP, MCZ, UASM). Jackson County: Kansas City (UK). Jefferson County: Kimmswick (UMMZ).
Miller County: Ozark Lake (CAS). Polk County: Aldrich (CU). St. Louis County: St. Louis (CAS, CM, USNM). Taney
County: Branson (CAS). County not determined: Bolival (CAS). NEBRASKA: Douglas County: Omaha, Child’s Point
(CAS). Lancaster County: Lincoln (CAS). OKLAHOMA: Cleveland County: (CAS); Norman (CAS). Latimer County:
(CAS). LeFlore County: Page (UMMZ). Lincoln County: east of Stroud (TCB). Marshall County : Lake Texoma State Park
(TE). Rogers County: Catdosa (CNHM). Tulsa County: Tulsa (CAS). Wagoner County: Cornell (UMMZ). PENNSYL-
VANIA: Westmoreland County: Jeanette (CM). TEXAS: Brazos County: College Station (INHS). WISCONSIN: Dane
County: (UW); Madison (CU, UW). Dodge County: Beaver Dam (CAS, KSU, MCZ, UMMZ). Milwaukee County: Mil-
waukee (UW), Vernon County: Westby (USNM). County not determined: Wingra Lake (UW).
Evarthrus alabamae Van Dyke, 1926
Figures 34, 107, 132
Evarthrus vagans alabamae Van Dyke, 1926:118. HOLOTYPE, male labelled as follows:
“Mobile, Ala. III. 08; Van Dyke Collection.” CAS. ALLOTYPE, labelled as follows:
“Mobile, Ala. 11.26.1901. H. P. Loding; Van Dyke Collection.” CAS. NEW STATUS. -
Csiki, 1930:673 ( Pterostichus ). - Leng and Mutchler, 1933:13 (Evarthrus). - Loding,
1945:16.
Evarthrus lodingi\ Loding, 1945: 16 (not Van Dyke).
Recognition. — The circular pronotum is diagnostic of alabamae and distinguishes it from
the closely related species seximpressus which has a more rectangular pronotum. Character-
istics of the male genitalia of alabamae and seximpressus also distinguish these species from
one another (fig. 106 cf. fig. 107).
Description. — Body length 14.6 — 18.7 mm. Form robust, particularly the pronotum.
Head between eyes and disc of pronotum with highly sinuous lines and amorphic meshes
composing microsculpture. Intervals of elytra with microsculpture formed by a combination
of raised amorphic and isodiametric meshes.
142
Freitag
Head slightly or moderately glossy; frontal grooves of average depth, distinct, with slight
middle bend, posterior halves directed laterally. Penultimate article of labial palpus with
five setae.
Pronotum slightly or moderately glossy; shape somewhat circular in outline as in fig. 34.
Disc of average convexity; sides strongly produced, moderately constricted anteriorly and
posteriorly, not sinuate in front of posterior angles; posterior angles not produced, obtuse
and broadly rounded; anterior transverse impression complete and deeply impressed; basal
foveae with sides continuous or not posteriorly; lateral bead much broader posteriorly.
Prosternal process with longitudinal groove distinct but not deep. Middle femur with Five or
six setae on anterior face.
Elytra dull or slightly glossy; slightly sinuate apically; intervals almost flat or strongly con-
vex; striae of average depth, distinctly punctate anteriorly indistinctly punctate posteriorly.
Male genitalia (fig. 107) with median lobe moderately arcuate, angle clearly obtuse; apical
blade rather broad, apex broadly rounded; apex with lateral edges deflected dorsally; right
paramere distinctly tapered apically, extended to approximately halfway point of median
lobe; internal sac with serrulate field apically; apical sclerite hght amorphic plate, with
darker elhptically-shaped basal portion. The genitalia of six males were examined in detail.
Stylus of female ovipositor slightly tapered apically.
Variation. — Body size and convexity of elytral intervals are somewhat variable, but there
is no apparent geographical clinal pattern in the variation.
Two specimens from Oakhurst, Texas and two from Livingston, Texas are smaller than
the average alabamae specimen (e.g. from Mobile, Alabama). They vaguely resemble engel-
manni. However the alabamae characters are evident, e.g. very round basal angles of the
elytra and broader apical blade of the male phallus. These appear to be variants of alabamae
and that is how I regard them.
Collecting notes. — Specimens of alabamae are found in pine-oak woods, in leaf litter.
G. E. Ball collected specimens of this species along with specimens of E. sinus in pine-oak
coastal forest.
Geographical distribution (fig. 132). - This species inhabits the Gulf Coastal Plain from
Alabama west to Texas. The Kansas and northern Arkansas records are probably wrong. I
have seen 163 specimens from the following localities.
United States - ALABAMA: Mobile County: Alabama Port (GEB); Grand Bay (ANSP); Mobile (AMNH, ANSP, AU,
CAS, CU, KSU, MCZ, UMMZ, USNM, UW). ARKANSAS: Bradley County: Pine Oak Woods (UA). Lawrence County:
Imboden (CNHM, MCZ, USNM). KANSAS: Clay County: (CNHM). LOUISIANA: Caddo County: Shreveport (CAS).
Jefferson Davis County: Lake Arthur (CAS). Natchitoches County: Natchitoches (UMMZ); Vowell’s Mill (USNM). County
not determined: Hart (CAS). MISSISSIPPI: Harrison County: Gulfport (CU); Handsboro (FDPI, UMMZ). Jackson County:
Pascagoula (USNM). TEXAS: Harrison County: Marshall (UMMZ). Polk County: five miles east of Livingston (AMNH);
Livingston (AMNH). San Jacinto County: two miles east of Oakhurst (AMNH). Travis County: Austin (MCZ).
Evarthrus engelmanni LeConte, 1852
Figures 35, 108, 132
Evarthrus engelmani LeConte, 1852:228. LECTOTYPE (here selected) a male, labelled as
follows: “Tex; engelmani 2.” MCZ. TYPE LOCALITY, Texas. - LeConte, 1858:28
(. Evarthrus ). - LeConte, 1863a:8. — LeConte, 1873:318. — Schaupp, 1880:49. - Leng,
1920:57. — Csiki, 1930:673 ( Pterostichus ).
Revision of Evarthrus
143
Evarthrus vagans\ Schaupp, 1880:49 (not LeConte). — Csiki, 1930:673 ( Pterostichus ). -
Loding, 1945:16 {Evarthrus).
Recognition. — The combination of the produced sides, produced posterior angles, and
posterior widening of the lateral bead of the pronotum is diagnostic of specimens of engel-
manni. These features distinguish engelmanni from nonnitens.
Description. - Body length 12.6 — 19.1 mm. Form average for the seximpressus group.
Head between eyes and disc of pronotum with highly sinuous lines and amorphic meshes
composing microsculpture. Microsculpture of intervals of elytra formed by bead-like isodia-
metric meshes.
Head slightly or moderately glossy; frontal grooves moderately deep, distinct, straight or
with slight middle bend, posterior halves directed laterally. Penultimate article of labial pal-
pus with five setae.
Pronotum slightly glossy; shape somewhat quadrate in outline as in fig. 35 with disc of
average convexity; sides strongly produced, moderately constricted anteriorly and posteri-
orly, distinctly sinuate in front of posterior angles; posterior angles small, more or less
produced, slightly or broadly obtuse; anterior transverse impression complete and deeply
impressed; basal lateral fovea with sides usually continuous posteriorly; lateral bead much
broader posteriorly than anteriorly. Prostemal process with longitudinal groove moderately
or deeply impressed. Middle femur with five or six setae on anterior face.
Elytra dull, slightly sinuate apically; intervals flat or slightly convex; striae not deep, in-
distinctly punctate anteriorly, obsoletely punctate or impunctate posteriorly.
Male genitalia (fig. 108) with median lobe moderately arcuate, angle clearly obtuse; apical
blade broad with broadly rounded almost truncate apex; lateral edges of apex not strongly
deflected dorsally; right paramere very narrow apically, not extended to halfway point of
median lobe; internal sac with serrulate field apically, apical sclerite light amorphic plate
with dark elliptical basal portion. The genitalia of eleven males were examined.
Stylus of female ovipositor moderately tapered apically.
Geographical variation. — The pronotum is variable, which sometimes makes it difficult to
separate engelmanni from the three other species in the seximpressus group. For example
several specimens from College Station, Texas are very much like seximpressus but they all
have a sinuation in front of the posterior angles of the pronotum which is characteristic of
engelmanni. Other specimens of engelmanni resemble individuals of nonnitens in pronotal
features but are distinguishable by their male genitalia.
Notes on synonymy . - I believe LeConte named this species after George Engelmann,
1809 — 1844, a physician in St. Louis and eminent botanist. This is probably why LeConte
changed the name engelmani (1852) to engelmanni in subsequent publications.
Collecting notes. — Specimens of engelmanni have been collected in forests (label data).
Geographical distribution (fig. 132). — This species inhabits eastern Texas. The El Paso
record is surely not correct. I have seen 69 specimens from the following localities.
United States - TEXAS: Bastrop County: Bastrop State Park (CNC). Brazos County: (MCZ); College Station (MCZ,
TAM). DeWitt County: Cuero (AMNH). El Paso County: El Paso (CM). Goliad County: (USNM). Grimes County: (TAM).
Harris County: Houston (CM). Madison County: (TAM). Tarrant County: east of Fort Worth (KSU). Travis County:
Austin (MCZ). Victoria County: Victoria (USNM). County not determined: Fedor (CAS, CM).
144
Freitag
Evarthrus nonnitens LeConte, 1873
Figures 36, 109, 132
Evarthrus nonnitens LeConte, 1873:320. LECTOTYPE (here selected) a female, labelled as
follows: “red disc; Red River; Type 5656; E. nonnitens LeC.” IV'CZ. TYPE LOCALITY,
Red River, Louisiana. - Schaupp, 1880:49 ( Evarthrus ). - Casey, 1918:362. - Leng,
1920:57. - Csiki, 1930:673 ( Pterostichus ).
Evarthrus enormis Casey, 1918:361. HOLOTYPE, female, labelled as follows: “Tex; CASEY
BEQUEST 1925; TYPE USNM 47125; enormis Csy.” USNM. TYPE LOCALITY, Hous-
ton, Texas. NEW SYNONYMY. - Leng, 1920:57 {Evarthrus). - Csiki, 1930:673
{Pterostichus).
Recognition. — Specimens of nonnitens are characterized by the combination of the ex-
tremely matte surface of the elytra, somewhat flattened disc of the pronotum, and relatively
narrow posterior portion of the lateral bead of the pronotum.
Individuals of nonnitens can be confused with specimens of seximpressus and engehnanni.
The duller elytra of nonnitens usually distinguishes it from both engelmanni and sexim-
pressus. In addition specimens of nonnitens and engelmanni can be distinguished from one
another by the difference in widths of the basal part of the lateral bead of the pronotum
(fig. 35 cf. fig. 36). Also seximpressus does not have a produced basal angle of the pronotum
while nonnitens has. The relative width of the apex of the median lobe of the genitalia is
also a reliable feature for separating these species (fig. 109 cf. figs. 106 and 108).
Description. - Body length 13.7 - 16.9 mm. Form average for the seximpressus group.
Head between eyes, disc of pronotum, and intervals of elytra with microsculpture com-
posed of highly sinuous entwined lines.
Head slightly glossy; frontal grooves deep, distinct, with middle bend, posterior halves
directed laterally. Penultimate article of labial palpus with five setae.
Pronotum dull, shape quadrate in outline as in fig. 36, disc somewhat flattened in middle;
sides slightly produced, slightly constricted anteriorly and posteriorly, sinuate in front of
posterior angles; posterior angles small and slightly produced, clearly obtuse; anterior trans-
verse impression complete and deeply impressed; basal lateral fovea with sides continuous or
not posteriorly; lateral bead slightly broadened posteriorly. Prosternal process with longi-
tudinal groove moderately or deeply impressed. Middle femur with five to seven setae on
anterior face.
Elytra very dull, matte; obsoletely sinuate apically; intervals flat or slightly convex; striae
not deep, distinctly punctate anteriorly, indistinctly punctate or impunctate posteriorly.
Male genitalia (fig. 109) with median lobe moderately arcuate, angle clearly obtuse,
evenly rounded; apical blade broad, apex broadly rounded almost truncate, lateral edges
deflected dorsally; right paramere very narrow apically extended to approximately halfway
point of median lobe; internal sac with serrulate field apically ; apical sclerite light amorphic
plate with dark oval basal portion. The genitalia of five males were examined.
Stylus of female ovipositor slightly tapered apically.
Notes on synonymy. - The type specimen of enormis Casey is an average specimen of
nonnitens in all respects.
Revision of Evarthrus
145
Geographical distribution (fig. 132). - This species is known from southern Arkansas and
the Gulf Coastal Plain in Mississippi, Louisiana and eastern Texas. I have seen 33 specimens
collected in the following localities.
United States - ARKANSAS: Bradley County: (UA); Crimson Clover (UA); Pine Oak Woods (UA). Clarke County:
(UA). Hempstead County: Hope (MCZ, UASM, UMMZ). LOUISIANA: Grant County: Grant Point, Dryprong (CNHM).
Lincoln County: five miles east of Ruston (AMNH); Ruston (MCZ). County not determined: Red River (MCZ). MISSIS-
SIPPI: Adams County: Natchez (CAS). TEXAS: Harris County: Houston (USNM). San Jacinto County: two miles cast
of Oakhurst (AMNH).
The hypherpiformis Group
Characteristics. — Penultimate article of labial palpus with five setae. Pronotum quadrate
in outline; anterior transverse impression complete, shallow medially. Prostemal process
without setae on apex. Three to five setae on third interval of elytron. Median lobe of male
genitalia with medial ventral bump; apex of apical blade flat and sharp, deflected ventrally.
Right paramere short and markedly tapered apically. This group is represented by one
species, hypherpiformis. It occurs in the northern Coastal Plain area of Alabama and
Mississippi.
Evarthrus hypherpiformis new species
Figures 37, 1 10, 132
Recognition. — The combination of body size, three to five setae on the third interval of
the elytron, flattened pronotum, and form of the male genitalia is diagnostic for this species.
Specimens of hypherpiformis resemble those of nonnitens but without setae on the apex of
the prostemal process.
Description. - HOLOTYPE, male, labelled as follows: “Marengo Co., ALABAMA Prairies
s. Demopolis June, 1935 A. F. Archer; loan from UMMZ; HOLOTYPE Evarthrus hypherpi-
formis R. Freitag(red label).” UMMZ.
Body length 18.0 mm, width 7.4 mm. Form somewhat flat, with parallel sides.
Microsculpture of head between eyes composed of isodiametric and amorphic meshes;
disc of pronotum with amorphic meshes; intervals of elytra with isodiametric bead-like
meshes. Integument of dorsum slightly glossy.
Head length 2.0 mm, width 4.1 mm; frontal groove deep and broadly impressed, posterior
halves slightly directed laterally. Penultimate article of labial palpus with five setae.
Pronotum length 5.3 mm, width 5.6 mm; form quadrate in outline as in fig. 37; disc
distinctly flattened in centre; sides not prominent, constricted moderately anteriorly slightly
posteriorly, slightly sinuate in front of posterior angles; posterior angles not prominent,
slightly obtuse, somewhat sharp ; anterior transverse impression complete; basal lateral fovea
with sides not continuous posterio-medially, central depressed portion flattened and some-
what rugose, extension from inner side to middle longitudinal line represented by a small
roughly sculptured area; lateral bead same width throughout length. Apex of prostemal
process extended far beyond middle coxae, longitudinal groove obsolete. Anterior faces of
middle femora with five setae on one, six setae on the other.
Elytra length 10.7 mm, width 7.4 mm; sides parallel, obsoletely sinuate apically ; intervals
distinctly convex; striae with small but distinct punctures anteriorly, obsoletely punctate
posteriorly. Third interval of each elytron with three setae.
146
Freitag
Male genitalia (fig. 110): median lobe moderately arcuate, with ventral medial bump;
apical blade with apex deflected dorsally at a sharp right angle; right paramere very short,
markedly tapered apically; internal sac serrulate apically, apical sclerite elongate, pale
colored plate.
Derivation of specific name. - The habitus of specimens of hypherpiformis is vaguely like
that of individuals of some species of the subgenus Hypherpes of the genus Pterostichus.
Variation among paratypes (four males, Alabama, Mississippi). — Body length 17.7 — 18.5
mm. The number of setae in the third interval of the elytron ranges from three to Five. The
genitalia of one male was examined. It resembled that of the holotype in all respects.
Disposition of type material. — The holotype and one paratype are in the collections of
the UMMZ, and the three other paratypes are in the collections of AMNH, CU, and UASM.
Geographical distribution (fig. 132). — This species is found in Alabama and Mississippi
only. I have seen Five specimens from the following localities.
United States - ALABAMA: Dallas County: Hazen (AMNH). Marengo County: south of Demopolis (UASM, UMMZ).
MISSISSIPPI: Oktibbeha County: Agriculture College (CU).
The sodalis Group
Characteristics. — Penultimate article of labial palpus with five to seven setae. Pronotum
subcordiform, sides moderately or strongly constricted posteriorly; anterior transverse im-
pression complete or incomplete. Prostemal process with shallow or obsolete longitudinal
groove, without setae at apex. Middle femur with 5 — 11 setae on anterior face. Male
genitalia with median lobe slightly or moderately arcuate; internal sac with apical sclerite
light amorphic plate usually with darker basal tooth, tooth sometimes unsclerotized. The
species sodalis, parasodalis, furtivus, altemans and iowensis are included in this group. They
occur across northeastern United States from New Jersey west to South Dakota, Nebraska,
and Kansas, and south to northern Alabama and Arkansas.
Evarthrus sodalis LeConte, 1848
Frontispiece and Figures 38-48, 64, 111, 133
Feronia (Molops) sodalis LeConte, 1848:349. LECTOTYPE (here selected) a male, labelled
as follows: “yellow disc; Type 5659; E. sodalis Lee. orbatus Lee.” MCZ. TYPE LOCAL-
ITY, Illinois. — LeConte, 1852:229 ( Evarthrus ). — LeConte, 1870:5. — LeConte, 1873:
318. - Schaupp, 1880:49. - Blatchley, 1910:101. — Casey, 1918:356 (Eumolops). -
Casey, 1920:197 ( Evarthrinus ). - Leng, 1920:57 (Eumolops). — Leonard, 1926:222. -
Csiki, 1930:672 (Pterostichus).
Feronia (Molops) colossus LeConte, 1848:343. LECTOTYPE (here selected) a male, labelled
as follows: “yellow disc; colossus 4.” MCZ. TYPE LOCALITY, Missouri. NEW COMBI-
NATION. - LeConte, 1852:233 (Evarthrus). - LeConte, 1863a:8. — LeConte, 1873:
318. - Schaupp, 1880:49. - Casey, 1918:356 (Eumolops). - Leng, 1920:57. - Csiki,
1930:672 (Pterostichus).
Feronia (Molops) corax LeConte, 1848:347. LECTOTYPE (here selected) a male, labelled as
follows: “green disc; Type 5661; E. corax Lee.” MCZ. TYPE LOCALITY, near Long’s
Peak. — LeConte, 1852:229 (Evarthrus). - Motschulsky, 1865:261. - LeConte, 1873:
318. - Schaupp, 1880:49. - Casey, 1918:357. - Leng, 1920:57. - Csiki, 1930:672
(Eumolops).
Revision of Evarthrus
147
Feronia (Pterostichus) vagans LeConte, 1848:349. LECTOTYPE (here selected) a male,
labelled as follows: “yellow disc; Type 5664; E. vagans Lee.” MCZ. TYPE LOCALITY,
Ohio. NEW SYNONYMY. - LeConte, 1852:229 {Evarthrus). - LeConte, 1863a:8. -
LeConte, 1873:320. - Leng, 1920:57.
Evarthrus fatuus LeConte, 1852:233. LECTOTYPE (here selected) a male, labelled as
follows: “yellow disc; Type 5060; E. fatuus LeC.” MCZ. TYPE LOCALITY, Iowa. —
LeConte, 1873:318 {Evarthrus). - Schaupp, 1880:49. - Casey, 1918:356 {Eumolops). -
Casey, 1920:197 {Evarthrinus). - Leng, 1920:57 {Eumolops). — Csiki, 1930:672
{Pterostichus). - Lindroth, 1966:474.
Evarthrus furtivus; Blatchley, 1910:101 (not LeConte).
Evarthrinus (Evarthrops) retractus Casey, 1920:197. HOLOTYPE, female, labelled as fol-
lows: “L. CASEY bequest 1925; TYPE USNM 47132; retractus Csy.” USNM. TYPE
LOCALITY, “probably Indiana.” NEW SYNONYMY. - Leng and Mutchler 1927:10
{Evarthrinus). — Csiki, 1930:673 {Pterostichus).
Evarthrinus inflatipennis Casey, 1924:78. HOLOTYPE, female, labelled as follows: “111.;
CASEY bequest 1925; TYPE USNM 47133; inflatipennis Csy.” USNM. TYPE LOCAL-
ITY, near Chicago, Illinois. NEW SYNONYMY. - Leng and Mutchler 1927:10 {Evar-
thrinus). - Csiki, 1930:673 {Pterostichus).
Eumolops sulcata Casey, 1918:355. HOLOTYPE, male, labelled as follows: “Fla; CASEY
bequest 1925; TYPE USNM 47134; sulcata Casy.” USNM. TYPE LOCALITY. FLORIDA
(this locality is probably incorrect). NEW SYNONYMY. — Casey, 1920:196 {Evarthrinus).
- Leng, 1920:57 {Eumolops). - Csiki, 1930:672 {Pterostichus).
Evarthrus lodingi Van Dyke, 1926:118. HOLOTYPE, male, labelled as follows: “Monte
Sano, Ala. Madison Co. 6.9.1 1.” H. P. Loding; Van Dyke Collection. CAS. TYPE LOCAL-
ITY, Monte Sano, Alabama. NEW COMBINATION. - Csiki, 1930:673 {Pterostichus). -
Leng and Mutchler, 1933:13 {Evarthrus).
Recognition. — The species altemans, furtivus, iowensis and parasodalis are remarkably
similar to sodalis in their external non-genitalic structures. Although some specimens of
sodalis can be distinguished from individuals of the first four species by external features,
the male genitalia are the only reliable diagnostic character of sodalis. The apical blade of
the median lobe is elongate and narrow in sodalis while it is shorter and broader in the
other species (fig. Ill cf. figs. 112, 113, 114, 115).
Description. - Body length 12.4 - 20 mm. Form robust.
Microsculpture: head between eyes with flattened amorphic meshes, disc of pronotum
with highly sinuous entwined lines, or transversely directed lines, or meshes transversely
stretched; intervals of elytra with isodiametric bead-like or flattened meshes, or highly sinu-
ous lines.
Head moderately glossy; frontal grooves of average depth, not sharply defined, straight or
with slight bend in middle, convexity directed medially. Penultimate article of labial palpus
with five to seven setae.
Pronotum moderately glossy; shape subcordiform in outline as in figs. 38 - 48; disc of
average convexity; sides moderately or strongly produced, constricted moderately anteriorly
strongly posteriorly, sinuation in front of posterior angles well marked, moderate, slight, or
148
Freitag
absent: posterior angles produced or not, broadly obtuse or slightly acute; anterior transverse
impression complete or incomplete; basal lateral fovea with sides usually but not always,
continuous posteriorly; lateral bead same width throughout. Prostemal process with shallow,
broadly excavated or obsolete longitudinal groove. Middle femur with five to eight setae on
anterior face.
Elytra of males moderately glossy or iridescent, females dull; apical sinuation slight,
obsolete, or absent; intervals flat, moderately convex, or highly convex; striae of average
depth, punctate anteriorly, impunctate or obsoletely punctate posteriorly.
Male genitalia (fig. Ill) with median lobe slightly arcuate, angle broadly obtuse; apical
blade elongate, right side and apex deflected dorsally; right paramere of average length
extending to halfway point of median lobe, rather broad, slightly tapered apically, apex
broadly rounded; internal sac with serrulate apical field, apical sclerite light apically with
darker basal tooth, or tooth not sclerotized. The genitaha of 21 males were examined in
detail.
Stylus of female ovipositor elongate, slightly tapered apically.
Geographical variation and subspecies. — There are three populations which are more or
less distinct in several structural features.
In Kansas and proximal areas specimens referred to the nominal species colossus are dis-
tinguished from typical sodalis by larger body size, more prominent posterior angles of the
pronotum, and the absence of an apical sclerite of the internal sac of the median lobe of the
male. Between these populations there are populations which have intermediate character-
istics and therefore all of these groups appear to be the same species. Because of the differ-
ences I recognize a subspecies s. colossus west of the Mississippi River, and mainly south of
the Missouri River, and an eastern subspecies mainly east of the Mississippi River and north
of Alabama and Tennessee.
The third distinct form, formerly the species lodingi, occurs in Tennessee, and northern
Alabama. It is distinguished from s. sodalis and s. colossus by highly convex and iridescent
elytral intervals of the males. This iridescence is because of the microsculpture which is com-
posed of numerous highly sinuous lines, placed very close to one another. Both s. sodalis
and s. colossus have isodiametric meshes forming the microsculpture on the elytral intervals
of the male. Furthermore, specimens from Alabama and Tennessee are generally larger than
s. sodalis specimens. They also differ from s. colossus individuals by having an apical sclerite
in the internal sac of the median lobe of the male. The few sodalis specimens which I have
seen from Kentucky appear intermediate in structural features between those of Tennessee
and Alabama, and Indiana, Illinois and Ohio. I believe the Alabama and Tennessee specimens
form a third subspecies, s. lodingi.
Notes on synonymy. — The lectotype of colossus LeConte is a sodalis specimen larger
than the average size and of the form which inhabits Kansas and Missouri.
The lectotype of corax LeConte, is an average sodalis specimen that inhabits western
areas of this species range; and it is characterized by the small prominent basal angles of the
pronotum.
The lectotype of vagans LeConte, is a sodalis specimen with very broadly rounded basal
angles of the pronotum, which is the common condition of specimens in northern Ohio.
Revision of Evarthrus
149
The lectotype of fatuus LeConte is a sodalis specimen with rectangular hind angles of the
pronotum as in fig. 43. This form is common in eastern Iowa.
The type specimen of inflatipennis Casey is a sodalis specimen that is average for the form
found in Illinois but slightly smaller in body size.
The type specimen of sulcata Casey is a sodalis specimen that is of the common form and
size found in Illinois.
The type specimen of lodingi Van Dyke is a sodalis specimen of the common form
inhabiting northern Alabama.
Collecting notes. — Specimens of s. sodalis and s. colossus are found in open grassy places
under cover. G. E. Ball collected specimens of s. sodalis beside railroad tracks under ties near
Ithaca, New York. I have taken s. colossus from under boards in pasture and abandoned
farmyards in Kansas. The subspecies s. lodingi is forest adapted and occurs in leaf litter.
The gut of a female s. colossus specimen, which I examined, was full of the remains of
ants.
Geographical distribution (fig. 133). — This species is widespread in northeastern United
States ranging from Pennsylvania west to Kansas and Nebraska, south to northern Mississippi
and Alabama, and north to Duluth, Minnesota.
E. s. sodalis LeConte
I have seen 664 specimens from the following localities.
Canada - ONTARIO: Point Pelee (Lindroth 1966).
United States - ILLINOIS: Adams County: five miles east of LaPrairie (CNHM). Bureau County: Princeton (UMMZ).
Champaign County: Champaign (CNC, INHS, RTB, UASM); Seymour (MCZ); Urbana (CNHM, INHS, RTB, RU, UMMZ,
UW). Cook County: Carle Woods (CNHM); Chicago (CAS, CNHM, CU, UASM, UMMZ, USNM, UW); Evanston (ASNP, CAS,
UMMZ); Glencoe (CNHM, UMMZ); Palos Park (CAS, CNHM, UMMZ); River Forest (CNHM); River Grove (USNM); Summit
(CNHM, INHS); West Northfield (MCZ); Willow Springs (CAS, CNHM, CNC, UMMZ). DuPage County: Glen Ellyn (CNHM).
Lake County: Fort Sheridan (UMMZ); Lake Zurich (RTB); Ravinia (UMMZ). LaSalle County: Ottawa (RTB). McHenry
County: Algonquin (INHS). McLean County: Bloomington (CNHM, CU, USNM); Normal (INHS). Mason County: Havanna
(INHS). Ogle County: Oregon (UMMZ). Peoria County: Peoria (INHS). Putnam County: (INHS). Richland and Lawrence
County: Wabash Valley (CM, MCZ, USNM). Rock Island County: Rock Island (UMMZ). Sangamon County: Springfield
(CNHM). Vermilion County: Danville (INHS); Kickapoo State Park (RTB); Oakwood (INHS). Washington County: Dubois
(INHS). Will County: Joliet (CNHM), Winnebago County: Rockford (CAS). Counties not determined: Edgebrook (UMMZ);
Somerset (INHS). INDIANA: Cass County: (MCZ). Gibson County: (UMMZ); Princeton (USNM). Jefferson County: Clifty
Falls State Park (GEB); Hanover (UMMZ). Knox County: Vincennes (USNM); Wheatland (UMMZ). Kosciusko County:
Winona Lake (UMMZ). Lagrange County: Lagrange (UMMZ). Marion County: (MCZ, UP). Monroe County: Bloomington
(UMMZ). Posey County: (CNHM); Mount Vernon (CNHM). Wells County: LaFayette (UMMZ). Warren County: Pine
(CNHM). Wells County: Blufton (UMMZ). County not determined: Indiana Dunes State Park: (RTB). IOWA: Benton
County: (UMMZ). Clayton County: McGregor (UMMZ). Des Moines County: Burlington (MCZ). Hamilton County: Randall
(CNC). Johnson County: Iowa City (CAS, MCZ, UMMZ, USNM); Solon (USNM). Linn County: Cedar Rapids (UMMZ).
Tama County: Traer (ISU). KENTUCKY; Edmonson County: Bee Spring (MCZ); Mammoth Cave National Park (TCB).
Fayette County: Lexington (TCB). Franklin County: Stony Creek, north of Frankfort (UMMZ). Hardin County: Summit
(CNHM). Harlan County: Cumberland Gap (MCZ). Henderson County (CNC). MICHIGAN: Jackson County: Jackson
(TH). Kalamazoo County: Climax (UMMZ); Gull Lake Biology Station (TH). Lenawee County: Adrian (TH). Munroe
County: (UMMZ). Oakland County: (UMMZ). Wayne County: Detroit (USNM). MINNESOTA: Houston County: Caledonia
(UMMZ). St. Louis County: Duluth (MCZ). MISSISSIPPI: Tishomingo County: Cook’s lodge near Iuka (UMMZ); six miles
east of Iuka (FDPI). MISSOURI: Buchanan County: St. Joseph (USNM). NEW JERSEY: Morris County: Lincoln Park
(CNHM). NEW YORK: Chautauqua County: Findley Lake (GEB); Mayville (GEB); Pendergast Creek, near Lake Chau-
tauqua (GEB). Erie County: Buffalo (GEB); Hamburg (CAS). Tompkins County: Ithaca (GEB, FDPI, UA); Turner Hill
150
Freitag
(GEB). Yates County: Dresden (UMMZ). Counties not determined: Van Cort’dt Park (CU); Windom (CU). OHIO: Allen
County: Lima (UMMZ, USNM). Columbiana County: Salineville (CAS, CU). Cuyahoga County: Cleveland (MCZ). Darke
County: (CAS); Beamville (UMMZ). Fairfield County: Millersport (CM). Franklin County: Columbus (CAS, MCZ, RU).
Hamilton County: (CNHM); Cincinnati (ANSP, CAS, UMMZ). Licking County: Alexandria (RTB). Mercer County: Mendon
(UMMZ). Ottawa County: Lakeside (UMMZ); Put-in-Bay, South Bass Island (UMMZ). Preble County: Eaton (UMMZ).
PENNSYLVANIA; Allegheny County: (CM, CU); Pittsburgh (CM, RU). Erie County: (CM). Forest County: Cook’s Forest
(CM). Warren County: (UMMZ). Westmoreland County: Jeanette (CM). TENNESSEE: Knox County: Knoxville (CNC);
30 miles west of Knoxville (AMNH). WISCONSIN: Bayfield County: (MCZ). Dane County: (UW); Madison (UASM, UW).
Dodge County: Beaver Dam (CAS, MCZ, TE, USNM). Green County: Albany (CAS, CNHM); Brodhead (UMMZ). Jefferson
County: Fort Atkinson (GEB). Milwaukee County: (UW); Milwaukee (CAS). Racine County: Burlington (CNHM). Wal-
worth County: Walworth (CNHM). County not determined: Rautubug (MCZ).
E. s. colossus LeConte
I have seen 151 specimens from the following localities.
United States - ARKANSAS: (UASM). IOWA: O’Brien County: four miles east of Sanborn (I SU). Woodbury County:
Sioux City (UMMZ). KANSAS: Chase County: (UK). Doniphan County: Wathena (UASM, USNM). Douglas County: (UK);
five miles north of Baldwin City (DL, RF); Lawrence (CAS, MCZ, UK, UMMZ). Franklin County: (UMMZ). Johnson
County: Mission (UK). Lawrence County: (UMMZ). Leavenworth County: Leavenworth (CAS, CNHM); Tonganoxie
(MCZ). Linn County: (UK). Pottawatomie County: Onaga (UK). Reno County: (ANSP). Riley County: (USNM); Man-
hattan (KSU, USNM). Shawnee County: Topeka (USNM). Wilson County: Benedict (CAS). Woodson County: two miles
east of Yates Centre (DL, RF). MISSOURI: Boone County: Columbia (CNHM). Buchanan County: St Joseph (USNM).
Clinton County: Cameron (CAS); Lathrop (CNC). Jackson County: Kansas City (UK). Pettis County: Sedalia (CNHM). St.
Louis (CAS, CNHM, USNM). County not determined: Pickle Springs (UMMZ). NEBRASKA: Cedar County: Randolph
(MCZ). Knox County: Creighton (CAS).
E. s. lodingi Van Dyke
I have seen 53 specimens collected in the following localities.
United States - ALABAMA: Jackson County: Point Rock (UMMZ). Madison County: Monte Sano (CAS, MCZ, UASM,
UK, UMMZ, USNM); Monte Sano State Park (CNHM). TENNESSEE: Cumberland County: Grassy Cove (CAS, UMMZ).
Davidson County: Madison (CU); Nashville (AMNH, USNM). Maury County: Columbia (ANSP). County not determined:
Cedar Glade (USNM).
Evarthrus parasodalis new species
Figures 49, 112, 133
Recognition. — The following combination of structures characterizes this species: prono-
tum with sides not sinuate or obsoletely sinuate in front of posterior angles, posterior angles
not prominent and broadly obtuse; apex of median lobe of male genitalia short and broad,
right paramere rather narrow apically; range, Arkansas. These features distinguish parasodalis
from the similar sodalis lodingi which has the following corresponding features: pronotum
with more produced sides, distinctly sinuate in front of posterior angles; apex of median
lobe elongate and narrow, right paramere broader apically; known from northern Alabama
and Tennessee.
Description. — HOLOTYPE, male, labelled as follows: “Washington Co., Ark. 1962 Trap
A 29— VI; HOLOTYPE Evarthrus parasodalis R. Freitag (red label).” MCZ.
Body length 16.9 mm, width 7.1 mm. Form typical of sodalis group.
Microsculpture: head between eyes and disc of pronotum with isodiametric and amorphic
meshes; intervals of elytra with transversely stretched meshes. Integument of dorsum moder-
ately glossy.
Revision of Evarthrus
151
Head length 1 .9 mm, width 3.7 mm; frontal groove deep, broadly impressed, middle bend
with convexity directed medially. Penultimate article of labial palpus with three medial and
three apical setae.
Pronotum length 4.8 mm, width 5.9 mm; form subcordate in outline as in fig. 49; disc of
average convexity; sides not prominent, constricted slightly anteriorly, moderately posteri-
orly, not sinuate in front of posterior angles; posterior angles not prominent, broadly obtuse;
anterior transverse impression complete, obsoletely impressed medially; basal lateral foveae
with sides not continuous posterio-medially, amorphic depression medially beside inner
side; lateral bead gradually broadened posteriorly. Prostemal process with obsolete longi-
tudinal groove. Anterior faces of middle femora with eight setae on one and 1 1 setae on the
other.
Elytra length 10.2 mm, width 7.1 mm; sides slightly produced, obsoletely sinuate apically;
intervals raised with flattened centres; striae deep with rather small indistinct punctures
throughout.
Male genitalia (fig. 1 12): median lobe moderately arcuate; apical blade short, broad, apex
evenly rounded; right paramere elongate, extended to apical half of median lobe, strongly
tapered apically; internal sac with serrulate field apically, apical sclerite light amorphic plate
with dark basal tooth.
ALLOTYPE, female, labelled as follows: “Washington Co. Ark. VII — 1 6 — 1960; Forest
leaf litter; Otis and Maxine Hite; ALLOTYPE Evarthrus parasodalis R. Freitag (green label).”
MCZ.
Body length 16.8 mm, width 7.1 mm. Form same as in holotype except pronotum with
sides more strongly constricted posteriorly.
Microsculpture on head between eyes and disc of pronotum same as in holotype; intervals
of elytra with raised bead-like isodiametric meshes.
Pronotum moderately glossy; length 4.5 mm, width 5.6 mm.
Elytra dull; length 10.5 mm, width 7.1 mm.
Stylus of ovipositor elongate and quite tapered apically.
Derivation of specific name. — This species is closely related to sodalis which is what the
name parasodalis connotes.
Variation among paratypes (41 males, 40 females, Arkansas). — Total length 15.6 — 19.3
mm. The genitalia of three males were examined in detail and in all respects resemble those
of the holotype. One female specimen collected at Hot Springs has produced and rather
sharp posterior angles of the pronotum.
Disposition of type material. — The holotype and allotype are in the collections of the
MCZ. The paratypes are in the collections of the following: CU, INHS, RF, RTB, UASM and
UA.
Collecting notes. — I collected a specimen of parasodalis in deciduous forest leaf fitter on
a hillside near Fayetteville, Arkansas.
Geographical distribution (fig. 133). — This species is known from Arkansas, only. I have
seen 83 specimens from the following localities.
United States - ARKANSAS: Conway County: (UA). Franklin County: (UA). Garland County: Hot Springs (UMMZ).
Montgomery County: north of Mount Ida (RTB). Washington County: (UA, UASM, CU);Cove Creek, 27 miles northwest
of Fayetteville (RF);Covc Creek Valley (CU).
152
Freitag
Evarthrus furtivus LeConte, 1852
Figures 50-51 , 113, 133
Evarthrus furtivus LeConte, 1852:234. LECTOTYPE (here selected) a male, labelled as
follows: “white disc; Type 5662; E. furtivus Lee.” MCZ. TYPE LOCALITY, here re-
stricted to Virginia. — LeConte, 1863a:8 ( Evarthrus ). - LeConte, 1873:319. — Schaupp,
1880:49. - Casey. 1918:355 ( Eumulops ). - Casey, 1920:195 ( Evarthrinus ). - Leng,
1920:57 ( Eumolops ). - Csiki, 1930:672 ( Pterostichus ).
Recognition. — Specimens of furtivus are extremely difficult to distinguish from indi-
viduals of s. sodalis by their external nongenitalic structures. The posterior angles of the
pronotum of furtivus are less broadly rounded than those of s. sodalis in southwestern
Pennsylvania where their geographical ranges overlap. For a certain identification however,
it is necessary to examine the male genitalia.
Description. - Body length 13.0 — 1 7.0 mm. Form typical of the sodalis group with sides
of pronotum and elytra somewhat convex.
Microsculpture: head between eyes and disc of pronotum with highly sinuous entwined
lines, often almost effaced; intervals of elytra with bead-like or more flattened isodiametric
meshes. Integument of dorsum moderately glossy; elytra slightly glossy in some specimens.
Head: frontal grooves not deep, not sharply defined, fairly straight, slightly oblique
toward one another. Penultimate article of labial palpus with five setae.
Pronotum subcordiform in outline as in figs. 50 — 51; disc of average convexity; sides
slightly or strongly produced, constricted moderately anteriorly, moderately or strongly
posteriorly, obsoletely or distinctly sinuate in front of posterior angles; posterior angles
produced or not produced, broadly obtuse ; anterior transverse impression incomplete; basal
lateral fovea with sides usually continuous posteriorly; lateral bead not broadened posteri-
orly. Prostemal process with shallow or obsolete longitudinal groove. Middle femur with five
or six setae on anterior face.
Elytra with apical sinuation obsolete; intervals slightly convex or flat; striae distinct, not
deep, indistinctly punctate anteriorly, obsoletely or impunctate posteriorly.
Male genitalia (fig. 1 13): median lobe slightly arcuate, angle broadly obtuse, apical blade
short, broad with apex evenly rounded; right paramere extended to halfway point of median
lobe, markedly tapered apically: internal sac with apical serrulate field, apical sclerite light,
broad, somewhat triangular with apical end graded into serrulate field around genital open-
ing. The genitalia of 1 1 males were examined.
Stylus of female ovipositor elongate, moderately tapered apically.
Geographical variation. - Specimens possessing pronota with somewhat sharp posterior
angles are common in southwestern Pennsylvania, but decrease in number southward where
specimens with more broadly rounded posterior angles are more numerous.
Geographical distribution (fig. 133). — This species ranges from southern Pennsylvania
south to Virginia. I have seen 1 1 3 specimens collected in the following localities.
United States - DISTRICT OF COLUMBIA: Washington (USNM). MARYLAND: Montgomery County: (USNM). NEW
JERSEY: Gloucester County: Malage (USNM). PENNSYLVANIA: Allegheny County: (CM); Pittsburg (CM). Cumberland
County: New Cumberland (CAS, CU, MCZ); Shippensburg (UMMZ). Dauphin County: Harrisburg (CU, MCZ). Fayette
County: Ohiopyle (CM). Philadelphia County: Germantown (ANSP); Philadelphia (CAS, MCZ). Westmoreland County:
Jeanette (CM). Counties not determined: Inglenook (CAS); Rockville (ANSP, CAS, MCZ). VIRGINIA: Arlington County:
Rosslyn (MCZ). Fairfax County: Mount Vernon (USNM). Henrico County: Richmond (AMNH). Nelson County: (USNM).
Spotsylvania County: Fredricksburg (MCZ). Counties not determined: Blackpond (MCZ, USNM); Edsall (USNM): Glen-
carlyn (USNM); Merdon (USNM). WEST VIRGINIA: Pocahontas County: Swamp Creek (TCB).
Revision of Evarthrus
153
Evarthrus altemans Casey, 1920
Figures 52, 114, 134
Evarthrinus (Evarthr ops) altemans Casey, 1920:196. HOLOTYPE, male, labelled as follows:
“la; CASEY bequest 1925; TYPE USNM 47 1 3 1 ; altemans Csy.” USNM TYPE LOCAL-
ITY, Keokuk, Iowa. PARA TYPE, female, labelled as follows: “Iowa, CASEY bequest
1925; altemans -2 PARATYPE 47131.” USNM. - Leng and Mutchler 1927:10 {Evar-
thrinus). - Csiki, 1930:673 ( Pterostichus ).
Recognition. — The similar sodalis colossus is distinguished from altemans by structural
details of the male genitalia and by the more laterally produced posterior angles of the
pronotum {figs. 45—47 cf. fig. 52). These species are largely allopatric.
Description. — Body length 13.4 — 18.4 mm. Form robust notably constricted at base of
pronotum.
Microsculpture: head between eyes with highly sinuous entwined lines, occasionally
amorphic meshes formed; disc of pronotum with highly sinuous lines or transversely
stretched meshes, often partially effaced; intervals of elytra with isodiametric meshes, bead-
like in females, flatter in males.
Head moderately glossy; frontal grooves of average depth, somewhat broad, generally
straight, slightly oblique toward one another. Penultimate article of labial palpus with six
setae.
Pronotum moderately glossy; shape somewhat cordiform in outline as in fig. 52; disc of
average convexity; sides markedly produced, moderately constricted anteriorly, strongly and
sharply constricted posteriorly, markedly sinuate in front of posterior angles; posterior
angles produced, almost right or slightly obtuse; anterior transverse impression incomplete;
basal lateral fovea with sides usually continuous posteriorly, medial side with anterior end
directed laterally; lateral bead not broad posteriorly. Prostemal process with shallow or
obsolete longitudinal groove. Middle femur with six to nine setae on anterior face.
Elytra of males slightly glossy, females dull; apical sinuation obsolete or absent; intervals
slightly convex, almost flat; striae of average depth, with small distinct punctures anteriorly,
obsoletely punctate or impunctate posteriorly.
Male genitalia (fig. 1 14): median lobe moderately arcuate, broadly obtuse; apical blade
short, broad, with apex very broadly rounded, almost truncate slightly deflected dorsally;
right paramere extended to apical half of median lobe, slightly tapered apically; internal sac
with serrulate field apically, preapical sclerite light near genital opening with darker basal
tooth. The genitalia of 20 males were examined.
Stylus of female ovipositor elongate, slightly tapered apically.
Collecting notes. — Members of this species are found under cover in open grassy places.
Geographical distribution (fig. 134). — Members of this species are common in a relatively
restricted range in Iowa and margins of peripheral states. I have seen 830 specimens from
the following localities.
United States - ILLINOIS: Adams County: five miles northeast of La Prairie (CNHM). Hancock County: Pilot Knob
State Park (ISU). Macoupin County: Chesterfield (RTB). Pike County: Rockport (CAS). IOWA: Appanoose County:
Moulton (UMMZ). Boone County: Boone (ISU); Ledges State Park (ISU). Crawford County: (MCZ); Denison (AMNH).
Dallas County: Perry (ISU). Davis County: (CAS). De Moines County: Burlington (MCZ). Dickenson County: Cayler
Prairie (ISU); Lakeside Laboratory (ISU). Hamilton County: Blairsburg (ISU); five miles south of Stanhope (ISU); Randall
154
Freitag
(ISU). Hardin County: Iowa Falls (CNHM). Henry County: Mount Pleasant (CAS, UASM, UMMZ). Johnson County: Iowa
City (ANSP, MCZ, UASM, USNM). Lee County: Fort Madison (MCZ). Linn County: Cedar Rapids (UMMZ); Palisades
(USNM). Lucas County: Chariton (USNM). Marshall County: State Centre (CU); ten miles west of Marshall Town (ANSP).
Montgomery County: Red Oak (ISU). O’Brien County: four miles east of Sanborn (ISU). Page County: Shenandoah (ISU).
Palo Alto County: Ruthuen (UMMZ). Plymouth County: Le Mars (ISU). Pochahontas County: Kaslow (ISU). Polk
County: Des Moines (ISU). Pottawattamie County: Council Bluffs (ISU). Sioux County: Hawarden (VMK). Story County:
Ames (AU, CAS, CU, ISU, MCZ, MSU, UMMZ, USNM, UW); four miles east of Gilbert (ISU); Maxwell (ISU); Nevada
(UASM); Soper’s Mill Dam near Gilbert (ISU). Tama County: Traer (ISU, USNM). Van Buren County: (ISU). Wayne
County: Lineville (ISU). Webster County: three miles west of Dayton (ISU). Winnebago County: Forest City (UMMZ).
Thompson (ISU). Counties not determined: Boonsboro (MCZ); Harold (CU); Lake Okoboji (USNM). MINNESOTA:
Lincoln County: Lake Benton (VMK). MISSOURI: Livingston County: six miles north of Chillocothe (ISU). St Louis
County: (CU); Overland (CAS); St. Louis (USNM). County not determined: Onandaga Cave (UMMZ). SOUTH DAKOTA:
Brookings County: Brookings (VMK); White (VMK). Deuel County: Gary (VMK). WISCONSIN: Bayfield County: (MCZ).
Evarthrus iowensis new species
Figures 53, 115, 134
Recognition. - The following combination of characteristics is diagnostic for the species
iowensis: small size; pronotum usually with complete anterior transverse impression; elytra
with first two umbilicate punctures of umbilicate series with normally raised areas around
them, and third puncture as large as either first or third; and form of male genitalia. Speci-
mens of alternans generally resemble individuals of iowensis but are distinguished by the
larger body size, sides of pronotum more strongly constricted posteriorly, incomplete
anterior transverse impression of pronotum, and relatively longer right paramere of male
genitalia.
The species substriatus and constrictus also strikingly resemble iowensis. In substriatus
and constrictus however the first and second anterior umbilicate punctures have areas be-
tween them which are flatter than the normal condition, and the third puncture is distinctly
larger than the first two. In addition the male genitalia are different (fig. 115 cf. figs. 1 16,
117 and 118).
Description. — HOLOTYPE, male, labelled as follows: “Iowa City, Iowa 5 — 15 Buchanan;
Loan from USNM: HOLOTYPE Evarthrus iowensis R. Freitag (red label).” USNM.
Body length 11.7 mm, width 5.0 mm. Form less robust than other species of sodalis
group.
Microsculpture of head between eyes, disc of pronotum, and intervals of elytra with
highly sinuous somewhat sparsely distributed lines, partially effaced. Integument of dorsum
glossy.
Head length 1.4 mm, width 2.8 mm; frontal grooves shallow and broad, slightly curved
with middle bend directed medially. Penultimate article of labial palpus with two medial
and three apical setae.
Pronotum length 3.2 mm, width 4.1 mm; form subcordate in outline as in fig. 53; disc of
average convexity; sides prominent, constricted moderately anteriorly and strongly posteri-
orly, distinctly sinuate in front of posterior angles; posterior angles prominent, sharp,
slightly obtuse; anterior transverse impression complete, distinctly impressed throughout;
basal lateral fovea with sides continuous posteriorly; lateral bead not broad posteriorly.
Prosternal process with longitudinal groove obsolete. Middle femur with five setae on
anterior face.
Revision of Evarthrus
155
Elytra length 7.1 mm, width 5.0 mm; sides slightly produced, not sinuate apically;
intervals almost flat; striae of average depth, indistinctly punctate throughout. Third inter-
val of each elytron with two setae.
Male genitalia (fig. 1 15): median lobe moderately arcuate; apical blade short, broad, apex
evenly rounded, almost truncate; right paramere not extended to apical half of median lobe,
strongly tapered apically; internal sac with apical serrulate field, apical sclerite light with
darker basal tooth.
ALLOTYPE, female, labelled as follows: “Iowa City, Iowa IV. 14 Wickham; Wickham
Collection 1933; loan from USNi/i; ALLOTYPE Evarthrus iowensis R. Freitag (green
label).” USNM
Body length 1 1 .5 mm, width 4.9 mm. Form same as in holotype.
Microsculpture on head between eyes and disc of pronotum same as in holotype; intervals
of elytra with raised, bead-like isodiametric meshes.
Pronotum glossy; length 3.0 mm, width 3.1 mm.
Elytra slightly glossy; length 7.1 mm, width 4.9 mm.
Stylus of ovipositor slightly tapered apically, apex broadly rounded.
Derivation of specific name. — This species is named iowensis because much of its range is
in Iowa.
Variation among paratypes (16 males, 15 females, Iowa, Minnesota, South Dakota). —
Total length 1 1 .2 — 13.9 mm. The genitalia of five males were examined and varied little or
not at all from that of the holotype. The elytra of females are duller than those of the
males. The number of setae in the third interval of an elytron varies from one to three.
Disposition of type material. — The holotype and allotype are in the collections of the
USNM. The paratypes are in the following collections: CAS, CU, ISU, KLE, MCZ, UASM,
UMMZ, USNM, and VMK.
Collecting notes. — V. M. Kirk has collected specimens of iowensis in corn fields in south-
eastern South Dakota.
Geographical distribution (fig. 134). — This species is confined to Iowa, Minnesota and
South Dakota. I have seen 33 specimens from the following localities.
United States - IOWA: Dickinson County: Cayler Prairie (ISU, UASM). Howard County: Elma (USNM). Johnson
County: Iowa City (USNM). Story County: Ames (ISU). Woodbury County: Sioux City (UMMZ). MINNESOTA: Olmsted
County: Rochester (CU). SOUTH DAKOTA: Brookings County: Brookings (VMK). Hutchinson County: Menno (VMK).
Yankton County: Yankton (VMK)
The substriatus Group
Characteristics. — Penultimate article of labial palpus with four or five setae. Pronotum
with sides strongly constricted posteriorly; anterior transverse impression complete or not
complete. Prosternal process with shallow or obsolete longitudinal groove without setae
apically. Middle femur with four to seven setae on anterior face. First two anterior punc-
tures of umbilicate series of elytra without normally raised ridges around them, third
umbilicate puncture distinctly larger than first two. Median lobe of male genitalia strongly
arcuate; internal sac with elongate narrow amorphic plate.
This group includes substriatus and cons trie tus whose aggregate geographical range in-
cludes the grasslands of central United States and northwestern Mexico.
156
Freitag
Evarthms substriatus LeConte, 1848
Figures 54. 77-78, 116-117, 134
Feronia (Molops) substricita LeConte, 1848:344. LECTOTYPE (here selected) a female,
labelled as follows: “green disc; Type 5616; E. substriatus Lee.” MCZ. TYPE LOCALITY,
near the Rocky Mountains. — LeConte, 1852:233 ( Evarthrus ). — LeConte, 1858:28. -
LeConte, 1863a:8. - LeConte, 1873:319. - LeConte, 1876:519. - Schaupp, 1880:49. -
Casey 1918:343 ( Anaferonia ). - Leng, 1920:56. — Csiki, 1930:671 ( Pterostichus ).
Evarthrus latebrosus LeConte, 1852:233. LECTOTYPE (here selected) a male, labelled as
follows: “green disc; Type 5617; E. latebrosus Lee.” MCZ. TYPE LOCALITY. Missouri
Territory. - LeConte, 1863a:8 ( Evarthrus ). - LeConte, 1873:319. - Schaupp, 1880:49.
- Leng, 1920:56 ( Anaferonia ). - Csiki, 1930:671 ( Pterostichus ).
Anaferonia evanescens Casey, 1918:343. HOLOTYPE, female, labelled as follows: “Mex;
CASEY bequest 1925;TYPE USNM 47100; evanescens Csy.” USNM. TYPE LOCALITY,
Colonia Garcia, Sierra Madre Mts., Chihuahua, Mexico. NEW SYNONYMY. — Csiki,
1930:671 ( Pterostichus ). - Leng and Mutchler, 1933:12.
Anaferonia pantex Casey, 1918:344. HOLOTYPE, female, labelled as follows: “Tex;
CASEY bequest 1925; TYPE USNM 47099; Anaferonia pantex Csy.” PARATYPES, two
males and three females, labelled as follows: “Tex; CASEY bequest 1925; pantex —2 to
pantex -6; PARATYPE USNM 47099.” USNM. NEW SYNONYMY. - Csiki, 1930:671
{Pterostichus). - Leng, 1920:56 {Anaferonia).
Recognition. — The species substriatus can be separated from the structurally similar
constrictus by the following characters: elytra with marked apical sinuation, plica large; last
external abdominal sternum with prominent dorsal lateral knob that articulates with plica
(more distinct in females); elytral striae almost effaced in some specimens; apex of apical
blade of median lobe of male evenly rounded. In contrast with these the structures of con-
strictus are as follows: elytra with slight apical sinuation: plica average for subgenus; last
external abdominal sternum with slightly raised mound that fits onto plica; elytral striae
always distinctly impressed; apex of apical blade of median lobe more truncate.
Specimens of iowensis also can be mistaken for those of substriatus. The structures that
distinguish them are recorded in the recognition section of iowensis.
Description. - Body length 9.5 — 14.5 mm. Form short with broad pronotum.
Microsculpture: head between eyes and disc of pronotum with sparsely distributed
sinuous lines and/or amorphic meshes, largely effaced; intervals of elytra with isodiametric
meshes.
Head slightly or moderately glossy; frontal grooves of average depth, bent medially with
convexity directed medially. Penultimate article of labial palpus with four or five setae.
Pronotum moderately or slightly glossy; shape in outline as in figs. 54 — 55; disc of average
convexity; sides markedly produced, moderately constricted anteriorly, strongly constricted
posteriorly, strongly sinuate in front of posterior angles; posterior angles prominent, approx-
imately right; anterior transverse impression complete or not complete; basal lateral fovea
with sides continuous posteriorly, shape as in fig. 54. Prostemal process with shallow or
obsolete longitudinal groove. Middle femur with four to six setae on anterior face.
Revision of Evarthrus
157
Elytra slightly glossy; apical sinuation sharply defined (fig. 78); intervals slightly convex
or completely flat; one to three setae on the third interval of elytron; striae very shallow
with impunctate obsolete dashes, or of average depth and indistinctly punctate; plica large
and prominent (fig. 77).
Last external abdominal segment with prominent dorsolateral knob which fits onto plica.
Male genitalia (fig. 116 — 117): median lobe strongly arcuate, angle approximately right;
apical blade elongate, apex evenly rounded, right paramere extended to halfway point of
median lobe or shorter, apically slightly curved or straight, tapered apically; internal sac
with apical serrulate field, apical sclerite light, elongate. The genitalia of 23 males were
studied.
Stylus of female ovipositor of average size, slightly tapered apically.
Geographical variation. — The pronotum is not markedly constricted posteriorly in speci-
mens from the southern end of the species range. In specimens from the state of Durango,
Mexico, for example, the sides of the pronotum are generally less prominent than those of
fig. 54. Further north in the state of Chihuahua, Mexico, specimens have a pronotum with
produced sides (fig. 54). The shape of the pronotum gradually changes northward until the
common condition is a pronotum with strongly produced sides and a marked posterior
constriction, which is like that of constrictus shown in fig. 55.
There is also a south-north cline in the depth of the striae of the elytra. In Mexico the
striae are obsolete and impunctate. These become gradually deeper and more distinctly
punctate northward.
The intervals of the elytra correspondingly change clinally from a completely flat condi-
tion in Mexico to distinctly convex in Kansas and Nebraska.
The male genitalia of Durango specimens differ from those of individuals from the rest of
the species range (fig. 117 cf . fig. 1 1 6). In Durango, the median lobe is usually strongly
arcuate, and the right paramere is slightly curved. Further north the median lobe is more
broadly rounded and the right paramere is straighter.
Notes on synonymy. — The lectotype of latebrosus LeConte is a substriatus specimen
with impressed elytral striae.
The type specimen of evanescens Casey represents an average substriatus specimen that
is found in northern Mexico. The type specimen of pantex Casey is a substriatus specimen
of the average kind found in Texas.
Collecting notes. — G. E. Ball collected specimens of this species in dry pine forest in
Mexico, and under tumbleweed in desert areas of New Mexico. Specimens are also found
under cover in open places such as pastures and com fields.
Geographical distribution (fig. 134). - This species ranges from Durango, Mexico, north
to southern Wyoming and Minnesota, and from eastern Arizona east to the eastern regions of
Nebraska, Kansas, Oklahoma and Texas. I have seen 525 specimens from the following
localities.
Mexico - CHIHUAHUA: Guerrero (USNM); Minaca (GEB); 31.9 miles south of Minaca (GEB). DURANGO: Arroyo
Hondo near LaFlor (DRW, GEB); Ciudad (AMNH); 18 miles east of El Salto (AMNH); J. Manuel 9300’, El Salto (CAS);
Otinapa (DRW, GEB); two miles east of La Ciudad (CNC).
United States - ARIZONA: Cochise County: Huachuca Mountains (GEB). Pima County: Madera Canyon, St. Rita
Mountains (GEB). COLORADO: Fremont County: Canon City (UASM). Huerfano County: Walsenburg (UMMZ). Las
Animas County: Trinidad (USNM). Logan County: Sterling (AMNH, USNM). Prowers County: Granada (CAS, MCZ).
158
Freitag
County not determined Regnier (AMNH). KANSAS: Chase County: Elmdale (KSU). Clark County: (CNC, UK). Dickinson
County: (CNHM). Douglas County: (UK). Ford County: (UMMZ); Dodge City (UK). Geary County: Fort Riley (CAS);
Junction City (UMMZ). Gove County: (UK). Hamilton County: (CNHM, UK). Harvey County: Sedgewick (CAS, USNM).
Keamy County: Lakin (MCZ). Kiowa County: Belvidere (UK). Meade County: (USNM). Ness County: Ness City (UK).
Pottawatomie County: Onaga (CAS). Reno County: (UMMZ, USNM). Riley County: (USNM); Manhattan (FDPI, KSU,
USNM). Scott County: Scott City (USNM). Sedgwick County: Mount Hope (USNM). Shawnee County: Topeka (CNHM,
USNM). Sheridan County: State Lake, near Studley (RF, UK). Sherman County: five miles west of Goodland (RF);
Goodland (CAS). Thomas County: Colby (RF). Wallace County: (CAS, UK, USNM); Sharon Springs (CNHM). Wilson
County: Benedict (UK). MINNESOTA: Hennepin County: Bloomington (AMNH). NEBRASKA: Furnas County: Cam-
bridge (MCZ). Lancaster County: Lincoln (CAS). Red Willow County: Indianola (MCZ). NEW MEXICO: Colfax County:
Raton (CAS). Lincoln County: Ruidoso (CAS). McKinley County: Coolidge (AMNH, MCZ, USNM); near Ramah (GEB).
Otero County: Cloudcroft (ANSP, CAS, CNHM, CNC, GEB, MCZ, UASM, UK, USNM); Mescalero Reservation (MCZ);
Sacramento (MCZ); Sacramento Mountains (CAS); 16 Springs Canyon, Sacramento Mountains (GEB). Quay County:
Tucumcari (MCZ, USNM). San Miguel County: Las Vegas (CAS); Las Vegas near Hot Springs (UK); Porvenir (CAS).
Santa Fe County: Sante Fe (ANSP, CAS). Torrance County: Tajique (UK). County not determined: Tajano Experimental
Station (CAS). OKLAHOMA: Beckham County: (CAS). Cleveland County: Norman (CAS). Comanche County: Fort Sill
Military Reservation (UMMZ); Wichita National Forest (CAS, UMMZ). Custer County (CAS). Garfield County: Enid
(AMNH). Harmon County: seven miles southwest of Hollis (UMMZ). Oklahoma County: Oklahoma City (CAS). Woods
County: (CAS). SOUTH DAKOTA: Yankton County: Yankton (VMK). TEXAS: Baylor County: eight miles south of
Seymour (CNHM). Bastrop County: Bastrop State Park (CNC). Bexar County: Somerset (CAS). Blanco County: (UMMZ);
Round Mountain (AMNH, CAS, CNHM, RU). Brazos County: College Station (RU). Brewster County: Alpine (USNM);
two miles south of Alpine (CAS). Brown County: Brownwood (RU). Comal County: New Braunfels (USNM). Coryell
County: Gatesville (AMNH). Crockett County: Ozona (AMNH). Dallam County: Rita Blanca Lake, Dalhart (AMNH).
Dallas County: Dallas (MCZ). DeWitt County: Cuero (AMNH). Edwards County: eight miles northeast of Rocksprings
(TCB). Gillespie County: Fredricksburg (RU). Hays County: San Marcos (CNHM). Howard County: Big Spring (CAS,
USNM). Jeff Davis County: Davis Mountains (CAS); Davis Mountains, six — ten miles west of Fort Davis (GEB); Davis
Mountains, 10 miles north of Fort Davis (DL); Fort Davis (CNC, MCZ); Limpia Canyon (DRW, GEB). Kerr County: Kerr-
ville (CNC); nine miles southwest of Kerrville (GEB); 20 miles southeast of Kerrville (CNC). Kleberg County: Riviera
(DRW). Llano County: Llano (AMNH). Pecos County: Blackstone Ranch, 16 miles south of Sheffield (GEB); Fort Stock-
ton (UMMZ). Runnels County: Ballinger (RU). San Patricio County: Lake Corpus Christi (UMMZ). Taylor County: Abiline
(AMNH, CAS); 25 miles southwest of Abiline (CNHM). Terrell County: Chandler Ranch (GEB); 16 miles north of Dryden
(GEB). Tom Green County: Christoval (AMNH, TAM); San Angelo (RU). Travis County: (UMMZ); Austin (CAS, MCZ,
USNM). Uvalde County: one mile south of Montell (TCB); Uvalde (CNHM). Wichita County: Wichita Falls (CAS).
Williamson County: Georgetown (TCB); Leander (TCB). County not determined: Camp Bullis (DRW). WYOMING:
Laramie County: Cheyenne (UMMZ).
Evarthrus constrictus Say, 1 823
Figures 55, 79-80, 118, 134
Feronia constricta Say, 1823b: 147. Type lost. TYPE LOCALITY, Arkansas River near the
Rocky Mountains. — LeConte, 1848:344 C Feronia ). - LeConte, 1852:233 {Evarthrus). —
LeConte, 1863a:8. — LeConte, 1873:319. — Schaupp, 1880:49. - Casey, 1918:345
{Anaferonia). — Leng, 1920:56. - Csiki, 1930:671 (. Pterostichus ). - Van Dyke, 1943:27
{Anaferonia). — Blackwelder and Blackwelder, 1948:2 {Evarthrus).
Feronia (Molops) ovipennis LeConte, 1848:345. LECTOTYPE (here selected) a female,
labelled as follows: “green disc; Type 5619;E. ovipennis Lee.” MCZ. TYPE LOCALITY,
near the Rocky Mountains. - LeConte, 1852:232 {Evarthrus). - LeConte, 1863a: 8. —
LeConte, 1873:319. — Schaupp, 1880:49. — Casey, 1918:343 {Anaferonia). — Leng,
1920:56. - Csiki, 1930:671 {Pterostichus). — Van Dyke, 1943:26 {Evarthrus). - Black-
welder and Blackwelder, 1948:2.
Revision of Evarthrus
159
Anaferonia vernicata Casey, 1918:344. HOLOTYPE, male, labelled as follows: “N. M.
CASEY bequest 1925; TYPE USNM 47105; vernicata Csy.” USNM. TYPE LOCALITY,
Alamogordo, New Mexico. NEW SYNONYMY. — Leng, 1920:56 ( Anaferonia ). — Csiki,
1930:671 ( Pterostichus ).
Anaferonia pimalis Casey, 1918:345. HOLOTYPE, female, labelled as follows: “Ari; CASEY
bequest 1925; TYPE USNM 47106; pimalis Csy.” USNM. TYPE LOCALITY, Southern
Arizona. NEW SYNONYMY. - Leng, 1920:56 (Anaferonia). - Csiki, 1930:671
( Pterostichus ).
Anaferonia latebrosus\C asey, 1918:346 (not LeConte).
Anaferonia pudica Casey, 1918:346. HOLOTYPE, female, labelled as follows: “Tex; CASEY
bequest 1925; TYPE USNM 47101 ; pudica Csy.” USNM. TYPE LOCALITY, Texas. NEW
SYNONYMY. - Leng, 1920:56 (Anaferonia). - Csiki, 1930:671 (Pterostichus).
Anaferonia papago Casey, 1918:346. HOLOTYPE, female, labelled as follows: “Ari; CASEY
bequest 1925; TYPE USNM 47102; papago Csy.” USNM. TYPE LOCALITY, Arizona.
NEW SYNONYMY. - Leng, 1920:56 (Anaferonia). - Csiki, 1930:671 (Pterostichus).
Anaferonia lixa\ Leng, 1920:56 (not LeConte).
Pterostichus ( Pterostichus ) (Sect Anaferonia) lixa\ Csiki, 1 930:671 (not LeConte).
Recognition. — Specimens of constrictus, substriatus and iowensis are structurally similar.
Their distinguishing features are described in the recognition sections of the two preceeding
species.
Description. — Body length 9.5 — 12.8 mm. Form relatively less robust than other species
of the subgenus Evarthrus.
Microsculpture: head between eyes, disc of pronotum with lines partially or completely
effaced; sparsely distributed sinuous lines; intervals of elytra with largely effaced to distinct
isodiametric meshes.
Head glossy; frontal grooves of average depth, curved with convexity directed medially.
Penultimate article of labial palpus with two median and four apical setae.
Pronotum glossy; shape as in fig. 55; disc strongly convex; sides markedly produced,
strongly constricted posteriorly with marked sinuation in front of posterior angles; posterior
angles produced, acute or nearly so; anterior transverse impression complete; basal lateral
fovea with sides continuous posteriorly. Prostemal process with shallow or obsoletely im-
pressed longitudinal groove. Middle femur with six or seven setae on anterior face.
Elytra of males moderately glossy; females slightly duller; apical sinuation not sharply
defined (fig. 80); intervals of average convexity or slightly flatter; striae distinctly impressed
with small punctures anteriorly; impunctate posteriorly; plica not prominent (fig. 79).
Last external abdominal segment with low dorsal convexity articulating with plica.
Male genitalia (fig. 118) with median lobe strongly arcuate, angle almost right, ventral
median hump present or absent; apical blade elongate, apex almost truncate, apical lateral
edges strongly deflected dorsally; right paramere extended to halfway point of median
lobe, slender apically; internal sac with serrulate field apically, apical sclerite dark, elongate.
The genitalia of 1 1 males were examined.
Stylus of female ovipositor of average size, slightly tapered apically.
160
Freitag
Notes on synonymy. — The original description of cons trie tus Say was used to identify
this species. The lectotype of ovipennis LeConte is an average specimen of constrictus. The
type specimens of vernicata Casey, pimalis Casey, pudica Casey, and papago Casey are all
average constrictus specimens.
Collecting notes. — This species has been collected in corn fields, and open pasture under
rocks.
Geographical distribution (fig. 134). — The range of this species extends as a relatively
narrow band from Arizona east to Kansas and Nebraska. I have seen 218 specimens from the
following localities.
United States - ARIZONA: Apache County: eight - 15 miles northeast of White River (AMNH); McKay’s Peak,
White Mts. (AMNH); White Mts. (AMNH, CAS, MCZ); Springville (UMMZ); White Mountain Reservation, east of McNary
(AMNH). Coconino County: five miles northwest of Flagstaff (AMNH); eight miles south of Flagstaff (GEB); Flagstaff
(CAS); 23 miles southwest of Heber (UMMZ); Williams (CAS, USNM). Gila County: Globe (MCZ). Navajo County: Heber
(UMMZ); Show Low (CAS). COLORADO: Denver County: Denver (CAS). El Paso County: Colorado Springs (AMNH,
CAS, MCZ, USNM). Huerfano County: Gardner (AMNH); LaVeta (CAS). Otero County: LaJunta (MCZ). Prowers County:
Granada (CAS, MCZ). Pueblo County: (MCZ). County not determined: Clayton (CNHM). IOWA: Woodbury County:
Sioux City (USNM). Clark County: (CAS, CNHM, MCZ, UASM, UK). Douglas County: (UK). Ford County: Dodge City
(UK). Grove County: (UK). Greeley County: (UK). Hamilton County: (UK). Harvey County: Sedgewick (CAS). Reno
County: (UMMZ, USNM); Hutchinson (CAS); Medora (UK). Scott County: Scott City (USNM). Wallace County: (UK);
Sharon Springs (CNHM); Wallace (USNM). County not determined: Fort Hayes (MCZ). NEBRASKA: Lancaster County:
Lincoln (CAS). NEW MEXICO: Catron County: Luna (UMMZ); seven miles south of Luna (AMNH). Colfax County:
Koehler (USNM); Prairie, near Koehler (CAS, USNM). Quay County: Tucumcari (USNM). San Doval County: Jemez
Mountains (CAS). San Miguel County: Beulah (ANSP, CAS); Las Vegas (INHS); Porvenir (CAS). Counties not determined:
Pinedale (GEB); Tres Ritos (CAS); Water Canon (UK). SOUTH DAKOTA: Hutchinson County: Menno (VMK). Yankton
County: Yankton (VMK). County not determined: Cedar Pass (USNM). TEXAS: Bell County: Belton Dam (CU). Bexar
County: 20 miles north of San Antonio (CAS). Hemphill County: Canadian (CAS). McLennan County: China Springs
(CNHM). Tarrant County: Fort Worth (MCZ). Travis County: Austin (MCZ).
The torvus Group
Characteristics. — Penultimate article of labial palpus with five to seven setae. Pronotum
more or less quadrate, not markedly constricted posteriorly, disc usually rugose, sides not
very prominent, distinctly sinuate in front of posterior angles; anterior transverse impression
complete. Prosternal process with shallow or obsolete longitudinal groove, without setae.
Middle femur with six — ten setae on anterior face. Male genitalia with apical blade of
median lobe relatively short and evenly rounded.
The species torvus and gravidus are included in this group. Both occur west of the
Mississippi River and occupy the Great Plains from Texas north to South Dakota.
Evarthrus torvus LeConte, 1863
Figures 56-57, 119-120, 135
Evarthrus torvus LeConte, 1863b:9. LECTOTYPE (here selected) a male, labelled as fol-
lows: “Col; Type 5657; E. torvus Lee.” MCZ. TYPE LOCALITY, Colorado. - LeConte,
1 863a: 8 (Evarthrus). - Schaupp, 1880:49. — Casey, 1918:356 (Ewnolops). - Leng,
1920:57. — Csiki, 1930:672 ( Pterostichus ).
Feronia (Evarthrus) acuminata Chaudoir, 1868:52. LECTOTYPE, male, one of two un-
labelled specimens of both sexes, beside which is a box label: “Tejas.” MHNP. NEW
SYNONYMY.
Revision of Evarthrus
161
Eumolops prominens Casey, 1918:353. HOLOTYPE, female, labelled as follows: “Fla;
CASEY bequest 1925; TYPE USNM 47128; prominens Csy.” USNM. TYPE LOCALITY,
Florida (this locality is probably incorrect). NEW SYNONYMY. — Leng, 1920:57
C Eumolops ). — Csiki, 1930:672 ( Pterostichus ).
Eumolops sexualis Casey, 1918:354. HOLOTYPE, male, labelled as follows: “N. M.; CASEY
bequest 1925; TYPE USNM 47124; Eumolops sexualis, Csy.” USNM. TYPE LOCALITY,
New Mexico. PARATYPE, one male and one female, labelled as follows: “N. M.; CASEY
bequest 1925; sexualis —2 and sexualis —3; PARATYPE USNM 47124.” NEW SYN-
ONYMY. - Leng, 1920:57 {Eumolops). — Csiki, 1930:672 {Pterostichus).
Eumolops inflatula Casey, 1918:354. HOLOTYPE, female, labelled as follows: “Col;
CASEY bequest 1925; TYPE USNM 47127; inflatula Csy.” USNM. TYPE LOCALITY,
Akron, Colorado. NEW SYNONYMY. - Leng, 1920:57 {Eumolops). - Csiki, 1930:672
{Pterostichus).
Eumolops (Evarthrinus) decepta Casey, 1918:357. HOLOTYPE, female, labelled as follows:
“Ind; CASEY bequest 1925; TYPE USNM 47356; Evarthrinus deceptus Csy.” USNM.
TYPE LOCALITY, Indiana (this locality is incorrect). NEW COMBINATION. - Casey
1920:194 {Evarthrinus). — Leng, 1920:57 {Eumolops). — Csiki, 1930:672 {Pterostichus).
Eumolops (Evarthrinus) impolita Casey, 1918:358. HOLOTYPE, male, labelled as follows:
“Tex; CASEY bequest 1925; TYPE USNM 47130; impolita Csy.” USNM. TYPE LOCAL-
ITY, Texas. PARATYPE, female, labelled as follows: “Tex; CASEY bequest 1925;
impolita -2 PARATYPE USNM 47130.” USNM. NEW SYNONYMY. - Casey, 1920:195
{Evarthrinus). — Leng, 1920:57 {Eumolops). - Csiki, 1930:673 {Pterostichus).
Evarthrinus (Evarthrinus) minax Casey, 1920:194. HOLOTYPE, male, labelled as follows:
“L; CASEY bequest 1925; TYPE USNM 47124; minax Csy.” USNM. TYPE LOCALITY,
Indiana (this locality is probably incorrect). PARATYPE, female, labelled as follows:
“Ind; CASEY bequest 1925; minax -2 PARATYPE USNM 47129.” USNM. NEW SYN-
ONYMY. — Leng and Mutchler 1927:10 (Evarthrinus). — Csiki, 1930:673 (Pterostichus).
Recognition. — The following characteristics are diagnostic for torvus: pronotum moder-
ately constricted anteriorly and posteriorly, subcordiform, almost quadrate, sides not
strongly produced (figs. 56—57), disc and basal fovea rugose north and west of Oklahoma;
male genitalia (figs. 1 19—120).
Specimens of the similar species gravidus are generally broader than are those of torvus.
In addition the pronotum of gravidus is more quadrate with distinctly crenulated sides in
front of the basal angles, and the sides of the basal foveae are not continuous posteriorly
(fig. 58). Further these two species can be separated by their different genitalic structures,
(figs. 119-120 cf. fig. 121).
Specimens of iowensis are also similar to southern torvus individuals. The species are
allopatric and are also distinguished by their genitalia (fig. 115 cf. figs. 1 19—120).
Description. — Body length 12.7 — 19.5 mm. Form robust or slender and elongate.
Microsculpture: head between eyes and disc of pronotum with sinuous lines distinctly
defined or effaced; intervals of elytra with isodiametric meshes occasionally slightly
stretched longitudinally.
Head moderately glossy; frontal grooves of average depth, somewhat broad, curved with
convexity directed medially. Penultimate article of labial palpus with five or six setae.
162
Freitag
Pronotum slightly or moderately glossy, more or less rugose or smooth; shape as in figs.
56—57; disc of average convexity; sides not strongly produced, slightly or moderately con-
stricted anteriorly, moderately constricted posteriorly, distinctly sinuate in front of poste-
rior angles; posterior angles prominent, approximately right; anterior transverse impression
complete; basal lateral fovea markedly rugose or not, sides almost always continuous poste-
riorly. Prostemal process with longitudinal groove slightly impressed or obsolete. Middle
femur with six to nine setae on anterior face.
Elytra glossy to dull; apical sinuations slight or obsolete; intervals distinctly convex to
flat; one to three setae on third interval; striae of average depth, with small distinct punc-
tures in anterior half, impunctate posteriorly.
Male genitalia (figs. 119—120) with median lobe moderately arcuate, angle broadly
obtuse; apical blade of average length or shorter, apex evenly rounded; right paramere
extended to apical half of median lobe or shorter, strongly tapered apically; internal sac
with apical serrulate field, apical sclerite light, elongate, basal and curled in or not. The
genitalia of 1 1 males were examined.
Stylus of female ovipositor of average size, slightly tapered apically.
Geographical variation and subspecies. — This species can be divided into two geo-
graphically separate groups of populations, one south and east of Oklahoma, the other
north and west of Oklahoma. In Oklahoma there are intermediate forms. Specimens of
western and northern populations are characterized by the following structures: body form
robust; pronotum with disc and basal lateral foveae deeply rugose; elytra of male glossy,
female slightly glossy, usually one seta in third interval; scutellar striae present; median lobe
of male with apical blade of average length; internal sac of median lobe with light apical
sclerite recurved basally. The corresponding structures of southern specimens differ as
follows: body more slender and elongate; pronotum with disc and basal lateral foveae with
normal sculpturing; elytra of males slightly glossy, females matte, two or three setae
normally in third interval; scutellar striae absent; apical blade of median lobe short; internal
sac with light apical sclerite not recurved basally.
I regard these populations as subspecies, torvus torvus north and west of Oklahoma, and
torvus deceptus further south and east.
Notes on synonymy. — The lectotype of torvus is similar to the common form that
inhabits Colorado. Tue type specimens of prominens Casey, sexualis Casey, and inflatula
Casey all look like the lectotype of torvus LeConte. The type specimens of decepta Casey,
impolita Casey, and minax Casey are the average specimens from southern portions of this
species range that have dull elytra and three punctures in the third interval of the elytron.
Collecting notes. — D. J. Larson and I collected t. torvus specimens under logs in a dried
out stream bed near Castle Rock, Colorado and under boards in a farmyard near Colby,
Kansas. G. E. Ball collected some specimens under rocks near the river, Cache la Poudre,
Colorado. Specimens have also been taken in corn fields.
Geographical distribution (fig. 135). — This species inhabits the Great Plains from south-
ern Texas north to South Dakota.
Revision of Evarthrus
163
E. torvus torvus
I have seen 307 specimens from the following localities.
United States - ARKANSAS: Hempstead County: Hope (UMMZ). COLORADO: Adams County: Bennett (CAS).
Boulder County: Boulder (CNC). Clear Creek County: Empire (CAS). Denver County: Denver (CAS, CNHM, CU, GEB,
USNM). Douglas County: ten miles north of Castle Rock (DL, RF). Jefferson County: Golden (CAS). Larimer County:
Fort Collins (CAS, KSU, RTB); Loveland (CAS); Poudre Canyon (GEB). Weld County: Greeley (CAS). Counties not
determined: Genessee Mountain Park (CAS); Lookout Mountain (CAS). IOWA: Louisa County: Oakville (ISU). O’Brien
County: four miles east of Sanborn (ISU). KANSAS: Chase County: Elmdale (KSU). Cheyenne County: (KSU). Clay
County: (ANSP, CAS, UMMZ, USNM). Doniphan County: Wathena (USNM). Douglas County: (CU, KSU, MCZ); five
miles north of Baldwin City (DL). Geary County: Junction City (UMMZ). Harper County: Harper (USNM). Kiowa
County: Belvidere (KSU). Marion County: Marion (CAS). Pottawatomie County: Onaga (CAS, MCZ, USNM, UW). Riley
County: (CNHM, USNM); Manhattan (KSU, USNM). Sedgwick County: Mount Hope (ANSP, UMMZ, USNM). Sherman
County: Goodland (CAS, CNHM). Sumner County: Wellington (USNM). Thomas County: Colby (RF). Wabaunsee
County: (KSU). Wallace County: Sharon Springs (CNHM); Wallace (KSU). Wilson County: (CNHM); Benedict (CAS).
Wyandotte County: Argentine (AMNH, MCZ). MISSOURI: Carter County: Van Buren (UMMZ). Greene County: Willard
(ANSP). St. Charles County: St. Charles (MCZ). Vernon County: (GEB). NEBRASKA: Cedar County: Randolph (MCZ).
Furnas County: Cambridge (MCZ). Hall County: Jet U. S. 34 and Platte River near Grand Island (GEB). Lancaster
County: Lincoln (CAS, CNHM, USNM). Phelps County: Holdredge (CAS). Platte County: near Platte (CAS). Red Willow
County: McCook (USNM). NEW MEXICO: Dona Ana County: near Rincon (MCZ). Otero County: Bent (CU); Cloudcroft
(CAS, GEB, MCZ, USNM); Mescalero Reservation (CU). San Miguel County: Gallinas Canon (MCZ); Las Vegas (CAS,
INHS, KSU); Sapello Creek (GEB). OKLAHOMA: Canadian County: Yukon (CAS). Cleveland County: (CAS). Kingfisher
County: Kingfisher (ANSP). Murray County: (CAS). Tulsa County: Tulsa (CAS, CNHM). County not determined: Wichita
National Forest (CAS). SOUTH DAKOTA: Custer County: (VMK). Fall River County: five miles south of Hot Springs
(AMNH). Lawrence County: Spearfish (VMK). TEXAS: Hidalgo County: Edinburg (UMMZ), (probably incorrectly
labelled).
E. torvus deceptus
I have seen 28 specimens collected in the following localities.
United States - TEXAS: Dallas County: Dallas (MCZ, RTB). DeWitt County: Cuero (AMNH). El Paso County: El
Paso (CM). Grayson County: Sherman (MCZ). Grimes County: Roans Prairie (AMNH). Lee County: Fedor (CM). Mon-
tague County: Forestburg (UMMZ). Tom Green County: Cristoval (AMNH).
Evarthrus gravidus Haldeman, 1853
Figures 58, 121 , 135
Evarthrus gravidus Haldeman, 1853:361. Type lost. TYPE LOCALITY, Texas (here se-
lected). - LeConte, 1858:28 {Evarthrus). — LeConte, 1863a:8. - LeConte, 1873:318. -
Schaupp, 1880:49. - Casey, 1918:354 ( Eumolops ). — Leng, 1920:57. - Csiki, 1930:672
( Pterostichus ).
Eumolops ampla Casey, 1918:353. HOLOTYPE, female, labelled as follows: “Tex.; CASEY
bequest 1925; TYPE USNM 47126; ampla Csy.” USNM. TYPE LOCALITY, Texas. NEW
SYNONYMY. - Leng, 1920:57 {Eumolops). — Csiki, 1930:672 (Pterostichus).
Recognition. — The robustness of the body, crenulate sides near the basal angles of the
pronotum, and lateral basal foveae of pronotum broad with sides not continuous posteriorly,
combined, distinguish members of gravidus from specimens of all other species of Evarthrus.
The pronotum of specimens of torvus is more constricted posteriorly than that of speci-
mens of gravidus. Also the sides of the basal lateral foveae of the pronotum are continuous
posteriorly in torvus but not in gravidus.
164
Freitag
Description. — Body length 15.5 — 21.7 mm. Form robust.
Microsculpture: head between eyes and disc of pronotum with sinuous lines and amorphic
meshes, sometimes partially effaced; intervals of elytra with isodiametric meshes. Integu-
ment of dorsum moderately glossy.
Head: frontal grooves of average depth, bent with convexity directed medially. Penulti-
mate article of labial palpus with six or seven setae.
Pronotum with surface more or less rugose; shape somewhat quadrate as in fig. 58; disc of
average convexity; sides not strongly produced, moderately constricted anteriorly, slightly
constricted posteriorly, slightly sinuate in front of posterior angles, crenulated posteriorly;
posterior angles sharp, approximately right; anterior transverse impression complete; basal
lateral fovea with sides usually not continuous posterio-medially. Prosternal process with
longitudinal groove shallow or obsolete. Middle femur with eight to ten setae on anterior
face.
Elytra slightly or obsoletely sinuate apically; intervals of normal convexity or completely
flat; one to three setae on third interval; striae of average depth or more shallow, apical two
thirds with small punctures, impunctate posteriorly.
Male genitalia (fig. 121): median lobe strongly arcuate, angle slightly acute; apical blade
short with apex evenly rounded; right paramere large, extended to apical third of median
lobe, moderately tapered apically; internal sac with apical serrulate field, apical sclerite
light, elongate, with basal tooth. The genitalia of four males were examined.
Stylus of female ovipositor slightly tapered apically with apex broadly rounded.
Notes on synonymy. — I selected Texas as the type locality because gravidus is known
from Texas only. The type specimen of ampla Casey is an average gravidus specimen.
Geographical distribution (fig. 135). — This species is confined to Texas. I have seen 137
specimens from the following localities.
United States - TEXAS: Collin County: Plano (USNM). Comal County: New Braunfels (CU, USNM). Dallas County:
Dallas (USNM). El Paso County: El Paso (CAS). Kerr County: Kerrville (CNC). Lee County: Fedor (CAS, CM). McLennan
County: China Springs (CNHM); Waco (MCZ). Montague County: Forestburg (UMMZ). Travis County: Austin (MCZ,
USNM). Victoria County: Victoria (USNM). County not determined: Therman (MCZ).
The gigas Group
Characteristics. — Body size large. Penultimate article of labial palpus with five or six
setae. Pronotum strongly constricted posteriorly, posterior angles prominent, anterior trans-
verse impression complete, prosternal process with obsolete longitudinal groove. Middle
femur with 7—11 setae on anterior face. Male genitalia with short, stub-like, right paramere
and apical blade of median lobe twisted 45° from horizontal plane, or right paramere long
and apical blade with left side deflected dorsally.
The species gigas, heros and sallei form this group. All three are known from eastern
Texas, southern Oklahoma, and Arkansas.
Revision of Evarthrus
165
Evarthrus sallei LeConte, 1873
Figures 59, 122, 136
Evarthrus sallei LeConte, 1873:319. LECTOTYPE (here selected) a male, labelled as fol-
lows: “9; red disc; Type 5663; E. sallei Lee.” MCZ. TYPE LOCALITY, Texas. - Schaupp,
1880:49 {Evarthrus). — Casey, 1918:356 ( Eumolops ). — Leng, 1920:57. — Csiki, 1930:
672 ( Pterostichus ).
Recognition. — The following combination of characteristics is diagnostic of this species:
basal lateral fovea of pronotum with sides continuous posteriorly forming straight base;
elytra with marked apical sinuation, intervals of males with transversely stretched meshes
comprising microsculpture; male genitalia with reduced stub-like paramere, apical blade of
median lobe twisted 45° from horizontal plane.
The subspecies sodalis colossus is similar to sallei in general appearance, but the basal
lateral fovea is U-shaped at the base. In addition these species are allopatric.
Description. — Body length 18 — 21 mm. Form robust, body with relatively parallel sides.
Microsculpture: head between eyes and disc of pronotum with sparsely distributed sinu-
ous lines often effaced or nearly so; intervals of males with transversely stretched meshes,
females with isodiametric meshes.
Head moderately to markedly glossy; frontal grooves of average depth, bent medially with
convexity directed medially. Penultimate article of labial palpus with five setae.
Pronotum moderately or slightly glossy; shape as in fig. 59; disc of average convexity;
sides strongly produced, moderately constricted anteriorly, strongly constricted posteriorly,
sharply sinuate in front of posterior angles; posterior angles very prominent, acute; anterior
transverse impression complete; basal lateral fovea with sides continuous posteriorly forming
straight base. Middle femur with seven or eight setae on anterior face.
Elytra of males moderately glossy iridescent in a few specimens, females less glossy; apical
sinuation sharply defined; intervals distinctly convex in males, slightly convex in females.
Striae of average depth, distinctly punctate anteriorly, apical parts indistinctly punctate.
Male genitalia (fig. 122) median lobe strongly arcuate, angle slightly obtuse, often with
low ventral medial bump; apical blade short with apex evenly rounded, twisted more than
45° from horizontal plane; right paramere very small, stub-like; internal sac with serrulate
field apically, apical sclerite dark elongate tooth. The genitalia of four males were examined.
Stylus of female ovipositor slightly tapered apically with broadly rounded apex.
Geographical distribution (fig. 136). — This species is known from Texas, only. I have
seen 48 specimens from the following localities.
United States - TEXAS: Comal County: New Braunfels (USNM). Dallas County: (CAS, INHS, UMMZ); Dallas (CAS,
INHS, KSU, MCZ, USNM). Jackson County: (USNM); Baroncuhua (USNM). Victoria County: Victoria (USNM).
Evarthrus gigas Casey, 1918
Figures 60, 123, 136
Megasteropus gigas Casey, 1918:350. HOLOTYPE, female, labelled as follows: “Tex;
CASEY bequest 1925; TYPE USNM 47123; Megasteropus gigas Csy.” USNM. TYPE
LOCALITY, Texas. PARATYPE, female, labelled as follows: “Tex; CASEY bequest
1925; gigas -2 PARATYPE USNM 47123.” USNM. - Leng, 1920:57 {Megasteropus). -
Csiki, 1930:672 {Pterostichus).
166
Freitag
Recognition. — The combination of large body size, elytra with flat intervals and very
shallow impunctate striae, and male genitalia (fig. 123), distinguishes gigas from all other
species of Evarthrus. Specimens of heros superficially resemble individuals of gigas but are
distinguished by their distinctly punctate elytral striae.
Description. — Body length 19.4 — 23.8 mm. Form robust.
Microsculpture: head between eyes and disc of pronotum with sinuous lines, almost
effaced in males, very dense in females; intervals of elytra with isodiametric, flat meshes in
males, markedly amorphic meshes in females.
Head markedly glossy in males, slightly so in females; frontal grooves short, of average
depth, sometimes bent with convexity directed medially. Penultimate article of labial palpus
with five setae.
Pronotum markedly glossy in males, slightly glossy in females; shape as in fig. 60; disc of
average convexity; sides strongly produced, moderately constricted anteriorly, strongly
constricted posteriorly, sharply sinuate in front of posterior angles; posterior angles acute
and very prominent; anterior transverse impression complete; basal lateral fovea with sides
not continuous posteriomedially. Middle femur with seven to ten setae on anterior face.
Elytra of males markedly glossy, those of females slightly glossy; distinctly sinuate
apically; intervals completely flat. Striae very shallowly impressed almost effaced, impunc-
tate, series of dashes rather than continuous lines in some specimens.
Male genitalia (fig. 123) with median lobe strongly arcuate, angle slightly obtuse, large
lobe-like evagination ventromedially; apical blade twisted more than 45° from horizontal
plane; right paramere very small, stub-like; internal sac with apical serrulate field, apical
sclerite dark elongate tooth. The genitalia of three males were examined.
Stylus of female ovipositor slightly tapered apically with broadly rounded apex.
Geographical distribution (fig. 136). — This species inhabits southeastern Texas. I have
seen 22 specimens from the following localities.
United States - TEXAS: Kleberg County: Kingsville (CU). Victoria County: Victoria (USNM).
Evarthrus heros Say, 1823
Figures 61, 65, 76, 124, 136
Feronia heros Say, 1823b: 145. Type lost. TYPE LOCALITY, “The Arkansa.” MCZ. —
LeConte, 1848:350 (Feronia). — LeConte, 1852:233 (Evarthrus). - Haldeman, 1853:
361. - LeConte, 1858:28. — LeConte, 1863a:8. - LeConte, 1873:318. — Schaupp,
1880:49. - Casey, 1918:352. — Leng, 1920:57. — Csiki, 1930:672 (Pterostichus). - Van
Dyke, 1943:27 (Eumolops). - Blackwelder and Blackwelder, 1948:2 (Evarthrus).
Feronia (Pterostichus) americana; LeConte, 1848:350 (not Dejean).
Me gas ter opus gigas ; Van Dyke, 1943:27 (not Casey).
Recognition. - The following combination of characteristics is diagnostic of heros: large
body size; scutellar stria of elytron long, separate from second stria; base of second stria
beginning near basal seta; intervals of elytra very shallow, usually indicated by rows of
distinctly impressed punctures; apex of median lobe of male with left side deflected
dorsally.
Revision of Evarthrus
167
Specimens of the similar species gigas are distinguished from heros by possession of very
shallow impunctate elytral striae.
Description. — Body length 18.7 — 27.1 mm. Form robust.
Microsculpture: head between eyes and disc of pronotum with sinuous lines, sometimes
almost effaced; intervals of elytra with isodiametric meshes.
Head glossy; frontal grooves of average depth, usually bent posteriorly with convexity
directed laterally. Penultimate article of labial palpus with five or six setae.
Pronotum glossy; shape as in fig. 61; disc of average convexity; sides strongly produced,
moderately constricted anteriorly, strongly constricted posteriorly, sharply sinuate in front
of posterior angles; posterior angles produced and acute; anterior transverse impression
complete; basal lateral foveae with sides not continuous posteriomedially. Middle femur
with 9—11 setae on anterior face (fig. 76).
Elytra of males markedly glossy, those of females moderately glossy, slightly sinuate
apically; intervals completely flat; striae very shallowly impressed, usually only rows of
punctures; scutellar stria long and separated from second stria which begins near basal seta
(fig. 65).
Male genitalia (fig. 124) with median lobe moderately arcuate, angle broadly obtuse;
apical blade with left side of apex deflected dorsally; right paramere extended to apical half
of median lobe; internal sac with apical serrulate field, apical sclerite elongate light plate.
The genitalia of five males were examined.
Stylus of female ovipositor slightly tapered apically with broadly rounded apex.
Collecting notes. — The original description of E. heros was used to identify this species.
Specimens of this species have been collected in cotton fields.
Geographical distribution (fig. 136). — This species is found in Arkansas, Oklahoma, and
eastern Texas. I have seen 69 specimens from the following localities.
United States - ARKANSAS: County not determined: Arkansa (MCZ). OKLAHOMA: McCurtain County: Milierton
(USNM). TEXAS: Collin County: Plano (USNM). Comal County: New Braunfels (CNHM, USNM). Cooke County:
Gainesville (USNM). Dallas County: Dallas (ANSP, CNHM, MCZ, UMMZ, USNM). Delta County: Cooper (USNM).
Ellis County: Waxahachic (USNM). Fannin County: Ladonia (USNM). Lee County: (UMMZ). McLennan County: Waco
(CAS). Montague County: Forestburg (UMMZ).
The gravesi Group
Characteristics. — Body size average. Penultimate article of labial palpus with three setae.
Pronotum constricted posteriorly, posterior angles prominent, sides of basal fovea not
continuous posteriorly, anterior transverse impression complete; posterior lateral setae on
bead. Middle femur with four setae on anterior face. Plica of elytron absent. Only one
species, gravesi, is included in this group.
Evarthrus gravesi new species
Figures 62, 136
Recognition. — The combination of the glossy dorsum, form of the pronotum, and
complete anterior transverse impression, absence of the plica of the elytron, and four setae
on the anterior face of the middle femur, distinguishes E. gravesi from all the other species
of Evarthrus. The general habitus of gravesi resembles that of substriatus or constrictus.
168
Freitag
Description. — HOLOTYPE, female, labelled as follows: “Pearl (Jackson) Rankin Co.
Miss. 23 — III — 1 959 R. C. & A. Graves; HOLOTYPE Evarthrus gravesi R. Freitag.” MCZ.
Body length 12.8 mm., width 5.5 mm. Form robust.
Microsculpture on head between eyes effaced; sparsely distributed, almost effaced,
sinuous lines on disc of pronotum; isodiametric meshes on intervals of elytra. Dorsum
glossy.
Head length 1.5 mm., width 3.0 mm.; frontal grooves of average depth, sharply defined,
straight, oblique. Penultimate article of labial palpus with three setae: two medial and one
apical.
Pronotum length 3.5 mm., width 4.5 mm.; form cordiform in outline as in fig. 62; disc
moderately convex; sides produced, constricted slightly anteriorly, strongly posteriorly,
distinctly sinuate in front of posterior angles; posterior angles produced, obtuse; anterior
transverse impression complete and deeply impressed. Middle femur with four setae on
anterior face; lateroventral margins of last article of tarsus with setae.
Elytra length 7.8 mm., width 5.6 mm.; not sinuate apically; plica absent; intervals
moderately convex; intervals deeply impressed and indistinctly punctate.
Stylus of ovipositor broad, slightly tapered apically.
This is the only specimen of this species seen by me. The position of the type locality
is indicated on fig. 136.
Derivation of specific name. — The type specimen was collected by Dr. Robert C. Graves,
Department of Biology, Bowling Green State University, Ohio. This species is named in
honour of the collector.
FOSSIL MATERIAL
Of the nominal fossil species Evarthrus tenebricus Scudder, only the head is preserved.
Scudder (1900) placed it in the genus Evarthrus “on account of the brevity of the last joint
of the labial palpus.” This structure is not diagnostic for the genus Evarthrus. Furthermore
the other characteristics given by Scudder in his description are common to many carabid
genera. I have not seen the specimen. It may or may not be a member of the genus
Evarthrus.
PHYLOGENY AND ZOOGEOGRAPHY OF THE SPECIES
OF E VAR THR US LECONTE
Phylogeny
My views concerning the phylogeny of Evarthrus are based on structural similarities and
differences of extant species, because a fossil record is not available. Fig. 137 is a time
divergence dendrogram of the history of Evarthrus. It is based on the principle that similar
organisms are related. Species that have many similar structures are closely related, while
those which are dissimilar are more remotely related. Since rate of divergence is unknown
the slopes of the branches of the dendrogram are not significant.
Revision of Evarthrus
169
To determine the relationships of the genus Evarthrus, I compared its characteristics with
those of a representative selection of Nearctic and Palaearctic genus-group taxa of the tribe
Pterostichini. I believe Evarthrus is most closely related to the Molops group (tribe Molopini
Jeannel, 1942 and 1948) of Europe and Africa. Among other structures the three that
Evarthrus and the Molops group share, and which 1 regard as patristic affinities are: pleura
and thoracic and abdominal sterna impunctate; basal portion of interval 7 of the elytron
raised and not deflected downward like other intervals; and antennae of larva with five
articles each (Van Emden, 1942).
Because the genera Evarthrus and Pterostichus have many structural similarities a word
about their relationships is necessary. A remarkably stable characteristic of Pterostichus is
the presence of punctures on the ventral sclerites, particularly in the groove of the mes-
episternum. Rarely the punctures are feebly developed but they usually can be found on the
mesepisternum. Also the known larvae of species of Pterostichus have antennae of four
articles each. Because adults of Evarthrus and of the Molops group have impunctate ventral
sclerites and the larvae have antennae of five articles, and are similar to Pterostichus in most
other features, I believe Pterostichus is a distant relative of these genera.
The progenitor of the genus Evarthrus probably had characteristics which are present in
extant species of the Molops group and Evarthrus, and some which are widespread in other
pterostichine genera. The hypothetical ancestor of Evarthrus probably possessed in combi-
nation the primitive states of characters presented in Table 1 . Advanced conditions of these
characters are also listed in Table 1, and illustrate the extent of evolutionary change. The
determination of the trends of change in these characters or morpho-clines (Maslin 1952)
provide a basis for establishing a phylogeny of Evarthrus.
One trend that is apparent is the increase of body size in all three subgenera and it is most
evident in the subgenus Evarthrus. The largest species of this subgenus are specialized, and
there is no doubt that large size is the specialized condition.
Another evolutionary trend is in the increase in the number of setae of the penultimate
article of the labial palpus. Most of the species in Fortax and Cyclotrachelus have two setae,
several species have three, or four, while E. unicolor has four, five, or six setae. In the sub-
genus Evarthrus most species possess five or six setae.
Two setae on the penultimate article of the labial palpus is the primitive condition
because it is common in pterostichine genera which are related to Evarthrus. Similarly the
setae on the anterior face of the middle femur tend to increase in number. Four setae is the
common condition in Fortax, Cyclotrachelus, and in the less specialized species of the
subgenus Evarthrus. This is probably the primitive condition. The setae range in number
from seven to eleven in specialized species of the subgenus Evarthrus.
There are two distinct trends in the direction of the eversion of the internal sac of the
median lobe. The internal sac everts dorsoapically in all species of Cyclotrachelus. In the
subgenus Fortax, the internal sac tends to evert to the left while in the subgenus Evarthrus
the eversion is to the right. The intermediate dorsoapical eversion is probably the primitive
state.
170
Freitag
TABLE I. Primitive and Specialized conditions of some characters of Evarthrus.
CHARACTER PRIMITIVE ADVANCED
BODY:
Size
Colour
Surface
Venter
MICROSCULPTURE:
Head, pronotum and elytra
HEAD:
Frontal grooves
Gula
Penultimate article of labial palpus
PRONOTUM:
Outline, dorsal aspect
Basal angles
Anterior transverse impression
Basal lateral fovea
Basal seta
PROSTERNAL PROCESS:
Basal and preapical portion
Apex
METEPISTERNUM:
ELYTRA:
Intervals
Interval 3
Interval 7
Striae
Umbilicate series
Plica
LEGS:
Colour
Middle femur
Basal article of middle and hind tarsus
Last tarsal article
ABDOMEN:
Last external sternum of female
MALE GENITALIA:
Median lobe
Right paramere
Eversion of internal sac
Apex of internal sac
FEMALE GENITALIA:
Stylus
Small
Black
Glossy
Impunctate
Almost effaced, sinuous,
sparsely distributed lines
Deep, straight, parallel
Without knobs on each side
With two medial setae
Quadrate
Not prominent
Complete with medial
portion feebly present
Monostriate
Near basal angle beside lateral bead
With obsoletely impressed
longitudinal groove
Not marginate, glabrous
Short
Low convexity
With one puncture
Raised at base
Shallow, Fmely punctate
Distinct ridges separating first
three anterior punctures
Small
Rufopiceous
With four setae on anterior face
With lateral groove
With ventral lateral setae
With two setae
Moderately arcuate
Extended to halfway point of
median lobe
Dorsoapical
With light apical serrulate field
Slightly tapered apically
Large
Unchanged
Dull
Unchanged
Amorphic meshes or
isodiametric meshes
Shallow, crescent-shaped, oblique
Flanked by raised knobs
With more than two medial setae
Cordate
Prominent
Medial portion absent or medial
portion clearly impressed
Bistriate or punctiform
In front of basal angle on lateral bead
With deep longitudinal groove
Marginate with setae
Unchanged
High convexity
With two or more punctures
Unchanged
Deeply impressed or coarsely punctate
Without distinct ridges between first
three anterior punctures
Large or absent
Red
With more than four setae on
anterior face
Without lateral groove
Without ventral lateral setae
Unchanged
Strongly arcuate or slightly arcuate
Very long or very short and stub-like
Right or left
With serrulate field and sclerite
Broadly rounded apically or
strongly tapered apically
Revision of Evarthrus
171
The progenitor of Evarthrus probably differentiated into the “ancestors” of the subgenus
Fortax and the Cyclotrachelus-Evarthrus lineage. The Fortax ancestor acquired a cordiform
pronotum and the basal foveae became deeper posteriorly and shorter. Most of the
structures of the ancestor of the Cyclotrachelus-Evarthrus group underwent slight modifica-
tions. The Fortax stock diverged into the morio group and obsoletus group lineages. The
basic stock of the morio group lost the ventral lateral setae of the last tarsal articles. It
differentiated and first gave rise to laevipennis which acquired sharp and somewhat
produced basal angles of the pronotum, but retained a relatively primitive eversion of the
internal sac. A later stock in which a left eversion of the internal sac developed, diverged
into morio and hernandensis . The primitive stock of the obsoletus group acquired a cordi-
form pronotum with punctiform basal lateral foveae and basal setae which were developed
anteriorly a short distance in front of the basal angles. It evolved into the extant species
obsoletus in which developed a modified median lobe of the male, and the progenitor of
iuvenis and approximatus. The latter two species inherited a more primitive median lobe,
but iuvenis acquired a derived eversion of the internal sac, to the left and when everted
curled around the left ventral side of the median lobe. This type of eversion is convergent
with that of hernandensis and morio because iuvenis is not closely related to them.
The basic stock of Cyclotrachelus acquired a modified pronotum with constricted
posterior sides and basal setae which became situated in the head. It retained the. primitive
monostriate basal lateral foveae of the pronotum, and dorsoapical eversion of the internal
sac of the median lobe.
The complementary stock which gave rise to the subgenus Evarthrus gained more setae on
the penultimate article of the labial palpus. It also acquired modified bistriate basal lateral
foveae of the pronotum and an internal sac that everted to the right.
The ancestral stock of the subgenus Cyclotrachelus differentiated into two stocks. One
gave rise to the spoliatus group. It retained the primitive internal sac with an apical sclerite.
The other evolved into the ovulum-faber complex, and it acquired a light-colored sclerite in
the internal sac. The species brevoorti is the earliest derivative of the spoliatus group because
the other members of the group are more closely related to one another than to brevoorti.
It evolved a modified truncate apex of the median lobe and lost the apical serrulate field of
the internal sac. The stock complementary to brevoorti gave rise to unicolor and the
ancestor of fucatus and spoliatus. It retained the primitive apex of the median lobe and
light serrulate apical field in the internal sac. The species unicolor gained a few extra setae
on the penultimate article of the labial palpus, and evolved a modified internal sac. The
progenitor of fucatus and spoliatus perhaps closely resembled the latter. The acquisition of
an extra seta on the penultimate article of the labial palpus of some fucatus individuals is
probably a recent modification.
The species vinctus evolved early in the history of the ovulum-faber complex for it is not
closely related to any of the other members of the group. It inherited the primitive sclerite
of the internal sac, and evolved produced, sharp, basal angles of the pronotum. The sister
stock of vinctus acquired a cleft apical sclerite characteristic of the ovulum group and the
faber group. The one which differentiated into alabamensis and the ancestor which gave rise
to texensis, macrovulum and ovulum evolved sharp, produced basal angles of the pronotum.
172
Freitag
The early derivative alabamensis acquired a deep longitudinal groove in the prosternal
process, and lost the glossiness of the elytra, while its diverging sister stock developed
crescent-shaped frontal grooves on the head before evolving into the ancestor of macrovu-
lum and texensis and ovulum. The species ovulum developed a deep longitudinal groove in
the prosternal process, but the ancestral stock, macrovulum-texensis retained the primitive
condition of that structure. Recently, macrovulum evolved a short paramere, while the
paramere of texensis has not changed from the primitive condition. The three members of
the faber group possess a deep groove in the prosternal process which was probably a feature
inherited from the ancestral stock of this group. It first gave rise to parafaber, a somewhat
distant relative of faber and levifaber, which retained the cleft apical sclerite of the internal
sac. Later the sister stock of parafaber acquired a modified C-shaped sclerite of the internal
sac and then differentiated into faber and levifaber.
Returning to the ancestor of the subgenus Evarthrus, this stock differentiated into two
stocks. One stock retained four setae on the anterior face of the middle femur, and then it
evolved the ancestor of the incisus group and the progenitor of the blatchleyi and sigillatus
groups. The other stock gained an extra seta on the anterior face of the middle femur, and
it gave rise to the seximpressus group ancestor and the forerunner of the rest of the lineages
in the subgenus Evarthrus.
The posterior constriction of the pronotum, short right paramere and median dorsal
hump on the median lobe of the male, and the dark elongate apical sclerite of the internal
sac, which are characteristic of incisus and whitcombi, were modifications acquired by their
common ancestor.
A change in the shape of the basal lateral foveae of the pronotum and formation of a
deep medial longitudinal groove in the prosternal process were modifications which
occurred in the stock that gave rise to the ancestors of the blatchleyi and sigillatus groups.
Both characteristics are present in all of the extant species of the two groups.
The ancestor of blatchleyi and floridensis probably evolved a slightly arcuate median lobe
and narrow apical blade; both extant species probably resemble their common ancestor in
most features.
The primitive stock that differentiated into sigillatus and the ancestor of sinus and
convivus acquired a light-colored apical sclerite in the internal sac but retained the primitive
type of median lobe and parameres of the male. Because the male genitalia of sigillatus are
unlike that of convivus and sinus, I believe sigillatus is an earlier derivative of this lineage.
The species sigillatus probably evolved as the sister stock of the ancestor of convivus and
sinus. Geographical variation in non-genitalic structures of sigillatus such as the shape of the
pronotum and elytra is probably evidence of recent differentiation. The ancestor of
convivus and sinus acquired modified genitalia which were inherited by both species.
One of the extraordinary structural modifications in the history of Evarthrus was the
acquisition of apical setae on the prosternal process by the basic stock of the seximpressus
group. This ancestor gave rise to two stocks: one which differentiated into alabamae and
seximpressus, gained rounded basal angles of the pronotum; and one that gave rise to
engelmanni and nonnit ens acquired distinct produced basal angles of the pronotum.
Revision of Evarthrus
173
The relationships of hypherpiformis are not clear because besides having its own distinct
appearance it shares structures with two distinctly related groups. Specimens of this species
have the general external habitus of the members of the seximpressus group, but lack the
setae of the prosternal process. They have 3 — 5 punctures on the third interval of the
elytron and the stump-like right paramere of the male genitalia which resembles that of sallei
and gigas of the gigas group. I have placed hypherpiformis close to the seximpressus group
on the dendrogram. This position requires that similar male genitalia were evolved in un-
related lineages.
The sister stock of the one which gave rise to the seximpressus group acquired a prono-
tum with a posterior constriction and small basal angles. It gave rise to the ancestors of two
stocks. One of them retained the primitive pronotal form, and differentiated into the basal
stocks of the sodalis group. Its sister group gained prominent angles of the pronotum and
diverged into the primitive stocks of the substriatus group, and the torvus and gigas groups.
The extant species furtivus, parasodalis and sodalis are descendants of a common form
which inherited and maintained a primitive pronotum with small basal angles. The promi-
nent angles of the pronotum of sodalis colossus are probably a recent modification. Also the
narrow apical blade of sodalis has probably evolved recently. The species alternans and
iowensis evolved from an ancestor that acquired prominent basal angles of the pronotum.
The three punctures on the disc of the elytron of some specimens of iowensis is probably a
recent change.
The species constrictus and substriatus are characterized by the absence of normal ridges
between the first three anterior punctures of the umbilicate series. This feature was
probably acquired by their common ancestor. Some specimens of substriatus have three
punctures on the third interval of the elytron which is probably a recent modification.
Only insignificant structural changes developed in the stock which gave rise to the
ancestors of the torvus and gigas groups. The derivative stock which evolved the torvus
group retained the primitive pronotal shape. The presence of three punctures in the third
interval of the elytron in som egravidus, torvus and deceptus individuals is probably a recent
modification.
A more pronounced posterior constriction of the pronotum was acquired by the ancestor
of the gigas group. It first gave rise to heros which inherited the primitive type of right para-
mere of the male genitalia. The sister stock of heros' evolved a stump-like right paramere
and differentiated into the extant species sallei and gigas.
The gravesi group is not closely related to any other Evarthrus group. Its members have
lost the plica and resemble the species of Cyclotrachelus in some detail, but generally look
like western species of the subgenus Evarthrus.
According to this phyletic scheme, the following characteristics have evolved more than
once in the Evarthrus lineage: pronotum with posterior constriction, four times; posterior
setae on lateral bead, twice; deep, sharply defined medial longitudinal groove of the
prosternal process, four times; three or more setae on the third interval of the elytron, five
times; short stump-like parameres, three times; and sclerite of the internal sac, at least four
times.
174
Freitag
Zoogeography
Introduction. — The chief objective of historical zoogeography is to interpret the geo-
graphical relationships of extant organisms in terms of past climate and physiography. These
events often provide evidence of former barriers, or alternatively, opportunities for dispersal
now no longer available. When a measure of concordance is found between such evidence
and present distributions of extant species, it may be assumed that these distribution
patterns are explained by changes which occurred in the past. In turn, geographical relation-
ships of species judged to be closely related phylogenetically can provide evidence about
past changes. The distribution of the species of Evarthrus is considered from both aspects.
In the sections which follow, we provide an account of the historical background, a descrip-
tion of the distribution pattern of the extant species, and an analysis of the distribution
pattern in historical terms.
The hypotheses we propose below are weakened by two defects in our data: we cannot
relate in detail the distribution of the species to different vegetation types, although the
patterns suggest such a relationship; and many of the species are rarely encountered so that
their ranges may be incompletely known. The following essay must, therefore be regarded
as only a first approximation to a description of the geographical distribution of Evarthrus.
Our analysis is thus preliminary, but we hope not too premature.
Historical background. - The history and dispersal of the genus Evarthrus must be
related to that of the biota of eastern North America, and to the geological history of that
area. Much has been written about this topic, and the following references have been con-
sulted: Auffenberg and Milstead, 1965; Ball, 1956, 1959; Berry, 1922, 1926; Blair, 1965;
Braun, 1950; Carlston, 1950; Clarke, 1896; Coleman, 1946; Davis, 1965; Flint, 1965;
Graham, 1964; Hibbard et al, 1965; Howden, 1963, 1969; King, 1959; Muller, 1965;
Richards and Judson, 1965; Ross, 1965; Schafer and Hartshorn, 1965; Selander, 1965;
Stebbins, 1951; and Whitehead, 1965. The most important events are: the history of the
distribution of the Arcto-Tertiary Geo flora; and Pleistocene events which relate to changes
in water level, and climate.
Briefly, the Arcto-Tertiary Geoflora in the early Tertiary was Holarctic in distribution,
occurring at higher latitudes than now. At present, its elements are concentrated in more
southern temperate regions, especially eastern Asia and eastern North America. Concerning
the Pleistocene, it is fairly well established that the glacial periods had two important
effects: there was a lowering of temperatures, resulting in a certain amount of faunal and
floral shifting; and second, sea level was lowered, resulting in the appearance of additional
land in coastal areas. During interglacial stages, the reverse changes occurred, and the
Mississippi River became greatly enlarged, as a result of glacial meltwater. Thus, this river
probably became a highly effective barrier to movement of flightless terrestrial animals.
The cyclical nature of these events during the Pleistocene should have led to alternation
of range contraction and expansion of organisms, with consequent geographical isolation
resulting from fragmentation of once-continuous ranges during unfavorable time, with the
reverse taking place during favorable times. Evolution occurred as a result of both sets of
circumstances.
The distribution pattern, in general. — This is simply illustrated by means of a grid
(Table 2), oriented like a map, with columns representing longitude and rows representing
1 . This section was written jointly by G. E. Ball and me.
THE NUMBER OF SPECIES OF Evarthrus PLOTTED IN 5° INTERVALS OF LONGITUDE AND LATITUDE*
Revision of Evarthrus
175
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176
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latitude. Each square represents 5° longitude and 5° latitude, as in figs. 125—136. The
number in each square represents the total number of species recorded from within that
space in figs. 125—136. The numbers are totalled horizontally and vertically to obtain a
“total interval value” for each 5° interval of longitude and latitude, and the “average 5°
interval value” is obtained by dividing the total interval value by the number of squares
relevant to the total interval value. In addition, the total number of species found in each 5°
interval is recorded. The absence of continental land is indicated by short horizontal lines in
the appropriate squares.
In general, the number of species is maximum in the southeast (30° — 34°N., east of
90°W.), and decreases in all directions. In part, the decrease is the result of restriction of
land area (for the squares 30 - 34; 79 — 75; and 20 — 24; 80 — 84, occupied mainly by
ocean, and the square 25 - 29; 80 - 84, including peninsular Florida). However, other
factors must be considered to explain the northern, western, and southern decreases.
The simplest explanation probably involves historical factors. The genus Evarthrus was
probably warm temperate and forest adapted, and has probably undergone most of its
evolution under these conditions. All three subgenera are represented in warm temperate
forest. With one exception, spread into peripheral areas has been accomplished by the most
highly evolved subgenus, Evarthrus s. str. The exception is£. (Cyclotrachelus) faber, which
is represented in the Florida keys, at the southeastern extremity of the range of the genus.
Howden (1969) suggests that the principal factor accounting for reduction of species
toward the north is Pleistocene glaciation and its associated drastic climatic and other
effects on species living in or near glaciated areas. We agree that these factors probably
influenced the diversity of Evarthrus toward the north, but we also believe that even if
glaciation had not taken place the genus would still have exhibited maximum diversity in
the southeast.
The Floridian Evarthrus fauna seems to be especially poor. Could this be the result of
climate encountered on the peninsula? Rohwer and Woolfenden (1969) provide historical
evidence that peninsular climate is one of the major factors accounting for Florida’s de-
pauperate breeding bird fauna, and reasoning by analogy we suggest that the impoverish-
ment of the Evarthrus fauna can be accounted for in similar fashion.
Simply expressed, the distribution pattern of Evarthrus is one of subtraction from a fauna
of maximum diversity on the eastern Gulf Coast. However, in detail, the distribution pattern
is much more complex, and the complexities must be described before it is possible to
present a more detailed account of historical zoogeography. For purposes of this analysis,
taxa of subspecific and specific rank are treated as equivalent. This more detailed analysis is
considered under four headings: extent of range, centers of concentration, effects of the
Pleistocene, and species pairs.
Extent of range. — One of the striking features of the distribution pattern is the relatively
small range of many of the species. This is illustrated by the data presented in Table 3. We
used as an index of range extent the linear distance between the most widely separated
localities because it was simply obtained and was sufficiently accurate for our purposes.
Because gravesi and parafaber are known from single localities only, they were excluded
from this analysis. Note that almost one-half of the taxa have ranges less than 500 miles
long, and that 80% of the taxa have ranges of 1 000 miles or less.
Revision of Evarthrus
177
TABLE 3
FREQUENCY DISTRIBUTION OF MAXIMUM LINEAR EXTENT OF
GEOGRAPHICAL RANGE IN MILES OF THE SPECIES OF Evarthrus.
CLASS N
2001 -2250 2
1751 -2000 1
1501 - 1750 3
1251 - 1500 1
1001 - 1250 2
751 - 1000 8
501 -750 5
251 - 500 10
0-250 12
Restricted ranges seem to suggest the existence of barriers to dispersal, or alternatively,
that the present ranges of many of the species are less extensive than they once were, and
that the surviving populations are relics of a once more extensive group of populations. This
second alternative is discussed below.
A simple test for the existence of barriers would be to compare the present known ranges
of species with physical features of the landscape suspected of being effective barriers to
flightless, lowland terrestrial animals. In the east are two obvious candidates: the Mississippi
River and the Appalachian Mountains. Data on distribution of species and subspecies with
reference to the Mississippi River are presented in Table 4.
TABLE 4
COMPARISON OF DISTRIBUTION OF THE SPECIES OF Evarthrus
IN RELATION TO THE MISSISSIPPI RIVER
178
Freitag
Two groups are compared: the total Evarthrus fauna and that segment of the fauna whose
taxa are either known to reach the banks of the Mississippi or can reasonably be expected to
be there. Both sets of data show that less than one-third of the total number of species are
represented on both sides of the river. Additional details are provided in the following
section.
The effectiveness of the Appalachian system as a barrier was measured in terms of the
distribution patterns of 1 1 taxa whose ranges flank the mountains (Table 5). Less than
one-third of these taxa occur on both sides. Of this group, two species are mainly to the east
and one is mainly to the west of the mountains.
TABLE 5
COMPARISON OF DISTRIBUTION OF THE SPECIES OF Evarthrus
IN RELATION TO THE APPALACHIAN MOUNTAINS
These data appear to support the view that topographical features are important in re-
stricting the ranges of the species of Evarthrus.
Centers of concentration. — Although the distribution patterns of the extant taxa of
Evarthrus appear to have been influenced by obvious barriers, these are not the limiting
factors for all of the species. This can be seen most clearly by a comparison of the patterns
of the species with more or less restricted ranges.
The object of this comparison was to attempt to locate centers of concentration, to be
used in conjunction with a consideration of historical zoogeography of the species of
Evarthrus. Excluded from the analysis initially were the wide-ranging species, and those
represented on both sides of the major barriers. The ranges of the remaining species were
examined for concordance, and centers of concentration were discovered. Subsequently, the
species with restricted ranges but occurring in two or more centers of concentration were
excluded. The species included in the analysis are called “centrant” species; those excluded
are “radiant” species (terminology after Hulten, 1937).
The distribution patterns of the centrant species are indicated in Table 6. The vertical
lines represent lines of longitude, the horizontal lines represent lines of latitude. Each
resulting square containing one or more species was lettered. The squares which contained a
number of species in common were combined to provide the centers of concentration, as in
Table 7.
Revision of Evarthrus
179
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TABLE 7
CENTERS OF CONCENTRATION OF THE SPECIES OF Evarthrus.
1. Letters are designations for intervals in Table 6.
2. species gravesi is included, from E.
3. species nonnitens is included, from E.
The locations of the centers are illustrated in Fig. 138. We are doubtful about the reality
of Center 6. It is based on the presence of two species there, known from nowhere else.
Center 7 contains no endemic species, but it seems central for a number of wide-ranging
western species.
These Centers of Concentration, several of which are at least partly independent of the
Appalachian-Mississippi barriers, suggest the existence of additional barrier systems. The
restricted ranges may be simply by-products of biotype impoverishment, discussed below.
The distribution of all species, in terms of these centers of concentration, is indicated in
Table 8.
Revision of Evarthrus
181
TABLE 8
DISTRIBUTION OF THE SPECIES OF Evarthrus IN RELATION TO THE
CENTERS OF CONCENTRATION
NAME OF SPECIES
morio
hernandensis
laevipennis
iuvenis
approximatus
obsoletus
brevoorti
spoliatus
fucatus
unicolor
vinctus
alabamensis
macrovulum
texensis
ovulum
parafaber
levifaber
faber
incisus
whitcombi
blatchleyi
floridensis
sigillatus
sinus
convivus
seximpressus
alabamae
engelmanni
nonnitens
hypherpiformis
s. sodalis
s. colossus
s. lodingi
parasodalis
furtivus
altemans
iowensis
substriatus
constrictus
t. torvus
t. deceptus
gravidus
sallei
gigas
heros
gravesi
TOTALS
CENTER NUMBER
4 5 6 7
8
182
Freitag
Degree of difference among these centers is indicated in Table 9, by means of an index of
difference (Greenslade, 1968). Of 28 comparisons, 12 were scored as 100, i.e., pairs of
centers shared no species in common. Five scored between 91 and 94.
TABLE 9
DISSIMILARITY VALUES AMONG CENTERS OF CONCENTRATION
OF THE GENUS Evarthrus.
CENTER CENTER NUMBER
11
4
^ x 100 64
1. Total number of species in each pair of centers.
2. Number of species in common between each pair of centers.
Revision of Evarthrus
183
In general, the peripheral centers differ strongly from one another. Center 7, for example,
shares no species with Centers 1, 2 and 3, and Center 8 shares a single species with Centers
2 and 3. Centers 4 and 8 are each connected with six other Centers, and, consequently, are
the least distinctive.
Because of its central position, one would expect Center 4 to be minimally distinctive.
Center 8, on the other hand, is peripheral, and for this reason, might be expected to hold a
more distinctive fauna. The expectation is not realized, probably because the area, although
suitable for a variety of species, has few endemics.
In striking contrast to Centers 4 and 8 is Center 1 . It shares species with only one other
Center, and consequently, has the most distinctive fauna of any Center.
A more interesting comparison involves the Centers of about the same latitude, but on
different sides of the Mississippi River. Centers 2 (southeastern) and 5 (southwestern), have
a combined fauna of 35 species, of which only two are shared (Index 94). Centers 4 (north-
eastern) and 8 (northwestern) have a combined fauna of 13 species of which three are
shared (Index 77). In each case, few species occur on both sides of the river. The difference
in Index values is the result mainly of the much more diverse southern fauna. However, this
fact is of less interest than is the thought that the effectiveness of the river as a barrier is
less in the north than in the south. This may be the result of one of two factors: the river,
being narrower, is easier to cross, and consequently a greater proportion of the northern
fauna has done so; or, the species occurring in the north are more adept at crossing water
barriers. Probably both factors are involved.
The barriers marking off other centers are not clear. Probably they are complexes of
direct climatic factors, and indirect ones, transmitted through conditions of soil and vege-
tation. Barriers of this type may have been as consequential in the development of diversity
in Evarthrus as have been the more obvious barriers. How did these barriers develop? We
suggest it was through loss of variability brought about as a by-product of the glacial stages.
Effects of the Pleistocene epoch. — The fact that Centers of Concentration of species
with limited ranges can be identified suggests the operation of some powerful factor. We
suggest that this factor was climatic fluctuation during the Pleistocene, which is known to
have caused range expansion and contraction of many species. Especially during glacial
stages, one might expect the ranges of more or less warm adapted species to have become
more or less restricted, with a consequent reduction in population size and number. During
interglacials and post-glacial time, the ranges of more broadly adapted species, the radiants,
might be expected to expand, whereas species which lost too much variability as a result of
reduction in population size and number would not be able to spread. This is essentially
Hulten’s Theory of Equiformal Progressive Areas (1937).
The position of the Centers of Concentration probably do not coincide precisely with the
former distribution of the centrant species during the Pleistocene epoch. The northern
species must have been south of their present ranges during the glacial stages, and the
coastal species must have been north of their present ranges during the interglacial periods.
However, at a more general level, a correlation is apparent. The centrant species on each
side of the Mississippi River and Appalachian Mountains were probably in the same longi-
tudinal zones in the past as they are now, and it seems unlikely that the ranges of centrant
184
Freitag
species from different centers overlapped. Can this be taken as evidence that the south-
eastern biota was not radically displaced by the cold glacial periods (Braun, 1950)? We
think it can be.
Thus the significance of the Centers is that they indicate in part positions of refugia
during the glacial stages. One can also infer from the locations that different species have
different ecological tolerances of rather narrow range, and that the evolution of diversity in
Evarthrus has involved development of these limited tolerances. Although evolution in
general seems to have led to the development of species of limited tolerances, especially in
the east, more broadly adapted species have evolved in the west. Some of the latter may be
of relatively recent origin, an inference based on their postulated phylogeny.
Species-pairs. - The analysis of the distribution of pairs of closely related species (sister
species) seems to be the most fruitful approach to historical zoogeography at the local level.
Their distributions are likely to provide the clearest evidence of location of former barriers
and of patterns of phylogeny. Of course, this is true only if the two sister species are largely
allopatric.
The two members of some species pairs are largely or completely sympatric, and conse-
quently, their present distribution gives no indication about their past geographical relation-
ships. These are: morio — hernandensis; iuvensis — approximate; alternans — iowensis;
substriatus — constrictus; and torvus — gravidus. These are excluded from the following
discussion. Also excluded are gravesi and hypherpiformis, both representing monotypic
groups and lacking close living relatives.
The distribution patterns are of several types: east — west; Floridian — Gulf Coast; and
north — south (Table 10). The last category is common among the western species of the
subgenus Evarthrus.
The range disjunctions of most of the east-west pairs of vicariant species are correlated
with the Mississippi River or with the Appalachian Mountains. In these cases, speciation has
taken place between present Centers of Concentration, and it is easy to imagine how the
isolation and consequent differentiation came about. There are also several instances of
vicariant distributions within Centers 2 and 5 . These involve species with coastal ranges, and
one can imagine the requisite range disjunctions arising as a result of changes in coast line
with fluctuation in sea level and sizes of rivers during the waxing and waning of the glacial
stages. The same explanation is relevant for vicariant distributions within Florida, and be-
tween Florida and the mainland.
The north-south disjunctions are more difficult to explain, because, for the most part,
obvious barriers do not occur between the northern and southern members of the pairs.
However, during glacial stages, one can imagine that range restriction occurred, as described
above, with isolates of once more wide-ranging species surviving and differentiating.
Vicariance relationships of these same kinds are evident within sister species-groups, and
are presumably the result of the same causes having been effective at an earlier time in the
phylogeny of the genus.
These distribution patterns suggest to us a model of how diversity has been generated in
the genus Evarthrus. Beginning with a species of relatively restricted range and tolerance
(Stage A), range expansion takes place as tolerances are expanded (Stage B). If a wide
Revision of Evarthrus
185
TABLE 10
DISTRIBUTION PATTERNS OF ALLOPATRIC SISTER SPECIES
OF THE GENUS Evarthrus
A. EAST - WEST RELATIONSHIPS
WESTERN VICAR EASTERN VICAR
186
Freitag
temperature tolerance evolves, the species may spread longitudinally into climatic zones
different from the ancestral one. If not, the species spreads latitudinally. Major physical
barriers may be crossed, or barriers may appear and become effective subsequently, breaking
the range of Stage B species. Differentiation of the resulting segregates takes place, leading
initially to divergence to the subspecies level (Stage C), and later to the species level (Stage
D). At this stage, two allopatric species are present in place of the original ancestral species.
Next, the ranges of these sister species may expand. The sister species may meet one
another, and eventually, the two sister species become sympatric (Stage E). Examples of all
of these stages are found among the extant species of Evarthrus. Stage A, Stage D and some
Stage E species form the nuclei of the Centers of Concentration — the centrants. Stage B
and Stage E species usually are widespread in more than one center — the radiant species.
It is suggested that the barriers are provided by permanent physiographical features, such
as mountains, rivers and the sea, or by temporary interruptions of ranges as a result of
Pleistocene climatic change.
Historical zoogeography. — In this section, an attempt is made to reconstruct the geo-
graphical history of the genus Evarthrus, based to a large extent on the considerations
presented above. The primitive ancestry of this genus became separated from its Palaearctic
sister stock (the ancestor of the Palaearctic molopine genera) sometime in the early Tertiary,
when the Bering Land Bridge was covered by the sea. This New World group inhabited the
Arcto-Tertiary forests in warm temperate regions during the middle Tertiary, possibly
ranging across the north from the west coast to the east. As this biota withdrew from the
west, so did Evarthrus, ultimately becoming restricted to the east. Probably representatives
of the genus arrived too late on the Gulf Coast to be able to enter eastern Mexico. This
suggests that they arrived in the area in post-Miocene time. Martin and Harrell (1957)
suggest that grasslands formed in this area in the Miocene. Had the early members of
Evarthrus been present before the grasslands formed they might be expected to be found
today in Mexico.
Because of the present extensive overlap in ranges of the extant subgenera, it is impossible
to guess the geographical relationships of these groups at the time of their origin. And, for
the same reason, it is impossible to reconstruct much of the subsequent history of the genus.
However, it is possible to suggest events which led to the development of many of the
extant species pairs.
The general assumption is that much of the evolution of markedly similar allopatric spe-
cies took place in the Pleistocene, a time span of several million years. Zoogeographers, such
as Blair (1958), Hubbell (1954) and Howden (1963), have suggested that such species have
arisen during the Pleistocene, because of the coincidence of the ranges of such species with
barriers that could have arisen during this time, only. However, on the basis of fossil
evidence, Shotton (1965) and Lindroth (1963) have denied that any stock diverged to
species level. The difference may be that in general, the more widespread species have been
collected as fossils, and they may be older, less rapidly evolving. In any event, it is clear that
a number of vertebrates have undergone evolution to the species level during the Pleistocene,
and we fail to see that insects should not have done the same. The taxa are discussed below,
in phylogenetic sequence.
Revision of Evarthrus
187
The morio and obsoletus groups of the subgenus Fortax are largely allopatric (Fig. 125,
cf. Fig. 126). The morio group occupies the Gulf Coastal Plain, while the obsoletus group
has a more northern distribution. Possibly the ancestral stock of these groups became
divided during early Pleistocene or late Pliocene time into a northern and southern stock.
Speciation of hernandensis and morio of the morio species group undoubtedly took place
in Floridian refuges, possibly during an interglacial period, when Florida was isolated by
water barriers from the mainland. Both species are presently almost completely confined to
Florida (Fig. 125). Because these are sympatric their geographical history is difficult to de-
termine. The species laevipennis and the ancestor of morio and hernandensis were probably
separated during an early interglacial. E. laevipennis evolved on the mainland coast just west
of Florida, and has moved northward in recent times.
The Appalachian Mountains are an effective barrier between the species obsoletus and the
other two members of the obsoletus group (Fig. 126). The obsoletus group ancestor prob-
ably became disjunct and gave rise to obsoletus west of the mountains while its sister stock
which ultimately give rise to approximatus and iuvenis speciated east of the mountains.
Since approximatus and iuvenis are very closely related and sympatric, little can be said
about the history of their geographical distribution.
Widespread sympatry among the species groups of the subgenus Cyclotrachelus masks
most of their distributional history. Within the species groups, however, the distribution
pattern among some species merits comment.
In the spoliatus group the more remotely related species brevoorti and unicolor are
sympatric on the Gulf Coastal Plain (Fig. 127). Little can be said about their geographical
history. The sister species fucatus and spoliatus flank the Appalachians. Since both extant
species are cool tolerant the separation of the ancestral population could have occurred as
follows: the ancestral population was distributed somewhere at the southern end of the
Appalachian System during a glacial period; with the coming of an interglacial the biota
began to shift northward and along with it went the ancestral population; as it moved north
it became divided at the base of the mountains; the northward shift continued and the two
populations, now completely separated, moved north one on either side of the mountains,
then differentiated into fucatus and spoliatus.
Each species of the ovulum group has a highly restricted geographical distribution (Fig.
128). The species vinctus is confined to high elevations of the southern Appalachians. It
may have become disjunct in the mountains during early stages of the Pleistocene. The
remaining species, alabamensis and the closely related ovulum, macrovulum, and texensis
are southern Gulf Coastal forms. The ancestor which gave rise to alabamensis and the basic
stock of ovulum, macrovulum and texensis was probably distributed along the coast from
southern Alabama down into Florida. During an early interglacial the Florida stock of the
species probably became isolated in a Florida refuge where it evolved, while the mainland
population evolved into alabamensis . With the advent of another glacial period and conse-
quent lowering of sea level, Florida was again united with the mainland. The ancestor of
ovulum, macrovulum, and texensis then moved north and became distributed from Florida
along the coast to Mobile, Alabama where it became sympatric with alabamensis. In the
following interglacial period this stock, just as its ancestor, became divided into a mainland
188
Freitag
population and one separated on a Floridian island. The mainland form differentiated into
the ancestor of macrovulum and texensis, and the other into ovulum. The macrovulum-
texensis stock crossed the Mississippi River, and differentiated into two species.
The zoogeography of the faber group probably parallels that of the ovulum group (Fig.
129). The ancestor of faber and levifaber , and parafaber probably ranged along the Gulf
Coast from Mobile, Alabama to peninsular Florida. During an interglacial it became divided
into a mainland population which speciated into parafaber and a Floridian isolate which also
speciated. During the following glacial period the Florida population radiated. When the
next interglacial occurred this population was separated. One portion became isolated in a
Florida refuge, while another part remained on the mainland northeast of Florida. Both
populations differentiated giving rise to faber in Florida and to levifaber on the mainland.
The subgenus Evarthrus is the only subgenus with extensive representation west of the
Mississippi River, although it is rich in species to the east of the river, as well. Clearly the
subgenus has been on both sides of the Mississippi for an extended period of time.
The two species of the incisus group are exclusively or largely west of the Mississippi
River (Fig. 130), and of these whitcombi is confined to Center 6, south of the Arkansas
River. To the north occurs the radiant species incisus. Possibly the Arkansas River consti-
tuted a partial barrier to dispersal, and the incisus stock differentiated to the north and
south of that barrier.
Figure 131 illustrates the distribution of the taxa of the blatchleyi and sigillatus groups.
The barrier effect of the Mississippi River and the Appalachian System is shown in remark-
able clarity. In addition Florida contains two more essentially endemic species. The ances-
tral stock of the blatchleyi group undoubtedly had a southern distribution perhaps near
northeastern Florida, while its sister stock which gave rise to the sigillatus group was isolated
further north. Possibly during an interglacial, the species floridensis evolved in a Floridian
refuge while blatchleyi evolved on the mainland near Florida; then during the following
glacial period blatchleyi moved south into Florida. Alternatively, the two could have evolved
from a discontinuously distributed Floridian stock.
Because of Pleistocene north-south biotic shifts, the ancestral stock of the sigillatus group
was divided into two populations one on either side of the Appalachians. That on the east
evolved into sigillatus. The western population then became disjunct. The species sinus must
have evolved on the Gulf Coast, while convivus evolved further north.
Extant species of the seximpressus group are mainly Gulf Coastal Plain species where they
are largely sympatric (Fig. 132). Only seximpressus ranges northward to Minnesota and Wis-
consin. Two species alabamae and nonnitens, have managed to cross the Mississippi River.
The ancestral stocks of the present species probably evolved on or near the Gulf Coastal
Plain. The species seximpressus probably evolved further north as a cool adapted form,
while the other three species arose on the Gulf Coastal Plain.
Because the relationships of the hypherpiformis group are not clearly understood there is
nothing to write concerning the zoogeography of it (Fig. 132).
All of the extant species of the sodalis group are relatively northern forms and as a group
cross the northern headwaters of the Mississippi River (Fig. 133). There is no doubt that the
primitive ancestors of this species group also had northern ranges and were cool adapted.
Revision of Evarthrus
189
Geographical variation in the species sodalis is probably recently acquired because at present
some distinct populations are partially isolated west of the Missouri River and some in
northern Alabama. The species parasodalis is confined to central and western Arkansas
while its sister species sodalis occurs further north, where it probably evolved.
The species furtivus may have been pinched off to the east of the Appalachians from its
immediate ancestral population during a general southward biotic glacial shift (Fig. 133).
The closely related species alternans and iowensis are sympatric (Fig. 134). They may
have evolved in the general region where they are now found, near the upper reaches of the
Mississippi River, but they could not have survived the glacial stages in this area. Therefore,
their present distribution is post-glacial.
The sympatric species substriatus and constrictus, which make up the substriatus group,
are two of the most widespread forms in the genus Evarthrus (Fig. 134). Together they
occupy the Great Plains from Durango, Mexico north to Minnesota and from central
Arizona east to the margin of the Mississippi River valley. They evolved in the Great Plains.
They, along with the species torvus and possibly gravidus are the only extant species of
Evarthrus which occur mainly on dry prairie.
E. gravidus occurs mainly in southeastern Texas but it has been collected as far west as El
Paso, Texas (Fig. 135). It is not as widespread in the Great Plains as its sister species torvus.
The subspecies of torvus occur north and south of one another in the Great Plains. This
suggests that southeastern Texas and the northern Great Plains could be centers of specia-
tion in which population became disjunct during the south-north Pleistocene biotic shifts.
E. gravidus could have speciated in isolation in southeastern Texas after a northward inter-
glacial shift while torvus speciated in the northern Great Plains. A southern shift during the
next glacial stage (Wisconsin?) brought torvus south into gravidus territory. During the next
interglacial (Recent?) again torvus moved north and southern and northern populations
which were partially separated subspeciated.
Because the members of the gigas group are largely sympatric the historical zoogeography
of the gigas group is almost impossible to reconstruct (Fig. 136). Speciation must have
occurred in the southeastern Great Plains area.
Phylogenetic relationships between the species gravesi and other Evarthrus species is
uncertain. We cannot comment on the geographical history of this species.
The present distributions of the species of Evarthrus indicate that Pleistocene events
profoundly affected the genus as a whole. At the present time forest species which live in
warm temperate regions have distinctly restricted distributions. Conversely, cold tolerant
and/or dry tolerant species are clearly more widespread and some may be undergoing further
divergence at the present time. Those species which were specifically suited for warm tem-
perate forest conditions, i.e. species on the Gulf Coastal Plain, probably moved southward
along with their biotic neighbourhood as a northern ice mass developed. Species which
could not tolerate even a slight depression in the temperature probably perished along the
Gulf Coastal Plain. Some moved into peninsular Florida. Other slightly more broadly adap-
ted forest species probably moved southward or eastward on the Gulf Coastal Plain as sea
level dropped and land emerged. At the height of the glacial periods, the ranges of these
species may have been greatly reduced, and their ranges have remained restricted to the
present time.
190
Freitag
On the other hand broadly adapted species made adaptive shifts during the Pleistocene.
Some became successful cold tolerant forms and others invaded dry regions west of the
Mississippi River. These species are dominant at the present time.
Because the species of Evarthrus are wingless it is not surprising that water is an effective
barrier to these species. The Mississippi drainage system and ocean around Floridian islands
were undoubtedly the two most important water barriers in the history of this genus. In
addition, another great obstacle was, and remains the Appalachian Mountains. These barriers
in conjunction with Pleistocene climatic changes forged the geographical patterns of the
present species of Evarthrus.
ACKNOWLEDGEMENTS
I am indebted to G. E. Ball for his guidance, aid, suggestions, encouragement and
constant interest throughout this study, and for reading and editing the manuscript.
I thank Dr. B. Hocking for reading the manuscript and offering suggestions concerning
this problem. I also thank C. H. Lindroth, W. G. Evans, and H. F. Clifford for reading and
criticizing the thesis on which this paper is based.
I thank the following institutions and persons for the loan of specimens: Norman
Anderson, Montana State University; T. C. Barr, Jr., University of Kentucky; R. T. Bell,
University of Vermont; H. D. Blocker, Kansas State University; H. R. Burke, Texas A & M
University; G. W. Byers, University of Kansas; L. Chandler, Purdue University; C. V. Covell,
Jr., University of Louisville; P. J. Darlington, Jr., Museum of Comparative Zoology at
Harvard University; T. L. Erwin, University of Alberta; K. L. Esau, Iowa State University;
Leonora K. Gloyd, State Natural History Survey Division, Urbana, Illinois; R. C. Graves,
Bowling Green State University; Kirby L. Hays, Auburn University; T. Hlavac, Michigan
State University; T. H. Hubbell, University of Michigan; V. M. Kirk, United States Depart-
ment of Agriculture; J. L. Laffoon, Iowa State University; D. J. Larson, University of
Calgary; Hugh B. Leech, California Academy of Sciences; J. E. H. Martin, Canada Depart-
ment of Agriculture; L. L. Pechuman, Cornell University; H. Radcliffe Roberts, Academy of
Natural Sciences, Philadelphia; P. Rouse, University of Arkansas; J. B. Schmitt, Rutgers
University; R. D. Shenefelt, University of Wisconsin; P. J. Spangler, Smithsonian Institution,
United States National Museum; P. Vaurie, American Museum of Natural History; G. E.
Wallace, Carnegie Museum; R. L. Wenzel, Chicago Natural History Museum; D. R. White
head, University of Alberta; R. Woodruff, State Plant Board of Florida; D. A. Young, North
Carolina State University.
I thank P. J. Darlington, Jr., (Museum of Comparative Zoology) and P. J. Spangler,
(United States National Museum) for permitting me to examine type specimens in then-
institutions, and for their kind hospitality.
R. B. Madge (British Museum of Natural History, London), and Jacques N£gre, (Museum
Nationale d’Histoire Naturelle, Paris) compared material with type specimens. I am grateful
to both gentlemen.
I would like to thank my wife Gayla for typing and reading the manuscript.
Revision of Evarthrus
191
I thank L. L. Kennedy of the University of Alberta, who kindly identified fungus zygotes
which were found in the gut of an Evarthrus specimen.
Thanks are also due to: D. R. Whitehead for criticizing the key and reading the manu-
script; David Larson for accompanying me on a collecting trip to southeastern United States
and for his criticisms of the key; J. Barron, T. Erwin, and R. E. Leech for criticizing the key
and for suggestions on other aspects of this problem; J. Scott for photographing specimens
of Evarthrus, and assisting with the preparation of distribution maps.
REFERENCES
Auffenberg, W., and W. W. Milstead. 1965. Reptiles in the Quaternary of North America,
pp. 557-568. In H. E. Wright, Jr., and D. G. Frey (ed.). The Quaternary of the United
States. Princeton University Press, Princeton.
Ball, G. E. 1956. A revision of the North American species of the genus Helluomorphoides
Ball, 1951 (Coleoptera, Carabidae, Helluonini). Proc. ent. Soc., Washington, 58(2) : 67-
91.
Ball, G. E. 1959. A taxonomic study of the North American Licinini with notes on the Old
World species of the genus Diplocheila Brulle (Coleoptera). Mem. Amer. ent. Soc., 16,
258 p.
Ball, G. E. 1960. Carabidae (Latreille, 1810), 55-181 pp. In Ross H. Arnett, Jr., The Beetles
of the United States. The Catholic University of America Press, Washington.
Ball, G. E. 1966. A revision of the North American species of the subgenus Cryobius
Chaudoir (Pterostichus , Carabidae, Coleoptera). Opusc. ent. Supp. 28, 166 pp.
Ball, G. E. 1967. Book review (of Lindroth, C. H. 1966. Part 4, Ground beetles of Canada
and Alaska). Coleopterists Bulletin, 21(2) : 40, 56, 58, 63, and 64.
Basford, N. L., J. E. Butler, C. A. Leone and F. J. Rohlf. 1968. Immunologic comparisons
of selected Coleoptera with analyses of relationships using numerical taxonomic methods.
Syst. Zool., 17(4) : 388-406.
Bates, H. W. 1882. Biologia Centrali-Americana, Insecta. Coleoptera, Carabidae. 1(1) : 40-
152.
Berry, E. W. 1922. Additions to the flora of the Wilcox group. U. S. Geol. Sur. Prof. Paper.
13 : 1-121.
Berry, E. W. 1926. Pleistocene plants from North Carolina. U. S. Geol. Sur. Prof. Paper.
140 : 97-119.
Blackwelder, R. R., and R. M. Blackwelder. 1948. Fifth supplement 1939-1947 (inclusive)
to the Leng catalogue of Coleoptera of America, north of Mexico. Mount Vernon, New
York. 87 p.
Blair, W. F. 1958. Distributional patterns of vertebrates in the southern United States in
relation to past and present environments. In Hubbs, Carl. L. (editor). Zoogeography.
Publication No. 5 1 of the American Association for the Advancement of Science, Wash-
ington, D. C. pp. 433-468, 1 1 figs.
Blair, W. F. 1965. Amphibian speciation, 543-556 pp. In H. E. Wright, Jr., and D. G. Frey,
(ed.). The Quaternary of the United States. Princeton University Press, Princeton.
192
Freitag
Blatchley, W. S. 1910. The Coleoptera or beetles of Indiana. Bulletin of the Indiana Depart-
ment of Geology and Natural Resources. No. 1 , 1386 p.
Boving, A. G. and F. C. Craighead. 1930. An illustrated synopsis of the principal larval
forms of the order Coleoptera. Ent. amer. 1 1 : 1-351.
Braun, E. L. 1950. Deciduous forests of eastern North America. Blakiston Company, Phila-
delphia, 596 p.
Brimley, C. S. 1938. The insects of North Carolina. North Carolina Department of Agricul-
ture. Division of Entomology. 560 p.
Carlston, C. W. 1950. Pleistocene history of coastal Alabama. Geol. Soc. Amer. BulL 61 :
1119-1130.
Casey, T. L. 1918. Amarinae and Pterostichinae. Memoirs on the Coleoptera. Lancaster
Press Inc., Lancaster, Pa. 8, 427 p.
Casey, T. L. 1920. Random studies among the American Caraboidea. Memoirs on the
Coleoptera. Lancaster Press Inc., Lancaster Pa. 9, 529 p.
Casey, T. L. 1924. Additions to the known Coleoptera of North America. Memoirs on the
Coleoptera. Lancaster Press Inc., Lancaster Pa. 11, 347 p.
Chaudoir, M. de. 1838. Tableau d’une nouvelle subdivision du genre Feronia Dejean suivi
d’une characteristique de trois nouveau genres de carabiques. Bull. Soc. Nat. Moscou,
1 1 : 3-32.
Chaudoir, M. de. 1868. Observations synonymiques sur les carabiques de l’Amerique septen-
trionale et descriptions d’especes nouvelles de ce pays. Extrait de la Revue et Magasin de
Zoologie. p. 1-64.
Clark. W. B. 1896. The Eocene deposits of the Middle Atlantic Slope in Delaware, Maryland,
and Virginia. U. S. Geol. Sur. Bull. 141, 167 p.
Coleman, A. P. 1946. The last million years. The University of Toronto Press, Toronto. 216
P-
Csiki, E. 1930. Pars 112. Carabidae:Harpalinae, 4(2) : 529-737. In W. Junk, and S.
Schenkling, (ed.). Coleopterorum catalogus auspiciis et auxilio. Berlin and ’s-Gravenhage.
Davis, M. B. 1965. Phytogeography and palynology of northeastern United States, p. 377-
402. In H. E. Wright, Jr., and D. G. Frey (ed.). The Quaternary of the United States.
Princeton University Press, Princeton.
Dejean, P. F. M. A. 1828. Species general des coleopteres de la collection de M. le compte
Dejean, Paris. 3, 556 p.
Flint, R. F. 1965. Introduction: Historical Perspectives, p. 3-11.7/2 H. E. Wright, Jr., and D.
G. Frey, (ed.). The Quaternary of the United States. Princeton University Press,
Princeton.
Germar, E. F. 1824. Insectorum species novae aut minus cognitae, descriptionibus illustra-
tae. Halae. 624 p.
Graham, A. 1964. Origin and evolution of the biota of southeastern North America:
evidence from the fossil plant record. Evolution 18(4) : 571-585.
Greenslade, P. J. M. 1968. Island patterns in the Solomon Islands bird fauna. Evolution,
22(4) : 751-761.
Haldeman, S. S. 1853. Descriptions of some new species of insects, with observations on
described species. Proc. Acad. nat. Sci. Philad. 6 : 361-365.
Revision of Evarthrus
193
Hibbard, C. W., D. E. Ray, D. E. Savage, D. W. Taylor, and J. E. Guilday. 1965. Quaternary
mammals of North America. In H. E. Wright, Jr., and D. G. Frey, (ed.). The Quaternary
of the United States. Princeton University Press, Princeton.
Horn G. H. 1875. Synonymical notes and descriptions of new species of North American
Coleoptera. Trans. Amer. ent. Soc. 5 : 126-156.
Horn, G. H. 1881. On the genera of Carabidae with special reference to the fauna of Boreal
America. Trans. Amer. ent. Soc. 9 : 91-196.
Horn, G. H. 1886. Notes on the Biologia Centrali-Americana. Trans. Amer. ent. Soc. 13 :
1-16 (Proc.).
Howden, H. F. 1963. Speculations on some beetles, barriers and climates during the
Pleistocene and pre-Pleistocene periods in some non-glaciated portions of North America.
Syst. Zool. 12 : 178-201.
Howden, H. F. 1969. Effects of the Pleistocene on North American insects. Ann. Rev. Ent.,
14 : 39-56.
Hubbell, T. H. 1932. A revision of the Puer Group of the North American Genus Melanoplus
(Orthoptera, Acrididae). Misc. Publ. Univ. Mich. Mus. Zool. 23 : 1-64.
Hubbell, T. H. 1952. Distribution of the Furcatus Group of Melanoplus , map and discus-
sion, 385 p. In Rodgers, Hubbell, and Byers, Man in the Biological World. McGraw-Hill,
New York.
Hulten, E. 1937. Outline of the history of arctic and boreal biota during the Quarternary
Period. Bokforlags Akteibolaget Thule, Stockholm. 168 pp., 42 plates.
Jeannel, R. 1942. Coleopteres Carabiques, 2me partie. Faune de France. Paris. 40 : 573-
1173.
Jeannel, R. 1948. Coleopteres Carabiques de la Region Malgache, 2me partie. Faune de
l’Empire Francais. 19 : 373-765.
King, D. B. 1959. The evolution of North America. Princeton University Press, Princeton.
189 p.
Lacordaire, J. T. 1854. Histoire naturelle des insectes. Genera des Coleopteres ou expose
methodique et critique de tous les genres proposes jusqu’ici dans cet ordre d’insectes.
Paris. 1 , 486 p.
LeConte, J. L. 1848. A descriptive catalogue of the geodaphagous Coleoptera inhabiting
the United States east of the Rocky Mountains. Annals of the Lyceum of Natural History
of New York, 4 : 173-474.
LeConte, J. L. 1852. Synopsis of the species of Pterostichus Bon and allied genera inhabiting
temperate North America. J. Acad. nat. Sci. Phil. 2 : 225-256.
LeConte, J. L. 1858. Catalogue of Coleoptera of the regions adjacent to the boundary line
between the United States and Mexico. Acad. Nat. Sci. Phil. 4 : 9-42.
LeConte, J. L. 1863a. List of the Coleoptera of North America. Prepared for the Smith-
sonian Institution. Smithson. Misc. Coll., No. 140, 78 p.
LeConte, J. L. 1863b. New species of North American Coleoptera. Prepared for the Smith-
sonian Institution. Smithson. Misc. Coll., No. 167, 92 p.
LeConte, J. L. 1870. Types of Mr. Newman (British Museum). Ann. Mag. nat. Hist. 4 : 5.
LeConte, J. L. 1873. The Pterostichi of the United States. Proc. Acad. nat. Sci. Phil. 302-
320 pp.
194
Freitag
LeConte, J. L. 1876. II. Coleoptera of southern Colorado and northern New Mexico,
collected in 1875, by Lieut. W. L. Carpenter, Ninth Infantry, 519-520 p. Report upon
new species of Coleoptera collected by expeditions for geographical surveys west of the
100 meridian, Lieut. G. M. Wheeler, Corps of Engineers, U. S. Army, in charge.
LeConte, J. L. and G. H. Horn. 1883. Classification of the Coleoptera of North America.
Smithson. Misc. Coll. No. 507, 567 p.
Leng, C. W. 1915. List of the Carabidae of Florida. Bull. American Mus. Nat. Hist., 34 :
555-601.
Leng, C. W. 1920. Catalogue of the Coleoptera of America, north of Mexico. Mount Vernon,
New York, 470 p.
Leng, C. W. and A. J. Mutchler. 1927. Supplement 1919-1924 (inclusive) to catalogue of the
Coleoptera of America, north of Mexico. Mount Vernon, New York, 78 p.
Leng, C. W. and A. J. Mutchler. 1933. Second and third supplements 1925-1932 (inclusive)
to catalogue of the Coleoptera of America, north of Mexico. Mount Vernon, New York,
1 1 2 p.
Leonard, M. D. 1926. A list of the insects of New York with a list of the spiders and certain
other allied groups. Cornell University agr. Expt. Sta. Mem. 101 : 1121 p.
Lindroth, C. H. 1955. Dejean’s types of North American Carabidae (Col.). Opusc. ent. 20 :
10-34.
Lindroth, C. H. 1963. The problem of late land connections in the North Atlantic area. In
Love, A. and D. Love. North Atlantic biota and their history. The MacMillan Company,
New York, pp. 73-85, 6 figs.
Lindroth, C. H. 1966. The ground-beetles of Canada and Alaska. Part 4. Opusc. ent. 29 :
409-648.
Loding, P. H. 1945. Monograph II. Catalogue of the beetles of Alabama. Geological Survey
of Alabama. University of Alabama. Wetumpka Printing Co., Wetumpka, Alabama, 172 p.
Martin, P. S. and B. E. Harrell. 1957. The Pleistocene history of temperate biotas in Mexico
and eastern United States. Ecology 38 : 468-480.
Maslin, T. P. 1952. Morphological criteria of phyletic relationships. Syst. Zool., 1 : 49-70.
Motschulsky, V. 1865. Enumeration des nouvelles especes de coleopteres rapportes de ses
voyages. Bull. Soc. Imp. Nat. Moscou, 38(4) : 227-313.
Muller, E. H. 1965. Quaternary Geology of New York, 99-1 12 p. In H. E. Wright, Jr., and D.
G. Frey, (ed.). The Quaternary of the United States. Princeton University Press, Princeton.
Newman, E. 1838. Entomological notes. Ent. Mag. 5 : 372-402.
Olson, A. L., T. H. Hubbell, and H. F. Howden. 1954. The burrowing beetles of the genus
Mycotrupes (Coleoptera :Scarabaeidae : Geotrupinae). Univ. Mich. Mus. Zool. Misc. Publ.
84, 59 p.
Richards, H. G. and S. Judson. 1965. The Atlantic Coastal Plain and the Appalachian
Highlands in the Quaternary. In H. W. Wright, Jr., and D. G. Frey, (ed.). The Quaternary
of the United States. Princeton University Press, Princeton.
Rohwer, S. A. and G. E. Woolfenden. 1969. Breeding birds of two Florida woodlands:
comparisons with areas north of Florida. Condor, 171(1) : 38-48.
Ross, H. H. 1965. Pleistocene events and insects. In H. E. Wright, Jr., and D. G. Frey, (ed.).
The Quaternary of the United States. Princeton University Press, Princeton.
Revision of Evarthrus
195
Ross, H. H., G. L. Rotramel, J. E. H. Martin, and J. F. McAlpine, 1967. Postglacial coloniza-
tion of Canada by its subboreal winter stoneflies of the genus Allocapnia. Can. Ent.
99(7) : 703-712.
Say, T. 1823a. Descriptions of insects of the families of Carabiei and Hydrocanthari of
Latreille, inhabiting North America. Trans. Amer. phil. Soc. 2 : 1-109.
Say, T. 1823b. Descriptions of coleopterous insects collected in the late expedition to
the Rocky Mountains, performed by order of Mr. Calhoun, Secretary of War, under the
command of Major Long. J. Acad. nat. Sci. Phil. 3 : 139-216.
Say, T. 1834. Descriptions of new North American insects and observations on some already
described. Trans. Amer. phil. Soc. 4 : 409-470.
Schafer, J. P., and J. H. Hartshorn. 1965. The Quaternary of New England, p. 1 13-128. In
H. E. Wright, Jr., and D. G. Frey, (ed.). The Quaternary of the United States. Princeton
University Press, Princeton.
Schaupp, F. G. 1880. Synoptic tables of Coleoptera. Evarthrus LeConte. Bull. Brooklyn ent.
Soc. 3 : 49.
Schuler, L. 1962. La Spermatheque des Chlaenius de France (Callistomorphi Jeannel).
L’Entomologiste. 18(4) : 77-86.
Schuler, L. 1963a. La spermatheque chez les Harpalidae et les Pterostichitae de France.
Rev. franc. Ent. 30(2) : 81-103.
Schuler, L. 1963b. Les organes genitaux femelles chez les Pterostichidae de France. Les
tribus Anchomenini et Sphodrini (suite). Le Cas des Patrobidae (Col. Carabiques). Bull.
Soc. ent. Fr. 68 : 13-26.
Scudder, S. H. 1900. Adephagous and clavicorn Coleoptera from the Tertiary deposits at
Florissant, Colorado with descriptions of a few other forms and a systematic list of the
non-rhynchophorous Tertiary Coleoptera of North America. Mon. U. S. Geol. Sur. 40,
148 p.
Selander, R. K. 1965. Avian speciation in the Quaternary, p. 527-542. In H. E. Wright, Jr.,
and D. G. Frey, (ed.). The Quaternary of the United States. Princeton University Press,
Princeton.
Shotton, F. W.1965. Movements of insect populations in the British Pleistocene. Geol.
Soc. Amer., Special Paper 84, pp. 17-33.
Simpson, G. G. 1961. Principles of animal taxonomy. Columbia University Press, New York.
247 p.
Stebbins, G. L., Jr., 1951. Variation and evolution in plants. Columbia University Press,
New York. 643 p.
Van Dyke, E. C. 1926. New species of Carabidae in the subfamily Harpalinae chiefly from
western North America. Pan-Pacific ent. 2 : 113-126.
Van Dyke, E. C. 1943. New species and subspecies of North American Carabidae. Pan-
Pacific ent. 19 : 17-30.
Van Emden, F. I. 1942. A key to the genera of larval Carabidae (Col.). Trans. R. ent. Soc.,
Lond. 92 : 1-99.
Whitehead, D. R. 1965. Palynology and Pleistocene phytogeography of unglaciated eastern
North America. In H. E. Wright, Jr., and D. G. Frey, (ed.). The Quaternary of the United
States. Princeton University Press, Princeton.
196
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1mm
Figs. 1-14. Pronotum, dorsal aspect. 1. E. hernandensis (Citrus County, Florida). 2. E. morio
(Alma, Georgia). 3. E. laevipennis (Mobile, Alabama). 4. E. laevipennis (near Spartensburg,
South Carolina). 5. E. approximatus (Rosslyn, Virginia). 6. E. iuvenis (near Roanoak,
Virginia). 7. E. obsoletus (near Tuscaloosa, Alabama). 8. E. unicolor (Umadilla, Georgia).
9. E. fucatus (Leesburg, Alabama). 10. E. spoliatus (Rock Creek, Washington, D. C.).
11-13. E. brevoorti (11. Mobile, Alabama; 12. Calvert, Alabama; 13. Clemson, South
Carolina). 14. E. vinctus (Clayton, Georgia).
Revision of Evarthrus
197
2mm
Figs. 15-36. Pronotum, dorsal aspect. 15. E. alabamensis (Mobile, Alabama). 16. E. ovulum
(Tallahassee, Florida). 17. E. macrovulum (Mobile, Alabama). 17a. E. texensis (Kirbyville,
Texas). 18. E. parafaber (Mobile, Alabama). 19. E. levifaber (Georgia). 20. E. faber (Punta
Gorda, Florida). 21. E. incisus (near Morrilton, Arkansas). 22. E. whitcombi (Hot Springs,
Arkansas). 23. E. blatchleyi (Jacksonville, Florida). 24. E. floridensis (Winter Park, Florida).
25-28. E. sigillatus (25. Easton, Pennsylvania; 26. Black Mountains, North Carolina; 27.
Climax, North Carolina; 28. Auburn, Alabama). 29. E. sinus (Alabama Port, Alabama).
30-32. E. convivus (30. Beamsville, Ohio; 31. Talladega, Alabama; 32. near Toomsuba,
Mississippi). 33. E. seximpressus (Le Flore County, Oklahoma). 34. E. alabamae (Gulfport,
Mississippi). 35. E. engelmanni (Cuero, Texas). 36. E. nonnitens (Bradley County, Arkansas).
198
Freitag
Figs. 37-62. Pronotum dorsal aspect. 37. E. hypherpiformis (near Demopolis, Alabama).
38-44. E. sodalis sodalis (38. near Lake Chautauqua, New York; 39. Cleveland, Ohio; 40.
Albany, Wisconsin; 41. near Frankfort, Kentucky; 42. Chicago, Illinois; 43. Dubois, Illinois;
44. near Luka, Mississippi). 45-47. E. sodalis colossus (45. near Yates Centre, Kansas; 46. St.
Joseph, Missouri; 47. St. Louis, Missouri). 48. E. s. lodingi (Monte Sano State Park, Ala-
bama). 49. E. parasodalis (Washington County, Arkansas). 50-51. E. furtivus (50. N. Cum-
berland, Pennsylvania; 51. Rosslyn, Virginia). 52. E. altemans (Ames, Iowa). 53. E. iowensis
(Iowa City, Iowa). 54. E. substriatus (Cloudcroft, New Mexico). 55. E. constrictus (Hamil-
ton County, Kansas). 56. E. torvus torvus (near Grand Island, Nebraska). 57. E. torvus
deceptus (Cuero, Texas). 58. E. gravidus (Austin, Texas). 59. E. sallei (Victoria, Texas).
60. E. gigas (Victoria, Texas). 61. E. heros (Dallas, Texas). 62. E. gravesi (Pearl, Mississippi).
Revision of Evarthrus
199
Fig. 63. Ventral aspect of head of E. unicolor. Figs. 64-65. Elytra, basal portion. 64. E.
sodalis colossus (St. Joseph, Missouri). 65. E. hews (Dallas, Texas). Figs. 66-69. Palpus of
labium. 66. E. hernandensis. 67. E. unicolor. 68. E. faber. 69. E. blatchleyi ■ Figs. 70-71.
Head, dorsal aspect. 70. E. alabamensis . 71. E. ovulum ■ Fig. 72. Ventral aspect of mandibles
of E. sigillatus. Fig. 73. Genitalia of female E. sigillatus. Figs. 74-76. Middle femur, anterior
face. 74. E. hernandensis. 75. E. seximpressus. 76. E. heros. Fig. 77. Plica of elytron and
dorsal hump of abdomen of E . substriatus (Kerrville, Texas). Fig. 78. Dorsal aspect of poste-
rior portion of elytra of E. substriatus (near Ft. Davis, Texas). Fig. 79. Plica of elytron and
dorsal hump of abdomen of E. constrictus (Denver, Colorado). Fig. 80. Dorsal aspect of
posterior portion of elytra of E. constrictus (Clark County, Kansas).
200
Freitag
Figs. 81-86. Male genitalia, median lobe and parameres: a, right lateral aspect; b, ventral
aspect; c, left lateral aspect; d, median lobe with internal sac everted. 81. E. hernandensis
(Juniper Springs, Florida), d, right apical aspect. 82. E. morio (Wellborn, Florida), d, left
lateral aspect. 83. E. laevipennis (Mobile, Alabama), d, left lateral aspect. 84. E. approxi-
matus (Fairfax County, Virginia), d, left lateral aspect. 85. E. iuvenis (near Roanoke,
Virginia), d, left lateral aspect. 86. E. obsoletus (Talladega, Alabama), d, left lateral aspect.
Revision of Evarthrus
201
Figs. 87-92. Male genitalia, median lobe and parameres: a, right lateral aspect; b, ventral
aspect; c, left lateral aspect; d, median lobe with internal sac everted. 87. E. unicolor (Lees-
burg, Alabama), d, ventral aspect. 88. E. fucatus (Leesburg, Alabama), d, right lateral aspect.
89. E. spoliatus (Rock Creek, Washington, D. C.), d, ventral aspect. 90. E. brevoorti (Mobile,
Alabama), d, left lateral aspect. 91. E. brevoorti (Lucedale, Mississippi). 92. E. vinctus
(Rabun County, Georgia), d, left lateral aspect.
202
Freitag
Figs. 93-98. Male genitalia, median lobe and parameres: a, right lateral aspect; b, ventral
aspect; c, left lateral aspect; d, median lobe with internal sac everted, similarily e-h. 93. E.
alabamensis (Mobile, Alabama), d, left lateral aspect. 94. E. ovulum (Toombs County,
Georgia), d, right lateral aspect. 95a-d. E. macrovulum (Mobile, Alabama), d, right dorso-
lateral aspect. 95e-h. E. texensis (Tyler Co., Texas). 96. E. parafaber (Mobile, Alabama),
d, left lateral aspect. 97. E. levifaber (Georgia), d, right lateral aspect. 98. E. faber (Punta
Gorda, Florida), d, right lateral aspect.
Figs. 99-105. Male genitalia, median lobe and parameres: a, right lateral aspect; b, ventral
aspect; c, left lateral aspect; d, median lobe with internal sac everted. 99. E. incisus (Wash-
ington County, Arkansas), d, ventral aspect. 100. E. whitcombi (Hot Springs, Arkansas), d,
right lateral aspect. 101. E. blatchleyi (Gainesville, Florida), d, right lateral aspect. 102. E.
floridensis (Winter Park, Florida), d, right lateral aspect. 103. E. sigillatus (Easton, Pennsyl-
vania), d, right lateral aspect. 104. E. sinus (Alabama Port, Alabama), d, right lateral aspect.
105. E. convivus (near Toomsuba, Mississippi), d, right lateral aspect; e, median lobe with
internal sac everted, ventral aspect.
204
Freitag
Figs. 106-1 12. Male genitalia, median lobe and parameres: a, right lateral aspect; b, ventral
aspect; c, left lateral aspect; d, median lobe with internal sac everted, right lateral aspect.
106. E. seximpressus (Washington County, Arkansas). 107. E. alabamae (Mobile, Alabama).
108. E. engelmanni (College Station, Texas). 109. E. nonnitens (Bradley County, Arkansas).
110. E. hypherpiformis (Manego County, Alabama). 111. E. sodalis (Mayville, N. Y.).
112 . E. parasodalis (Washington County, Arkansas).
Revision of Evarthrus
205
Figs. 1 13-120. Male genitalia, median lobe and parameres: a, right lateral aspect; b, ventral
aspect; c, left lateral aspect; d, median lobe with internal sac everted, right lateral aspect.
1 13. E. furtivus (Mt. Vernon, Virginia). 1 14. E. alternans (Adams County, Illinois). 115.
iowensis (Iowa City, Iowa). 116. E. substriatus (Minaca, Chihuahua, Mexico). 117. E. sub-
striatus (Durango, Mexico). 118. E. constrictus (Austin, Texas). 1 19. E. torvus torvus (near
Castle Rock, Colorado). 120. E. torvus deceptus (Texas).
206
Freitag
d
Figs. 121-124. Male genitalia, median lobe and parameres: a, right lateral aspect; b, ventral
aspect; c, left lateral aspect; d, median lobe with internal sac everted, right lateral aspect.
121. E. gravidus (Forestburg, Texas). 122. E. sallei (Victoria, Texas). 123. E. gigas (Kings-
ville, Texas). 124. E. heros (Gainesville, Texas).
Revision of Evarthrus
207
Figs. 125-130. Maps of geographical distribution. 125. E. hemandensis (•),£. morio (A),
E. laevipennis (■). 126. E. approximatus (■), E. iuvenis (A), E. obsoletus (•). 127. E.
unicolor (■), E. fucatus (★), E. spoliatus (A), E. brevoorti (•). 128. E. vinctus (■), E. alaba-
mensis (A), E. ovulum (•), E. macrovulum (★), E. texensis <▼), 129. E. parafaber (■), E.
levifaber (A), E. faber (•). 130. E. incisus (•), E. whitcombi (A).
208
Freitag
Figs. 131-132. Maps of geographical distribution. 131. E. blatchleyi (★), E. floridensis (•),
E. sigillatus (•), E. sinus (■), E. convivus (±). 132. E. seximpressus (•),£. alabamae (A), E.
engelmanni (★), E. nonnitens (■), E. hypherpiformis (•).
Revision of Evarthrus
209
Figs. 133-134. Maps of geographical distribution. 133. E. sodalis sodalis (•), E. sodalis
colossus (A), E. sodalis lodingi (■), E. parasodalis (★), E. furtivus (*). 1 34. E. altemans (•),
E. iowensis (A), E. substriatus (■), E. constrictus (★).
210
Freitag
Figs. 135-136. Maps of geographical distribution. 135. E. torvus torvus (•), E. torvus decep-
tus (A), E. gravidus (■). 136. E. sallei (■), E. gigas (A), E. heros (•), E. gravesi (•). Fig. 137.
Hypothetical phylogeny of the species of the genus Evarthrus. Species groups are numbered
as follows: 1 - the mono group, 2 - the obsoletus group, 3 - the spoliatus group, 4 - the
ovulum group, 5 - the faber group, 6 - the incisus group, 7 - the blatchleyi group, 8 - the
sigillatus group, 9 - the seximpressus group, 10 - the hypherpiformis group, 1 1 - the sodalis
group, 1 2 - the substriatus group, 1 3 - the torvus group, 1 4 - the gigas group, 1 5 - the gravesi
group.
Revision of Evarthrus
21 1
Fig. 138. Geographical distribution of the subgenera Fortax (• • •), Cyclotrachelus (- - -),
and Evarthrus ( ), and centers of speciation (1-8).
Freitag
ERRATA - Quaest. ent. 1969: 5(1):
“New distributional records for Canadian soldier flies (Diptera:Stratiomyidae. Part I.
Beridinae and Sarginae”. By Max W. McFadden.
page 5. The species Allognosta brevicomis Johnson is known from Quebec, not Quebic.
page 7. The title of the paper by James (1951) is “The Stratiomyidae of Alaska”, not
Alberta.
“A revision of the genera Philophuga Motschoulsky and Tecnophilus Chaudoir with notes
on the North American Callidina (Coleoptera:Carabidae)”. By David J. Larson.
page 29. Page references in key:
Tecnophilus Chaudoir, p. 44, not page 38.
Philophuga Motschoulsky, page 29, not page 24.
Infemophilus new genus, page 43, not page 37.
page 67 - Table 17. The unnamed branch arising from the Philophuga stem was accidentally
inserted by the Temporary Editor, and should be deleted.
For all of these errors the Temporary Editor (G. E. Ball) apologizes to the authors and to
the readers of Quaestiones entomologicae .
E.q -Q f}(/
Quaestiones
en
tomologi
icae
MUS. COMP. ZOOll
library
FEB fc \
S >v •
• |M^
A periodical record of entomological investigations,
published at the Department of Entomology, Uni-
versity of Alberta, Edmonton, Canada.
VOLUME V
NUMBER 3
JULY 1969
QUAESTIONES ENTOMOLOGICAE
A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.
Volume 5 Number 3 July 1969
CONTENTS
Book Review 213
Graham — A comparison of sampling methods for adult mosquito populations in central
Alberta, Canada 217
Chiang — Some pharmacological properties of the nerve cord of the cockroach;
Periplaneta americana (L.) 263
Errata 307
BOOK REVIEW
SMITH, DAVID, S. 1968. Insect Cells. Their Structure and Function. Oliver and Boyd Ltd.,
Edinburgh, Scotland, xvii + 372 pp., cxviii plates. Cloth bound. $16.00 U. S. A.
With the publication of this volume Smith has done for Entomology what D. W. Fawcett
in his An Atlas of Fine Structure, The Cell (1966) did for Vertebrate Zoology. He has pro-
duced a concise and beautifully illustrated summary of our current knowledge of the fine
structure and function of insect cells.
Each tissue and tissue product of the insect body is illustrated by one or more carefully
chosen transmission or scanning electron micrographs, many of which originated in Smith’s
laboratory. The cells and cell products treated are: integument, muscle, neuromuscular
junctions, nervous system, corpus cardiacum, corpus allatum, compound eye, tracheal sys-
tem, dorsal vessel, haemocytes, pericardial cells, oenocytes, fat body, mycetocytes, salivary
glands, silk glands, fore-gut, mid-gut, hind-gut, peritrophic membrane, rectal papillae, Mal-
pighian tubules, anal papillae, testis, vas deferens, accessory glands, ovary and spermatheca.
Although 24 species of insect are represented in the book, the majority of the plates contain
micrographs prepared from only five: Oncopeltus fasciatus, Ephestia kiihniella, Carausius
morosus, Calliphora erythrocephala, and Periplaneta americana. In all the plates abbrevia-
tions are minimal.
The text of the book includes up-to-date background summaries of the basic morphology
and physiology of each of the tissues illustrated. In each summary the reader is referred, for
further information, to the relevant chapter in the sixth edition of V. B. Wigglesworth’s The
Principles of Insect Physiology (1965). Reference to each of the micrographs is made in the
text and the main features contained in these illustrations are mentioned in the captions.
Each summary is supplied with a list of references covering the principal contributions made
in that area. Throughout the text reference is made to ultrastructural studies of comparable
214
tissues in vertebrate animals, a consistency which will undoubtedly enhance the value of the
book for general biologists as well as broadening the perspective of entomologists. A selected
list of references to papers published after the submission of the manuscript (June 1967) is
added as an appendix. Throughout the text Smith is careful also to mention avenues worthy
of further investigation.
No experience is required, on the part of the reader, in the interpretation of electron
micrographs. The necessary background is provided in the introduction in the form of a
foldout illustration. On this is centered a diagram of the “generalized cell” and its organelles.
Each organelle is also illustrated by a small electron micrograph prepared from insect materi-
al. Each of these micrographs is numbered and, in a key on either side of the diagram, its
role in cell physiology is summarized.
Although the manufacture of the book is good, the book is very heavy (2.75 lbs.) for its
linear dimensions and it is possible that the strength of the binding will prove insufficient to
support the weight of the pages over a long period of time.
The editing of the book is excellent. I list here the few typographical errors noticed. On
page 220, mention is made in the last line of the caption to a plate which is not included.
On page 232 it is the outer not the inner surface of the crop that bears a lattice work of vis-
ceral muscle fibres. On page 256 the magnification of the plate and the insert are reversed.
Exception is taken also to the statement on page 183 that insects possess no sex hormones.
Beginning in 1963 Jacqueline Naisse published a series of papers proving convincingly that
androgens are produced by the developing testes of the lampyrid beetle Lampyris noctiluca
L.
The addition of a conventional photomicrograph of the tissue under discussion in each
section would have eased the reader’s difficulty in positioning the electron micrographs that
follow in their relative positions in the tissue.
These are minor criticisms of a text that will probably become a classic. This book should
be present in the personal collection of every insect morphologist and physiologist. The rea-
son is that most of the papers cited on insect ultrastructure have been published in journals
few of which are generally perused by entomologists — an enlightening observation on the
origin of most contributions in this field.
B. S. Heming
Department of Entomology
University of Alberta.
■J
A COMPARISON OF SAMPLING METHODS FOR ADULT MOSQUITO
POPULATIONS IN CENTRAL ALBERTA, CANADA
PETER GRAHAM Quaestiones entomologicae
Department of Biology 5 : 21 7-261 1969
Lindenwood College
St. Charles, Missouri 63301
u. s : a.
Nine sampling methods for adult mosquitoes were compared: Malaise traps, Malaise traps
baited with carbon dioxide, light traps, visual attraction trap, rotary sweep net, animal bait,
human bait and captures of resting mosquitoes in a trailer. The position of the trap as well
as its type was found to affect both the size and composition of the catch. Rotary sweep
nets were found to have a definite attraction for mosquitoes and this may be selective for
some species . Light traps caught a relatively larger proportion of parous mosquitoes than
other methods, but other physiological stages showed no differences between methods.
In recent years a vast literature on sampling methods for mosquito populations has
accumulated but this deals mainly with size and species composition of the catches.
In any study of trapping methods for insects it must be realized that the catch depends
on three sets of factors: those in the trap, those in the environment and those in the insect.
The catch depends on the population density of the insect, its “availability” and its
activity. Corbet (1961) considered that light traps in Uganda sampled only those mosquitoes
engaged in non-specific activities and did not catch those engaged in feeding, swarming,
or oviposition. Biddlingmayer (1967) has published a study of the effects of environment
and species composition on different trap types but has not considered the effects of the
physiological state of the mosquito.
Apart from Corbet (1961) I know of only one study relating the physiological state of an
insect of medical importance to survey methods, the work of Burse 11 (1961). Barr (1958)
mentions that the age and physiological state of mosquitoes affects the captures in light
traps, but the citation he gives for this, Nielsen and Nielsen (1953), is incorrect, as this paper
makes no mention of factors affecting light trap captures. Russian workers have paid con-
siderable attention to the physiological age of mosquitoes, have elucidated many factors in
the biology of the insects and have provided methods for determining age (Detinova, 1962)
but have not related age to sampling procedure. I have attempted to fill part of this need in
relation to woodland mosquitoes in central Alberta.
THE STUDY AREA
The study area is on the west shore of George Lake, 53°57’N and 1 14°06’W, about 40
miles northwest of Edmonton, Alberta. The area lies at the southern margin of the boreal
mixed forest subzone (LaRoi, 1968). All traps were within 300 yards of the campsite, more
or less in the centre of a square mile field site operated by the Department of Entomology,
University of Alberta, (Fig. 1 ).
FIGURE I
SKETCH MAP OF STUDY AREA AND CAMPSITE
GEORGE LAKE FIELD SITE
218
Graham
Sampling Methods
219
Away from the lake shore the vegetation of the field site consists of almost untouched
mature poplar forest, with small areas of spruce on the northern and western boundaries.
Prior to 1930 some trees were removed by neighbouring farmers, but otherwise the forest
has not been disturbed. The principal trees are Populus tremuloides Michx. and P. balsam-
ifera L.. Other trees are Picea glauca (Moench.), Betula papyrifera Marsh., Alnus tenuifolia
Nutt, and Salix species. Larix laricina Koch, is common in neighbouring wetlands but rare
on the field site. The understory is more diverse, consisting of a large number of shrub and
herb species: Amelanchier alnifolia Nutt., Viburnum edule (Michx.), Rosa acicularis Lindl.,
Cornus stolonifera Michx. and Ribes lacustre (Pers.) are common shrubs. Cornus canadensis
L., Solidago species, Epilobium angustifolium L., and Aster species are common herbs.
Ledum groenlandicum Oeder. forms more or less oval bogs in a few places, usually on
clumps of sphagnum moss. On the northern boundary there is an area of sedge ( Carex
species) meadow which contains a number of permanent water holes. A stream flows out of
the lake just south of the campsite and is blocked by several beaver dams. There is a fringe
of Carex bordering the lake and a floating mat of Typha species round the lake edge.
About half the surrounding country is cleared for cultivation and grazing, mainly on the
east, south and northwest, resulting in a patchwork of woodland, pasture and cultivation
which allows a rich mosquito fauna (Graham in prep.).
In the winter of 1964-1965 above normal snow falls were recorded and melt water
remained well into summer. Also nearly six inches of rain fell in the last two weeks of June
1965. Thus the majority of spring larvae were able to complete their development and
second broods of many species developed. In the winter of 1965-1966, below normal snow
falls occurred and most melt water had dried up by late spring, so many larvae did not
complete development. Heavy rains did not fall till late July and August and the resulting
pools soon dried up, so second broods were not prominent. Heavy snow fell in the winter
of 1966-1967, but did not melt till the end of April. In 1966, snow had almost disappeared
by 21 April, but in 1967 it was still deep on this date. Break up of ice on the lake had
occurred on 21 April 1966 but did not take place till the end of the first week of May in
1967. According to Mr. E. Donald, a neighbouring farmer, the 1967 spring was ten days to
two weeks behind the long term average at George Lake. Table 1 presents names of the
major species of mosquitoes taken and their abundance in 1 966.
An example of the difference in mosquito populations in the 1965 and 1966 seasons is
given by the captures in a light trap operated at the Victoria Golf Course in the City of
Edmonton. In 1965 this trap was run from 9 July to 30 August and caught 2826 mosqui-
toes, an average of 75 per night. In 1966 the same trap was run from early May to the end
of August and caught six mosquitoes. Control measures in the urban area were the same in
both years. In the spring of 1967 traps at George Lake caught approximately four times as
many mosquitoes as in the spring of 1 966, though I had the impression that the mosquito
nuisance in the field site was worse in 1 966.
METHODS
Sampling
Nine methods of sampling adult mosquitoes — Malaise traps, New Jersey light traps, a
220
Graham
Table 1 Numbers of mosquito species identified at George Lake in the spring and in
summer of 1966.
visual attraction trap, a rotary sweep net, chicken bait, rat bait, human bait, carbon dioxide
bait, and collections of resting mosquitoes inside a trailer — were tested in this study. The
localities of these traps are shown in Fig. 1 . Traps used in different years were operated in
the same places.
All meteorological data were obtained from a recording thermohygrograph in a Stevenson
screen at the campsite.
Malaise traps. - This type of trap was first described by Malaise (1937), but its impor-
tance in ecological studies has only recently become apparent. I chose the modification of
Townes (1962) as it is operational from all four directions. Breeland and Pickard (1965) and
Smith et al. (1965) have recently demonstrated the value of this type in mosquito studies.
One trap was used in 1965 and two in 1966 and 1967. These had four entrances four feet
high and six feet wide and the catching head was eight feet above the ground. The traps
were erected over old tracks, MI near the lake shore and Mil on the top of a low rise about
300 yards into the forest. Calcium cyanide, in the form of Cyanogas G, was used as a killing
agent. Three teaspoonfuls in a manila envelope remained lethal for five days. The traps are
shown in Plates 1 , 2 and 3.
Light traps. — Two battery-operated standard New Jersey light traps (Lt) were used. This
type was chosen as it is perhaps the trap most frequently used by mosquito workers. I decid-
ed not to use ultra violet light since the standard model is more often used and some studies
(Zhogolev 1959; Downey 1962) have indicated that while U. V. greatly increases the catch
Sampling Methods
221
Plate 2
Malaise trap II in April 1967.
222
Graham
of some biting flies it might be less attractive to some mosquito species. It was not possible
to compare U. V. with the standard model.
One of the two, Lt I, was situated at the forest edge on the lake shore and the other,
Lt II, a short distance into the forest. A six volt car battery operated one of these traps for
three nights. Air flowed at 121.5 cubic feet per minute through Lt I and at 120 cubic feet
per minute through Lt II. These figures were determined with a Biram’s anemometer. The
light traps are shown in Plates 4, 5 and 6.
Visual attraction trap. — A visual attraction trap of the type described by Haufe and
Burgess (1960) was used, with a net, instead of the hourly timing device described, for
collecting the catch (Plate 7). Unfortunately, only one trap was available and the power
supply permitted only restricted hours of operation.
The cylinder made one complete revolution every two seconds and the air flowed through
the trap at 741 cubic feet per minute. Complete engineering blue prints of this trap are
obtainable from the Canada Department of Agriculture, Medical and Veterinary Entomol-
ogy Branch, Lethbridge, Alberta.
This type of trap was originally developed for use in the north, where short summer
nights make light traps inefficient.
Rotary sweep net. — A group of four electrically driven rotary sweep nets was used. Two
nets were at 45.5 inches and two at 58 inches above the ground. At each level, the base of
one was 23 inches and the other was 34 inches from the shaft, so that no two nets swept the
same volume of air. The nets were 12.5 inches in diameter and the trap made one complete
revolution per second, thus the trap swept 1 1 67 cubic feet of air per minute (Plate 8). The
insects caught were removed with an aspirator. Operation of this trap was restricted by the
power supply.
Rotary traps have been used by several workers (Chamberlin and Lawson, 1945; Stage
and Chamberlin, 1945; Stage et al., 1952; Love and Smith, 1957) who assumed the traps
have no attraction for insects and take unbiased random samples.
Chicken baited traps. — Two small traps were constructed in 1965; each held one chicken.
These were not very successful.
Two large traps each capable of holding several birds were used in 1966. They were six
feet long, by four feet wide, 12 inches high at the corners and 22 inches high at the centre.
One end was closed in with a roosting box 17 inches by four feet in dimension. The floor
was one inch mesh wire cloth. Four egress traps protruded from the roosting box, two on
each side. The rest of the sides were made of 14 x 18 inch mesh galvanized wire gauze.
Ingress traps were tried but the birds sat on them and broke them. The mosquitoes entered
by the wire cloth floor and were caught in the egress traps as they left. The traps did not
catch many mosquitoes, possibly because the birds ate them.
The two traps were designated CBI and CBII (Plates 9 and 10). CBI was baited with white
leghorns and CBII with bantams. The latter often escaped so that the number in the trap
varied from two to five. These traps were only operated in 1966.
Rat baited traps. - The two traps used with chickens in 1965 were modified and used
with rats in 1966. Each was of a different design. They were designated RBI and RBII.
RBI was a modified Magoon (1935) trap 19 inches long and 17 inches wide and 12 inches
high. The animal chamber was closed in by 1 4 x 18 inch mesh galvanized wire gauze so that
Sampling Methods
223
Plate 3 Catching head of Malaise trap I.
Plate 4 Light trap I as seen from the Lake shore.
224
Graham
Plate 5 Light trap I showing proximity to the Lake.
Plate 6
Light trap II showing position in the forest.
Sampling Methods
225
Plate 7
The visual attraction trap.
Plate 8 The Rotary trap.
Plate 9 Chicken baited trap I.
Plate 10 The catching cages of chicken baited trap II.
226
Graham
Sampling Methods
227
the mosquitoes could not reach the bait. The mosquitoes entered a collecting cage 19
inches long, six inches high and six inches wide by means of a no-return baffle and were
removed with an aspirator. RBII was a circular trap, 15 inches in diameter with the bait
cage coming to a cone 1 2 inches high. This cone projected through a hole three inches in
diameter in the top plate into a cone three inches high in the bottom collecting cage, the
two forming a no-return baffle. The catching cage was a removable cylinder nine inches high
and 10 inches in diameter. Flanges six inches wide in RBI and 4V2 inches wide in RBII were
added in 1966 to direct the mosquitoes into the trap. These traps are shown in Plates 1 1 and
12.
Originally three rats were used in each cage, but births often modified this. These two
traps were run close to each other in the middle of the campsite. RBI was operated in June,
July and August 1966 and May and June 1967; RBII was only operated in July and August
of 1966.
Many designs have been suggested for animal bait traps using large animals, the so-called
stable traps of Magoon (1935), Bates (1944), Roberts (1965), but relatively little attention
has been given to the use of small animals as mosquito bait (Southwood, 1966). These
would appear to offer certain advantages because of their smaller size and the fact that they
need less attention than cattle or horses.
Human bait. — On one day in most weeks from late May to the end of August, 1966, 1
sat quietly with trouser legs rolled up and caught any mosquitoes which alighted in a fifteen
minute period. In May these collections were made in the afternoon and after that at 1800
hours.
Carbon dioxide baited traps. — Several authors (Brown, 1951 ; Bellamy and Reeves, 1952;
Newhouse et al. , 1966 and others) have shown that carbon dioxide used alone or in con-
junction with another attractant is good bait for mosquitoes. I decided to try the release of
carbon dioxide from a cylinder in Malaise traps. The gas was released at from one to six
litres per minute with an average rate of five litres per minute (approximately equivalent to
the amount of carbon dioxide expired by 20 men at rest). It proved difficult to control the
flow accurately in the field with changing conditions of temperature and barometric pres-
sure. The gas was released through a flowmeter and led up into the catching head by means
of a plastic tube (Plate 13). On the night of 12/13 July, 1966 releasing the gas direct from
the cylinder without a flowmeter was tried. Both 25 lb. and 50 lb. cylinders of carbon diox-
ide were tried; the former proved better as they were more portable. One 25 lb. cylinder
lasted approximately 16 hours. The traps were run from 1700 hours to 0900 hours the fol-
lowing morning.
Carbon dioxide was used on alternate nights, the other Malaise trap being used as a con-
trol. Carbon dioxide traps were operated in July and August 1966 and May and June 1967.
Collections of resting mosquitoes and miscellaneous collections. — Also on one day per
week in June, July and August, 1966 and from 16 May to 15 June 1967, all mosquitoes
found resting in one of the trailers at the camp were collected early in the afternoon. In
June 1966 this trailer was used as a kitchen and dining room, thereafter as. a store. In 1966
a carton of dry ice was kept in it and rats were kept there over weekends in a screened cage.
In 1967 rats were kept in the trailer in a unscreened cage.
228
Graham
Plate 12
Rat baited trap II.
Sampling Methods
229
Plate 13 Malaise trap I with CO2 cylinder in place.
230
Graham
At various times during the summer of 1966, mosquitoes were caught with a sweep net,
when biting at times other than when human bait captures were in progress and in a C. D. C.
(Communicable Diseases Center) miniature light trap. These collections have been included
only in total catch figures.
Handling and Dissection
Weekend catches of Malaise traps, human bait, and miscellaneous collections were identi-
fied and counted. All other collections were frozen and taken to the laboratory. There they
were identified, counted and dissected or, if numerous, subsampled and dissected. In 1967
the very large collections in carbon dioxide traps were subsampled for dissection and sub-
sampled again for identification. The number of specimens in each sample was estimated
from these two subsamples. This estimated number was used to obtain the proportion of
each species in the total carbon dioxide trap catch. The number identified varied from 1 00%
to 20% of the total, being proportionately smaller in the larger samples. This is probably an
accurate estimate of the numbers of the more numerous species but not of the rarer ones.
Specimens for dissection were first assigned to a stage of Sella (Detinova, 1962) and to
one of five arbitrary categories of external wear. They were dissected in distilled water
under XI 2 of a Wild M5 stereomicroscope. The contents of the ventral oesophageal diver-
ticulum and the mid gut were noted. The ovaries were then examined under X50 for the
stage of Christophers (Clement, 1963).
The ovaries were removed to a drop of water on a microscope slide, allowed to dry and
stored till they could be examined for parity or nulliparity by Detinova’s method of ovarian
tracheation (Detinova, 1962). The ovaries were then examined in a drop of distilled water
under XI 00 of a Propper compound microscope.
All dissections were done within one week of capture and the specimens were kept frozen
in dry ice until dissected. Corbet (1961) showed that mosquitoes were suitable for dissec-
tion after being kept frozen for three months. I found that it was possible to use Detinova’s
method on ovaries which had been stored dry on a slide for a year.
Males were counted and identified to genus only.
Except for Malaise trap captures all mosquitoes were killed by freezing with dry ice.
RESULTS AND DISCUSSION
Malaise traps, light traps, visual attraction trap, rotary trap, rat bait, chicken bait and
Malaise and carbon dioxide were compared both quantitatively and qualitatively. Two
methods — human bait and collections in the trailer — were not standardized enough for
quantitative comparison.
Note on statistics used
The statistical analysis of the data obtained in this study presented certain difficulties,
since the nature of the study did not allow the randomization of catches. All traps had to
be operated in the same place and technical difficulties as well as the nature of some of the
traps prevented simultaneous operation. Therefore the statistical tests applied are not all
strictly applicable to the data obtained, though I believe they assist in the interpretation of
the results.
Sampling Methods
231
Quantitative comparison of the trap types was obtained by converting the catch into
catch per 100 trap hours to standardize and to allow for the fact that the different traps
were run for different lengths of time. This test is not strictly applicable but does help to
confirm conclusions reached by other methods. An index of trap “effectiveness” was ob-
tained by dividing the catch per 100 trap hours in the trap or trap type under consideration
by the catch per 100 trap hours in the combined Malaise traps over the same period. The
Malaise traps were chosen as standards as they appear to be passive and to have no attraction
for mosquitoes. The combined Malaise traps were used in an attempt to minimize the
effects of trap position. This “index of effectiveness” permits a ranking of traps and trap
types in order of effectiveness.
A modified geometric mean, the Williams mean (Haddow, 1960), was used for studying
the effects of the addition of carbon dioxide to Malaise traps. It is obtained by the expres-
sion Mw = antilog ^x+ * -1 where x is the value of each sample and N is the
number of samples. The addition of one to each sample value allows the inclusion of zero
catches which cannot be included in a normal geometric mean. Williams has shown that
where there is a large variation in the size of samples or one sample is very different from
the others, this mean gives a better measure of central tendency than the arithmetic mean.
The test was used in the qualitative comparison of the trap types. Simpson et al.
(1960) was the principal reference used for statistical methods.
Relative effectiveness — catch per unit time
Table 2 shows the relative effectiveness of the traps in 1966 and 1967 and Table 3, the
effectiveness in June of 1966 and 1967. Table 4 shows the catch per 1000 trap hours of the
five most abundant species and Culex territans which is believed to feed on cold blooded
vertebrates and so differs from the other species caught which are believed to feed mainly
on warm blooded vertebrates.
The results in May and June 1967 are similar to those in 1966 except that light traps
caught less than either the visual attraction or the rotary traps and the catch per 1 00 trap
hours in all traps averaged four times larger than in 1966.
Perhaps the most interesting result was the high catch in the rotary trap which is gener-
ally believed to have no attractive influence and to take a random sample of flying insects
(Stage and Chamberlin, 1945; Love and Smith, 1957; Juillet, 1963; Southwood, 1966). This
trap had an index of effectiveness of 5.1 in 1966 and 9.7 in 1967 and was well within the
range of traps with an attractive influence, namely light, bait, and visual attraction traps.
This indicates that the rotary trap does in fact exert some attractive influence on mosqui-
toes. It was impossible to observe the approach of mosquitoes to a trap of this size so that
the nature of this influence could not be elucidated. It is well known that many biting
Diptera including mosquitoes (Clement, 1963) are attracted to moving objects. Tabanids
and tsetse flies ( Glossina sp.) are attracted to moving motor vehicles (Duke, 1955; Glascow,
1963). The stimulus of a rotary trap may be similar to that of a moving vehicle.
Though the light traps caught nearly three times as many mosquitoes in June 1967 as in
June 1966, their relative effectiveness was nearly halved. This is probably due to the absence
of Culiseta inornata in 1967 as this species formed a large proportion of the 1966 catch.
232
Graham
Table 2 The numbers of adult female mosquitoes taken per 100 trap hours in differ-
ent traps at George Lake 1966 and 1967.
Trap 1966 (June July August)
* August only.
** Adjusted using Malaise trap figures per equivalent nights.
*** The ‘Index of Effectiveness’ is the no. of mosquitoes per 100 trap hours taken in trap,
divided by the no. of mosquitoes per 100 trap hours in the combined Malaise traps.
Sampling Methods
233
Table 3 Comparison of mosquito captures per 100 trap hours in June 1966 and June
1967 at George Lake.
Trap 1966 1967 Index of Comparative
Effectiveness Increase
in catch
1966 1967 1966/67
Fig. in brackets = no. mosquito caught. Av. increase 4.30
Apart from the rotary and light traps the “indices of effectiveness” for the two years are
very similar. This indicates that the relative effectiveness of a trap does not change much
with population size; but it may be changed considerably if the species composition changes.
The number of mosquitoes per unit volume of air filtered in June 1967 was calculated
for the rotary, visual attraction, and light traps. Only the volume of air flowing through the
trap was used; no estimate of “area of influence” was made. This gives an estimate of the
actual efficiency of these traps. The rotary trap captured 2.4 mosquitoes, the visual attrac-
tion trap 0.3 and the light traps 1.3 mosquitoes per 10,000 cubic feet of air. If it is assumed
that rotary traps have no attraction to mosquitoes but capture only those which come
within range, then the efficiency of rotary and Malaise traps should be approximately the
same. I calculated the air flow needed to give a catch of 34.1 mosquitoes per 100 trap hours
(the figure in Malaise traps) if the efficiency in Malaise traps is the same as that of the
rotary trap. This was 23.5 cubic feet per minute, which means that the average wind speed
through these traps would have been 0.9 feet per minute. That is, these traps would have
to have been standing in virtually still air during June 1967; since this was not so, I infer
that the efficiency of the Malaise traps was below that of the rotary trap.
The high actual efficiency of the rotary trap, above both light and visual attraction
traps, is additional evidence that this type of trap does provide an attractive stimulus for
mosquitoes.
Proportion of males
Comparatively few males were taken and relatively little attention was paid to them as
they formed only about 1% of the total catch in 1966 and 1967. Table 5 shows the pro-
portion of males taken in 1966 and 1967.
Table 4 Numbers of adult females of selected species of mosquito caught per 1 000 trap hours and actual numbers caught in different trap
types at George Lake from 1 June to 1 September 1966.
234
Graham
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Sampling Methods
235
Table 5 Proportions of male mosquitoes in traps at George Lake, 1966 and 1967.
Trap 1966 (1st June — 1st Sept.) 1967 (15 May — 30 June)
No. 66 Total catch 6 : 6 + 9 No. 66 Total catch 6 :6 + 9
X2 2 1966 - 33.033 P = 0.005 (Light against rest)
X ^2 1967 — 1.457 P = 0.5 (Light against rest)
* August only
In both years light traps took the largest proportion of males, but the statistical signifi-
cance of this is doubtful. The position of the trap was important; Lt II took a much great-
er proportion in both years than Lt I. Light traps are known to take a larger proportion of
the males of some insects than are in the population (Southwood, 1966) and to take large
numbers of male mosquitoes (Barr, 1958). Belton and Galloway (1965) found 50% of light
trap captures of nearly 6000 mosquitoes were males at Belleville in Ontario. Breeland and
Pickard (1965), however, found 22% in Malaise trap captures were males but only 12% in
light traps.
Species composition
Diversity. — The index of diversity a was introduced by Fisher et al. (1943) as a measure
of the diversity of a population. It is obtained from the expression S = aloge(l+N/a) where
S is the number of species and N the number of individuals. An approximation, adequate
for most needs, can be obtained from nomograms in Williams (1964) and Southwood
(1966). This index is dependent on the size of the sample as well as its diversity but is
useful for comparing traps operated the same period and has been successfully used to
compare methods of catching Heteroptera by Southwood (1960).
The indices of diversity for the trap types in 1966 are shown in Table 6. There were no
significant differences between trap types, which indicates that the smaller catches were due
to lower effectiveness rather than to the unavailability of certain species.
236
Graham
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Sampling Methods
237
Proportions of different species. — The proportions of the major species in the catches of
the different trap types are shown in Table 6. Table 7 shows the proportions in paired traps.
* July and August only
X 2 12 Malaise 170.65 P = <0.001
X26 Mal+C02 15.11 P = <0.01
X26 Light 26.23 P = <0.005
The low catch of Culiseta inornata in the rotary trap and the absence of this species from
the visual attraction trap in 1966 is hard to explain since this species formed 27% of the
catch in the nearby Malaise I.
The animal bait traps showed great similarity. The rat and chicken bait traps did not
differ significantly while the human bait and rat bait traps differed only at the 5% level.
Aedes canadensis formed over 20% of the catches in chicken bait traps but was scarce in
other traps and A. cinereus was most abundant in human bait catches.
Discussion. — The results of Breeland and Pickard (1965) are of interest. They found 52%
of Malaise trap captures were Aedes compared to 54% in light traps and 50% in biting
catches. Forty seven percent of the Aedes in their Malaise traps and 52% in their light
traps were A. vexans, which indicates that the preponderance of this species in light traps
238
Graham
is often more due to its preponderance in the population rather than to any specific attrac-
tion to light, though this species is often stated to be greatly attracted to light (Huffacker
and Bach, 1943; Love and Smith, 1957). Although Love and Smith found a high “index of
attractivity” to light for this species, the proportion of A. vexans was actually higher in
their sweep nets than in their light traps (53% and 50% respectively). Breeland and Pickard
found light traps gave a significantly lower diversity, 3±0.3, than the Malaise traps, 5±0.5
(my calculations). At George Lake only Culiseta inornata and Anopheles earlei were above
the numbers expected in light traps if there was no difference between trap type.
Haufe and Burgess (1960) compared a visual attraction trap to a suction trap, which
like a Malaise trap presumably takes a random sample of the flying insect population. They
found that though a visual attraction trap caught ten times as many mosquitoes as a suc-
tion trap, there was no significant difference between the proportion of band-legged and
black-legged Aedes between the two traps. At George Lake the main difference between
Malaise I and the visual attraction trap was the low number of Culiseta inornata and the
high number of Culex territans in the visual attraction trap and there were no significant
differences in the proportions of Aedes species, which indicates that this trap takes a
random sample of the mosquitoes which approach it. Haufe and Burgess (1960) found
this trap caught all mosquitoes approaching to within about 30 inches of it and observations
at George Lake support this.
The rotary trap catch was significantly different from the Malaise trap catch but this
applies mainly to the catch of Culiseta inornata which was lower than expected and that of
Aedes punctor which was higher than expected. In 1967 this species formed 58% of the
catch. The evidence shows that this trap exerts an attractive stimulus to mosquitoes and
this may be selective for some species, possibly A. punctor.
The animal bait traps differ from the others in that their attraction depends on the
feeding habits of the adult female mosquitoes. Captures on chickens, rats and humans
differed very little. Differences between human bait and the others were probably due to
position.
Effects of carbon dioxide on the catch in Malaise traps
Rudolfs (1922) suggested that mosquitoes were attracted to carbon dioxide and since
then some controversy has arisen over whether this is merely an activating agent (Willis,
1947; Laarman, 1955) or whether it also has an orienting effect (Reeves, 1953), but
Clement (1963) states the importance of carbon dioxide as an aid in host finding by mos-
quitoes has yet to be determined.
Several workers have found that the addition of dry ice (solid carbon dioxide) to light
traps greatly increases the catch (Reeves and Hammon, 1942; Huffacker, 1942; Huffacker
and Bach, 1943 and Newhouse et al., 1966). Carestia and Savage (1967) found that the
catch in a C. D. C. miniature light trap was greatly increased by the addition of carbon
dioxide from a cylinder and that the catch increased as the rate of flow was increased.
Reeves (1953) used carbon dioxide as bait in a stable trap and caught large numbers of
Culex tarsalis at 26 ml. CO2 per minute (equivalent to one chicken) and the catch increased
as the rate of flow increased. Bellamy and Reeves (1952) designed a portable trap, from a
twenty pound lard can, which used dry ice as bait.
Sampling Methods
239
Hayes et al (1958) and Dow (1959) have compared carbon dioxide bait with other
mosquito attractants and find it compares very favorably as an attractant for adult females
and Brown (1951) and Brown et al. (1951) have found carbon dioxide an effective attrac-
tant in the field.
Table 8 shows the catch of mosquitoes in Malaise + CO2 traps and in Malaise traps over a
period of six nights in 1966 and seven in 1967. The traps were run from 1700 hours to 0900
hours the following morning.
Table 8 Numbers of mosquitoes caught in Malaise and Malaise + CO2 traps on equiv-
alent nights at George Lake in July, August 1966 and May and June 1967.
* Mw = Williams mean = antilog
-1
x = Number per sample
N = Number of samples
240
Graham
Table 9 shows the proportions of species in the Malaise and the Malaise + CC>2 traps.
The complete Malaise trap captures are used, rather than only those on equivalent nights for
these latter were too low for accurate analysis. These figures show that the addition of
carbon dioxide to a Malaise trap greatly increases its catch and the numbers of nearly all
species caught are increased. The increase for some species is greater than for others, Aedes
species appearing to be more attracted to carbon dioxide than non -Aedes species. Three
Aedes species showed significantly higher proportion in Malaise + CO2 traps; these were
Table 9 Proportion of female mosquito species in Malaise and Malaise + CO2 traps at
George Lake.
* Estimated total
** above expected in CO2 trap
t below expected in CO2 trap
Fig. in brackets = no. caught
Sampling Methods
241
A. vexans in 1966 and A. intrudens and A. punctor in 1967. A punctor showed no signifi-
cant difference in 1966 and in fact the proportion was slightly higher in the Malaise traps,
possibly because this species is relatively less abundant in August than in the spring. The
greatly increased proportion of A. vexans in carbon dioxide traps is interesting as Huffacker
and Bach (1943) took a lower proportion of this species in light traps with carbon dioxide
than in light alone, but Carestia and Savage (1967) and Newhouse et al. (1966) took slightly
higher proportions of A. vexans in light traps with carbon dioxide than with light alone.
Both Carestia and Savage and Newhouse et al. found the proportions of Culex species were
greatly increased when carbon dioxide was added to light traps. This did not occur at
George Lake as the only common Culex was C. territans which feeds mainly on amphibians.
Table 10 shows the Williams mean catch per trap night in Malaise, Malaise + CO2 and
light traps. To obtain some idea on how carbon dioxide attracts mosquitoes I watched both
traps on several evenings in May and June in 1967. The traps had to be observed through
binoculars from at least twenty yards distance; otherwise the mosquitoes left the trap for
the observer. About half an hour after the traps were started, a swarm of mosquitoes formed
over the catching head of Malaise I + CO2, which was on low ground. At Malaise II + CO2,
which was on the top of a low ridge, no swarm formed but large numbers of mosquitoes
settled on the baffles of the trap. I saw very few settling on these in Malaise I + CO2. Many
of the settled mosquitoes crawled or flew upwards and were caught.
In both traps many mosquitoes remained settled very close to the carbon dioxide outlet
for periods of up to fifteen minutes.
The formation of a swarm over the carbon dioxide outlet and the very large numbers
caught show that carbon dioxide probably exerts a considerable orienting stimulus to adult
female mosquitoes; but it is easily overridden by the approach of a host animal such as man.
Table 10 Comparison of mosquito captures per night in Malaise, Malaise + CO2 and
Light traps at George Lake, July and August 1966 and May and June 1967.
* Mw = Williams mean
242
Graham
Physiological state
The contents of the ventral diverticulum. — Trembley (1952) and Hocking (1953) have
shown that sugar solutions and nectar normally pass into the ventral oesophageal diverticu-
lum and not into the stomach. Hocking (1953) has shown the importance of nectar as an
energy source for mosquitoes. Thus, the contents of the ventral diverticulum are a partial
measure of the energy resources available to the mosquito. The amounts of liquid in the
ventral diverticulum in female mosquitoes caught in different trap types are shown in Table
11. In most mosquitoes the ventral diverticula were either empty or only partially filled;
in only 30 out of 650 mosquitoes were they full.
There were no significant differences between trap types.
Table 1 1 Comparison of the contents of the ventral diverticulum of mosquitoes
caught in the trap types at George Lake in 1966.
Contents of Ventral Diverticulum
0 = empty
1 = partially full
2 = full
Ovarian development and stage in gonotrophic cycle. - Tables 12 and 13 show the stages
of Sella and Tables 14 and 15 show the stages of Christophers. Table 16 shows the occur-
rence of gravid females in the traps. In 1966 light traps caught a significantly higher propor-
tion of the higher stages than the other traps but this was not so in 1967. A striking differ-
ence between the two years was the large number of resting mosquitoes which had ovaries
in stages III— V of Christophers in 1967, which has been discussed above.
Three gravid females were taken in animal bait traps and seven in carbon dioxide traps,
but it is unlikely they were attracted to the bait.
Corbet (1961) found that in Mansonia fuscopennata (Theobald), in Uganda, light traps
sampled only those specimens engaged in “non-specific activity” i.e., those not engaged in
Sampling Methods
243
Table 12 Comparison of the stages of Sella of mosquitoes caught in the different trap
types at George Lake over the periods 1st June to 1st September 1966 and
1 6th May to 30th June 1 967.
1966
Stage of Sella
swarming, biting, or ovipositing. Standfast (1965) confirmed this for Culex annulirostris
Skuse but he believed this indicated activity in the intermediate stages of the gonotrophic
cycle, that is females in stages III and IV of Christophers or III to VI of Sella. Corbet
(1961), on the other hand, found 90% of M. fuscopennata in light traps were in stages I and
II of Christophers and none were gravid. George Lake results do not support this since a
number of gravid females and individuals in intermediate stages of the gonotrophic cycde
were taken in light traps. Both Corbet and Standfast based their conclusions on the fact
that peak light trap captures did not coincide with peaks of biting, swarming or oviposition
activity. Captures were not recorded at hourly intervals at George Lake, but mosquitoes
were often found biting round light traps in the evenings and in the mornings of nights when
none were caught. Corbet and Standfast worked on tropical mosquitoes, which may ex-
plain some of the differences.
244
Graham
Table 13 Comparison of the stages of Sella of Aedes species in the different trap types
at George Lake 1st June to 1st September 1966 and 16th May to 30th June
1967.
1966
Physiological age of adult females as shown by the proportion of parous females. - In
three years of study 1683 pairs of ovaries were examined for parity. The proportions of pars
in the traps are shown in Tables 17—21. Except in August 1966, light traps caught a higher
proportion of pars than other traps. In August 1966 the greater part of the light trap catch
was Culiseta inornata and Anopheles earlei, most of which were probably about to over-
winter and these species appear to overwinter as nullipars. All the Aedes taken in light traps
in August 1966 were parous. The statistical significance of the higher proportion of pars in
light traps is doubtful. In 1965 and 1966 the proportion of pars in light traps was signifi-
cantly higher at the 5% level when tested against the rest combined but not significant when
the traps were tested individually. In the spring of 1967, however, the proportion of pars in
light traps was significantly higher at the 1 % level when the traps were tested individually.
In all cases the parity rate in light traps was higher for Aedes species than for the total catch.
Sampling Methods
245
Table 14 Comparison of the stage of Christophers of mosquitoes in different trap
types at George Lake from 1st June to 1st September 1966 and 15th May
to 30th June 1967.
Table 21 shows the parity rate in five species taken in different trap types. With the ex-
ception of Anopheles earlei the parity rate in light traps was higher than in other traps.
It is clear that light traps have a slightly higher attraction for older female mosquitoes
than the other trap types tested.
Damage to mosquitoes by different collection methods
I noticed that the condition of specimens caught in different traps varied considerably. I
investigated this using six arbitrary damage categories. The results are shown in Table 22.
The specimens taken in Malaise traps both with and without carbon dioxide are in much
better condition then those taken in other traps, probably because the insects were dead
before falling into the collecting bottle and there were no moving parts in the traps. These
traps also involved a minimum of handling both during and after capture. The rotary trap
246
Graham
Table 15 Comparison of the stage of Christophers of Aedes species in different trap
types at George Lake, 1st June to 1st September 1966 and 16th May to 30th
June 1967.
1966
Stage of Christophers
damaged specimens more than any other and the mean shown is possibly too low as the
catch of this trap included a high proportion of unidentified specimens which were not
assigned to any damage category.
General discussion
Southwood (1966) has reviewed methods of sampling insects, including mosquito popu-
lations. Though a great deal of ingenuity has been expended on the design of methods for
sampling adult mosquito populations and on the refinement of these methods, they are
mainly aimed at the largest possible catch. A few methods have been designed for special
purposes such as window traps (Muirhead-Thompson, 1951) which are designed to catch
mosquitoes entering or leaving buildings, a trap to catch mosquitoes emerging from cesspits
(Saliternik, 1960) and several traps designed to catch resting mosquitoes (Russell and San-
Occurrence of gravid female mosquitoes in the different traps at George Lake in 1965, 1966 and 1967.
Sampling Methods
247
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248
Graham
Table 17 Proportion of parous mosquitoes in trap types at George Lake 1st June to
1st September 1966.
* Operated 27th July to 1st September only
** significant at 5% level for those traps run for the whole period.
X2 2 = 5.447 (Light vers, rest) P = <0.05
Table 18 Comparison of like proportion of parous mosquitoes in trap types operated
in August 1965 and August 1966 at George Lake.
Trap type 1965 1966
not significant not significant
all 1 3 Aedes species caught were parous.
Sampling Methods
249
Table 19 Comparison of the proportion of parous mosquitoes taken in various trap
types at George Lake in June 1966 and 1967.
Trap type June 1966 May, June 1967
P N P:P+N P N P:P+N
x25 = 20.14 P = <0.001
250
Graham
Table 21 Comparison of the proportion of parous females of selected mosquito species
in trap types at George Lake (Results for 1965, 1966 and 1967 combined).
Trap Species
* Key
0. Pristine, unrubbed, very fresh appearance.
1. Very good, unrubbed, but not so fresh.
2. Good, some mesonotal scales missing but pattern clearly discernible.
3. Fair, mesonotum rubbed, but species still identifiable by scale pattern.
4. Rubbed, mesonotum with most scales missing, species not identifiable by scale pattern.
5. Bald, almost all scales missing, black legged Aedes sp. rarely identifiable.
Sampling Methods
251
tiago, 1934; Smith, 1942; Snow, 1949; and Muirhead-Thompson, 1958). These methods
are usually biased towards a few species. Many resting-site methods have been designed to
catch anopheline vectors of malaria. Methods designed for general survey work should catch
as wide a spectrum of species as possible and should not be selective for any species or
physiological state.
A large number of factors affect the efficiency of sampling methods. One of these is geo-
graphical location. The pit shelter (Muirhead-Thompson, 1958), was designed and worked
well in Rhodesia and in Java (D. A. Muir, pers. comm.), two areas very different in climate
and with very different mosquito species but failed when tried in Sarawak, an area similar in
climate and with many mosquito species in common with Java. The methods I tested were
all designed to be of use in general survey work. The findings may apply only to central
Alberta but should also apply to much of the southern part of the boreal forest in which
similar conditions and species occur. The mosquito fauna of this area is peculiar for the
great preponderance both in the numbers and in the number of species of Aedes and the
relative unimportance of all other genera except Culiseta. At George Lake five genera were
found but Aedes comprised 72% of the species and 85% of the individuals. This can be com-
pared to Kentucky, where eight genera were found and Aedes comprised 31% of the species
and 52% of the individuals (Breeland and Pickard, 1965). It is thus improbable that findings
in one area will apply in to to to the other.
The importance of the physiological state of the mosquitoes has been ignored in most
studies on sampling adult mosquito populations. Bursell (1961) showed that the physio-
logical state of tsetse flies ( Glossina swynnertoni Austen) varies according to the sampling
method used and this greatly affected interpretation of the results. Differences in the physi-
ological age of mosquitoes taken by different sampling methods could affect the results in
disease transmission studies as mosquitoes only become infected with disease-causing micro-
organisms after they have fed on an infected host. Thus a trap which takes a higher propor-
tion of physiologically older females than occur in the population will give an exaggerated
infection rate and if the method is selective for a few species may cause the vectorial impor-
tance of some species to be overrated. Though such a method may be useful where infected
females are rare or a pool of mosquitoes is used and an exact infection rate is not required.
The low activity of the adult females in stages III and IV of Christophers is important and
must be taken into account in population studies using sampling methods which catch active
mosquitoes, as this means that a significant proportion of the population is inactive and so
unavailable for sampling. Most studies, including this one, which show population peaks of
mosquitoes are actually showing peaks of activity rather than actual population peaks and
though the activity and population peaks are probably similar, this is by no means certain.
Methods of capturing resting adult female mosquitoes are inaccurate, biased towards a few
species and almost impossible to correlate with methods of taking active mosquitoes. If the
length of the gonotrophic cycle at different temperatures and the average number of gono-
trophic cycles passed through by the females in a population were known, an estimate of
the proportion in the stages III and IV of Christophers could be obtained. Polovodova’s
method enables the number of gonotrophic cycles passed through by a female mosquito to
be accurately determined and work on this in North America has been started both on
252
Graham
Culex tarsalis (Nelson, 1964; Burdick and Kardos, 1963) and on univoltine Aedes species
(Carpenter and Nielsen, 1965), but almost nothing appears to be known of the length of the
gonotrophic cycle in most North American species of mosquito. This is a fruitful field for
future research. The use of Polovodova’s method in determining the life history, vectorial
importance and population dynamics of Anopheles maculipennis Meigen is shown by Deti-
nova (1962).
The contents of the ventral diverticulum provide a partial measure of the energy reserves
available to the mosquito. Theoretically it should be possible to distinguish the proportion
of mosquitoes which have recently migrated into an area; these should have empty or nearly
empty ventral diverticula, having used up most of their energy resources on the flight and
the resident population should have full or nearly full diverticula. However, it would be
necessary to conduct a thorough investigation of the nectar resources available and of the
plants frequented by mosquitoes before the contents of the ventral diverticulum could be
used to distinguish migrants from resident mosquitoes. At George Lake the majority of mos-
quitoes had empty or only partially filled ventral diverticula in 1966. Since flowers were
abundant during the whole of the investigation, it is possible that many of the mosquitoes
caught had migrated in from outside the field site; this is supported by the very few larvae
found near the study area and the few males caught. Males are believed to be more seden-
tary than females, seldom moving more than a few miles from breeding sites, while adult
female Aedes in temperate regions may undertake long distance migrations (Clement,
1963).
Since male mosquitoes do not take blood meals, relatively little attention has been paid
to them in the past, as is shown by the few references to males in Bates (1949) and Clement
(1963). I paid little attention to male mosquitoes in this study. In the last few years the
development of sterile male methods of insect control has resulted in considerable interest
in male mosquitoes. Males probably give a better idea of the population breeding in the
vicinity of the study because they are more sedentary than the females and the adult males
of many species provide more reliable characters for specific determination than do the
adult females. It is unlikely that any one method will be equally effective for sampling both
adult females and adult males because their biologies differ considerably. The swarming
habits of the males of many species of mosquito will make the siting of traps even more
critical for males than females and this, coupled with the fact that females appear to be
longer lived than males and as they only mate once, it is unlikely that any one sampling
method will give a true sex ratio. The important sex ratio, the number of males to unmated
females, can probably be best estimated from rearing experiments.
Methods believed to take a random sample of the active population. — Malaise traps exert
no recognizable attraction to mosquitoes and so I believe they take a random sample of the
active mosquitoes and that this is unbiased both towards species and towards physiological
state. There is one possible area of bias, that is, against blood meal seeking mosquitoes. If
the generally accepted theory of host finding in mosquitoes, which is, that the biting cycle
represents the frequency with which a population in random flight comes within the range
of attraction of a host (Mattingly, 1949), is correct then there is no bias. However, Corbet
(1961) has shown that there may be a definite urge to bite and it is possible that some mos-
Sampling Methods
253
quitoes with this urge may rest on the vegetation until activated by the presence of a pos-
sible host as do tsetse flies ( Glossina species) (Glascow, 1963). If this is so then not many
hungry mosquitoes will be caught in Malaise traps which would introduce an element of
physiological bias.
At George Lake the proportion of the rarer species in Malaise traps was less than in traps
which used an attractive element. This indicates that these traps are unlikely to catch large
samples of these species, though they took a larger number of species than any other trap
type probably because of the longer operating time. The four species not taken in Malaise
traps were all rare, no more than two specimens of each being taken by all methods in 1966.
No species were taken by Malaise traps alone. Breeland and Pickard (1965) found that
Malaise traps caught a higher proportion of rare species and several species they recorded
were only taken in Malaise traps.
The Malaise trap has certain advantages over other traps; it has no moving parts; it can
work with a minimum of servicing, and needs to be emptied only once or twice a week,
which allows it to be operated in remote places; it operates twenty-four hours a day and the
catch is preserved in good condition. A few disadvantages are important; if left for any
length of time spiders spin webs across the entrances; it is very vulnerable to vandalism;
low efficiency and large size make it necessary to operate this trap for a prolonged period in
a single site and the position of the trap is more critical than for other trap types; many
mosquitoes which enter the base of the trap get attracted out before they are caught, so
that the number seen round the traps is no indication of the catch; and it is very difficult to
obtain a meaningful estimate of the volume of air filtered and so obtain an absolute density
figure. These disadvantages may preclude the use of Malaise traps for some studies.
Smith et al. (1965) found Malaise traps alone were capable of predicting outbreaks of
biting flies in Kentucky. The advantages listed above make this trap superior to most other
presumably unbiased methods of collection where absolute density figures are not required.
Suction traps (Southwood, 1966) require a motor or permanent electric supply and re-
quire regular servicing. The position of these traps is critical as in Malaise traps. The volume
of air filtered can be easily obtained so this trap can give an absolute density figure for fly-
ing mosquitoes.
A fairly recent innovation is a net attached to a car. These have been used by Stage and
Chamberlin (1945), Biddlingmayer (1964, and 1967) and Sommerman and Simmet (1965).
Provided the car is driven fast enough to eliminate attraction to moving bodies these nets
probably give a random sample of active insects. The period of operation is limited but this
method can cover a wide area. Sommerman and Simmet (1965) have provided a design
that enables the catching container to be changed at distance or time intervals, which makes
the results easier to interpret. If driven along the same route at the same time of day at regu-
lar intervals this method should produce useful results, but a series of Malaise traps at strate-
gic intervals would probably provide as useful if not more useful information at less cost.
The rotary trap is generally considered to take a random sample of flying insects and the
volume of air filtered can be easily determined, so if this trap had no attraction for mos-
quitoes an absolute density figure would be given. But the randomness of the sample taken
in these traps is open to doubt as the traps almost certainly exert some attractive stimuli to
254
Graham
mosquitoes and these may be selective for some species. Also, Maw (1964) has shown that
the nets can become charged with static electricity and repel some small flying insects. This
trap is bulky, requires considerable attention and damages the catch. I do not believe rotary
traps can supply data that cannot be obtained equally well by Malaise traps.
Much the same is true of the visual attraction trap, though this trap probably does take a
random sample of active mosquitoes.
If a random sample of active mosquitoes over a prolonged period is required and absolute
density figures are not necessary, Malaise traps can obtain this with less trouble than any
type of mechanically operated trap and are just as capable of detecting variations in popula-
tion level. Where a large sample is required or only a limited time is available some other
method such as visual attraction or car trap will possibly be better.
Methods known to take biased samples of the population. — Methods which use light, a
bait or which capture mosquitoes in resting places are considered here.
A large number of designs of light traps for insects have been described (Southwood,
1966), but relatively few are suitable for mosquitoes. Many small insects including mos-
quitoes are repelled by strong light (Verheijen, 1960; Barr et al. , 1963) and so traps designed
to catch these usually include a suction fan, like the New Jersey trap. Loomis (1959) con-
sidered that the air flow through a New Jersey trap must be standardized if two traps are
being compared. At George Lake in 1966 the number of mosquitoes per 10,000 cubic feet
of air filtered was 0.73 in Lt. I and 0.25 in Lt. II which are in the same ratio as the catches
per 100 trap hours are (2.9). This shows that the difference between the catches in the two
traps was not due to difference in air flow through them.
The species composition of light trap captures is usually considerably different from that
in the natural population. This has been shown by Southwood (1960) for Iieteroptera and it
has also been found in mosquitoes. The attraction to light may vary within a single species
over its geographical range and under different environmental conditions. Anopheles earlei
was one of the species in which a high proportion of the catch was taken in light traps at
George Lake, but McLintock et al. (1966) found a much lower proportion of this species
was taken by light traps than in collections by other methods in Saskatchewan.
Light traps can only provide data on relative changes in mosquito populations and are
probably of little use in life-table studies or in studies in which the true species composition
of the mosquito fauna is required. In spite of many drawbacks, light traps are useful survey
tools for mosquitoes and, if the attraction for older (parous) mosquitoes is found to be
widespread, will prove especially useful in some disease transmission studies. Light traps are
easily standardized and if run in the same place over a long period give an indication of
population changes. Clark and Wray (1967) used light traps in studies which enabled accu-
rate prediction of Aedes vexans invasions of Illinois cities. Few modern workers would go
as far as Mulhern (1953), who stated that since light traps were mechanized they gave better
results than methods involving collection by hand, such as human biting rate collections,
since these have a human element in them.
Light traps are particularly efficient for male mosquitoes (Belton and Galloway, 1965;
Southwood, 1966) and they are probably the most efficient method of sampling males.
Light traps were not very effective at George Lake because it is near the northern limit at
which light traps are useful.
Sampling Methods
255
Animal bait traps are often used and collections of mosquitoes settling on man have fre-
quently been used to obtain density figures for mosquitoes. Human baited traps are des-
cribed by Gater (1935) and Klock and Biddlingmayer (1953), and traps baited with large
animals by Magoon (1935), Shannon (1939), Bates (1944) and Roberts (1965). Wharton
et al. (1963) modified a Malayan trap (Gater, 1935) for use with monkeys as bait. In recent
years interest has arisen in mosquitoes which attack birds, in connection with studies on
arbor viruses, and several traps baited with chickens or other birds have been described
(Flemings, 1954 and Rainey et al., 1962). Lumsden (1958) and Worth and Jonkers (1962)
have designed traps which use small vertebrates as bait. The Lumsden trap uses a fan and can
give timed captures; portable versions of it are described by Snow et al. (1960) and Minter
(1961).
The advantages of small mammal traps seem to be great. At George Lake captures in rat
baited traps were very similar in composition to human bait captures, so white rats appear
to have great value as bait in mosquito surveys. They are hardy, can survive outdoors if
given shelter and can be left unattended for several days. As they give an approximation of
the human biting rate they will give an idea of the nuisance value of the mosquito species
present. Human bait captures while very useful can only be done for limited periods and are
more expensive. Birds require more attention and large traps and, while essential in some
virus transmission studies, they may give a false impression of the species composition, in
studies where nuisance to man is important.
The only animal-substitute bait tested was carbon dioxide. When added to a Malaise trap
this greatly increased both the catch and the number of species per unit time, but it appears
to be especially attractive to some species, destroying the random nature of Malaise trap
captures. For most purposes this is probably not too great a disadvantage and the increased
catch will offset this, as the species attracted will probably be pest species.
The erratic behaviour of the dry ice trap of Bellamy and Reeves (1952) was possibly due
to the small entrance area since the orienting stimulus of carbon dioxide appears to be weak.
The main disadvantage of the addition of carbon dioxide from a cylinder to Malaise traps is
that it is expensive and cannot be operated in remote places. However, dry ice is reasonably
cheap and when in large enough blocks and suitably packed can last for several days while
emitting large quantities of carbon dioxide. I believe that a useful mosquito trap for general
survey purposes in places which cannot be visited every day would be a Malaise type trap
modified to use dry ice as a bait. In central Alberta 25 pounds of dry ice lasts two to three
days, which would make it possible to service this trap at bi-weekly intervals. Alone or com-
bined with small mammal bait traps these traps could be operated in places several miles
from a city, close to major mosquito breeding areas, and provide accurate data for fore-
casting the need for control measures in the city.
Olkowski et al. (1967) found Malaise traps baited with dry ice caught significantly more
tabanids (Diptera; Tabanidae) than did unbaited traps in California. At George Lake taba-
nids were too scarce and erratic in occurrence for any conclusions to be drawn. Anderson
et al. (1967) found the same proportion of the population of Symphoromyia (Diptera;
Leptidae) were taken in Malaise traps baited with dry ice as were attracted to their natural
hosts. They considered that these traps could give equivalent information on factors in-
256
Graham
fluencing attack rates at a lower cost, as could direct observation and collection of the flies
from the hosts.
Collections in resting sites are useful for a few limited purposes, especially for obtaining
blood fed females for host determination (World Health Organization, Division of Malaria
Eradication and Lister Institute of Preventive Medicine, 1 960) or for obtaining detailed in-
formation on the resting habits of mosquitoes for control with residual insecticides (Muir-
head-Thompson, 1951). These methods have proved very valuable in studies on anophelines
but less successful for northern Aedes.
Results from central Alberta (Graham in Prep, and Happold, 1965) indicate that no local
Aedes species exhibit any special tendency to enter buildings, but Anopheles earlei, Culiseta
inornata and Culex tarsalis are known to hibernate in basements. Shemanchuk (1965) has
shown that the principal hibernation sites of these species are in animal burrows. Further
studies on the resting habits of Alberta mosquitoes are required.
Conclusions
The principal conclusions drawn from this study are that the position as well as the trap
type greatly affects the size and composition of the catch. Malaise traps both with and
without carbon dioxide are probably the most useful types of traps for general mosquito
survey work as they can be operated away from sources of power and need relatively little
attention. Light traps will continue to be very useful in mosquito surveys, especially inside
urban areas, as long as their limitations are clearly understood. Animal bait and resting-site
captures are useful for specific purposes. Human bait captures are particularly useful for
assessing mosquito nuisance and should always be used by urban mosquito control organiza-
tions in conjunction with other sampling methods but are of little value alone for forecast-
ing the need for adult control measures in cities.
Although not tested in this study, car trap captures will probably be very useful in areas
where vandalism or other factors prevent the use of Malaise or animal bait traps outside the
city limits, but they should not be used instead of these.
ACKNOWLEDGEMENTS
I wish to express my thanks to B. Hocking, G. E. Ball, D. A. Boag, W. G. Evans, K. Smillie
and J. A. Downes for reading and criticizing the manuscript; to N. Wood for assistance in
the field and to W. Proctor for help with equipment. I also wish to thank the City of
Edmonton and the University of Alberta for the provision of grants and the Canada Depart-
ment of Agriculture, Lethbridge, Alberta, for the loan of traps.
Sampling Methods
257
REFERENCES
Anderson J.R., W. Olkowski and J.B. Hoy. 1967. Relationship between host attack rate and
CC>2 baited Malaise trap catches in certain Symphoromyia species. Proc. 35th Conf. Calif.
Mosquito Control Assoc.: 77.
Barr A.R. 1958. The mosquitoes of Minnesota. Univ. Minn. Agric. exp. Sta. Tech. Bull. 228.
154 pp.
Barr A.R., T.A. Smith, M. Boreham and K.E. White. 1963. Evaluation of some factors
affecting the efficiency of light traps in collecting mosquitoes. J. econ. Ent. 56: 123-127.
Bates M. 1944. Notes on the construction and use of stable traps for mosquito studies. J.
nat. Mai. Soc. 3: 135-145.
Bates M. 1949. The natural history of mosquitoes. New York, Harper and Row, 1965 re-
print: x + 378 pp.
Bellamy R.E. and W.C. Reeves. 1952. A portable mosquito bait trap. Mosquito News 12:
256-258.
Belton P. and M. Galloway. 1965. Light trap collections of mosquitoes near Belleville,
Ontario. Proc. ent. Soc. Ont. 96: 90-96.
Biddlingmayer W.L. 1964. The effect of moonlight on the flight activity of mosquitoes.
Ecology 45: 87-94.
Biddlingmayer W.L. 1967. A comparison of trapping methods for adult mosquitoes: Species
response and environmental influence. J. Med. Ent. 4: 200-220.
Breeland S.G. and E. Pickard. 1965. The Malaise trap — an efficient and unbiased mosquito
collecting device. Mosquito News 25 : 87-94.
Brown A.W.A. 1951. Studies on the responses of the female Aedes mosquito; Part IV; field
experiments on Canadian species. Bull. ent. Res. 42: 575-582.
Brown A.W.A. , D.S. Sarkaria and R.P. Thompson. 1951. Studies on the responses of the
female Aedes mosquito; Part I; the search for attractant vapours. Bull. ent. Res. 42:
105-114.
Burdick D.J. and E.H. Kardos. 1963. The age structure of fall, winter and spring populations
of Culex tarsalis in Kern County, California. Ann. ent. Soc. Amer. 56: 527-535.
Bursell E. 1961. The behavior of tsetse flies (Glossina swynnertoni) in relation to problems
of sampling. Proc. R. ent. Soc. Lond. A. 36: 69-74.
Carestia R.R. and L.B. Savage. 1967. Effectiveness of carbon dioxide as a mosquito attrac-
tant in the C.D.C. miniature light trap. Mosquito News 27: 90-92.
Carpenter M.J. and L.T. Nielsen. 1965. Ovarian cycles and longevity in some univoltine
Aedes species in the Rocky Mountains of the western United States. Mosquito News 25:
128-135.
Chamberlin J.C. and F.R. Lawson. 1945. A mechanical trap for the sampling of aerial mos-
quito populations. Mosquito News 5: 4-7.
Clark J.C. and F.C. Wray. 1967. Predicting influxes of Aedes vexans into urban areas. Mos-
quito News 27: 156-163
Clement A.N. 1963. The physiology of mosquitoes. Oxford, Pergamon Press, ix + 393 pp.
258
Graham
Corbet P.S. 1961. Entomological studies from a high tower in Mpandga forest, Uganda, VI:
Nocturnal flight activity of Culicidae and Tabanidae as indicated by light traps. Trans. R.
ent. Soc. Lond. 113: 301-314.
Detinova T.S. 1962. Age grouping methods in Diptera of medical importance, with special
reference to some vectors of malaria. Monogr. Wld. Hlth. Org. No. 47: 216 pp.
Dow R.P. 1959. A method of testing insect traps and attractants and its application to stud-
ies of Hippelates pusio and Culex tarsalis. J. econ. Ent. 52: 496-502.
Downey J.E. 1962. Mosquito catches in New Jersey mosquito traps and ultra violet light
traps. Bull. Brooklyn ent. Soc. 57: 61-63.
Duke B.O.L. 1955. Studies on the biting habits of Chrysops, III. Ann. trop. Med. Parasit.
(Liverpool). 49: 362-367.
Fisher R.A., A.S. Corbet and C.B. Williams. 1943. The relation between the number of indi-
viduals and the number of species on a random sample of an animal population. J. anim.
Ecol. 12: 42-58.
Flemings M.P. 1954. An altitude biting study of Culex tritaeniorhyncus and associated
mosquitoes in Japan. J. econ. ent. 52: 275-285.
Gater B.A.R. 1935. Aids to the identification of Anopheline imagines in Malaya. Govt. S.S.
and Mai. Adv. Board. F.M.S. 242 pp.
Glascow J.P. 1963. The distribution and abundance of tsetse. Oxford, Pergamon Press, xi +
241 pp.
Graham P. in prep. Observations on the biology of adult female mosquitoes at George Lake,
Alberta.
Haddow A.J. 1960. Studies on the biting habits and medical importance of East African
mosquitoes in the genus Aedes. Bull. ent. Res. 50: 759-779.
Happold D.C.B. 1965. Mosquito ecology in central Alberta. II. Adult populations and activi-
ties. Can. J. Zool. 43: 821-846.
Haufe W.O. and L. Burgess. 1960. Design and efficiency of mosquito traps based on visual
response to patterns. Can. Ent. 92: 124-140.
Hayes R.O., R.E. Bellamy, W.C. Reeves and M.J. Willis. 1958. Comparison of four sampling
methods for measurement of Culex tarsalis adult populations. Mosquito News 18: 218-
227.
Hocking B. 1953. The intrinsic range and speed of flight of insects. Trans. R. ent. Soc.
Lond. 104: 223-345.
Huffacker C.B. 1942. Tests with carbon dioxide and light as an attractant for mosquitoes
with especial emphasis on the malaria mosquito, Anopheles quadrimaculatus . Mosquito
News 2: 561-569.
Huffacker C.B. and R.C. Bach. 1943. A study of methods of sampling mosquito popula-
tions. J. econ. Ent. 36: 561-569.
Juillet J.A. 1963. A comparison of four traps used for capturing flying insects. Can. J. Zool.
41: 219-223.
Klock J.W. and W.L. Biddlingmayer. 1953. An adult mosquito sampler. Mosquito News 13:
157-159.
Sampling Methods
259
Laarman J.J. 1955. Host seeking behavior of the malaria mosquito Anopheles maculipennis
atroparvus. Acta Leidensia 25: 1-144.
LaRoi G. 1968. Taiga. In W.G. Hardy, Editor. Alberta, a natural history. Edmonton, M.G.
Hurtig.
Loomis E.C. 1959. A method for the more accurate determination of air volume displace-
ment in light traps. J. econ. Ent. 52: 342-345.
Love G.J. and W.W. Smith. 1957. Preliminary observations on the relation of light traps to
mechanical sweep net collections in sampling mosquito populations. Mosquito News 17:
9-14.
Lumsden W.H.R. 1958. A trap for insects biting small vertebrates. Nature, Lond. 181:
819-820.
Magoon E.H. 1935. A portable stable trap for capturing mosquitoes. Bull. ent. Res. 26:
363-372.
Malaise R.A. 1937. A new insect trap. Ent. Tidskr. 58: 148-164.
Mattingly P.F. 1949. Studies on West African forest mosquitoes. Part II. The less common
species. Bull. ent. Res. 40: 387-402.
Maw M.G. 1964. An effect of static electricity on captures in insect traps. Can. Ent. 96:
1482.
McLintock J., A.N. Burton, H. Dillenberg and J.G. Rempel. 1966. Ecological factors in the
1963 outbreak of western equine encephalitis in Saskatchewan. Can. J. Publ. Hlth. 57:
561-575.
Minter D.M. 1961. A modified Lumsden suction trap for biting insects. Bull. ent. Res. 52:
233-238.
Muirhead-Thompson R.C. 1951. Mosquito behavior in relation to malaria transmission in
the tropics. London, Arnold viii + 219 pp.
Muirhead-Thompson R.C. 1958. A pit shelter for sampling outdoor mosquito populations.
Bull. Wld. Hlth. Org. 19:1116-1118.
Mulhern T.D. 1953. Better results with mosquito light traps through standardizing mechani-
cal performance. Mosquito News 13: 130-133.
Nelson R. 1964. Parity in winter populations of Culex tarsalis Coquillet in Kern County,
California. Amer. J. Hyg. 80: 242-253.
Newhouse V.F., R.W. Chamberlain, J.G. Johnson and W.D. Sudia. 1966. Use of dry ice to
increase mosquito catch in the C.D.C. miniature light trap. Mosquito News 26: 30-35.
Nielsen L.T. and A.T. Nielsen. 1953. Field observations on the habits of Aedes taeniorhyn-
chus. Ecology 34: 141-156.
Olkowski W., J.R. Anderson and J.B. Hoy. 1967. Relationship between host attack rates
and CC>2 baited Malaise trap catches of certain Tabanid species. Proc. 35th Conf. Calif.
Mosquito control Assoc.
Rainey M.B., G.V. Warren, A.D. Hess and J.S. Blackmore. 1962. A sentinel chicken shed
and mosquito trap for use in encephalitis field studies. Mosquito News 22: 337-342.
Reeves W.C. 1953. Quantitative field studies of a carbon dioxide chemotropism of mosqui-
toes. Amer. J. trop. Med. Hyg. 2: 233-246.
260
Graham
Reeves W.C. and McD. Mammon. 1942. Mosquitoes and encephalitis in Yakima Valley,
Washington. IV. A trap for collecting live mosquitoes. J. inf. Dis. 70: 275-277.
Roberts R.H. 1965. A steer baited trap for sampling insects affecting cattle. Mosquito News
25: 281-285.
Rudolfs W. 1922. Chemotropisms of mosquitoes. New Jersey Agr. Exp. Sta. Bull. 367. 22
pp.
Russel P.F. and D. Santiago. 1934. An earth-lined trap for anopheline mosquitoes. Proc.
ent. Soc. Wash. 35: 1-21.
Salitemik Z. 1960. A mosquito light trap for use on cesspits. Mosquito News 20: 295-296.
Shannon R.G. 1939. A method for collecting and feeding mosquitoes in jungle yellow fever
studies. Amer. J. trop. Med. 19: 131-138.
Shemanchuk J.A. 1965. On the hibernation of Culex tarsalis Coquillet, Culiseta inornata
(Williston) and Anopheles earlei Vargas (Diptera: Culicidae) in Alberta. Mosquito News
25: 456-462.
Simpson G.G., A. Roe and R.C. Lewontin. 1960. Quantitative Zoology, 2nd edition. New
York, Harcourt-Brace. vii + 440 pp.
Smith G.E. 1942. The keg shelter as a diurnal resting place for Anopheles quadrimaculatus.
Amer. J. trop. Med. 22: 257-269.
Smith G.E., S.G. Breeland and E. Pickard. 1965. The Malaise trap, a survey tool in medical
entomology. Mosquito News 25: 389-400.
Snow W.E. 1949. Studies on portable resting stations for Anopheles quadrimaculatus in the
Tennessee Valley. J. nat. Mai. Soc. 8: 336-343.
Snow W.E., E. Pickard and R.E. Sparkman. 1960. A fan trap for collecting biting insects
attacking avian hosts. Mosquito News 20: 315-316.
Sommerman K.M. and R.P. Simmet. 1965. Car top trap with terminal cage in auto. Mos-
quito News 25: 172-182.
Southwood T.R.E. 1960. The flight activity of Heteroptera. Trans. R. ent. Soc. Lond. 112:
173-220.
Southwood T.R.E. 1966. Ecological methods. London, Methuen, xviii + 291 pp.
Stage H.H. and J.C. Chamberlin. 1945. Abundance and flight activity of certain Alaskan
mosquitoes as determined by means of a rotary trap. Mosquito News 5: 8-16.
Stage H.H., C.M. Gjullin and W.W. Yates. 1952. Mosquitoes of the north-western states.
U.S.D.A. Handbook No. 45. 95 pp.
Standfast H.A. 1965. A miniature light trap which automatically segregates the catch into
hourly samples. Mosquito News 25: 48-53.
Townes H. 1962. Design for a Malaise trap. Proc. ent. Soc. Wash. 64: 253-264.
Trembley H.L. 1952. The distribution of certain liquids in the oesophageal diverticulae and
stomach of mosquitoes. Amer. J. trop. Hyg. 1 : 693-710.
Verheijen F.J. 1960. The mechanism of the trapping effect of artificial light sources upon
animals. Arch. Neerland. Zool 13: 1-107.
Wharton D.H., D.E. Eyles and M.C.W. Warren. 1963. The development of methods for trap-
ping vectors of monkey malaria. Ann. trop. Med. Parasit. Liverpool. 57: 32-46.
Sampling Methods
261
Williams C.B. 1964. Patterns in the balance of nature. London, Academic Press, vii + 324
pp.
Willis E.R. 1947. The olfactory responses of female mosquitoes. J. econ. Ent. 40: 769-778.
World Health Organization, Division of Malaria Eradication and Lister Institute of Preven-
tive Medicine. 1960. A study of the blood feeding patterns of anopheline mosquitoes
through precipitin tests. Bull. Wld. Hlth. Org. 22: 685-720.
Worth C.B. and A.H. Jonkers, 1962. Two traps for mosquitoes attracted to small verte-
brates. Mosquito News 22: 15-21.
Zhogolev D.T. 1959. (Light traps as a method for collecting and studying vectors of disease
organisms.) Ent. Obozr. 38: 766-773. (In Russian).
SOME PHARMACOLOGICAL PROPERTIES OF THE NERVE CORD
OF THE COCKROACH ; PERIPLANETA AMERICANA (L.)
PETER K. CTIIANG
Department of Entomology
Quaestiones entomologicae
5 : 263-306 1969
University of Alberta
Edmonton, Alberta
Fluctuations in the endogenous activity and synaptic transmission were observed in in-
tact, in situ nerve cords o/ Periplaneta americana (L.). The endogenous activity and synaptic
transmission were inhibited by hemicholinium; choline reversed this inhibition. Carbachol
inhibited electrical activity entirely. Nicotine stimulated immediately upon application,
followed by block. Neither dimethylphenyl piperazinium nor methacholine produced any
observable effect. Pilocarpine depressed endogenous activity and synaptic transmission.
Acetylcholine at 10~ 3 M produced no observable effect, but at 10~ 2 M caused a progressive
blocking without apparent stimulation. Choline blocked synaptic transmission in some prep-
arations. Eserine caused synaptic facilitation, followed by block ; tetraethylpyrophosphate
produced similar effects. Pyridine-2-aldoxime methiodide reactivated synaptic transmission
and endogenous activity in TEPP-treated nerve cords. Choline apparently did not reactivate
the electrical activity of TEPP-treated nerve cords. Phenoxybenzamine and tranylcypromine
blocked both the endogenous activity and synaptic transmission. Spectrofluorometric assay
did not reveal the presence of noradrenaline in the nerve cords. The acetylcholinesterase
activity of individual 6th abdominal ganglion varied considerably.
In Periplaneta americana (L.), a system of neurons has been described which is called the
giant fiber system (Hess, 1958; Roeder, 1948a, 1953, 1962). Afferent fibers from mechano-
receptors on the caudal segments converge on the last abdominal (6th) ganglion and synapse
with a small number of giant fibers. The giant axons ascend the nerve cord through the ab-
dominal ganglia, and there appears to be no synapse at any point in the giant fibers of the
abdominal nerve cord (Hess, 1958; Roeder, 1948a). Some of the giant fibers apparently
ascend to the brain without interruption, and others synapse in the thoracic ganglia with
motor neurons supplying the leg muscles (Hess, 1958). The giant fiber system mediates
evasive response of the insects (Roeder, 1948a, 1962, 1963). In the intact roach, puffs of
air applied to the sensilla on the cerci induce an alarm reaction causing evasive response
(Roeder, 1948a, 1962).
In vertebrates, acetylcholine plays a vital role in preganglionic transmission in the auto-
nomic nervous system, skeletal neuromuscular junctions, and the central nervous system
(Koelle, 1963, 1965; and others). Acetylcholine may also have a role in adrenergic transmis-
sion (Burn and Rand, 1965; Koelle, 1963). Acetylcholine (ACh) and the enzymes, choline
acetylase (ChA) and acetylcholinesterase (AChE) which facilitate ACh synthesis and hydrol-
ysis respectively, have been demonstrated in the insect tissues including nervous tissue, yet
the physiological role of ACh in insects remains to be demonstrated (Colhoun, 1963b;
Smith, 1965).
264
Chiang
Acetylcholine has been suggested as a transmitter substance in the cockroach nervous
system (Smith and Treherne, 1965; Treherne, 1966; Yamasaki and Narahashi, 1960). An
increase of ACh content was demonstrated in nerve cords of P. americana treated with
eserine (Colhoun, 1958b; Mikalonis and Brown, 1941). Colhoun (1963b) summarized the
amounts of ACh content in various species of insects. ACh content in the 6th abdominal
ganglion was 63 ju gm./gm. (Colhoun, 1958a), and is located in “structural compartments.”
Cholinacetylase, the enzyme for the synthesis of ACh, was demonstrated in blowfly heads
(Smallman, 1956). True acetylcholinesterases have been demonstrated in or isolated from
the central nervous system (CNS) of various species of insects (Treherne, 1966).
Investigating synaptic and axonic transmission inP. americana, Roeder (1948b) first con-
cluded that there was no evidence showing that ACh was a synaptic transmitter of nerve
activity, although anticholinesterases disrupted synaptic transmission in the 6th abdominal
ganglion. The conclusion was also based on the failure of high external concentrations of
ACh, other choline esters, and cholinergic blocking agents to interfere with synaptic con-
duction.
Later, Roeder and Kennedy (1955) proposed that organophosphates could, besides in-
hibiting AChE, block ACh receptors at high concentrations. To circumvent the ineffective-
ness of externally applied ACh, Twarog and Roeder (1956, 1957) desheathed the last ab-
dominal ganglion, and observed that ACh, between 10-2 M and 10-3 M exerted a rapid
and pronounced effect on the synaptic transmission of the ganglion. In two of a series of
seventeen experiments, only a moderate decrease in synaptic response was noted. In all
others, within one to five minutes, bursts of synchronous action potentials were followed by
synaptic depression and block. Desheathing also reduced the concentrations at which other
pharmacological agents affected synaptic transmission. This led to the postulation of an ion
barrier theory, which held that the connective tissue sheath investing the roach nervous sys-
tem retards the passage of ions from the bathing medium to the interior of ganglion and
connectives (Twarog and Roeder, 1956, 1957). But the authors also pointed out the possi-
bility of well protected synapses in insect ganglia (Twarog and Roeder, 1957).
Based on his observations on the electrical activity of Locusta migratoria (L.), Hoyle
(1952, 1953) also suggested that the sheath surrounding peripheral nerves and ganglia of
insects may act as a selectively permeable barrier, separating the molecules and ions of the
haemolymph from those of the nervous system. O’Brien (1957, 1959a, b) reported that
ionized pharmacological compounds had low toxicity to insects. Histological studies demon-
strated that methylthiocholine was completely prevented from penetrating the intact nerve
sheath (Winton, Metcalf, and Fukuto, 1958).
Treherne (1961a, b, c; 1962a, b), and Treherne and Smith (1965a, b) demonstrated that
inorganic ions and ACh do penetrate the nerve tissue of insects. The influx of 14C-labelled
ACh into the extracellular system of the roach nerve cord occurred extremely rapidly, with
a half-time of approximately 50 seconds (Treherne and Smith, 1965a). The insect nervous
system is not virtually isolated beneath an impermeable nerve sheath, but is in a dynamic
equilibrium with some smaller ions and molecules in the haemolymph (Treherne, 1965b).
Intense AChE activity was demonstrated in the neuropile, sheaths encapulating the neu-
ron perikarya, and the perineurium of the nerve sheath of the insect CNS (Iyatomi and
Cockroach Nerve Cord
265
Kanehisa, 1958; Wigglesworth, 1958). Electron microscopy has revealed the following
distribution of eserine-specific esterase activity in the 6th abdominal ganglion of the roach,
P, americana (Smith and Treherne, 1965): (1) in the glial sheaths of the axons in the con-
nective and cereal nerves; (2) in the glial folds encapsulating the neuron perikarya in the
ganglion: (3) in localized areas along the membrane of axon branches with the neuropile,
frequently in association with local clusters of synaptic vesicles.
Bisset et al. (1960) found (3, j3-dimethyl acrylcholine in the prothoracic gland of the gar-
den tiger moth Arctia caja. But Chang and Kearns (1961), and Colhoun and Spencer (1959)
failed to demonstrate the presence of any free choline esters, other than ACh in the nervous
system of the cockroach. Colhoun (1963b) pointed out the difficulty in accepting ACh as a
synaptic transmitter first because of the failure to prove that ACh accumulated in nervous
tissue as a free ester following stimulation in the presence of anticholinesterases, and second,
the lack of evidence for antidromic stimulation or the blocking effects of some pharmaco-
logical agents, such as curare and atropine.
The role that biogenic amines may play in synaptic transmission in insects is largely un-
certain, though catecholamine-like substances have been extracted from insects. Frontali
(1968) demonstrated the presence of catecholamines in the roach brain by means of a
fluorescence histochemical method.
EXPERIMENTAL METHODS
Pumphrey and Rawdon-Smith (1937), and Roeder (1948a) have shown that stimulation
of the cereal sensilla of P. americana elicits a volley of impulses from the cereal sensory
fibers which pass to the 6th abdominal ganglion, and the impulses subsequently pass to the
nerve cord. Colhoun (1958b, 1960) studied synaptic transmission in the 6th abdominal
ganglion in situ when the cerci were exposed to air puffs. Adaptation to air puffs was not
shown in the 6th abdominal ganglion (Colhoun, 1960). There was an increase in total ACh
content in eserinized, isolated nerve cords, which were stimulated by air puffs (Colhoun,
1960). But there was no detectable change in total ACh content in the in situ nerve cords or
the 6th abdominal ganglia, treated the same way (Colhoun, 1960).
Since each step involved in neurohumoral transmission represents a potential point of
drug attack, Koelle (1965) proposed the following four possible approaches for identifying
a cholinergic synapse by considering the prototype drugs that affect processes concerned in
each step:
(1) Interference with the release of the transmitter:
Hemicholinium (HC-3) can block synaptic transmission by blocking the transport system
by which choline accumulates in the terminals of cholinergic fibers, and thus it limits the
synthesis of ACh.
(2) Promotion of the transmitter release:
Carbachol is supposed to act by releasing ACh at the synapse. It also probably acts
directly at postsynaptic cholinergic receptors.
(3) Combination with postsynaptic receptor sites:
266
Chiang
When a drug combines with a receptor, two effects may be observed: the same effect as
that of ACh (i.e., cholinomimetic); or no apparent direct effect but, by occupying the recep-
tor site, the drug prevents the action of endogenous ACh (i.e., cholinergic blockade).
(4) Interference with the destruction or dissipation of the transmitter:
The primary action of anticholinesterases, such as organophosphates and carbamates, is
the inhibition of AChE, with the consequent accumulation and action of endogenous ACh
at sites of cholinergic transmission. All drugs in this class probably have in addition, direct
actions at cholinoreceptive sites, and elsewhere.
Electrophysiological Studies
Materials
The chemicals were obtained as follows: acetylcholine chloride, nicotine, and pyridine-2-
aldoxime methiodide (2-PAM) from City Chemical Corporation; acetyl-beta-methyl choline
chloride, and eserine sulfate from Nutritional Biochemical Corporation; choline chloride
from Eastman Kodak Company; hemicolinium-3(HC-3) from Aldrich Co.; carbachol (carba-
mylcholine) and pilocarpine from British Drug House; phenoxybenzamine hydroxide (di-
benzyline), and tranylcypromine from Smith Kline and French. Tetraethylpyrophosphate
(TEPP) was donated by California Chemical Company, and 1, l-dimethyl-4-phenyl pipera-
zinium (DMPP) was donated by Dr. Graham Chen of Parke, Davis & Company.
Methods
Rearing of P. americana. - The roaches were reared in glass battery jars in the culture
room at 30°C and 50 to 60% R.H. The insects were supplied with water and rabbit pellets as
food. The roaches were kept in the laboratory first for at least 1 2 hours before an experi-
ment. Only male, adult roaches were used for the electrophysiology experiments.
In situ nerve cord preparation. — A male cockroach was lightly anaesthetized with carbon
dioxide. After the wings, antennae, and legs were cut off, the roach was pinned on a wax
block, slit dorsally and eviscerated. The nerve cord was cleared of connectives on both sides
from the last abdominal ganglion up to the thorax, and moistened with insect saline: NaCl
9.0 gm., KC1 0.2 gm., CaC^ 0.2 gm. per liter of distilled water, pH 7.0 (Pringle, 1938). A
fresh wax block was used for each roach. Mounted on a micro-manipulator, a pair of fine-
tapered platinum electrodes was hooked under the nerve cord between the 5th and 6th ab-
dominal ganglia. A period of 1 5 minutes was allowed for the nerve cord to achieve its steady
state (Weiant, 1958). Stimulation of the anal cerci by air puff was accomplished by pressing
a rubber bulb connected to a 16 cm. long Pasteur pipette, the tip of which was 0.5 cm.
from, but directed toward, the left cercus. Modified after Colhoun (1958b, 1960), this
method eliminated the possible production of neuroactive agents by electrical stimulation
reported by Sternburg, Chang and Kearns (1959). Potentials evoked by drugs or by pregang-
lionic stimulation could be detected conveniently. The analysis of synaptic response was
according to that of Prosser (1940).
Before recording the electrical activity of a nerve cord, the bathing solution was carefully
withdrawn with a Pasteur pipette, leaving the nerve cord resting on the electrodes. Wires
from the two platinum electrodes were connected push-pull to a Tektronix Type 122 pre-
amplifier, from which single-ended output was connected to a Tektronix Type 502 dual-
beam oscilloscope. The upper beam was used to display the electrical activity.
Cockroach Nerve Cord
267
A flexible copper plate was placed above, and a second one was placed below the rubber
bulb which did the air puffing. These plates were held in place by a piece of lucite. One
piece of wire was soldered to each of the copper plates; one led directly to a terminal of a
six volt radio battery and the other led to an input of the lower beam of the oscilloscope.
From the other terminal of the battery, a wire was connected to the grounding input of the
lower beam. Whenever the bulb was pressed, a current occurred between the two copper
plates, resulting in a short visible trace on the oscilloscope screen. This indicated the begin-
ning of the effect of air puffing the cerci of roaches, not the duration of the air puff.
Recording of the electrical activity was made with a Grass C-4 oscilloscope camera, using
Kodak Kind-1732 photographic paper.
The spikes were counted under a pair of binocular microscopes by the use of a tabulator.
Each set of records was counted twice, and the average was used. Since the synaptic
response varied in duration, the activity is expressed as spikes/air puff, whereas the endo-
genous activity in spikes/sec.
Eserine was first dissolved in acetone, and then diluted to the desired concentrations in
insect saline with no more than 1% acetone in the final solutions. Up to 8% acetone pro-
duced no detectable effect on the AChE activity (Dauterman, Talens, and van Asperen,
1962). Each solution was applied gently by a Pasteur pipette to the 6th abdominal ganglion.
After the initial observation was made, the entire nerve cord was bathed in the solution.
Since the solution gradually leaked out of the roach abdomen, additional solution was
applied as required.
Determination of AChE Activity
The AChE was assayed by a modification of van Asperen’s method (1962) which was
based on Gomori’s technique (1953). This sensitive method is primarily a colorimetric deter-
mination of naphthol produced by the enzymatic hydrolysis of naphthylacetate.
Materials
Naphthol was obtained from Fisher Scientific Company, a-naphthylacetate from Eastman
Kodak Company, and diazoblue-B from Edward Gurr, Limited.
Methods
Enzyme Preparation. — The 6th abdominal ganglion was dissected from an adult roach,
male or female, and rinsed in insect saline solution. The ganglion was then homogenized in
ice-cold 0.01 M phosphate buffer pH 7 in a Potter and Elvehjem homogenizer. The homo-
genate was centrifuged at 30,000 x g for 20 minutes and the supernatant diluted to one-half
ganglion per ml.
Substrate Solutions. — Substrate solution was prepared by diluting a stock solution of a-
naphthylacetate (0.03 M) in acetone with 0.01 M phosphate buffer pH 7 to give a final sub-
strate concentration of 3 x 10-4 M.
Diazoblue-B , Sodium Laurylsulfate Solution. — This solution, used for the quantitative
determination of the amount of naphthol produced, consists of 3 parts of a 1% diazoblue-B
solution and 7.5 parts of a 5% sodium laurylsulfate solution. The a-naphthol reacts with
diazoblue-B to give a strong blue color. Sodium laurylsulfate immediately stops all esterase
activity, and solubilizes the naphthol-diazoblue complex (van Asperen, 1962).
268
Chiang
Assay of AChE. - Cockroach nervous system contains AChE, ali-esterase (AliE), and
aryl-esterase (ArE) (Chadwick, 1963). Eserine is believed to inhibit the AChE activity com-
pletely, but not the other two esterases (Chadwick, 1963).
The assay was started by pipetting 0.5 ml. of enzyme solution (% ganglion) into 5 ml. of
the substrate solution. The reaction was stopped by the addition of 1 ml. of diazoblue
laurylsulfate solution (DBLS). In each assay, enzymatic hydrolysis was permitted in 1 tube
for 0 minutes and a second tube for 10 minutes at 40°C. The resulting color was read in a
Beckman DU-2 spectrophotometer at 600 m pi 5 minutes after addition of DBLS. The total
esterase activity (EAO.D.) was equal to O.D. of material incubated 10 minutes - O.D. of
material incubated 0 minutes. To determine the nonspecific esterase activity, the same pro-
cedure was followed, except that the substrate solution contained eserine at a particular
concentration. This gave an O.D. after AChE inhibition (IAO.D.). The AChE activity (N
O.D.) which was calculated as N O.D. = EAO.D. - I O.D.
The same procedure was followed to assay the AChE inhibited by TEPP when electrical
activity of the roaches ceased, except that the ganglia were homogenized with a phosphate
buffer (pH 7) containing 0.3% acetylcholine chloride (Colhoun, 1959a).
Spectrofluoremetric Determination of Noradrenaline
Modified after that of Shore and Olin (1958), this procedure has been successful in assay-
ing catecholamines of mammalian hearts (C. W. Nash, personal communication). The cate-
cholamines are first oxidized to red indole derivatives which then become strongly fluores-
cent hydroxyindoles in the alkali.
Materials
The distilled water used throughout this experiment was demineralized twice.
(1) Salt-saturated butanol: 49 g. of sodium chloride, and 2 ml. of HC1 were added to 946
ml. of reagent grade n-butanol.
(2) 4% versene (pH 6. 3-6.5): 8 g. of EDTA were dissolved in 200 ml. of distilled water,
and the pH adjusted to 6. 3-6. 5 using sodium hydroxide
(3) 0.1 M Iodine: 0.75 g. of iodine and 14.40 g. of potassium iodide were dissolved in 300
ml. of distilled water.
(4) Alkaline sulfite solution: 0.63 g. of anhydrous sodium sulfite were dissolved in 5 ml.
of distilled water; to this 20 ml. of 5 N NaOH were added.
(5) Standard noradrenaline was obtained from New England Nuclear Corportation.
Method
The abdominal nerve cords were dissected from adult roaches, and transferred to ice-cold
saline. The nerve cords were then frozen in lots of 10, and stored overnight in a deep freeze.
Before the extraction procedure began, the nerve cords were thawed, taken out of the vials
and frozen again by liquid nitrogen. The total weight of the nerve cords was 0.493 g. The
nerve cords were homogenized in Potter and Elvehjem homogenizer with 4 ml. of cold, acid,
salt-saturated butanol for 5 minutes. The homogenate was transferred to a 15 ml. screw-cap
centrifuge tube, shaken for 5 minutes and centrifuged for 10 minutes at 540 x g. The super-
natant was transferred to another 1 5 ml. screw-cap centrifuge tube, and mixed with 6 ml. of
heptane and 1.5 ml. of 0.01 N HC1. The mixture was shaken and centrifuged at 540 x g for
5 minutes. The organic layer was discarded.
Cockroach Nerve Cord
269
The acid layer was then pipetted in 0.5 ml. aliquots into three test tubes containing 1 ml.
of 4% versene each, and 0.2 ml. of 0.1 M iodine was added to two of these tubes. After 2
minutes, 0.5 ml. of alkaline sulfite solution was added. The third tube was used as a tissue
blank, which had its order of oxidation reversed, i.e., alkaline sulfite preceding iodine. After
another 2 minutes, 0.6 ml. of 5 N acetic acid was added. The contents of the tubes were
heated in a boiling water bath for 5 minutes, and cooled rapidly in cold water to room
temperature. The fluorescence for noradrenaline was read in an Aminco Bowman spectro-
fluorometer with activation and fluorescence wavelength at 385 m /x and 485 m n respec-
tively.
Two noradrenaline standards, and two reagent blanks were run together with the sample.
The noradrenaline standards would give a reading sensitive to about 0.5 /x g./g. of roach
nerve tissue, comparable to those of vertebrate concentrations in the CNS (Udenfriend,
1964).
RESULTS
The Normal Endogenous Activity
Four roach nerve cords were treated with saline alone, and the endogenous activity was
observed during an 8-hour period (Fig. 1 ). The synaptic transmission was generally good
even at the 8th hour. The endogenous activity and synaptic transmission varied from hour
to hour. The patterns of endogenous activity and synaptic transmission were almost identi-
cal although the peaks did not always coincide. The endogenous activity showed a tendency
to decrease during the first hour. This might be due to the decomposition of arginine phos-
phate during dissection, attendant stimulation and injury of the nerve cord (Engel and
Gerard, 1935).
Tobias et al. (1946) demonstrated that roach nerve cords could synthesize ACh. Though
excised, nerves of lobster leg were able to synthesize ACh (Dettbarn and Rosenberg, 1966).
The Effect of Hemicholinium (HC-3)
Hemicholinium (10 ~3 M) blocked synaptic transmission of two roaches (Fig. 2C, D), and
incompletely blocked the synaptic activity of two other roaches (Fig. 2A, B). Both the
endogenous activity and synaptic transmission showed steady decline. Starting after an hour
or two, the amplitudes of the endogenous activity and synaptic response decreased (Fig. 3).
In order to facilitate the depletion of endogenous ACh, the cerci were exposed to puffs
for a period of ten minutes each hour. The endogenous activity of the nerve cord of four
roaches was completely abolished after one to five hours.
Three other roaches were treated first with hemicholinium ( 1 0 "3 M), and then choline
chloride (10-3 M) was applied when both the endogenous activity and synaptic transmission
appeared to reach the lowest level (Figs. 4, 5). Again, the activity pattern showed a decline
in the presence of HC-3 alone. After the choline was applied, the endogenous activity was
increased, and the synaptic transmission was greatly improved.
No previous work has been done on the effect of HC-3 on the electrical activity of insect
nervous system. HC-3 is reported to interfere with the synthesis of ACh by preventing the
270
Chiang
Fig. 1 . Electrical activity in the ventral nerve cord of P. americana treated with insect saline.
Each graph represents the data from a different roach; • spikes/sec., □ spikes/air puff.
SP'K« - SPIKES
Cockroach Nerve Cord
271
Fig. 2. Electrical activity in the ventral nerve cord of P. americana treated with 10-3 M
HC-3. Each graph represents the data from a different roach; • spikes/sec., □ spikes/air puff.
In Fig. 2A, the data from 4 controls treated with saline are also presented; * average spikes/
sec., + average spikes/air puff.
272
Chiang
A
I MMI (Hi » t '■"»
B
m. <fiM * Mu
C
*WiMi ' i b» ' 4*
■UOU
/■■
D
E
F
G
Fig. 3. Electrical activity in the ventral nerve cord of P. americana treated with 10 3 M
hemicholinium. (A) before treatment; (B) 0 hour; (C) 1 hour; (D) 2 hours; (E) 3 hours;
(F) 4 hours; (G) 5 hours. Solid line air puff applied. Film speed 10 cm/sec.
SPIKES
Cockroach Nerve Cord
273
access of choline to choline acetylase (Gardiner, 1961), or by blocking the transport system
by which choline accumulates in the terminals of cholinergic fibers (Macintosh, 1961).
HC-3 inhibited the ACh synthesis in mammalian sympathetic ganglia (Birks and Macintosh,
1961; Macintosh, Birks and Sastry, 1956). HC-3 also inhibited the synthesis of ACh in
minced mouse brain (Gardiner, 1961). Postganglionic sympathetic transmission was blocked
by HC-3 (Burn and Rand, 1960). The block of ACh synthesis by HC-3 could be reversed by
large doses of choline (Birks and Macintosh, 1961 ; Gardiner, 1961).
The degree of inhibition of ACh synthesis by HC-3 depended upon the concentration of
choline present (Gardiner, 1961). The variation in the inhibitory effect on the electrical
activity in the present study could probably be explained on such basis.
</>
*
cO
160-
Fig. 4. Reactivation of electrical activity by 10 3 M choline in the ventral nerve cord of P.
americana treated with 10-3 M HC-3. Each graph represents the data from a different roach;
• spikes/sec., □ spikes/air puff, i choline added. In Fig. 3A the data from 4 controls treated
with saline are also presented; * average spikes/sec., + average spikes/air puff.
274
Chiang
A
»'■ tf
B
4* - kWS
4***
E
+
G
Hf#
+
Fig. 5. Reactivation of electrical activity by choline in the ventral nerve cord of P. americana
treated with 10 ~3 M hemicholinium (HC-3). (A) before treatment; (B) 0 hour; (C) 1 hour;
(D) 2 hours; (E) 3 hours; (F) 4 hours; (G) 6 hours. Solid line air puff applied. Film speed
10 cm/sec.
Cockroach Nerve Cord
275
The Effect of Carbachol (Carbamylcholine)
Carbachol at 10 ~3 M had no observable effect on the endogenous activity or the synaptic
transmission of three roaches (Fig. 6A, B, C). But at 10 ~2 M, carbachol blocked both the
endogenous activity and synaptic transmission in three nerve cords after approximately two
hours (Fig. 6D, E, F). Half an hour after the application of carbachol, the endogenous
activity decreased almost to zero in one nerve cord (Fig. 6E). The spikes of synaptic trans-
mission were barely visible. In the other two nerve cords, there was some indication that
some nerve cells were acted upon. The spikes of these cells were of low amplitudes.
The results were contrary to Roeder’s finding (1948b): carbachol at 10-2 M had no effect
on synaptic transmission in the roach’s 6th abdominal ganglion. Perhaps his period of obser-
vation was different.
Ginsborg and Guerrero (1964) reported that carbachol depolarized the sympathetic gang-
lion cells of the frog, and the amplitude of the spontaneous synaptic potential was de-
pressed five minutes after addition of carbachol. It was concluded that the depolarization
was apparently a direct result of the drug acting on the receptor, and not due to a “drug
induced” release of transmitter from the presynaptic fibers (Ginsborg and Guerrero, 1964).
Carbachol can release ACh only for brief periods before desensitization ensues (McKinstry
and Koelle, 1967). Carbachol itself is hydrolyzed much more slowly than ACh, and is com-
pletely unaffected by AChE (Barlow, 1964; Koelle, 1965).
The Effect of Nicotine
Since nicotine can mimic ACh in cholinergic synapses and junctions (Albert, 1965), it was
used so that the stimulatory and blocking effect of other drugs of the same category could
be compared. As reported by others (Roeder and Roeder, 1939; Welsh and Gordon, 1947),
nicotine (10-3 M) caused an immediate stimulation upon application, and then blocked syn-
aptic and endogenous activity irreversibly. Since the effect was immediate, a graph is not
presented.
The Effect of Dimethylphenylpiperazinium (DMPP)
DMPP produced no observable effect even at 10~2 M (Fig. 7). The electrical activity pat-
tern was similar to that of the saline controls.
No previous work has been done on the effect of DMPP on the electrical activity of in-
sects. DMPP is a selective stimulant of autonomic ganglion cells in vertebrates, although its
blocking effect is less potent than nicotine (Chen and Portman, 1954; Chen, Portman, and
Wickel, 1951; Leach, 1957).
The Effect of Methacholine (Acetyl-j3-methylcholine)
Methacholine at 10 2 M did not produce any observable effect on the electrical activity
of the roach nerve cords within a period of five hours (Fig. 8). Methacholine is equiactive as
ACh in vertebrates at the postganglionic parasympathetic nerves (Albert, 1965; Bebbington
and Brimblecombe, 1965). Geber and Voile (1965) observed that methacholine was even
more potent than ACh as a depolarizing agent in the sympathetic (superior cervical) ganglion
of the cat. Methacholine may also produce ganglionic hyperpolarization by a direct action
on the sympathetic ganglion cells (Voile, 1965).
276
Chiang
Hr. hr.
Fig. 6. Electrical activity in the ventral nerve cord of P. americana treated with carbachol;
A, B, & C 10 "3 M; D, E, & F 10 ~2 M. Each graph represents the data from a different
roach; • spikes/sec., □ spikes/air puff. In Fig. 6A, the data from 4 controls treated with
saline are also presented; * average spikes/sec., + average spikes/air puff.
Cockroach Nerve Cord
277
Fig. 7. Electrical activity in the ventral nerve cord of P. americana treated with 10 2 M
dimethylphenyl piperazinium (DMPP). Each graph represents the data from a different
roach; • spikes/sec., □ spikes/air puff. In Fig. 7A, the data from 4 controls treated with
saline are also presented; * average spikes/sec., + average spikes/air puff.
SPIKES _ SPIKES
278
Chiang
Fig. 8. Electrical activity in the ventral nerve cord of P. americana treated with 10-2 M
methacholine. Each graph represents the data from a different roach; • spikes/sec., □ spikes/
air puff. In Fig. 8A, the data from 4 controls treated with saline are also presented; * average
spikes/sec., + average spikes/air puff.
Cockroach Nerve Cord
279
Since methacholine is less readily hydrolyzed than ACh (Grollman, 1960), the failure of
methacholine to affect synaptic transmission is probably not due to its destruction by
hydrolysis. Roeder (1948b) also showed that methacholine was ineffective in stimulating or
blocking synaptic transmission in the roach.
The Effect of Pilocarpine
Pilocarpine at 10 "3 M showed a definite depressing effect on both the endogenous activ-
ity and synaptic transmission (Fig. 9). The amplitudes of the spikes decreased with time.
The endogenous activity and synaptic transmission were both blocked in one preparation
(Fig. 9D). Some low amplitude spikes appeared after one hour, indicating some cells were
acted upon. These low amplitude spikes resembled those induced by carbachol. Twarog and
Roeder (1957) also observed such effects from nerve cords, the ganglia of which were
desheathed.
Although pilocarpine is chiefly a powerful parasympathomimetic agent, stimulating the
organs innervated by postganglionic fibers, it can also exert ganglionic stimulation (Koelle,
1965; Grollman, 1960).
Pilocarpine is a weak competitive inhibitor of fly-head AChE in vitro, and does not exert
any progressive inhibition (Chadwick, 1 964). The results of the present investigation support
the conclusion of Chadwick (1964) that inhibition of AChE is not involved in the poisoning
of roaches by pilocarpine.
Pilocarpine injected into the heads of praying mantis produced a state of great excitation.
But when injected into the roaches, it produced immobility and apparent paralysis (Roeder
and Roeder, 1939).
The Effect of Acetylcholine
ACh (10 ~3 M) produced no observable effect within a period of five hours. But at 10-2
M ACh blocked both the endogenous activity and synaptic transmission in five out of six
intact preparations (Fig. 10). The stimulatory effect of ACh observed in mammalian ganglia
(Geber and Voile, 1965; Takeshige and Voile, 1962, 1963; Voile, 1965) was not observed
in the roaches. ACh exhibited a progressive depressing effect (Fig. 11). Three eserinized
nerve cords (eserine 10 ~5 M) were washed with saline when synaptic after-discharge occur-
red, and then 10-4 M, 10-3 M, 10~2 M ACh was applied to them immediately. No apparent
stimulation was observed. The endogenous activity and synaptic transmission approached
normal within a period of 30 minutes.
Roeder and Roeder (1939) reported that 10~3 M ACh produced a definite increase in the
level of endogenous activity in isolated nerve cords. But when the nerve cords were in situ,
ACh 10 "2 M had no effect (Twarog and Roeder, 1957). If the ganglia were desheathed and
eserinized, 10 "3 M ACh caused a partial synaptic block, and in concentrations between 3 x
10 "3 M and 5 x 10-3 M, most preparations showed brief after-discharge followed by block;
some preparations were incompletely blocked (Twarog and Roeder, 1957). Yamasaki and
Narahashi (1960) claimed that 10 ~2 M ACh was effective in depolarizing isolated roach
ganglia. They also found that if the ganglia were desheathed and eserinized, ACh was effec-
tive at concentrations as low as 10 ”4 M; but 10 "5 M had little effect. However, Takeshige
and Voile (1964) observed that when small doses of ACh were applied to eserinized cat’s
280
Chiang
Fig. 9. Electrical activity in the ventral nerve cord of P. americana treated with 10 “3 M
pilocarpine. Each graph represents the data from a different roach; • spikes/sec., □ spikes/air
puff. In Fig. 9A, the data from 4 controls treated with saline are also presented; * average
spikes/sec., + average spikes/air puff.
Cockroach Nerve Cord
281
A
Fig. 10. Electrical activity in the ventral nerve cord of P. americana treated with 10 "2 M
acetylcholine. Each graph represents the data from a different roach; • spikes/sec., □ spikes/
air puff. In Fig. 10A, the data from 4 controls treated with saline are also presented;
* average spikes/sec., + average spikes/air puff.
282
Chiang
A
i|4»" »
F
t— i mm* » *
G
Fig. 1 1. Electrical activity in the ventral nerve cord of P. americana treated with 10-3 M
ACh. (A) before treatment; (B) 0 hour; (C) 1 hour; (D) 2 hours; (E) 3 hours; (F) 4 hours;
(G) 5 hours. Solid line air puff applied. Film speed 10 cm/sec.
Cockroach Nerve Cord
283
sympathetic ganglia, hyperpolarization occurred also, in addition to depolarization caused
by a larger dose of ACh (Takeshige and Voile, 1962).
The Effect of Choline
Choline (10 “2 M) blocked synaptic transmission in four out of six preparations (Fig. 12).
Both the endogenous activity and synaptic transmission of one preparation were blocked.
The endogenous activity of the other five preparations was unimpaired. In some prepara-
tions, regular medium-high and low amplitude spikes occurred. In doses three to four times
greater than ACh, choline caused a low amplitude, but sustained, depolarization of the cat’s
sympathetic ganglia (Geber and Voile, 1965). Choline accelerated the rate of failure of post-
ganglionic firing in cat’s sympathetic ganglion (Voile and Koelle, 1961).
The Effect of Eserine
Eserine at 10 “7 M produced no observable effect. But anticholinesterase effects were
observed with 10 “5 M eserine about one hour after treatment; these effects consisted of
periodic bursts of low amplitude spikes and synaptic after-discharge in response to a single
puff, i.e., facilitation. Washing the preparation with saline abolished these effects. These
observations are in agreement with those of Roeder (1948b), and Yamasaki and Narahashi
(1960). The low amplitude spikes increased in frequency and magnitude with time. A syn-
aptic block sometimes developed after the onset of after-discharge caused by a single puff.
Since the effect of eserine is well established, a graph is not presented.
When the in situ ganglion was desheathed, Twarog and Roeder (1957) found that eserine
at 10 ~5 M was effective within five minutes, compared with 10 to 40 minutes in intact but
isolated nerve cords (Yamasaki and Narahashi, 1960).
The Effect of Tetraethylpyrophosphate (TEPP)
TEPP (10 "4 M) exerted a rapid and pronounced effect on the nerve cords. Within a min-
ute of application, repetitive discharge occurred. The duration of the discharge varied from
22 to 60 seconds. A synaptic block and an electrical quiescent period followed the initial
discharge. Four out of six nerve cords did not show any recovery from the completely inac-
tive state when washed with saline. The “endogenous” activity of one preparation reap-
peared for four hours after an initial quiescent period, but air puffs could not elicit a synap-
tic response (Fig. 13A). However, in one preparation, both the “endogenous” activity and
the synaptic transmission persisted, despite the alternation of electrical quiescence and syn-
aptic block (Fig. 13B).
TEPP was reported to block synaptic transmission in nerve cords that were in situ (Col-
houn, 1960; Roeder, 1948b), and in nerve cords with desheathed ganglia (Yamasaki and
Narahashi, 1960). Sternburg et al. (1959) proposed that TEPP caused a neuroactive sub-
stance to be released as a result of synaptic facilitation, and this rather than TEPP was
responsible for blocking the electrical activity. They also suggested that the same substance
participates in neurotransmission within the central nervous system of roaches. The number
of replicable observations was not indicated.
284
Chiang
Fig. 12. Electrical activity in the ventral nerve cord of P. americana treated with 10-2 M
choline. Each graph represents the data from a different roach; • spikes/sec., □ spikes/air
puff. In Fig. 12 A, the data from 4 controls treated with saline are also presented; * average
spikes/sec., + average spikes/air puff.
Cockroach Nerve Cord
285
Fig. 13. Electrical activity in the ventral nerve cord of P. americana treated with 10~4 M
TEPP. Each graph represents the data from a different roach; • spikes/sec., □ spikes/air puff.
In Fig. 13B, the data from 4 controls treated with saline are also presented; * average spikes/
sec., + average spikes/air puff.
The Effect of Pyridine- 2-aldoxime Methiodid (2-PAM) upon TEPP-Treated Nerve Cords
When complete electrical blockade was observed in four nerve cords, two treated with
10"4 M TEPP, two with 10~3 M TEPP, 2-PAM (10-3 M) was applied to them (Figs. 14, 15).
Both the endogenous activity and synaptic transmission reappeared at the first hour, with
the occurrence of facilitation and quiescence. With the exception of one preparation, synap-
tic after-discharge and electrical quiescence stopped at the third hour. The electrical activity
of all four appeared normal at the fourth hour.
When applied alone, 10 "3 M 2-PAM had no apparent effect on the electrical activity of
the nerve cords (Fig. 1 6). In mammals, 2-PAM is a potent neuromuscular blocking agent
(Holmes and Robins, 1955).
Phosphorylated AChE can be reactivated by 2-PAM effectively (Hobbiger, 1963). The
reactivation of phosphorylated AChE by 2-PAM involves the bonding of 2-PAM’s quarter-
nary ammonium group to the anionic site of the phosphorylated AChE, thus allowing the
hydrolysis of the phosphorylated-oxime-enzyme complex to occur, giving AChE and phos-
phorylated-oxime (Loomis, 1956; Wilson, 1960; Wilson and Ginsburg, 1958).
286
Chiang
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UJ
*
£
1/5
2,800-1
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Fig. 1 4. Reactivation of electrical activity by 2-PAM in the ventral nerve cord of P. ameri-
cana treated with TEPP; A & B 10 "3 M TEPP, C & D 10 ~4 M TEPP; 10-3 M. 2-PAM was
added at 0 hour when complete block was observed; • spikes/sec., □ spikes/air puff. In Fig.
14A, the data from 4 controls treated with saline are also presented; * average spikes/sec.,
+ average spikes/air puff.
Cockroach Nerve Cord
287
i >
B
1 m i*m*m >< mm
c
" 4' H
D
E
Fig. 15. Reactivation of electrical activity by 2-PAM in the ventral nerve cord of P. ameri-
cana treated with 1CT4 M TEPP. (A) before the nerve cord was treated with TEPP. Electrical
activity ceased at 0 hour. (B) 1 hour after 10 "3 M 2-PAM added; (C) & (D) 2 hours; (E) 4
hours. Solid line air puff applied. Film speed 10 cm/sec.
288
Chiang
Fig. 16. Electrical activity in the ventral nerve cord of P. americana treated with 10-3 M
2-PAM. Each graph represents the data from a different roach; • spikes/sec., □ spikes/air
puff. In Fig. 16A, the data from 4 controls treated with saline are also presented; * average
spikes/sec., + average spikes/air puff.
Phosphorylated roach AChE was reactivated by 2-PAM in vitro (Colhoun, 1959b; Brady
and Sternburg, 1966). The dose of 2-PAM necessary to reverse neuromuscular block in the
rat nerve-diaphragm was close to the toxic dose for TEPP (Holmes and Robins, 1955), and
the time taken for the reversal of block by 2-PAM was the same whether the anticholines-
terase had been in contact with the preparation for a few minutes or for many hours.
The Effect of Choline upon TEPP-Treated Nerve Cords
After immersion in 10 "3 M TEPP for 30 minutes, three nerve cords were then treated
with 10 "3 M choline (Fig. 17). One preparation remained completely blocked. The endo-
genous activity of another preparation appeared at the first hour, but became blocked again
at the fourth hour. Both the endogenous activity and synaptic transmission of a third prepa-
ration resumed at the third hour, with typical signs of AChE inhibition. But the endogenous
activity stopped again an hour later, with synaptic block to follow two hours later. Choline
is a weak reactivating agent of phosphorylated AChE (Hobbiger, 1963).
Fig. 17. Effect of 10-3 M choline on the electrical activity of the ventral nerve cord of P.
americana treated with 10 ~3 M TEPP. Each graph represents the data from a different
roach; • spikes/sec., □ spikes/air puff; i choline added. In Fig. 17 A, the data from 4 controls
treated with saline are also presented; * average spikes/sec., + average spikes/air puff.
Determination of AChE Activity
Fig. 18 shows the effect of eserine concentration upon the rate of a-naphthylacetate
hydrolysis by the esterases in the roach 6th abdominal ganglion. About 50% of the total
activity was readily inhibited by 1 0 "8 M eserine. The addition of 1 0 "5 M eserine apparently
blocked the AChE completely after 10 minutes of incubation. This is in agreement with the
results of van Asperen (1962) when fly-head AChE was used. Using manometric technique,
Chadwick and Hill (1957) reported that 96% of the roach nerve cord AChE was inhibited by
10-5 M eserine.
The activity of the individual ganglia varied from 1.520 /x M naphthol to 2.96 /x M naph-
thol produced per minute, with a mean of 1 .947 /x M naphthol per minute (Table 1 ). The
mean activity of three pooled homogenates was 2.1 1 1 /x M naphthol produced per minute.
But the difference between the two means is not statistically significant (P < 0.05) by
Student-t test. Yamasaki and Narahashi (1960) observed considerable variation of AChE
activity in roach nerve cords.
After an electrical block became apparent in three roach nerve cords treated with 1 0 “4 M
TEPP, the AChE activity was assayed immediately (Table 2). The mean AChE activity was
11.16% of that of the control. Inactivation of AChE ran parallel with the synaptic after-
discharge (Yamasaki and Narahashi, 1960).
290
Chiang
OPTICAL DENSITY
Fig. 18. Inhibition of roach ganglion AChE activity by eserine. Temperature 40°C. Substrate
3 x 10-4 M a-naphtyl acetate. Homogenate concentration % ganglion per tube. Incubation
time 10 minutes.
The Effect of Adrenergic Drugs
Phenoxybenzamine (dibenzyline) blocks the a-receptors of the adrenergic nerves effec-
tively and persistently (Nickerson, 1965). When phenoxybenzamine (10 ~3 M) was applied
to the nerve cords of roaches, a blocking effect was observed (Fig. 19A, B, C). Washing with
saline did not eliminate the blocking effect. Being a /3-haloalkylamine that can cause tissue
damage as nitrogen mustard (Nickerson, 1949), phenoxybenzamine may not be specific in
its action. Phenoxybenzamine increased the spontaneous efflux of H3 -noradrenaline by 50%
in cat’s colon (Costa et al., 1966).
Monoamine oxidase (MAO) is the enzyme chiefly responsible for the physiological inac-
tivation of 5-hydroxytryptamine (5-HT), and endogenous catecholamines (Kopine, 1964;
Page, 1958; Shore et al., 1957). Tranylcypromine, a MAO inhibitor, blocked the endogenous
activity and the synaptic transmission of the nerve cords of roaches at 10-3 M (Fig. 19E, F,
G). MAO inhibitors can produce an irreversible inactivation of MAO by forming stable
complexes with the enzyme, causing an elevation of biogenic amines (Javik, 1965; Pletscher,
1966). But MAO inhibitors can also inhibit other enzymes as well (Javik, 1965).
Table 1: AChE activity in 6th abdominal ganglion of P. americana.
Cockroach Nerve Cord
291
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phate buffer
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AChE activity = EA0.D.cnn less eserine - IAO.D. with eserine
600 600
One unit activity = lyM. of naphthol produced/min.
Table 2: AChE activity in 6th Abdominal ganglion of
292
Chiang
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(pH 7) containing 0.3%
ACh
Control = Mean AChE activity of individual ganglia in Table
Cockroach Nerve Cord
293
D
120-
160
80
N.
Fig. 19. Electrical activity in the ventral nerve cord of P. americana treated with 10~3 M
phenoxybenzamine (A, B, C), and 10"3 M tranylcypromine (D, E, F). Each graph represents
the data from a different roach; • spikes/sec., □ spikes/air puff. In Fig. 18A, the data from
4 controls treated with saline are also presented; * average spikes/sec., + average spikes/air
puff.
294
Chiang
Spec tro flu orometric Determination of Noradrenaline
There was no indication of the presence of noradrenaline in the sample (Fig. 20). Owing
to the large number of roaches required to yield sufficient amount of nerve cords, the exper-
iment was run only once. Unger (1957) believed that adrenaline, noradrenaline, and hista-
mine are not among a number of cardiac accelerators obtained from the abdominal nerve
cords, corpora allata, corpora cardiaca and haemolymph.
Fig. 20. Fluorescence spectra and light scatter of 0.25 /a g of standard noradrenaline, a
sample of extraction of roach abdominal nerve cords, and a reagent blank assayed by the
iodine method. Excitation was at 385 m p.
DISCUSSION
The physiological role of ACh in nerve tissues has been reviewed extensively by Koelle
(1963). In the synaptic and neuroeffector sites, there are three possible roles that can be
attributed to ACh. (1) On arrival of an impulse in an axon terminal, ACh is liberated and dif-
fuses across the synaptic cleft to combine with a receptor to bring about a localized depolar-
izartion, the postsynaptic potential (PSP). The latter in turn initiates electronically a nerve
action potention (AP) in the second neuron. (2) ACh acts first at an axon terminal from
which it is liberated to cause the release of additional quanta of ACh, which produce the
Cockroach Nerve Cord
295
PSP. (3) In non-cholinergic neurons, AP liberates ACh from presynaptic terminals, which
acts at the same terminals to effect release of another synaptic transmitter. The latter pro-
duces PSP, which initiates AP.
The demonstration that hemicholinium blocked both the endogenous activity and the
synaptic transmission either entirely or partially (Fig. 4), and the restoration of activity by
choline, indicates that the neural activity of the roach nerve cord is dependent upon ACh.
Drugs that act at cholinergic sites are generally divided into two categories: nicotinic and
muscarinic (Albert, 1965; Goth, 1966). The muscarinic drugs act mainly on the peripheral
autonomic nervous system, stimulating the post-ganglionic parasympathetic receptors of
organs such as smooth muscles, cardiac muscle, endocrine glands, etc., without having an
effect upon ganglionic transmission or skeletal muscles (Goth, 1966). The muscarinic recep-
tors are atropine sensitive (Goth, 1966). The nicotinic drugs can stimulate autonomic gang-
lion and end-plates of skeletal muscles (Goth, 1966). The nicotinic receptors are divided
into the ganglionic receptors which are hexamethonium sensitive, and the skeletal muscle
receptors which are sensitive to curare (Goth, 1 966). Nicotine has effects on both the gang-
lionic and neuromuscular transmission (Albert, 1965; Goth, 1966). However, these are only
working hypotheses for the mammalian peripheral nervous system. The terms “nicotinic”
and “muscarinic” are not applicable to the central nervous system (Curtis, Ryall and Wat-
kins, 1965; Feldberg, 1950).
Khromov-Borisov and Michelson (1966) pointed out that the invertebrate muscles are
mainly nicotinic. This invites the speculation that the roach cereal synapse does not have a
muscarinic receptor. The failure of atropine and methacholine to produce any blocking or
killing effect (Roeder, 1948b; Tobias et al., 1946) supports such a hypothesis.
Ambache (1955) pointed out the limitations of using atropine as a pharmacological cri-
terion for cholinergic nerves: (1) atropine might be destroyed by an esterase; (2) true atro-
pine resistance in cholinergic systems might be due to “proximity”, (3) secondary formation
of atropine-resistant pharmacological agents.
The failure of DMPP, a nicotinic drug, to produce any observable effects points to the
uniqueness of the roach cereal ganglion.
It is of interest to compare the in vivo effects of injecting carbachol, ACh, and methacho-
line into roaches with the results of the present investigation. The toxic dose of carbachol
for the roaches was 0.5 to 0.1 g./kg., and 7 to 10 g./kg. for the ACh; methacholine even at
20 g./kg. produced no killing effect (Tobias et al., 1 946).
Treherne (1966) postulated a “biochemical barrier” theory, which is partly based on the
histochemical evidence that AChE is situated “strategically” on the glial membranes border-
ing extracellular channels. Despite the rapid influx of ACh, the concentration of ACh within
the roach ganglion was reduced to 8.1 x 10"5 M when the nerve cord was bathed with a
solution of 10 "2 M ACh (Treherne and Smith, 1965b). The relative inability of externally
applied ACh to affect insect nerves would thus seem to result from the presence of this
“biochemical barrier” rather than to the “ion barrier” (Treherne, 1966).
Although choline could block synaptic transmission, the overall effect of the ACh in the
present investigation could not be explained entirely by the hydrolysis of ACh to choline
and acetate. In addition to its depolarizing property, it is possible that ACh also hyperpolar-
296
Chiang
izes the nerve cells in the roach cereal ganglion. When the nerve cords were isolated, or de-
sheathed, ACh was allowed to reach a cholinoreceptive site to cause depolarization. This
would be completely in accord with the finding that there are multiple cholinoreceptive
sites within the mammalian ganglia, and that ACh can be excitatory or inhibitory depending
on the receptor (Eccles and Libet, 1961; Geber and Voile, 1965; Koelle, 1965; Takeshige
and Voile, 1963, 1964). Since the ganglionic potentials represent an algebraic summation of
simultaneous processes, it is conceivable that the temporally related hyperpolarization and
postganglionic firing resulted from monitoring certain populations of cells and fibers (Take-
shige and Voile, 1 964). Thus, while the ganglion as a whole appeared to be in a hyperpolar-
ized state, the postganglionic fibers monitored might have been partially depolarized (Take-
shige and Voile, 1964). Unfortunately, this possibility could not be tested with the present
experimental technique. The results of carbachol and pilocarpine could also be explained by
this postulation. It is also possible that ganglionic hyperpolarization caused by ACh was
mediated by means of an inhibitory transmitter liberated within a ganglion (Voile, 1965).
Simultaneous presynaptic inhibition could also occur (McLennan, 1963).
It was recently found that ACh acts both as an excitatory and inhibitory transmitter in
the same abdominal ganglion of two species of marine mollusc of the genus Aplysia (Kandel
and Frazier, 1967; Tauc and Gershenfeld, 1962). Two types of ganglion cells have been
demonstrated by the actions of ACh. The response of one group of cells to ACh was charac-
terized by depolarization and acceleration of the rate of firing. Conversely, ACh produced
hyperpolarization and the blockade in the second group.
The possible role ACh may play in the sodium pump mechanism (Hokin, Hokin and
Shelp, 1960) should not be overlooked. A given drug, such as ACh itself, may produce either
cholinomimetic or cholinergic blocking action at certain sites, depending upon the dose,
rate of combination with the receptors, and other factors (Koelle, 1962; Paton, 1961). Also
of equal importance is the role the inhibitory system may play when the insect CNS is either
intact or semi-intact (Milburn, Weiant, and Roeder, 1960; Roeder, 1962).
Koelle (1963) surmised that the earliest function of ACh and its associated enzymes in
primitive organisms was probably the modification of the passage of various substances
across the cell membranes. With the subsequent development of different types of structural
complexity of cellular membranes in accordance with their various functions, the ACh-
AChE system itself has been achieved in nervous tissues, where its components and function
are concentrated predominantly at synapses and junctional sites. At such regions, the ACh
may serve both as the transmitter and as the agent for the liberation of other chemical
mediators.
Catecholamines have been demonstrated by fluorescence microscopy in the autonomic
ganglia of mammals (Jacobowitz and Koelle, 1963; Hamberger, Norberg and Sjoqvist, 1965;
Hamberger, Norberg and Unguestedt, 1965; Norberg and Sjoqvist, 1966; Owen and Falck,
1965). Adrenergic sensory neurons were demonstrated in two genera of molluscs (Dahl et
al., 1963). Noradrenaline and adrenaline have both inhibitory and facilitatory actions on
sympathetic ganglia of the cat (De Groat and Voile, 1966a, b). Eccles and Libet (1961)
proposed that ACh can release an adrenergic substance through a chromaffin cell to produce
an action potential. It was postulated that ACh released from sympathetic fibers could
Cockroach Nerve Cord
297
liberate noradrenaline from a peripheral store, in the same way as injected ACh (Burn, 1966;
Burn and Rand, 1965; Chang and Rand, 1960).
The possibility that catecholamines may participate in roach synaptic transmission was
suggested by Twarog and Roeder (1957), who recorded asynchronous bursts of low voltage
spikes following application of adrenaline and noradrenaline to desheathed roach ganglion.
After-discharge and synaptic blocking were observed at concentrations between 10-3 M and
10~2 M. But when applied to intact ganglion, noradrenaline and adrenaline failed to show
any stimulatory or blocking effect (Colhoun, 1959a). Application of 5 x 10-5 M dopamine
to the roach cereal ganglion induced bursts of activity which propagated along the nerve
cord; however, this substance did not have any effect on the synaptic transmission (Gahery
and Boistel, 1965). Since not all the regions in the roach cereal ganglion, as shown by elec-
tron micrograph, are associated with membrane-bound cholinesterase activity (Smith and
Treherne, 1965), such non-cholinergic synapses could be regarded as possible candidates for
transmission mechanisms involving catecholamines (Treherne, 1966).
Injection of adrenaline into roaches paralyzed them temporarily, but injection of nora-
drenaline produced no recognizable response (Barton Brown et al., 1961). All adrenergic
neuron blocking agents examined in detail show a variety of actions at cholinergic sites
(Boura and Green, 1965).
Colhoun (1963a) showed the presence of 5-HT in the nerve cords of roaches, though the
amount is extremely low in comparison with ACh. Grollman (1960) believed that 5-HT is a
primitive synaptic transmitter. It is a chemical transmitter in some invertebrates and verte-
brates (Brodie and Shore, 1957; Owen and Falck, 1965; Welsh, 1957), and can potentiate
and buffer ganglionic transmission (Page, 1958). If some of the noncholinergic regions are
occupied by 5-HT, then the data for the adrenergic drugs in the present investigation be-
come interpretable. But alpha adrenergic blocking agents can also react with receptors for
histamine, 5-HT and ACh (Nickerson, 1965).
Catecholamines and adrenaline-like substances have been extracted from insects (Camer-
on, 1953; Gregerman and Wald, 1952; Ostlunde, 1954; von Euler, 1961). The neurosecre-
tion from corpus cardiacum of P. americana and Rhodinus sp. is related to adrenaline (Bar-
ton Browne et al., 1961; Wigglesworth, 1954). The present investigation conforms to the
belief of Unger (1957) that noradrenaline is not present in the nerve cord of roaches. The
exact nature of the catecholamines present in the roach brain (Frontali, 1968) has yet to be
determined chemically.
Four possible functions have been proposed for the AChE in cholinergic nervous systems
(Koelle, 1963; Voile and Koelle, 1961): (1) temporal or spatial limitation of the transmitter
action of ACh at the postsynaptic site; (2) rapid hydrolysis of ACh to provide an immediate
source of choline for uptake by the presynaptic terminals and synthesis to ACh; (3) protec-
tion of the presynaptic terminals against reactivation by self-liberated ACh; (4) prevention
of accumulation of activating concentrations of ACh during the resting stage.
Confirmation of the first and fourth proposal can be obtained by the results of antichol-
inesterases (Figs. 13, 14; Tables 1, 2), only on the ground that ACh is a transmitter that
diffuses across the synaptic cleft to act on a postsynaptic receptor site in the roach cereal
ganglion. The synaptic after-discharge, facilitation and electrical quiescence were probably
298
Chiang
due to the accumulation of endogenous ACh on the postsynaptic sites. It is also possible
that ACh caused another transmitter substance to be released and act on the postsynaptic
sites, and since the ACh was unable to be hydrolyzed, it blocked all the sites that would
release the stimulatory transmitter.
No evidence was obtained to support the second proposal. When choline was applied to
the nerve cords with or without AChE-inactivation by TEPP, it either could not restore the
electrical activity to its normal state or it only accelerated the rate of failure of postgang-
lionic firing. Nothing can be said about the third proposal until more is known about the
synapses in insects.
The present investigation brought out the effectiveness of the “biochemical barrier”.
Indeed, such a barrier may be highly advantageous for the insects, because of their open
circulatory system. Ginetsinskii (1947) concluded that in the course of evolution, reduction
occurs in the cholinoreceptive zone, and in the number of drugs to which the cholinore-
ceptive zone is sensitive, i.e., an increase in the precision and specificity of cholinoreception.
Such may be the case for the insects. The “biochemical barrier” and the cholinoreceptive
specificity may explain the well protected synapses in insect ganglion.
The experiments with 2-PAM (Fig. 15) point to the significance of AChE in synaptic
transmission, although 2-PAM was also shown to reactivate axonal transmission (Dettbarn,
Rosenberg and Nachmansohn, 1964). The concentration of AChE in a given neuron reflects
the extent of the participation of ACh in the synaptic transmission (Koelle, 1962). Ehren-
preis (1967) pointed out the possible function of AChE as part of the receptor for ACh and
other cholinergic compounds.
The slow in vivo inhibition of AChE by carbamates indicates the effective concentrations
of ACh at the active sites of enzymes are normally very low (Winteringham, 1966). Stimu-
lated central activity may, accordingly, result in relatively large increases in effective sub-
strate concentrations at the enzyme sites, but this would not be reflected in detectable
changes in total ACh content of the central nervous system, and may only involve a small
fraction of the total ACh (Winteringham, 1966).
Colhoun (1959a), and Smallman and Fisher (1958) observed an increase of ACh content
in roach nerve cords treated with TEPP. Treating the roach with DDT also resulted in a
marked accumulation of ACh although the AChE was not inhibited (Colhoun, 1959a, b;
Tobias et al., 1946). But the elevation of ACh content in the nerve cords treated with TEPP
could be resolved into 2 peaks, an initial peak occurring at the first half hour, and a second
peak at 24 hours (Colhoun, 1959a). The initial peak was absent in the DDT-treated nerve
cords. It was postulated that the ACh increases in late TEPP and DDT poisoning were the
result of ACh synthesis in a form not available to AChE (Colhoun, 1959a), and that DDT
might accelerate the rate of ACh synthesis (Lewis, 1953).
A neuroactive substance was released into the blood of the roach during the course of
DDT poisoning (Colhoun, 1959b; Sternburg et al., 1959). It was suggested that the source
of the toxicant was apparently the CNS itself during periods of great nervous activity,
whether initiated by electrical stimulation or by constant bombardment of sensory-central
synapses due to excessive afferent impulses generated in the sensory nerves by direct action
of DDT (Sternburg et al., 1959). It seems possible that ACh itself could act as a releaser of
the neuroactive substance.
Cockroach Nerve Cord
299
Spontaneous recovery of AChE activity in organophosphate-treated insects has been
demonstrated, but the recovery was a result of AChE synthesis rather than a reversal of
inhibited AChE activity (Brady and Sternburg, 1966).
The considerable variations of AChE concentration in the roach ganglion (Table 1) may
explain the occasional ineffectiveness of ACh and some other related drugs at the same
dosage. But that the electrical activity of one roach nerve cord could continue in the pres-
ence of high concentration of TEPP was perplexing, unless there is an enzyme in the nerve
cord that can hydrolyze TEPP. It was demonstrated that diisopropyl fluorophosphate (DEP)
was hydrolyzed readily by a phosphatase to diisopropyl phosphoric acid inside a squid axon
(Hoskin, Rosenberg and Brazin, 1966).
Since the whole field of insect neuropharmacology is “conditioned” by rather well docu-
mented experiments with mammals (E. H. Colhoun, personal communication), there are
limitations in interpreting these data.
ACKNOWLEDGMENTS
I am greatly indebted to R. H. Gooding for advice and supervision, and to B. Hocking for
his encouragement and suggestions. I am grateful to F. R. Calaresu, and C. Heath, Depart-
ment of Physiology; and to W. G. Evans, Department of Entomology, for their advice. I
sincerely thank C. W. Nash, Department of Pharmacology, for his advice and facilities for
the spectrofluorometric work; and R. J. Reiffenstein, Department of Pharmacology, for his
advice and gift of some drugs. Thanks are also due to E. H. Colhoun, Department of Pharma-
cology, University of Western Ontario, for his helpful criticism of the manuscript, G. Chen,
Parke, Davis & Company, for the gift of DMPP, and Cyanamid of Canada for partial support
of the research.
REFERENCES
Albert, A. 1965. Selective toxicity. Methuen, London. 394 p.
Ambache, N. 1955. The use and limitation of atropine for pharmacological studies on
autonomic effectors. Pharmacol. Rev. 7: 467-500.
Barlow, R. B. 1964. Introduction to chemical pharmacology. Methuen, London. 455 p.
Barton Brown, L., L. F. Dobson, E. S. Hodgson and J. K. Kiraly. 1961. Adrenergic proper-
ties of the cockroach corpus cardiacum. Gen. comp. Endocrinol. 1 : 232-236.
Bebbington, A. and R. W. Brimblecombe. 1965. Muscarinic receptors in the peripheral and
central nervous systems, p. 143-172. In N. J. Harper and A. B. Simmonds (eds.). Advances
in drug research, vol. 2. Academic Press, New York.
Birks, R. and F. C. Macintosh. 1961. Acetylcholine metabolism of a sympathetic ganglion.
Can. J. Biochem. Physiol. 39: 787-827.
Bisset, G. W., J. F. D. Frazer, M. Rothschild, and M. Schachter. 1960. A pharmacologically
active choline ester and other substances in the garden tiger moth, Arctia caja (L.). Proc-
R. Soc.,B. 152:255-262.
300
Chiang
Boura, A. L. A. and A. F. Green. 1965. Adrenergic neurone blocking agent. Ann. Rev.
Pharmacol. 5: 183-212.
Brady, U. E. and J. Sternburg. 1966. Recovery of the cholinesterase activity in organo-
phosphate treated insects. J. Ins. Physiol. 12: 1171-1185.
Brodie, B. B. and P. A. Shore. 1957. A concept for a role of serotonin and norepinephrine
as chemical mediators in the brain. Ann. N. Y. Acad. Sci. 66: 631-642.
Burn, J. H. 1966. Adrenergic transmission. Pharmacol. Rev. 18: 459-470.
Burn, J. H. and M. J. Rand. 1960. Transmission failure in sympathetic nerves produced by
hemicholinium. Brit. J. Pharmacol. 15: 588-600.
Burn, J. H. and M. J. Rand. 1965. Acetylcholine in adrenergic transmission. Ann. Rev.
Pharmacol. 5: 163-182.
Cameron, M. C. 1953. Secretion of an orthodiphenol in the corpus cardiacum of the insect.
Nature, Lond. 172: 349-350.
Chadwick, L. E. 1963. Actions on insects and other invertebrates, p. 741-748. In G. B.
Koelle (ed.). Cholinesterases and anticholinesterases. Springer-Verlag, Berlin.
Chadwick. L. E. 1964. Inhibition of fly-head cholinesterase in vitro by pilocarpine and
atropine. J. ins. Physiol. 10: 573-585.
Chadwick, L. E. and D. L. Hill. 1957. Inhibition of cholinesterase by di-isopropyl fluoro-
phosphate, physostigmine and hexamethyl tetraphosphate in the roach. J. Neurophysiol.
10: 235-246.
Chang, S. C. and C. W. Kearns. 1961. The occurrence of acetylcholine in the nervous tissue
of American cockroaches. Ann. ent. Soc. Amer. 54: 711-715.
Chang, V. and M. J. Rand. 1960. Sympathetic postganglionic cholinergic fibers. Brit. J.
Pharmacol. 15: 56-66.
Chen, G., and R. Portman. 1954. Effect of 1, 1 -dimethyl-4-phenyl piperazinium iodide on
peristaltic reflexes of isolated guinea pig ileum. Proc. soc. exp. Biol. N. Y. 85: 245-248.
Chen, G., R. Portman and A. Wickel. 1951. Pharmacology of 1, l-dimethyl-4-phenyl pipera-
zinium iodide, a ganglionic stimulating agent. J. Pharmacol. 103: 330-336.
Colhoun, E. H. 1958a. Physical release of acetylcholine from the thoracic nerve cord of
Periplaneta americana L. Nature, Lond. 181 : 490.
Colhoun, E. H. 1958b. Acetylcholine in Periplaneta americana L. II. Acetylcholine and
nervous activity. J. Ins. Physiol. 2: 1 17-127.
Colhoun, E. H. 1959a. Acetylcholine in Periplaneta americana L. III. Acetylcholine in
roaches treated with tetraethyl pyrophosphate and 2, 2-bis(p-chlorophenyl)-l , 1, 1-tri-
chloroethane. Can. J. Biochem. 37: 160-272.
Colhoun, E. H. 1959b. Physiological events in organophosphorus poisoning. Can. J. Bio-
chem. 37: 1127-1134.
Colhoun, E. H. 1960. Acetylcholine in Periplaneta americana L. IV. The significance of
esterase inhibition in intoxication, acetylcholine levels, and nervous condition. Can. J.
Biochem. 38: 1363-1367.
Colhoun, E. H. 1963a. Synthesis of 5-hydroxytryptamine in the American cockroach.
Experientia 19: 9-10
Cockroach Nerve Cord
301
Colhoun, E. H. 1963b. The physiological significance of acetylcholine in insects and obser-
vations upon other pharmacological substances, p. 1-46. In J. W. L. Beament, J. E.
Treherne, and V. B. Wigglesworth (eds.). Advances in insect physiology, vol. 1. Academic
Press, New York.
Colhoun, E. H. and E. Y. Spencer. 1959. Acetylcholine effects of gamma-carbomethoxy-
propyltrimethyl-ammonium bromide. Science 130: 504-505.
Costa, E., D. J. Boullin, W. Hammer, W. Vogel, and W. B. Brodie. 1966. Interactions of
drugs with adrenergic neurons. J. Pharmacol. 18: 577-597.
Curtis, D. R., R. W. Ryall and J. C. Watkins. 1955. Cholinergic transmission in the mamma-
lian central nervous system, p. 137-145. In G. B. Koelle, W. W. Douglas, A. Carlsson and
V. Trcka (eds.). Pharmacology of cholinergic and adrenergic transmission. Macmillan,
New York.
Dahl, E., B. Falck, C. von Mecklanburg, and H. Myhrberg. 1963. Adrenergic sensory neurons
in invertebrates. Gen. comp. Endocrinol. 3: 693.
Dauterman, W. C., A. Talens and K. van Asperen. 1962. Partial purification and properties
of fly-head cholinesterase. J. Ins. Physiol. 8: 1-14.
De Groat, W. C. and R. L. Voile. 1966a. The actions of the catecholamines on transmission
in the superior cervical ganglion of the rat. J. Pharmacol. 154: 1-13.
De Groat, W. C. and R. L. Voile. 1966b. Interactions between the catecholamines and gang-
lionic stimulating agents in sympathetic ganglia. J. Pharmacol. 154: 200-215.
Dettbarn, W. and P. Rosenberg. 1966. Effect of ions on the efflux of acetylcholine from
peripheral nerve. J. gen. Physiol. 50: 447-460.
Dettbarn, W., P. Rosenberg and D. Nachmansohn. 1964. Restoration by a specific chemical
reaction of “irreversibly” blocked axonal electrical activity. Life Sci. 3: 55-60.
Eccles, R. M. and B. Libet, 1961. Origin and blockade of synaptic responses of curarized
sympathetic ganglia. J. Physiol. 157: 487-503.
Engel, L. G. and R. W. Gerard. 1935. The phosphorus metabolism of invertebrate nerve. J.
biol. Chem. 112: 370-392.
Ehrenpreis, S. 1967. Molecular aspects of cholinergic mechanisms, p. 1-78. In A. Burger
(ed.). Drugs affecting the peripheral nervous system. Marcel Dekker, New York.
Euler, U. S. von. 1961. Occurrence of catecholamines in Acrania and invertebrates. Nature,
Lond. 190: 170.
Feldberg, W. 1950. The role of acetylcholine in the central nervous system. Brit. med. Bull.
6: 312-321.
Frontali, N. 1968. Histochemical localization of catecholamines in the brain of normal and
drug-treated cockroaches. J. Ins. Physiol. 14: 881-886
Gahery, Y. and J. Boistel. 1965. Study of some pharmacological substances which modify
the electrical activity of the sixth abdominal ganglion of the cockroach, Periplaneta
americana, p. 73-78. In J. E. Treherne and J. W. L. Beament (eds.). Insect central nervous
system. Academic Press, New York.
Gardiner, J. E. 1961. The inhibition of acetylcholine synthesis in brain by a hemicholinium.
Biochem. J. 81: 297-303.
Geber, G. L. and R. L. Voile, 1965. Ganglionic stimulating properties of aliphatic esters
of choline and thiocholine. J. Pharmacol. 150: 67-74.
302
Chiang
Ginetsinskii, A. G. 1947. Quoted by Magazanik et al. (1965).
Ginsborg, B. L. and S. Guerrero. 1964. On the action of depolarizing drugs on sympathetic
ganglion cells of the frog. J. Physiol. 172: 189-206.
Gomori, G. 1953. Human esterases. J. Lab. clin. Med. 42: 445-453.
Goth, A. 1966. Medical pharmacology: principles and concepts. Mosby, Saint Louis. 668 p.
Gregerman, R. I. and G. Wald. 1952. The alleged occurrence of adrenaline in the mealworm.
J. gen. Physiol. 35: 489-493.
Grollman, A. 1960. Pharmacology and therapeutics. Lea and Febiger, Philadelphia. 1079 p.
Hamberger, B., K.-A. Norberg and F. Sjoqvist. 1965. Correlated studies of monoamines and
acetylcholinesterase in sympathetic ganglia, illustrating the distribution of adrenergic and
cholinergic neurons, p. 41-54. In G. B. Koelle, W. W. Douglas, A. Carlsson, and V. Trcka
(eds.). Pharmacology of cholinergic and adrenergic transmission. Macmillan, New York.
Hamberger, B., K.-A. Norberg and U. Ungestedt. 1965. Adrenergic synaptic terminals in
autonomic ganglia. Acta Physiol. Scand. 64: 285-286.
Hess, A. 1958. Experimental anatomical studies of pathways in the severed central nerve
cord of the cockroach. J. Morph. 103: 479-507.
Hobbiger, F. 1963. Reactivation of phosphorylated acetylcholinesterase, p. 921-988. In G.
B. Koelle (ed.). Cholinesterases and anticholinesterase agents. Springer- Ver lag, Berlin.
Hokin, M. R., L. E. Hokin and W. D. Shelp. 1960. The effects of acetylcholine on the turn-
over of phosphatidic acid and phosphoinositide in sympathetic ganglia and in various
parts of the central nervous system in vitro. J. Gen. Physiol. 44: 217-226.
Holmes, R. and E. L. Robins. 1955. The reversal by oximes of neuromuscular block pro-
duced by anticholinesterase. Brit. J. Pharmacol. 10: 490-495.
Hoskin, F. C. G., P. Rosenberg and M. Brazin. 1966. Re-examination of the effect of DFP
on electrical and cholinesterase activity of squid giant axon. Proc. Nat. Acad. Sci. Wash.
55: 1231-1235.
Hoyle, G. 1952. High blood potassium in insects in relation to nerve conduction. Nature,
Lond. 169: 281-282.
Hoyle, G. 1953. Potassium ions and insect nerve muscle. J. exp. Biol. 30: 121-135.
Iyatomi, K. and K. Kanehisa. 1958. Localization of cholinesterase in the American cock-
roach. Jap. J. appl. Ent. 2: 1-10.
Jacobowitz, D. and G. B. Koelle. 1963. Demonstration of both acetylcholinesterase (AChE)
and catecholamines in same nerve trunk. The Pharmacologist 5: 270.
Javik, M. E. 1965. Drugs used in the treatment of psychiatric disorders, p. 159-214. In L.
Goodman and A. Gilman (eds.). The pharmacological basis of therapeutics. Macmillan,
New York.
Kandel, E. R. and W. T. Frazier. 1967. Opposite synaptic actions mediated by different
branches of an identifiable interneuron in Aplysia. Science 155: 346-349.
Khromov-Borisov, N. V. and M. J. Michelson. 1966. The mutual dispositions of locomotor
muscles and the changes in their disposition in the course of evolution. Pharmacol. Rev.
18: 1051-1090.
Koelle, G. B. 1962. A new general concept of the neurohumoral functions of ACh and
AChE. J. Pharm. Pharmacol. 14: 65-90.
Cockroach Nerve Cord
303
Koelle, G. B. 1963. Cytological distribution and physiological functions of cholinesterases,
p. 187-298. In G. B. Koelle (ed.). Cholinesterase and anticholinesterases. Springer- Verlag,
Berlin.
Koelle, G. B. 1965. Neurohumoral transmission and the autonomic nervous system; anti-
cholinesterase agents; parasympathomimetic agents, p. 399-476. In L. Goodman and A.
Gilman (eds.). The pharmacological basis of therapeutics. Macmillan, New York.
Kopin, I. J. 1964. Storage and metabolism of catecholamines: the role of monoamine
oxidase. Pharmacol. Rev. 16: 179-191.
Leach, G. D. H. 1957. Ganglionic blocking action of dimethylphenyl piperazinium (DMPP).
J. Pharm. Pharmacol. 9: 747-751.
Lewis, S. E. 1953. Acetylcholine in blowflies. Nature, Lond. 172: 1004-1005.
Loomis, J. 1956. The effect of an aldoxime on acute serin poisoning. J. Pharmacol. 18:
123-128.
Macintosh, F. C. 1961. Effect of HC-3 on acetylcholine turnover. Fed. Proc. 20: 562-568.
Macintosh, F. C., R. I. Birks and P. B. Sastry. 1956. Pharmacological inhibition of acetyl-
choline synthesis. Nature, Lond. 178: 1181.
Magazanik, L. G., N. R. Fruentov, E. K. Roshkova, R. S. Rybolovlev and M. Ya. Mikhel’son.
1965. On the evolution of cholinoreceptive sites of locomotor muscle, p. 1 13-127. In G.
B. Koelle, W. W. Douglas, A. Carlsson and V. Trcka (eds.). Pharmacology of cholinergic
and adrenergic transmission. Macmillan, New York.
McKinstry, D. N. and G. B. Koelle. 1967. Acetylcholine release from the cat superior cervi-
cal ganglion by carbachol. J. Pharmacol. 157: 319-327.
McLennan, H. 1963. Synaptic transmission. W. B. Saunders, Philadelphia. 134 p.
Mikalonis, S. J. and R. H. Brown. 1941. Acetylcholine and cholinesterase in the insect
central nervous system. J. celLcomp. Physiol. 18: 401-402.
Milburn, N., E. A. Weiant, and K. D. Roeder. 1960. The release of efferent activity in the
roach, Perplaneta americana, by extracts of the corpus cardiacum. Biol. Bull. 118: 111-
119.
Nickerson, M. 1949. The pharmacology of adrenergic blockade. Pharmacol. Rev. 1 : 27-101.
Nickerson, M. 1965. Adrenergic receptor mechanism, p. 303-315. In G. B. Koelle, W. W.
Douglas, A. Carlsson and V. Trcka (eds.). Pharmacology of cholinergic and adrenergic
transmission. Macmillan, New York.
Norberg, K.-A. and F. Sjoqvist. 1966. New possibilities for adrenergic modulation of gang-
lionic transmission. Pharmacol. Rev. 18: 743-751.
O’Brien, R. D. 1957. Esterases in the semi-intact cockroach. Ann. ent. Soc. Amer. 50:
223-229.
O’Brien, R. D. 1959a. Comparative toxicology of some organophosphorus compounds in
insects and mammals. Can. J. Biochem. Physiol. 37: 1113-1 122.
O’Brien, R. D. 1959b. Effect of ionization upon penetration of organophosphates to the
nerve cord of the cockroach. J. econ. Ent. 52: 812-816.
Ostlunde, E. 1954. The distribution of catecholamines in lower animals and their effects on
the heart. Acta physiol. Scand. 31 : suppl. 112: 1-65.
Owen, C. and B. Falck. 1965. Localization of neuronal monoamines at the cellular level, p.
165-183. In G. B. Koelle, W. W. Douglas, A. Carlsson and V. Trcka (eds.). Pharmacology
of cholinergic and adrenergic transmission. Macmillan, New York.
Page, I. H. 1958. Serotonin (5-hydroxy tryptamine), the last four years. Physiol. Rev. 58:
277-335.
304
Chiang
Paton, W. D. M. 1962. A theory of drug action based on the rate of drug-receptor combina-
tion. Proc. roy. Soc., B. 154: 21-69.
Pletscher, A. 1966. Monoamine oxidase inhibitors, Pharmacol. Rev. 18: 121-129.
Pringle, J. W. S. 1938. Proprioception in insects. I. A new type of mechanical receptor from
the palps of the cockroach. J. exp. Biol. 15: 101-113.
Prosser, C. L. 1940. Action potentials in the nervous system of the crayfish. Effects of drugs
and salts upon synaptic transmission. J. cell. comp. Physiol. 16: 25-38.
Pumphrey, R. J. and A. F. Rawdon-Smith. 1937. Synaptic transmission of nerve impulses
through the last abdominal ganglion of the cockroach. Proc. R. Soc., B. 122: 106-118.
Roeder, K. D. 1948a. Organization of the ascending giant fiber system in the cockroach
{Periplaneta americana). J. exp. Zool. 108: 243-261.
Roeder, K. D. 1948b. The effect of anticholinesterases and related substances on nervous
activity in the roach. Johns Hopk. Hosp. Bull. 83: 587-603.
Roeder, K. D. 1953. Electric activity in nerves and ganglia, p. 423-462. In K. D. Roeder
(ed.). Insect physiology. John Wiley and Sons, Inc., New York.
Roeder, K. D. 1962. Neural mechanisms of animal behavior. Am. Zoologist 2: 105-115.
Roeder, K. D. 1963. Nerve cells and insect behavior. Harvard University Press, Massachu-
setts. 189 p.
Roeder, K. D. and N. K. Kennedy. 1955. The effect of certain trisubstituted phosphine
oxides on synaptic conduction in the roach. J. Pharmacol. 114: 21 1-230.
Roeder, K. D. and S. Roeder. 1939. Electrical activity, nerve cord, roach. J. cell. comp.
Physiol. 14: 1-12.
Shore, P. A. and J. Olin, 1958. Identification and chemical assay of norepinephrine in brain
and other tissues. J. Pharmacol. 122: 295-300.
Shore, P. A., J. A. R. Mead, R. G. Kuntzman, S. Spector and B. B. Brodie. 1957. On the
physiological significance of monoamine oxidase in brain. Science 126: 1063-1064.
Smallman, B. N. 1956. Mechanisms of acetylcholine synthesis in the blowfly. J. Physiol.
132: 343-357.
Smallman, B. N. and R. W. Fisher. 1958. Effect of anticholinesterases on acetylcholine
levels in insects. Can. J. Biochem. Physiol. 36: 575-594.
Smith, D. S. 1965. Synapses in the insect nervous system, p. 39-57. In J. E. Treherne and
J. W. L. Beament (eds.). Insect central nervous system. Academic Press, New York.
Smith, D. S. and J. E. Treherne. 1965. The electron microscopic localization of cholinester-
ase activity in the central nervous system of an insect, Periplaneta americana L. J. cell.
Biol. 26: 445-465.
Stemburg, J., S. C. Chang and C. W. Kearns. 1959. The release of a neuroactive agent by the
American cockroach after exposure to DDT or electrical stimulation. J. econ. Ent. 52:
1070-1076.
Takeshige, C. and R. L. Voile. 1962. Bimodal response of sympathetic ganglia to acetyl-
choline following eserine or repetitive preganglionic stimulation. J. Pharmacol. 138:
66-73.
Takeshige, C. and R. L. Voile. 1963. Cholinoreceptive sites in denervated sympathetic gang-
lia. J. Pharmacol. 141: 206-213.
Cockroach Nerve Cord
305
Takeshige, C. and R. L. Voile. 1964. A comparison of the ganglion potentials and block pro-
duced by acetylcholine and tetramethylammonium. Brit. J. Pharmacol. 23: 80-89.
Tauc, L. and H. M. Gershenfeld. 1962. A cholinergic mechanism of inhibitory synaptic
transmission in a molluscan nervous system. J. Neurophysiol. 25: 236-262.
Tobias, J. M., J. J. Kollros and J. Savit. 1946. Acetylcholine and related substances in cock-
roach, fly and crayfish and the effect of DDT. J. cell. comp. Physiol. 28: 159-182.
Treheme, J. E. 1961a. Sodium and potassium fluxes in the abdominal nerve cord of the
cockroach, P. americana L. J. exp. Biol. 38: 315-322.
Treheme, J. E. 1961b. The movement of sodium ions in the isolated abdominal nerve cord
of the cockroach, Periplaneta americana. J. exp. Biol. 38: 629-636.
Treherne, J. E. 1961c. The efflux of sodium ions from the last abdominal ganglion of cock-
roach, P. americana L. J. exp. Biol. 38: 729-736.
Treherne, J. E. 1962a. Transfer of substances between the blood and central nervous system
in vertebrate and invertebrate animals. Nature, Lond. 196: 1181-1183.
Treherne, J. E. 1962b. The distribution and exchange of some ions and molecules in the
central nervous system of Periplaneta americana. J. exp. Biol. 39: 193-217.
Treherne, J. E. 1965b. The chemical environment of the insect central nervous system, p.
21-30. In J. E. Treherne and J. W. L. Beament (eds.). Insect central nervous system.
Academic Press.
Treherne, J. E. 1966. The neurochemistry of arthropods. Cambridge University Press, Cam-
bridge. 149 p.
Treherne, J. E. and D. S. Smith. 1965a. The penetration of acetylcholine into the central
nervous system of an instect C Periplaneta americana L.). J. exp. Biol. 43: 13-22.
Treherne, J. E. and D. S. Smith. 1965b. The metabolism of acetylcholine in the intact cen-
tral nervous system of an insect (. Periplaneta americana L.). J. exp. Biol. 43:441-454.
Twarog, B. M. and K. D. Roeder. 1956. Properties of the connective tissue sheath of the
cockroach abdominal nerve cord. Biol. Bull. Ill: 278-286.
Twarog, B. M. and K. D. Roeder. 1957. Pharmacological observations on the desheathed
last abdominal ganglion of the cockroach. Ann. ent. Soc. Amer. 50: 231-237.
Udenfriend, S. 1964. Fluorescence assay in biology and medicine. Academic Press, New
York. 517 p.
Unger, H. 1957. Untersuchunger zur neurohormonalen Steuerung der Herzatigkeit bei
Schaben. Biol. Zbl. 76: 204-225.
Van Asperen, K. 1962. A study of housefly esterase by means of a sensitive colorimetric
method. J. Ins. Physiol. 8: 401-416.
Voile, R. L. 1965. Interactions of cholinomimetic and cholinergic blocking drugs at sympa-
thetic ganglia, p. 85-94. In G. B. Koelle, W. W. Douglas, A. Carlsson and V. Trcka (eds.).
Pharmacology of cholinergic and adrenergic transmission. Macmillan, New York.
Voile, R. L. and G. B. Koelle. 1961. The physiological role of acetylcholinesterase (AChE)
in sympathetic ganglia. J. Pharmacol. 133: 223-240.
Weiant, E. A. 1958. Control of spontaneous activity in certain efferent nerve fibers from the
metathoracic ganglion of the cockroach Periplaneta americana. Proc. \0th Int. Cong.
Ent. 2: 81-82.
306
Chiang
Welsh, J. H. 1957. Serotonin as a possible neurohumoral agent: evidence obtained in lower
animals. Ann. N. Y. Acad. Sci. 66: 618-630.
Welsh, J. H. and H. T. Gordon. 1947. The mode of action of certain insecticides on arthro-
pod axon. J. cell. comp. Physiol. 30: 147-172.
Wigglesworth, V. B. 1954. Neurosecretion and the corpus cardiacum of insects. Pub. Staz.
Zool. Napoli 24: 41-45.
Wigglesworth, V. B. 1958. The distribution of esterase in the nervous system and other
tissues of the insect Rhodnius prolixus. Quart. J. micro. Sci. 99: 441-450.
Wilson, I. B. 1960. Acetylcholinesterase, p. 501-517. In P. Boyer, H. Lardy and K. Myrback
(eds.). The enzymes, vol. 4. Academic Press, New York.
Wilson, I. B. and S. Ginsburg. 1958. Reactivation of alkylphosphate inhibited acetylcholin-
esterase by bis quaternary derivatives of 2-PAM and 4-PAM. Biochem. Pharmacol. 1 :
200-206.
Winteringham, F. P. W. 1966. Metabolism and significance of acetylcholine in the brain of
the adult housefly, M usca domestica L. J. Ins. Physiol. 12: 909-924.
Winton, M. Y., R. L. Metcalf and T. R. Fukuto. 1958. The use of acetylthiocholine in the
histochemical study of action of organophosphorus insecticides. Ann. ent. Soc. Amer.
51: 436-448.
Yamasaki, T. and T. Narahashi. 1960. Synaptic transmission in the last abdominal ganglion
of the cockroach. J. Ins. Physiol. 4: 1-13.
307
ERRATA Quaestiones entomologicae. 1969: 5(2)
“A revision of the species of the genus Evarthrus Le Conte (Coleoptera: Carabidae)” by
Richard Freitag.
Page 96. Key, couplet 10. The species Evarthrus vinctus Le Conte is known from Ten-
nessee, Georgia, North and South Carolina, not from Mississippi.
Page 207. Locality records for Evarthrus texensis Freitag are indicated on Fig. 128 by
filled half-circles, not by inverted triangles.
Page 211. On Fig. 138, the numbers 1 - 8 represent centers of concentration, not centers
of speciation.
e. d, -Q<n<t
v ' J
Quaestiones
entomologicae
MUS. COMP. 200L.
FES »
A periodical record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.
NUMBER 4
VOLUME V
OCTOBER 1969
QUAESTIONES ENTOMOLOGICAE
A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.
Volume 5 Number 4 27 October 1969
CONTENTS
Graham - Observations on the biology of the adult female mosquitoes
(Diptera:Culicidae) at George Lake, Alberta, Canada 309
Fredeen — Outbreaks of the black fly Simulium arcticum Malloch in Alberta 341
OBSERVATIONS ON THE BIOLOGY OF THE ADULT FEMALE MOSQUITOES
(DIPTERA :CULICIDAE) AT GEORGE LAKE, ALBERTA, CANADA
PETER GRAHAM
Department of Biology
Thomas More College
Covington, Kentucky 41017
U.S.A.
The seasonal distribution of the more important mosquito species is discussed. No species
was found to be particularly abundant inside buildings and mosquitoes did not appear to
enter buildings to digest their blood meals, but appear to digest these near the feeding site.
A significant difference was found between the occurrence of certain species at the lake
shore and in the forest. Mosquitoes were found to be relatively inactive when in stages I I -IV
of Christophers and 3-5 of Sella of the gonotrophic cycle. Retention of eggs by parous
females was found to be widespread and to occur in 7% of the parous females.
A key to the adult female mosquitoes of central Alberta is given.
During studies comparing the effectiveness of different mosquito sampling methods at the
George Lake field site, in 1965, 1966 and 1967, a number of observations on the biology of
the adult female mosquitoes was made. As these observations were incidental to the main
study, they are somewhat superficial, but I believe they are worth recording as relatively
little is known about the biology of mosquitoes in this area.
The methods of collection and the study area are described elsewhere (Graham, 1969).
NOTES ON THE IDENTIFICATION OF AND KEY TO THE
ADULT FEMALES OF CENTRAL ALBERTA SPECIES OF MOSQUITOES
No key to mosquitoes was found to be completely satisfactory for the identification of
the adult female mosquitoes taken at George Lake. I, therefore, constructed a key, based
largely on the works of Barr (1958), Carpenter and LaCasse (1955), Rempel (1953) and
Quaestiones entomologicae
5: 309-339 1969
310
Graham
Vockeroth (1954b), which includes all species of mosquito recorded from Edmonton,
George Lake and Flatbush. It includes most if not all species of mosquito likely to be found
in the parkland and boreal forest regions of the province. Fig. 1 illustrates characteristics
used in the key. This key may prove useful until such time as a complete taxonomic study
of Alberta species of mosquitoes is carried out. Such a study is required before a full under-
standing of this mosquito fauna can be achieved.
I
FIG.
KEY A.
KEY B.
1. A.
B.
Lateral view of a generalized mosquito thorax.
(After Steward and McWade, 1961).
Claw characters of Aedes species (from Vockeroth, 1954b).
Biology of Mosquitoes
311
Keys to the adult female mosquitoes of central Alberta
Key to Genera. —
1 . Palps almost as long as proboscis; scutellum rounded . . . Anopheles earlei, p. 3 1 6
— Palps short, less than 1/3' length of proboscis; scutellum trilobed 2
2. ( 1) Spiracular bristles present Culiseta, p. 316
— Spiracular bristles absent 3
3. ( 2) Post spiracular bristles present; tip of abdomen pointed Aedes, p. 322
— Post spiracular bristles absent; tip of abdomen rounded 4
4. ( 3) Wings with many pale scales, wing scales broad
Coquillettidia perturbans, p. 322
— Wing scales all dark and narrow Culex, p. 322
Key to Culex species (from Rempel, 1953)
1 . Tarsal segments ringed with white tarsalis, p. 322
Tarsal segments not ringed with white 2
2. ( 1) White bands on apices of abdominal terga territans, p. 322
— White bands on bases of abdominal terga restuans, p.322
Key to Culiseta species
1 . Hind tarsal segments ringed with white 2
— Hind tarsal segments not ringed with white 5
2. ( 1) Wing scales forming conspicuous spots 3
— Wings without conspicuous spots 4
3. (2) Tarsal white rings broad; very large species alaskaensis, p. 3 1 6
— Tarsal white rings narrow incidens, p.320
4. ( 2) Abdominal pale bands on bases of terga only, white in color
C. morsitans dyari, p.321
— Abdominal pale bands on both apices and bases of terga, usually pale yellow
brown in color C. sylvestris minnesotae, p. 3 2 1
5. ( 1) Costa of wing with mixed pale and dark scales inornata, p.320
— Costa with dark scales only impatiens, p.320
Key to Aedes species (based on Vockeroth, 1954b)
1 . Hind tarsal segments ringed with white 2
— Hind tarsal segments not ringed with white 11
2. (1) Tarsal white rings on both apices and bases of tarsal segments 3
— Tarsal white rings on bases of tarsal segments only 5
3. ( 2) Wings with both dark and light scales on most veins 4
— Wing scales entirely dark canadensis, p.325
4. ( 3) Dark and light scales equally distributed on veins campestris, p.324
Third vein (R4+5) with more dark scales than 2nd (R2+3) or 4th(M)
dorsalis, p.325
5. (2) Tarsal white rings very narrow, lA or less than length of segment
vexans, p.324
312
Graham
Tarsal white rings broader, at least 1/3 of length of segment 6
6. ( 5) Large yellow species; abdominal terga almost completely yellow scaled; tarsal
claw as in Fig. 1 B 4 flavescens, p. 326
Not as above, abdominal terga with abundant dark scales 7
7. ( 6) Tarsal claw large, main claw almost parallel to accessory tooth and slightly sinuate,
Fig. IB 1 excrucians, p.325
Tarsal claw smaller, not sharply bent beyond tooth 8
8. ( 7) Mesonotum with some contrasting markings; tarsal claw with long .accessory
tooth, Fig. 1 B 2 9
Mesonotum almost uniform yellow brown; tarsal claw with short accessory tooth,
Fig. IB 3 riparius, p.327
9. ( 8) Palps and torus with some white scales; lower mesepimeral bristle 1, 2 or absent
fitchii*, p.325
Palps and torus usually without white scales; lower mesepimeral bristles 3 or
more 10
10. ( 9) Palps lacking hairs on basal half of apical segment at inner edge
increpitus*, p. 326
Palps with hairs on basal half of apical segment at inner ventral edge
stimulans*, p.327
11. ( 1 ) Fore coxa with a patch of brown scales on anterior surface; small species
cinereus, p.324
Fore coxa with patch of white scales on anterior surface 12
12. (11) Wing scales distinctly bicoloured 13
Wing scales all dark or with pale scales restricted to base of costa 14
13. (12) Wings with pale and dark scales intermixed, dark predominating; lower mese-
pimeral bristles usually present niphadopsis
Wing veins alternating black and white scaled; lower mesepimeral bristles absent
spenceri, p.327
14. (13) Post-coxal scale patch present 15
— Post-coxal scale patch absent 21
15. (14) Hairy species, postpronotum with setae scattered over posterior half impiger
Less hairy species, postpronotal setae restricted to a single or irregular double row
along posterior margin 16
16. (15) Sides of mesonotum silvery grey; base of costa with numerous white scales in a
conspicuous patch 17
Sides of mesonotum yellow or dark; base of costa with only a few or no white
scales 19
17. (16) White scales on costa covering basal 1/7 cataphylla, p.325
White scales on costa restricted to extreme base 18
18. (17) Sternopleuron with scales extended to anterior angle; mesonotum with numerous
white scales giving a “frosted” appearance, medium strip indistinct
trichurus, p.327
Sternopleuron with scales extending half way to anterior angle; mesonotum with
distinct median brown stripe implicatus, p.326
* Adult females of these species cannot be distinguished with certainty.
Biology of Mosquitoes
313
19. (16) Bristles of scutellum and mesonotum black; postmetasternal membrane with 15
or more scales pionips, p. 326
Bristles of scutellum and mesonotum yellow or bronze; postmetasternal mem-
brane bare or with less than 1 2 scales 20
Base of costa with distinct patch of white scales hexodontus, p. 326
Base of costa with no or few scattered white scales at most punctor, p. 327
Hypostigial scale patch present pullatus, p. 327
Hypostigial scale patch absent 22
Scales on sternopleuron extended to anterior angle; mesepimeron scaled to near
lower margin; mesonotum with contrasting dark lines 23
Scales of sternopleuron extended half way to anterior angle; lower 1/3 of mese-
pimeron bare; mesonotum usually uniform yellow brown intrudens, p. 326
Bristles of scutellum and mesonotum bright yellow; abdominal white bands indis-
tinct or absent diantaeus, p. 325
Bristles of scutellum and mesonotum black or bronze; abdominal white bands
distinct 24
Lower mesepimeral bristles present communis, p. 325
Lower mesepimeral bristles absent sticticus, p. 327
Notes on identifications of adult females. — The characters given in the key should enable
most specimens of adult female mosquitoes from central Alberta to be identified, provided
they are not badly rubbed, but some qualifications and explanations are necessary.
Central Albertan species of Anopheles, Culex, and Coquillettidia present no problems and
even badly rubbed specimens of these genera can usually be determined. The only confusion
likely in Culiseta is that rubbed specimens of C. sylvestris minnesotae Barr may be mistaken
for C. morsitans dyari (Coquillett). However, a close examination will usually reveal a few
pale scales on the apices of the terga.
The genus Aedes presents most of the identification problems and all specimens unidenti-
fied in this study were in this genus. Aedes cinereus Meigen is best distinguished by the
brown patch of scales on the fore coxa. A character often given in keys, the absence of
white bands on the abdominal terga is unreliable in central Alberta; Carpenter and LaCasse
(1955: 266) state that the abdomen of this species is as follows: “First tergite with a median
patch of brown scales, a few pale scales intermixed; remaining tergites brown without pale
bands or with narrow partial or complete bands”; the specimen they figure has complete
bands. Forty four specimens of A. cinereus collected at George Lake in 1965 and 1966 were
examined. Of these 14 had complete bands, 14 had no bands, and 16 had incomplete bands.
Specimens of A. cinereus are likely to be confused with small specimens of Aedes intrudens
(Dyar) which have white fore coxal scale patches, and specimens of Aedes vexans (Meigen)
which have pale bands on the tarsi.
In the subgenus Ochlerotatus the species with pale bands on both apices and bases of the
tarsal segments are no problem. The “ stimulans ” group, those with bands only on the bases
of the tarsal segments, are difficult to separate. Claw characters distinguish A. excrucians
(Walker) and A. riparius Dyar and Knab from the others. A. flavescens (Muller) can only be
20. (19)
21. (14)
22. (21)
23. (22)
24. (23)
314
Graham
confused with A. riparius and then only when rubbed. Specimens of A. fitchii (Felt and
Young), A. stimulans (Walker) and A. increpitus Dyar are easily separated from other band-
legged species by claw characters, but are impossible to distinguish from each other with any
certainty. The characters in the key are all unreliable. In this study all doubtful specimens
were referred to A. fitchii since larval surveys and other work (Happold, 1965a and 1965b;
Wada, 1965) have shown this to be by far the most abundant species of the three in central
Alberta and the larvae of A. fitchii only have been found at George Lake.
Black legged Ochlerotatus are a difficult group. A. punctor (Kirby) and A. hexodontus
Dyar occur in two forms: “tundra” type with uniform yellow brown mesonota and “ punc-
tor” type which have contrasting lines on their mesonota. Wada (1965) considered the best
way to distinguish these two species was the presence of a white spot on the base of the
costa of A. hexodontus, but Jenkins and Knight (1950) state that A. punctor “tundra” type
may also have a few white scales on the costal base. The criterion I used was — a conspicu-
ous white spot on the base of the costa — A. hexodontus; none or only a few scattered white
scales on the base of the costa — A. punctor.
Vockeroth (1952, 1954b) has discussed the separation of A. pionips Dyar from A. com-
munis (De Geer). The presence of a postcoxal scale patch in A. pionips appears to be the
best character. I could find no completely satisfactory way of distinguishing A. pionips
from A. punctor or A. hexodontus. The black mesonotal bristles are usually adequate but
some specimens of A. punctor and A. hexodontus have dark bronze bristles. These may be
separated with difficulty by the characters given by Beckel (1954).
Specimens of A. intrudens, A. communis, and A. sticticus (Meigen) present some prob-
lems of differentiation. A. intrudens is the only species whose members lack postcoxal scale
patches which normally lack contrasting lines on the mesonotum, but a few individuals have
indistinct lines on the mesonotum and closely resemble A. communis, and a few of these
may lack lower mesepimeral bristles and resemble A. sticticus. The scale patch on the stemo-
pleuron which only reaches half way to the interior angle in A. intrudens will distinguish
these. The presence of lower mesepimeral bristles distinguishes specimens of A. communis
from those of A. sticticus and the bright yellow mesonotal bristles and incomplete abdom-
inal bands distinguish specimens of A. diantaeus Howard, Dyar and Knab from other species
without a postcoxal scale patch.
DIVERSITY
Twenty nine species of mosquitoes have been recorded from the George Lake field site.
Nineteen species were taken in 1965, 27 in 1966 and 25 in 1967. The species found and
their relative abundance are shown in Table 1. Pucat (1965) records 38 species of Culicinae
from Alberta.
Seven species made up 80% of the 1966 collections. These were: Culiseta inomata (Willis-
ton), Aedes excrucians, A. fitchii, A. communis, A. punctor, A. riparius and A. vexans. This
preponderance of a few species was expected from the work of Williams (1964) and has
been recorded in northern mosquitoes by Happold (1965a, 1965b) in Alberta, and by
Skiersca (1965) in Poland, as well as by other authors elsewhere.
Biology of Mosquitoes
315
Table 1. Mosquito species collected at George Lake in 1965, 1966 and 1967, with their
relative abundance.
Genus Anopheles Meigen, 1818
A. earlei Vargas, 1 943 c
Genus Culiseta Felt, 1904
Subgenus Culiseta Felt, 1904
C. alaskaensis (Ludlow), 1906 fc
C. inornata (Williston), 1893 a
Subgenus Culicella Felt, 1904
C. sylvestris minnesotae Barr, 1957 fc
C. morsitans dyari (Coquillett). 1901 p
Genus Culex Linnaeus, 1758
Subgenus Culex Linnaeus, 1758
C. tarsalis Coquillett, 1896 p
Subgenus Neoculex Dyar, 1905
C. territans Walker, 1856 c
Genus Coquillet tidia Dyar, 1905
C. per turbans (Walker), 1856 c
Genus Aedes Meigen, 1818
Subgenus Aedes Meigen, 1818
A. cinereus Meigen, 1818c
Subgenus Aedimorphus Theobald, 1903
A. vexans (Meigen), 1 830 Va
Subgenus Ochlerotatus Lynch Arribalzaga, 1891
A. canadensis (Theobald), 1901 r
A. cataphylla Dyar, 1 9 1 6 p
A. communis (De Geer), 1776 c
A. diantaeus Howard, Dyar & Knab, 1913 p
A. dorsalis (Meigen), 1830 p
A. excrucians (Walker), 1856 Va
A. fitchii (Felt & Young), 1904 a
A. flavescens (Muller), 1764 fc
A. hexodontus Dyar, 1916 r
A. implicatus Vockeroth, 1954 c
A. intrudens Dyar, 1919 fc
A. pionips Dyar, 1919 fc
A. pullatus (Coquillett), 1904 r
A. punctor (Kirby), 1837 a
A. riparius Dyar & Knab, 1907 c
A. spencerii (Theobald), 1901 p
A. sticticus (Meigen), 1838 fc
A. stimulans (Walker), 1 848 p
A. trichurus (Dyar), 1904 p
Key
316
Graham
The coefficient of diversity (Fisher et al., 1943) was 5 ± 0.04 in 1966 and 3 ± 0.02 in
1967, even though the number of species found was almost the same. It appears that this
coefficient is a function of sample size as well as population size in a population with a large
number of individuals, but a limited number of species. The coefficient of diversity is of use
in comparing different areas or traps sampled at the time, as has been shown by Williams
(1964) for mosquitoes taken in light traps in several cities in Iowa. It is also useful for com-
paring captures in the same trap in different years, provided the samples are similar in size or
the number of possible species is very large, as in the Lepidoptera studied by Williams
(1964).
Genus Anopheles Meigen, 1818
Anopheles earlei Vargas, 1943. — One species of Anopheles is recorded from Alberta. This
is a member of the widespread “ Anopheles maculipennis ” complex and until recently was
confused with a Pacific Coast species, A. occidentalis Dyar and Knab. It is distributed over
much of the northern U. S. A. and Canada north to Labrador and Alaska. Probably all
records of A. occidentalis from east of the mountains refer to this species. It is fairly com-
mon at George Lake.
There is believed to be only one generation per year in Alberta (Happold, 1965a). Over-
wintering appears to be by nulliparous females, which hibernate in basements and animal
burrows (Shemanchuk, 1965). They leave their hibernation sites in early spring and oviposit
soon after. At George Lake adults emerged in late July in 1966 but in late June in 1967.
The adult females are long lived and overwintered females could still be found in late July
1966. Dissections for parity rate confirmed the presence of only one generation in 1966
(Fig. 2) but the situation in 1967 was not clear as nullipars predominated in late June. Fig. 3
shows the parity rate in 1967 compared with Culiseta alaskaensis (Ludlow) which has only
one generation. No hibernation sites were found at George Lake but a series of females
taken at animal burrows by Shemanchuk at Rimbey, Alberta in October, 1966, were all
nulliparous.
Larvae of this species were found in a sedge meadow along with those of Aedes vexans in
August 1966 at George Lake.
Genus Culiseta Felt, 1904
This genus is often referred to as Theobaldia Neveu-Lemaire, 1902, but this name was not
available for this group of insects.
Culiseta alaskaensis (Ludlow), 1906. — This circumboreal species is a typical mosquito of
the boreal forest. It overwinters as an adult female and is one of the earliest species to leave
hibernation, becoming active as soon as the snow thaws. Though Happold (1965a) states “It
is common in Alaska . . . but rare in Alberta and Saskatchewan” it is far from rare at George
Lake, and was numerous enough to be a nuisance in May of both 1966 and 1967. In 1966
the emerging generation was seldom encountered, only two adults being taken in July and
August. The overwintered generation had disappeared by late June. The females which leave
hibernation are voracious biters, but there is no information on whether the newly emerged
females bit or not.
Number of Specimens
Biology of Mosquitoes
317
Fig. 2. Seasonal changes in the numbers of parous and nulliparous female mosquitoes at
George Lake. June to August 1966.
Shaded area = no. nulliparous E = early; L = late
Number of Specimens
318
Graham
Fig. 2. Continued
Biology of Mosquitoes
319
Fig. 2. Continued
320
Graham
Fig. 2. Continued
Culiseta impatiens (Walker), 1848. — This species has not been found at George Lake,
though it occurs at Flatbush (Happold, 1965a).
Culiseta incidens (Thompson), 1869. — This is a western North American species. It has
not been found at George Lake but occurs at Flatbush.
Culiseta inomata (Williston), 1893. — Unlike the other Central Albertan species of Culi-
seta this species, which is confined to North America, appears to be a southern rather than a
northern or mountain form. It occurs in both forest and open country and has successfully
adapted to urban conditions, forming a large proportion of the mosquitoes breeding in
Edmonton. It was common at George Lake in 1966, when it was most abundant near the
lake shore, but was not taken after the second week in May in 1967. Happold found it to be
rare at Flatbush; he found no larvae and very few adult females.
This species overwinters as an adult female, but has several generations in a summer. Wada
(1965) found that oviposition in Edmonton continued into mid August. Unfortunately most
of the females taken at George Lake were gravid so no confirmation of generation number
was possible. Malaise trap captures in 1966 show three peaks, which suggests two generations:
one peak in early May, when overwintered females leave hibernation sites, one peak in early
August, when the adults resulting from eggs laid by the overwintered females emerge and
one in September, which may represent a second generation. Larvae have been found in
woodland pools in Edmonton but none have been found at George Lake.
321
Biology of Mosquitoes
o.
Fig. 3. Changes in the number of parous and nulliparous females of Culiseta alaskaensis
and Anopheles earlei during the spring and early summer of 1967 at George Lake.
Shaded area = no. nulliparous E = early; L = late
Culiseta sylvestris minnesotae Barr, 1957. — Specimens of this species are often confused
with specimens of C. morsitans dyari (Coquillett). I did not distinguish between the two
species in 1965. All 1966 specimens which I preserved were C. sylvestris minnesotae but it is
possible that a few C. morsitans dyari (Coquillett) may have been taken.
C. sylvestris minnesotae hibernates as an adult female, which leaves winter quarters early
in the spring, being recorded in the first week of May 1967, before C. alaskaensis. The dis-
tribution of this species is not yet well known. Stone (1965) records it from Minnesota,
Utah, Ontario, New Jersey and Massachusetts. Curtis (1967) states it has been recorded
from the borders of British Columbia but not yet in that province. Probably many records
of C. morsitans dyari (Coquillett) will prove to be C. sylvestris minnesotae. Barr (1958)
states that females of C. sylvestris minnesotae do not appear to take human blood.
Culiseta morsitans dyari (Coquillett), 1901. - This is a holarctic species with a more or
less northern distribution. It is not common at George Lake, where only five specimens have
been definitely identified. Elsewhere in Alberta I have seen specimens of this species from
the Cypress Hills, Edmonton, and Flatbush.
Very little is recorded of the biology of this species in North America. In Europe, Marshall
(1938) and Wesenberg-Lund (1921) found that it overwinters as a larva, and unlike other
Culiseta species it oviposits on wet mud, laying eggs singly instead of in rafts. Howard et al.
(1915) considered it hibernated as an egg in North America but this does not seem to be so,
and there is no study of its life history. Stone et al. (1959) suggest that the North American
form may be different from the palearctic one. At George Lake in 1967 females appeared in
late June, and one dissected was nulliparous.
322
Graham
Genus Culex Linnaeus, 1758
Culex restuans Theobald, 1901. — This species has not been found at George Lake but
was taken by Klassen (1959) in Edmonton.
Culex tarsalis Coquillett, 1896. — Two specimens of this common prairie species were
taken in 1966. They had probably migrated into the field site from surrounding open lands.
This species may be extending its range north in Alberta. Hocking (pers. comm.) informs me
that it has increased in abundance in Edmonton in the last ten years.
Culex territans Walker, 1856. — This is a holarctic species which until 1949 was confused
with C. apicalis Adams in North America. It is probable that all records of C. apicalis east of
the mountains and north of Utah refer to C. territans. Possibly because it rarely bites man, it
is recorded as rare in most regional mosquito records (Happold, 1965a, 1965b; Curtis, 1967;
Steward and McWade 1961). It is believed to feed mainly on cold blooded vertebrates but
Means (1965) has recorded it feeding on man. It was fairly common at George Lake in 1966
and 1967, and was one of the first mosquitoes to leave winter quarters. In 1967 C. territans
was the first mosquito taken, one specimen being taken in a Malaise trap in April before the
snow had completely melted.
From seasonal distribution data it appears that there were two generations in 1966 (Fig.
4), but dissections were too few to confirm, though they do support this.
Genus Coquillettidia Dyar, 1 905
This is a mainly tropical genus; two species penetrate into northern regions, one in
Eurasia and one in North America.
Coquillettidia perturbans (Walker), 1856. - This widespread North American species was
first taken in Alberta at Flatbush by Happold in 1961, (Happold, 1965a) and this is still the
only published locality (Pucat, 1964, 1965). I have seen specimens from Edson, and P. Shera
(unpublished report) recorded it from Elk Island National Park. At George Lake it was not
found in 1965, but was fairly common in 1966 and had just appeared in 1967 when work
was stopped. It is probably common and widespread over the forested parts of Alberta.
There appears to be only one generation per year and this is the only Canadian mosquito
definitely known to overwinter as a larva (Happold, 1965a). As yet no larvae have been
found in Alberta.
Adult females of this species are reputed to be fierce biters, but were not numerous
enough to be a nuisance at George Lake. Happold (1965a) at Flatbush and Burgess and
Haufe (1960) in Ontario found this species abundant in the forest canopy.
Genus Aedes Meigen, 1818
This is the predominant mosquito genus in Alberta and in most northern regions. It in-
cludes 70% of the species and 80% of the specimens caught at George Lake. The genus is
distributed from the equator to the limit of land in the north and is found on many oceanic
islands.
Three subgenera, Aedes Meigen, Aedimorphus Theobald, and Ochlerotatus Lynch Arribal-
zaga, are found in Alberta.
No. tok«n
Biology of Mosquitoes
323
W
10 .
0 .
Aedes
intrudens
Fig. 4. Seasonal changes in the numbers of some of the less common mosquito species
taken by all methods at George Lake. 21 April to 29 September 1966.
E = early
L = late
324
Graham
All Alberta Aedes species overwinter as eggs and with the possible exceptions of A. vexans
and A. dorsalis , there is only one generation per year. Brust (pers. comm.) informs
me that diapause is not obligatory in the eggs of A. sticticus, so this species may also be
multivoltine if the season permits.
Aedes (Aedes) cinereus Meigen, 1818. — This small holarctic species is common in central
Alberta. It is a woodland species which is also common in Poland (Skiersca, 1965). It is
common at George Lake, where it is a late-appearing species reaching an adult population
peak in July and early August in 1966. 1 noticed that it is a low flier and bites mainly below
the knee and around the ankles.
Aedes (Aedimorphus) vexans (Meigen), 1830. — This appropriately-named species has one
of the widest and most unusual ranges of any mosquito, being found in the Palearctic,
Nearctic, and Oriental regions and also in the Transvaal, Fiji, Samoa and New Caledonia
(Stone et al., 1959). It was very common at George Lake in 1965 and 1966 but was not
found there in 1967, though it may have appeared after work stopped. It was taken in June
1966.
Rempel (1953) suggested there were two forms of A. vexans in Saskatchewan, a large
form in the prairies and a smaller one in the parklands and forest.
A. vexans is a migratory species, capable of flying long distances. Horsfall (1955) records
many instances of migration in temperate regions and de Meillon and Khan (1965) have
recorded a large migration in Burma. A result of this migratory tendency is that this species
is often a major pest species in cities like Edmonton with efficient control programs, as it
migrates in from breeding sites which may be several miles from the city. Clark and Wray
(1967) have studied the influx of A. vexans into Des Plaines, Illinois, and provided a method
of predicting invasions.
A. vexans is a late emerging species with a 1966 population peak in late summer. The
adult females continued well into September 1966 and larvae were taken in late August.
Stage et al. (1938) recovered a marked female 55 days after release.
In the southern part of its range the species is multivoltine, but the number of generations
in northern parts is not clear, as the eggs do not hatch simultaneously. Some may require
several floodings before hatching (Horsfall, 1955), giving rise to broods. Gjullin et al (1950)
found that eggs remained viable for three to four years if kept moist.
Nullipars were found from June to the end of August 1966 and predominated up to late
August, indicating that emergence took place in June, July, and early August, but there was
no evidence of more than one brood.
Subgenus Ochlerotatus Lynch Arribalzaga, 1891
This is the dominant subgenus in northern regions, but also occurs in the tropics. In
North America this subgenus extends further north than any other culicid. Two species,
A. nigripes, and A. impiger, occur within 500 miles of the pole. Nineteen species have been
recorded at George Lake, and Pucat (1965) records 25 from Alberta.
Aedes campestris Dyar and Knab, 1907. — This species has been recorded at Edmonton
(Klassen, 1959; Wada, 1965), but not yet at George Lake.
Biology of .VJosquitoes
325
A. canadensis (Theobald), 1901. — This species is widespread in the forested parts of
North America. It is not common at George Lake. Rempel (1950) found it to be fairly
common in the aspen grove parkland of Saskatchewan. It is a late emerger. At George Lake
it appeared at the end of June in both 1966 and 1967.
Aedes cataphylla Dyar, 1916. — This holarctic species is confined to western Northern
America in the Nearctic Region (Stone, 1965). It is not common at George Lake, but is
often a major pest species in Edmonton (Klassen, 1959; Klassen and Hocking, 1963). It is
one of the first species to emerge and Klassen found emergence completed by 19 May, 1958,
at Edmonton.
Aedes communis (De Geer), 1776. — This is an important woodland species with a cir-
cumboreal distribution and is a well known pest species in Europe and North America.
Chapman and Barr (1964) have described a subspecies A. communis nevadensis, from the
western U. S. A. and two larval forms of the typical subspecies. All larvae examined from
Alberta belonged to the eastern form. Hocking (1954) described an autogenous form from
Churchill on Hudson Bay in which autolysis of the flight muscles takes place. The species is
common at George Lake.
It is a fairly early emerging species; in 1966 a population peak of adult females occurred
in early June. Nullipars predominated to late June, indicating that some emergence took
place up to then.
Aedes diantaeus Howard, Dyar and Knab, 1917. - This species has a wide distribution in
the boreal forests of the old and new worlds, but it is seldom abundant (Vockeroth, 1954b).
In Alberta it has only been recorded from Flatbush (Happold, 1965a; Pucat, 1964). A single
specimen was found at George Lake in 1966.
Aedes dorsalis (Meigen), 1830. — This species occurs in the grasslands of the Palearctic
and Nearctic regions. It is abundant in southern Alberta but uncommon north of Edmontoa
One specimen was taken at George Lake in 1965.
Khelevin (1958) has shown that diapause in the egg of this species is facultative and it can
have several generations per year. Like A. vexans it is a migratory species (Horsfall, 1955).
Aedes excrucians (Walker), 1856. — This is a holarctic woodland species with a somewhat
more southerly distribution than A. communis. Together with A. vexans , it was one of the
two most abundant species at George Lake in 1965 and 1966. It was also abundant in 1967,
but work stopped before it reached population peak. It is a fairly early emerger, first appear-
ing in the last week in May 1966 but not till the second week in June 1967. The emergence
period appears to be prolonged as nullipars predominated until mid July and could still be
found in early August 1966. The adult females were found up to the end of August but
none were taken in September 1966. Matheson (1944) records a single instance of larvae
being found in September in New York. It is a persistent biter, and is a major pest species in
central Alberta.
Aedes fitchii (Felt and Young), 1904. — This species is confined to North America, where
it has the same range as A. excrucians. It is common in central Alberta. At George Lake it
appeared in early June 1966. Nullipars predominated until mid July and were found till mid
August, indicating a prolonged emergence period. Adult females were taken up to the end of
August but were not taken in September either in 1965 or in 1966.
326
Graham
It is an important pest species in central Alberta.
Larvae were found in a pool in a spruce grove on the western fence of the George Lake
field site along with those of A. implicatus in May 1966.
Aedes flavescens (Muller), 1764. — This species occurs in grassland areas of Eurasia and
North America. It is common in central Alberta and fairly common at George Lake. Speci-
mens taken on the field site had probably migrated in as it is an open country species.
Hearle (1929) considered it to be the most numerous mosquito on the Canadian prairies.
Happold (1965a) did not find it in the forest at Flatbush but found it common in an alfalfa
field.
Adult females were taken from early June to mid August 1966 at George Lake. Nullipars
were found till early July. A single male was taken in June 1967, when females were more
abundant than in 1966.
Aedes hexodontus Dyar, 1916. — This tundra and open country species was first recorded
from Alberta by Wada in 1964 at Edmonton (Wada, 1965). It was not common at George
Lake. It is an early species and was not found after June in either year.
Aedes implicatus Vockeroth, 1954. — This is a boreal forest species, confined to North
America, and occurs north almost to the tree line. It is locally common in central Alberta,
being common at George Lake but rare at Flatbush. In late May and early June 1967 this
species was the most abundant Aedes at George Lake.
It is an early emerger, the adult females reaching a population peak in mid June in both
1966 and 1967. There was some indication that a second brood occurred in late July 1966,
as pars predominated in late June and early July, but the three specimens dissected in late
July and early August were nulliparous. In 1967 only nullipars were taken in May but pars
predominated in late June and the numbers taken began to decline in mid June. Nielsen and
Rees (1961) believed this to be a short lived species; if so, then second broods are needed to
account for nullipars occurring in late summer.
In early May 1 966 larvae were found in pools in ruts in the road near the campsite and in
a pool on the western boundary along with those of A. fitchii.
Aedes increpitus Dyar, 1916. — This species has been taken at Edmonton by Klassen and
Wada, but has not yet been found at George Lake.
Aedes intrudens Dyar, 1919. - Another species with a circumboreal distribution, it was
common at Flatbush and relatively rare at George Lake in 1966, but common there in 1967.
It is an early species, with a population peak in late spring and only pars were taken in
July 1966. Happold recorded it as the most important Aedes species in buildings at Flat-
bush. At George Lake only one specimen was taken in the trailer in 1966 but it was promi-
nent indoors in spring 1967. Matheson (1944) writes of A. intrudens : “Dyar states that this
species readily invades houses, but I have never taken them in houses though I have found
them abundantly in wooded areas throughout the season.”
Aedes pionips Dyar, 1919. — This Nearctic species is found in Canada and the western
U. S. A. It is rare at George Lake.
It is generally considered to be a late emerger (Carpenter and LaCasse, 1955) but at
George Lake in 1966 the population peak appeared to be in early summer, adult females
continued into August, nullipars were found in late July.
Biology of Mosquitoes
327
Aedes pullatus (Coquillett), 1904. — This species is confined to the western mountains and
to the Ungava peninsula and South Baffin Island in North America. It also occurs in Europe,
where it is not a mountain species. In Alberta it is common in the Rocky Mountains. Klassen
and Wada did not find it at Edmonton but in some years it is common there (Hocking, pers.
comm.). It was rare at George Lake, and only found there in late May and early June.
Jenkins and Knight (1950) found it to be one of the most abundant species at Great
Whale River on the east coast of Hudson Bay and Vockeroth (1954b) states that the eastern
forms may be different from the western, though no morphological differences could be
found.
Aedes punctor (Kirby), 1837. — This is an important woodland species with a holarctic
distribution. It is a major pest species in both North America and northern Eurasia. It is
common in central Alberta, being the major black-legged Aedes species at both George Lake,
Edmonton and Flatbush.
In 1966 adult females were taken from late May into September, the population peak
being in early June. In 1967 this species reached its peak somewhat later than A. implicatus,
becoming the most abundant species in the third week in June. In 1966 there was a second
peak in August, which may have signified a second brood, but the few dissections done at
this time did not confirm this, the individuals examined being mainly parous. Nullipars
predominated until early July and were found until mid August; probably some emergence
took place in each summer month in 1966.
Aedes riparius Dyar and Knab, 1907. — This species occurs in western North America and
northern Eurasia. It is mainly an open country species but also occurs in woodlands. It is
common in central Alberta.
In 1966 it was not found until late June when it occurred in fairly large numbers, which
possibly indicates it did not breed in the area. Nullipars predominated in June and July and
some were found in August. In 1967 this species was taken in late May and was the most
abundant band-legged Ochlerotatus species till the end of June when A. excrucians began to
reach peak emergence.
Aedes spencerii (Theobald), 1901. — This is a prairie species, confined to western North
America; it is among the most numerous species in southern Alberta but rare north of Ed-
monton. One specimen was taken at George Lake in August 1966 and several in late May
1967.
A. sticticus (Meigen), 1838. — This is a holarctic species which extends its range south to
the Gulf of Mexico in North America. It is locally common in central Alberta being fairly
common at George Lake and rare at Flatbush.
Adult females were taken from late May to the end of July and nullipars to mid July in
1966.
Aedes stimulans (Walker), 1848. - This species has much the same range as A. fitchii. It
is common over much of the forest country of Canada, but appears to be rare in central
Alberta. Only four specimens have been definitely identified at George Lake and Happold
did not find it at Flatbush.
Aedes trichurus (Dyar), 1904. — This is a woodland species found in the northern U. S. A.
and southern Canada. In Alberta it occurs as far north as Beaverlodge (Rempel, 1950). It
was rare at George Lake.
328
Graham
SEASONAL DISTRIBUTION
Happold (1965b) discussed the seasonal distribution of mosquitoes at Flatbush, where
most of the important species found at George Lake occur, and the George Lake findings
are in agreement with his.
Fig. 5 shows the 1966 distribution of the commoner species in Malaise trap catches
corrected to 100 hours running time.
The mosquito population showed three peaks in 1966: the first in early May when the
overwintering females left hibernation; one in early July, representing the peak population
of Aedes species; and the third in early August formed by the emergence of those females
which enter hibernation and second broods of Aedes. This peak was probably much more
pronounced in 1965, when weather conditions favoured second broods.
In 1965, adults of Anopheles intrudens and pars of Anopheles earlei were taken in
August. This was almost certainly the result of weather conditions in 1965. Happold (1965a)
noted that in 1961 adults of Anopheles earlei survived for a longer period than in 1962.
During 1961 the weather conditions were like those in 1965. The adults of other species
may exhibit increased longevity as a result of favorable climatic conditions.
In all three years Aedes mosquitoes became abundant each spring approximately at the
same time as the poplars flushed.
MOVEMENT INTO BUILDINGS
Unlike tropical areas where disease transmission inside human dwellings is important,
there is relatively little information from North America on the movement of mosquitoes
into buildings. Matheson (1944) mentions a few species as persistent house enterers and
Happold (1964b) has recorded the species he collected indoors at Flatbush.
During June, July, and August 1966 and in May and June 1967 special captures were
carried out in a partially screened trailer at George Lake. In 1966, 1 12 specimens of 16 spe-
cies and in 1967, 96 specimens of 12 species were taken in this trailer (Table 2).
In 1966 70% of the indoor catch was composed of A. punctor, A. excrucians, A. pionips
and A. communis , while in 1967 Anopheles earlei, Aedes punctor and A. intrudens made up
the body of the catch. A. hexodontus and A. sticticus were also prominent in both years.
Happold (1965b) found A. earlei and A. intrudens made up 80% of the indoor catch at
Flatbush. In 1967 all A. earlei found indoors at George Lake were taken in May. This may
indicate that the overwintered females enter buildings to feed but that the trailer did not
provide a good hibernation site. Elsewhere individuals of this species are known to hibernate
in buildings, especially basements. The most striking difference between the George Lake
and Flatbush results was the absence of A. punctor from indoor captures at Flatbush,
although it was one of the most abundant local species. It formed over 30% of the indoor
catch at George Lake.
In central Alberta, the entry of mosquitoes into buildings depends more on the environ-
ment rather than on specific innate tendency to enter buildings as has been recorded in the
tropics, where exophilous and endophilous genera and species are recognized (Holstein,
1954; Muirhead-Thompson, 1951).
No./IOO Trop hours Na/IOO Trap hour* Na/IOO Trap hours
329
Fig. 5.
Black legged
Aedes
I — ■ I — l
Seasonal changes in the numbers of mosquitoes caught per 100 trap hours, in
Malaise traps, at George Lake. May to September 1966.
E = early
L = late
5
10
5
5
15
10
5
15
10
5
70
00
50
40
30
Graham
Aedes
punctor
A. implicatus
CZ71
A. communis
A.fitchii .
A.riparius
Fig. 5. Continued
Biology of Mosquitoes
331
Table 2. Mosquito species taken indoors at George Lake in 1966 and in 1967 and
numbers taken.
Species Number Caught
Total 112
96
332
Graham
DISTRIBUTION WITHIN THE STUDY AREA
Fig. 6 shows the relative abundance of the major species at the lake shore and in the
forest in 1 966. Culiseta inomata and Aedes vexans were more abundant in the lake shore
traps and A. excrucians and A. fitchii in the forest traps. The other species were more or less
evenly distributed. In 1967 A. implicatus was far more abundant in the forest traps.
Fig. 6. Relative abundance of mosquito species taken in Malaise traps at the lake shore
and in the forest at George Lake in 1966. Shaded - genera other than Aedes
1 = Anopheles earlei
3 = other Culiseta
5 = Coquillettidia per turbans
7 = A. communis
9 = A. fitchii
11= ^4. punctor
13 = A. vexans
2 = Culiseta inomata
4 = Culex territans
6 = Aedes cine reus
8 = A. excrucians
10 = A. implicatus
1 2 = A. riparius
14 = other Aedes
PROPORTION PAROUS
Fig. 2 shows the seasonal distribution of pars and nullipars* in the species taken more
abundantly in 1966 and Fig. 3 shows the parity rate for Culiseta alaskaensis and Anopheles
earlei, in the spring of 1967. The distributions in Aedes excrucians and A. fitchii exactly fit
those expected for a univoltine species which overwinters as eggs, i.e. there are two over-
lapping curves, one in early summer with nullipars predominant and one in late summer
with pars predominant. The curve for Anopheles earlei in 1966 fits the expected for a uni-
voltine species which overwinters as an adult female, though the number dissected was
small. The 1967 curve, however, indicates that considerable emergence took place in late
*Nullipars = those individuals which have not yet laid eggs.
Pars = those individuals which have laid at least one batch of eggs.
Biology of Mosquitoes
333
June and as this appears rather early for females to enter hibernation it is possible that there
was a second generation. The 1 967 curve for Culiseta alaskaensis is as expected for a species
which overwinters as an adult nulliparous female. The picture for the other species is not so
clear.
Shelenova (1959) found that in Russia, members of Aedes were generally shorter lived
than were members of Anopheles in the same area, few Aedes individuals having passed four
gonotrophic cycles, while anophelines often had passed 12 to 29. Carpenter and Nielsen
(1965) also found few individuals of Aedes with more than four dilations in the ovarian
ducts in Utah.
Corbet (1961) and Hamon et al. (1964) have studied differences in activity between pars
and nullipars, especially with regard to biting activity. Neither found any significant differ-
ences, though Hamon et al. found a modification of the biting cycle after spraying with
insecticide had removed the older portion of the population. They concluded that extrinsic
factors, such as light intensity and temperature are more important in controlling activity of
adult female mosquitoes than the age of the insect.
OVARIAN DEVELOPMENT AND ACTIVITY
Table 3 shows the stages of Sella and of Christophers in the total catch, in total Aedes
species and in the commoner species of mosquitoes. By far the greater proportion of all
Table 3. Distribution of the stages in the gonotrophic cycle (stages of Sella and of
Christophers) in the more important species of mosquito taken at George Lake
in 1966.
Stage of Sella
334
Graham
Table 3. continued.
Stage of Christophers
females caught were in stages 1 and 7 of Sella and I, II or V of Christophers. Thus they were
either unfed or gravid, with few in the intermediate stages. Christophers (1911) and Clement
(1963) have shown that the eggs normally develop to stage II of Christophers and then
development stops until a blood meal is taken. This agrees with the findings of Carpenter
and Nielsen (1965) in Utah, where 89% of the active females examined were in stages I and
II of Christophers.
A surprising part of the 1966 findings at George Lake was that very few of the resting
older females caught in the trailer were gravid or in intermediate stages. In 1 967 rats were
kept in unscreened cages in the trailer and many specimens of Anopheles earlei as well as
Aedes species were found in the intermediate stages of Sella and Christophers. This indicates
that these mosquitoes enter houses in search of food and if they find it remain to digest
their blood meals, but otherwise leave fairly soon. It also indicates their digestion of blood
meals is done very close to the feeding site. Carpenter and Nielson (1965) found about half
the resting females taken in their study were gravid but found very few intermediates.
Biology of Mosquitoes
335
These findings indicate that there is very little flight activity during intermediate stages of
the gonotrophic cycle. This is most probably because the blood fed or gravid female mos-
quito is a relatively inefficient flying machine as the weight of the blood meal may exceed
the weight of the insect. This little flight activity may apply mainly to woodland species as
individuals resting in grassland or tundra are more likely to be disturbed by passing animals.
Corbet (1961) found that most individuals engaging in “non-specific activity” were in the
early stages of Christophers but Standfast (1965) believed that light trap captures in Austra-
lia showed that a considerable part of the population was engaging in “non-specific activity”
during the night, interpreted this to mean that there was considerable activity in the inter-
mediate stages of the gonotrophic cycle, but did not record any dissections to support this.
Carpenter and Nielson (1965) report that a fair number of biting mosquitoes had ovaries
in stage II of Christophers and were nulliparous. They suggest that this is an indication of
autogeny and mention that some of the species in which this occurred have been shown to
be capable of autogeny by Chapman (1962).
At George Lake in 1966 and 1967 records were kept of specimens in stage I of Sella and
stage III of Christophers. Twenty three were found of which seven were nulliparous, forming
0.007% of all nullipars. These seven consisted of five species: A. communis, A. fitchii, A.
intrudens, A. punctor, and A. riparius. Two of these, A. communis and A. punctor were
included in those found by Carpenter and Nielson (1965) and are mentioned as capable of
autogeny by Chapman (1962) but the phenomenon appears to be rarer at George Lake
than in Utah where 4% of the biting females had ovaries in stage III of Christophers.
Eighty percent of the females of Culiseta inornata taken at George Lake in 1965 and in
1966 were gravid. The reason for this is not known but it could indicate some degree of
autogeny. No indication of autogeny in this species had been found in the laboratory.
RETENTION OF EGGS BY PAROUS FEMALE MOSQUITOES
AND OTHER OVARIAN ABNORMALITIES
During dissections for parity rate I noticed that several parous females with ovaries of
otherwise normal appearance (i.e., they did not have the sac-like appearance of those in
females which have just oviposited) had retained eggs in one ovary. The number of eggs
retained was usually one but six were found in one Aedes excrucians.
Table 4 shows the species and the number in each with retained eggs. This appears to be a
widespread phenomenon as it was recorded in ten species of three genera. It is also inter-
esting to note that the percentage of pars with retained eggs in each year was approximately
7%. The slightly higher proportion in August over June, may be due to older multiparous
females having a greater tendency to retain eggs over younger pauciparous females; but this
will need further study.
In one parous Aedes punctor taken in a Malaise trap in late June 1967 there was differ-
ential development in the ovaries, one being in stage II and one in stage IV of Christophers.
336
Graham
Table 4. Parous female mosquitoes at George Lake with eggs retained in one ovary.
ACKNOWLEDGEMENTS
I wish to express my thanks to B. Hocking, G. E. Ball, W. G. Evans, D. A. Boag, K. Smillie
and J. A. Downes for reading and criticizing the manuscript and to N. Wood for much help
in the field.
I should also like to thank the City of Edmonton and the University of Alberta for the
provision of grants to enable this work to be carried out.
REFERENCES
Barr, A. R. 1958. The mosquitoes of Minnesota. Univ. Minn. Agric. exp. Sta. Tech. Bull.
228. 154 pp.
Beckel, W. E. 1954. The identification of adult female A edes mosquitoes (Diptera:Culicidae)
of the blacklegged group, taken in the field at Churchill, Manitoba. Can. J. Zool. 32:
324-358.
Burgess, L. and W. O. Haufe. 1960. Stratification of some prairie and forest mosquitoes in
the lower air. Mosquito News 20: 341—346.
Carpenter, S. J. and W. J. LaCasse. 1955. The mosquitoes of North America (north of
Mexico). University of California Press, Berkeley. 360 pp; 127 pi.
Biology of Mosquitoes
337
Carpenter, S . J. and L. T. Nielsen. 1965. Ovarian cycles and longevity in some univoltine
Aedes species in the Rocky Mountains of the western United States. Mosquito News 25:
128-135.
Chapman, H. C. 1962. A survey for autogeny in some Nevada mosquitoes. Mosquito News
22: 134-136.
Chapman, H. C. and A. R. Barr. 1964. Aedes communis nevadensis, a new subspecies of
mosquito from western North America. Mosquito News 24: 440-447.
Christophers, S. R. 1911. The development of egg follicles in Anopheles. Paludism 2: 73-88.
Clark, J. C. and F. C. Wray. 1967. Predicting influxes of Aedes vexans into urban areas.
Mosquito News 27: 156-163.
Clement, A. N. 1963. The physiology of mosquitoes. Pergamon Press, Oxford. 393 pp.
Corbet, P. S. 1961. Entomological studies from a high tower in Mpanga forest, Uganda,
VIII: The age composition of biting mosquito populations, according to time and level.
Trans. R. ent. Soc. Lond. 1 13: 336-345.
Curtis, L. C. 1967. The mosquitoes of British Columbia. Occasional papers, British Columbia
Provincial Museum No. 15. 90 pp.
de Meillon, B. and Z. H. Khan. 1965. Examples of the use of simple age grading in the
assessment of Culex fatigans populations. W.H.O./V.C. /Sem./W.P./25. 64. (mimeographed).
Fisher, R. A., A. S. Corbet and C. B. Williams. 1943. The relation between the number of
individuals and the number of species on a random sample of an animal population. J.
anim. Ecol. 12: 42-58.
Gjullin, L. M., W. W. Yates and H. H. Stage. 1950. Studies on Aedes vexans (Meig.) and
Aedes sticticus (Meig.), flood water mosquitoes in the lower Columbia River valley. Ann.
ent. Soc. Am. 43: 262-275.
Graham, P. 1969. A comparison of sampling methods for adult mosquito populations in
central Alberta, Canada. Quaest. ent. 5: 217-261.
Hamon, J., S. Sales, J. P. Adam and P. Grenier. 1964. Age physiologique et cycles d’agres-
servite chez Anopheles gambiae et A. funestus dans la region de Bobo-Dioulasso (Haute
Volta). Bull. Soc. ent. Fr. 69: 110-121.
Happold, D. C. B. 1965a. Mosquito ecology in central Alberta. I. The environment, the
species and studies of the larvae. Can. J. Zool. 43: 795-819.
Happold, D. C. B. 1965b. Mosquito ecology in central Alberta. II. Adult populations and
activities. Can. J. Zool. 43: 821-846.
Hearle, E. 1929. The life history of Aedes flavescens (Muller). Trans. Roy. Soc. Can. Sect.
V. Ser. 3. 23: 85-101.
Hocking, B. 1954. Flight muscle autolysis in Aedes communis (de Geer). Mosquito News 14:
121-123.
Holstein, M.H. 1954. Biology of Anopheles gambiae. Monogr. Wld. Hlth. Org. No. 9: 1 72 pp.
Horsfall, W. R. 1955. Mosquitoes; their bionomics and relation to disease. Ronald, New
York. 723 pp.
Howard, L. O., H. G. Dyar and F. Knab. 1915. The mosquitoes of North and Central
America, Vol. III. Carnegie Institute, Washington, Publ. No. 1 59. 523 pp.
Jenkins, D. W. and K. L. Knight. 1950. Ecological survey of the mosquitoes of Great Whale
338
Graham
River, Quebec. Proc. ent. Soc. Wash. 52: 209-223.
Khelevin, N. W. 1958. The effects of environmental factors on the embryonic diapause and
on the number of generations in a season of Aedes caspius dorsalis Mg. (Diptera: Culi-
cidae). Effect of temperature on embryonic diapause in Aedes caspius dorsalis Mg. Ent.
Rev. 37: 19-35.
Klassen, W. 1959. The influence of the North Saskatchewan river on the dispersion of Aedes.
Thesis. University of Alberta, Edmonton. 120 pp.
Klassen, W. and B. Hocking. 1963. Control of the dispersion of Aedes along a deep river
valley. Mosquito News 23: 23-26.
Marshall, J. F. 1938. The British mosquitoes. British Museum (Nat. Hist.) London. 341 pp.
Matheson, R. 1944. Handbook of the mosquitoes of North America. Comstock Publishing
Co., Ithaca, N. Y. 314 pp.
Means, R. G. 1965. Culex territans Walker biting man in nature. Mosquito News 25: 489.
Muirhead-Thompson, R. C. 1951. Mosquito behavior in relation to malaria transmission in
the tropics. Arnold, London. 219 pp.
Nielsen, E. T. and D. M. Rees. 1961. An identification guide to the mosquitoes of Utah.
University of Utah, Biological Series 12(3), 63 pp.
Pucat, A. 1964. Seven new records of mosquitoes from Alberta. Mosquito News 24: 419-
421.
Pucat, A. 1965. List of mosquitoes from Alberta. Mosquito News 25: 300-302.
Rempel, J. G. 1950. A guide to the mosquito larvae of western Canada. Can. J. Res. D.
28: 207-248.
Rempel, J. G. 1953. The mosquitoes of Saskatchewan. Can. J. Zool. 31: 433-509.
Shelenova, M. F. 1959. The biology of the principal series of Aedes in the forest zone
of the European part of the U. S. S. R. Med. Parazit. (Mosk.) 28 : 456-462. (In Russian,
English summary).
Shemanchuk, J. A. 1965. On the hibernation of Culex tarsalis Coquillett, Culiseta inornata
(Williston) and Anopheles earlei Vargas (Diptera:Culicidae) in Alberta. Mosquito News
25: 456-462.
Skiersca, B. 1965. Ecological studies on the occurrence and distribution of the Culicine
fauna in the coastal forest belt. Ekologia Polska A. 3: 527-573.
Stage, H. H., C. M. Gjullin and W. W. Yates. 1938. Flight range and longevity of flood water
mosquitoes in the lower Columbia River valley. J. econ. Ent. 30: 940-945.
Standfast, H. A. 1965. A miniature light trap which automatically segregates the catch into
hourly samples. Mosquito News 25: 48-53.
Steward, C. C. and J. W. McWade. 1961. The mosquitoes of Ontario (Diptera:Culicidae),
with keys to species and notes on distribution. Proc. ent. Soc. Ont. 91 : 121-188.
Stone, A. 1965. Family Culicidae, pp. 105-120. In A. Stone, C. W. Sabrosky, W. W. Wirth,
R. H. Foote, and J. R. Coulson (Editors). Agriculture Handbook No. 276. 1696 pp.
Stone, A., K. L. Knight and H. Starke. 1959. A synoptic catalogue of the mosquitoes of the
world. Thos. Say Foundation. 358 pp.
Vockeroth, J. R. 1952. The specific status of Aedes pionips Dyar (Diptera: Culicidae). Can.
Ent. 84: 243-247.
Biology of Mosquitoes
339
Vockeroth, J. R. 1954. Notes on the identification and distributions of Aedes species of
northern Canada, with a key to the adult females (Diptera:Culicidae). Can. Ent. 86:
241-245.
Wada, Y. 1965. Population studies of Edmonton mosquitoes. Quaest. ent. 1: 187-222.
Wesenberg-Lund, C. 1921. Contributions to the biology of Danish Culicidae. Mem. Acad.
Roy. Sci. et Lettres, Copenhagen, Sec. Sci. 8th ser. No. 1. 210 pp.
Williams, C. B. 1964. Patterns in the balance of nature. Academic Press, London. 324 pp.
OUTBREAKS OF THE BLACK FLY
SIMULIUM ARCTICUM MALLOCH IN ALBERTA*
F. J. H. FREDEEN Quaes tiones Entomologicae
Entomologist 5: 341-372 1969
Canada Agriculture Research Station
University Campus
Saskatoon, Saskatchewan
The subgenus Gnus Rubzov is represented in Alberta by at least four species. One of
these, Simulium arcticum Mall., is the only species of black fly known to kill large farm
animals in the province. It is widely distributed, the immature stages occurring in most or all
streams and rivers originating on the slopes of the Rocky Mountains and other elevated areas
such as the Swan Hills. Outbreaks that have killed animals are known from only two areas.
Near Minburn, east of Edmonton, a few cattle were killed and people suffered serious effects
from bites in June of 1956 and in late May of 1961. Near Athabasca, north of Edmonton,
outbreaks occur throughout a period of several weeks every summer, disrupting grazing and
breeding activities and reducing production of milk and beef. A few cattle were killed there
in 1955 or 1956, and in June of 1963 and 1964.
Although bulls are most affected, no farm animals are exempt from attack. Initial attacks
are usually so sudden and violent that animals are damaged before they can be taken in-
doors. Humans are relatively immune to attack but when bitten sometimes require medical
attention or even hospitalization.
Other species of the subgenus Gnus known to occur in Alberta are S. corbis Twinn and
S. malyshevi D. R. and V. in west-central and northern areas and S. defoliarti S. and P. in
the extreme south-west. S. nigricoxum S. is not known outside of the Northwest Territories
and Alaska. Damaging outbreaks of S. defoliarti occur in south-central British Columbia but
not in Alberta.
DISTRIBUTION OF 5. ARCTICUM IN ALBERTA
Simulium arcticum Malloch is widely distributed in Alberta (Table 1 and Fig. 1 ). In addi-
tion to the records in Table 1 , Hearle (1932) reported that S. arcticum was abundant in the
Athabasca River at Jasper (no date), Strickland (1938) listed it as occurring on July 19 in
the Jasper area (possibly Hearle’s collection) and Abdelnur (1968) reported on some aspects
of its life history and habits in the Pembina and Athabasca Rivers at Flatbush.
Although larvae and pupae were encountered throughout the entire collecting season
(mid-May to early September) the period of greatest abundance on the plains was June 5 to
15, and in the mountains, early July. Future surveys in greater detail will undoubtedly prove
that these periods of peak abundance vary from year to year. In Saskatchewan the immature
stages have been collected in almost every month of the year although the species normally
overwinters as eggs (Fredeen et al., 1951) and larvae and pupae attain greatest abundance
between mid-May and mid-June each year.
* Contribution No. 349, C. D. A. Research Station, Saskatoon.
342
Fredeen
TABLE 1 . Streams and rivers in Alberta and the Peace River District in British Columbia, from which the immature stages of Simulium arcticum
Mall, were collected.
Black Fly
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Site* Location Date Abundance Stages of development
344
Fredeen
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346
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Black Fly
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The subgenus Gnus (Rubzov, 1940) to which S. arcticum belongs is holarctic. S. arcticum,
however, is restricted to western North America, occurring throughout the mountainous
regions from Alaska south to California. Specimens have been collected as far east as
Churchill, Manitoba (Twinn et al. , 1948). In Saskatchewan, farm animals have been killed in
numerous outbreaks originating in the Saskatchewan River system (Cameron, 1922; Rempel
and Arnason, 1947; Fredeen, 1958). The most destructive series of outbreaks in Saskatche-
wan occurred 1944 to 1947 inclusive when more than 1100 animals were killed. Some
aspects of the life cycle of this black fly other than these outbreaks are described by
Fredeen (1958, 1 963), Fredeen et al. (1951) and Abdelnur ( 1 968). In Alberta, livestock has
been killed in at least five outbreaks (Table 2).
The immature stages of three other species of the subgenus Gnus were also collected in
Alberta and northern British Columbia but these species have never been implicated in
damaging outbreaks in Alberta. Specimens of S. corbis Twinn were obtained from a few
streams and rivers in northern areas and the west-central foothills as follows:
Assineau Creek at Canyon Creek (July 3, 1950, May 31 and June 23, 1961); Fawcett River, Smith (May 31, 1961);
Sucker Creek, High Prairie (May 31, 1961); Sweeney Creek, 10 miles S.W. of Clear Prairie (June 16, 1961); Clear Creek,
Clear Prairie (June 16, 1961); Wagner Creek, Widewater (June 23, 1961); Sakwatamau River, Whitecourt (June 5, 1963);
Ksituan River, Gordondale (June 7, 1963); Chambers Creek and Shunda Creek, Horburg (June 7, 1963); Haven Creek,
Nordegg (July 4, 1963); Kiskatinaw River, Dawson Creek, B. C. (June 15, 1961); Buffalo Creek, Progress, B. C. (June 6,
1963).
S. defoliarti was collected only in the foothills in the S. W. corner of the province:
Belly River, W. of Mountain View (August 28, 1962; August 10, 1953); Highwood River, W. of Longview (August 8,
1954); Castle River, W. of Pincher Creek (May 7, 1955); Crowsnest River, N. of Lundbreck (May 7, 1955).
In the Shuswap River in south-central British Columbia this species breeds in such large
numbers that chemical control of the larvae is occasionally required. Severe outbreaks in
1951 affected gains in beef animals, causing losses in excess of 24,000 dollars (Curtis, 1954).
However, in Alberta it is not known to occur in nuisance numbers. S. malyshevi D. R. and
V. was collected as follows:
Clearwater River, Waterways (June 18, 1948); E. Prairie River, Enilda (June 3, 1961); Goose Creek, Calais (June 6,
1961); Beaton River, Cecil Lake, B. C. (June 16, 1961).
A fifth species of the subgenus Gnus, S. nigricoxum S. may also occur in the north end of
the province as it is widely distributed in the Northwest Territories and Alaska.
Samples of the immature stages of these black fly species were obtained by wading into
the margins of rapids to pick up rocks and tree branches by hand. Equipment has not yet
been devised that will allow the river bed to be quantitatively sampled in deep, fast-flowing
rapids. Since the levels of mountain-fed rivers are relatively unstable, even at distances of
1000 miles or more from their sources, and since the margins of these rivers advance and
retreat irregularly according to rates of snow melt and precipitation in the watersheds, our
marginal samples served only to indicate the presence or absence of S. arcticum. Seldom
were water levels low enough to allow examination of infestations of larvae and pupae in the
relatively permanent mid-river sites. However, on June 5, 1963, low levels on the McLeod
River at Whitecourt and the Waskahigan and Little Smoky Rivers at Little Smoky, exposed
dense infestations of larvae and pupae, approximately similar to those sometimes seen in
both branches of the Saskatchewan River in Saskatchewan prior to damaging outbreaks.
Moderately dense infestations were discovered in several other large rivers including the
TABLE 2. Damaging black fly outbreaks in Alberta attributed to Simulium arcticum Mall.
348
Fredeen
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Black Fly
349
North Saskatchewan River at Rocky Mountain House on June 12, 1949 and the Athabasca
at Fort Assiniboine on July 14, 1964. Eight per cent of the S. arcticum larvae in the Atha-
basca River in mid-July of 1964 were parasitized with mermithid nematodes, but nowhere
else were these parasites observed.
SOME CHARACTERISTICS OF THE
NORTH SASKATCHEWAN AND ATHABASCA RIVERS
These two rivers and their major tributaries flow in vegetation-free beds consisting mainly
of boulders, gravel and clean sand. The growth of vegetation is effectively prevented by the
eroding effect of the ice during the spring breakup and by frequent flood crests during the
summer.
In the upper reaches, gradients are relatively steep and boulders and gravel predominate;
on the plains the gradient is flatter, often averaging only about one foot per mile, and here
the river bed consists mainly of sand. In some regions of the plains, however, the gradients
are much steeper than this and the rivers flow in series of rapids over gravel and boulders.
These boulders provide favoured attachment sites for enormous numbers of S. arcticum
larvae and pupae. An outstanding example is a rock-filled weir across the North Saskatche-
wan River at Prince Albert, Saskatchewan which on June 9 and 10, 1947 was estimated to
contain more than 7 x 109 pupae (Fredeen, 1958).
On the North Saskatchewan River in Alberta there are rapids north of Willingdon and
Myrnam, north to north-west of the outbreak area near Minburn (Fig. 1). On the Athabasca
River there are numerous rapids between Whitecourt and Smith, a few scattered rapids from
Smith down to about the Pelican Portage Settlement and very numerous rapids and falls
downriver from there to Fort McMurray. This latter section of the river has an average
gradient of about 5.4 feet per mile and, although it was not sampled during these investiga-
tions, was considered likely to be a major breeding site for S. arcticum. It is 65 miles and
more, directly north of the center of the annual outbreak area in the county of Athabasca.
The average ice-free period in these two rivers lasts from 6.5 to 7.0 months (Table 3). Ice
breakup in the spring generally begins when the river level rises sharply as a result of melting
snow on the plains and foothills. Following this crest the level generally subsides irregularly
until a new major crest occurs in late June or early July as a result of rapid thawing and pre-
cipitation in the mountains. Maximum volume discharges ranging up to about 200,000 cu
feet/sec have been recorded on both rivers and minimum discharges of less than 2000 cu
feet/sec generally occur in the winter.
Water temperatures during the summer in these two rivers are likely to be a few degrees
lower than those on the North Saskatchewan at Prince Albert in Saskatchewan where
maxima of 70 to 75°F are attained in July and August. There the larvae and pupae of S.
arcticum attain greatest abundance in May and June when water temperatures range from
45 to 65°F.
Water turbidity increases with increases in the water level. Values greater than 3000 ppm
have been measured on the South Saskatchewan River. This turbidity has been used to
advantage during black-fly larviciding (Fredeen et al. , 1953).
350
Fredeen
TABLE 3. Some characteristics of the North Saskatchewan River at Edmonton, Alberta.
and the Athabasca River at Athabasca, Alberta. (Canada Department of Energy,
Mines and Resources, 1967).
Drainage area (sq. miles)
Average annual discharge
(ft3 /sec)
Maximum discharge
(ft3 /sec)
Minimum discharge
(ft3 /sec)
Average ice-free period
(months)
North Saskatchewan River
10,500
7,660 (54 years)
204,500 (June 28, 1915)
220 (Jan. 1, 1940)
7.0 (April 13 to Nov. 10)
Athabasca River
29,600
15,100 (30 years)
199,600 (June 10, 1954)
1 ,610 (December 14, 1956)
6.5 (April 21 to November 8)
SOME FACTORS INVOLVED IN THE
DEVELOPMENT OF DAMAGING OUTBREAKS
Rempel and Arnason (1947) suggested that the appearance of a massive swarm of S.
arcticum was due simply to mass emergence plus fortuitous winds. Additionally there is
recent evidence that the first ovarian cycle may be completed autogenously; in some in-
stances at least, the females do not attack animals for blood until after the eggs from the
first ovarian cycle have been laid (Fredeen, 1963). This means that emerging females can
accumulate, perhaps throughout a period of several days, awaiting weather conditions suit-
able for oviposition. This could result in the release of enormous numbers of blood-thirsty
black flies within a period of a very few hours. Perhaps both factors have been responsible
in the development of damaging outbreaks.
Winds play a very important part in the distribution of adult black flies. Variable-direc-
tion winds will scatter them over a wide sector, thinly enough that they will obtain their
blood meals unnoticed. However, livestock owners should be wary when the wind blows
steadily from proven breeding grounds for even a few hours. In one Saskatchewan outbreak,
livestock was killed 140 miles downwind from such a breeding site. In a few other instances
an abrupt shift in the wind direction has been known to transfer black flies from one area
into another, with resultant fatalities in both (Rempel and Arnason, 1947).
Black Fly
351
The appearance of large numbers of black flies in a district is generally sudden and
furthermore often occurs in the early morning or late evening so that a severe attack can
begin before livestock owners are aware of their presence. Also, reactions to massive injec-
tions of the toxin are swift, and the animals can become fatally ill within a few hours after
the black flies commence their attack.
The nature of the toxin is poorly known. Hutcheon and Chivers-Wilson (1953) showed
that the salivary extract contained an anticoagulant and a substance that gave reactions
similar to those from histamine. A detailed account of symptoms in cattle is given by Millar
and Rempel (1944) who investigated an outbreak of S. arcticum in the Macdowall area in
Saskatchewan. They reported that residents in the district noticed the appearance of black
flies in the district on the evening of May 30 but did not consider them to be unusually
abundant until the following morning. Animals subjected to heavy attack throughout May
31 developed fluid-filled swellings along the underlines. Some additionally developed a
heavy, jerky breathing accompanied by a strong trembling of the muscles. Animals with
such symptoms either died within 1 5 minutes to two hours, or made a complete recovery
within 48 hours. Deaths were attributed to acute toxemia but anaphylactic shock was not
ruled out.
TYPES OF LOSSES TO BE EXPECTED
DURING OUTBREAKS OF 5. ARCTICUM
Fatalities
The numbers of fatalities in livestock that have occurred in Alberta as a result of black fly
outbreaks are small when compared with losses during the large outbreaks in Saskatchewan
in the 1 940s. However, about half of the losses were those of herd sires (generally the most
expensive animals in the herds). Mature bulls are often more attractive to blood-sucking flies
than are cows or calves. Furthermore, imported bulls show less resistance to attack and
damage than do native bulls, especially those allowed to spend most of their time out-of-
doors. However, whether imported or native, at the first signs of an outbreak the herd sires
require special attention and should be stabled or at least kept under close observation.
Some veterinarians were reluctant to agree that these livestock fatalities in Alberta re-
sulted from black fly attacks. Although I did not personally observe any of the fatalities, I
agree with the conclusions reached by the owners, none of whom were preconditioned in
their thinking by being aware of similar occurrences in Saskatchewan.
Suspension of breeding activities
During black fly outbreaks, breeding activities are interrupted when the cattle become
involved in protective activities, i.e. seeking shelter in buildings, dense brush and sloughs.
This suspension of breeding activities results in an irregular and delayed calf crop the follow-
ing year, thus decreasing net returns from the enterprise. Also, individuals of S. arcticum
normally attack along the underline of an animal and the bull’s sheath thus becomes in-
352
Fredeen
flamed and may develop secondary infections requiring treatment by a veterinarian. Some
bulls remain permanently sterile despite treatment.
Declines in the production of milk and beef
These phenomena are frequently observed by livestock owners but losses are difficult to
measure. These losses naturally occur when normal pasture grazing is interrupted and physi-
cal activities are increased. Additionally, however, a milking cow’s udder is a favourite
target for black flies and this makes extraction of milk a more difficult process, either for
the nursing calf or for man. Reports from owners such as “black flies drove cattle out of the
pasture”, “black flies kept cattle in shed all day”, “cows’ udders red with blood”, “cows
difficult to milk because of black flies”, “milk production down something terrible”, are
commonplace during outbreaks.
General losses resulting from repeated threats of outbreaks
A loss that is seldom recognized in an area subjected to frequent, severe black fly out-
breaks is the general shift from livestock to alternate farm enterprises. In the Athabasca
region of Alberta livestock enterprises should predominate to ensure the healthiest eco-
nomic development of the region because of the relatively short frost-free season, the rough
terrain with much marginal land suitable only for pasture and forage crops, and soils that
require crop rotations for best productivity. However, some residents have either reduced or
eliminated their livestock enterprises and certain highly skilled breeders have even emigrated
to other districts where they will not be threatened by black flies. These shifts do not seem
to be warranted in terms of measurable losses that have occurred but rather can be attrib-
uted to the general suspense created by unpredictable and uncontrolled outbreaks.
The moderate numbers of black flies that occur almost daily during the summer can be
tolerated by men and animals without too much discomfort. However, increases in the
numbers of black flies can generally be expected whenever the wind blows steadily for a
few hours from breeding grounds in the river and occasional outbreaks have also occurred
when the wind has blown from some direction other than the river (Fig. 2). Since winds
often shift during the night, and black fly flight activity is generally greatest in early morn-
ing and late evening hours, many severe outbreaks have begun with surprising suddenness.
Sometimes the first indication of an outbreak has been the stampede of cattle from pasture
to barn. Less fortunate operators have their livestock scatter into the brush where they are
less likely to be protected. Thus a strong element of suspense prevails throughout the out-
break season.
Effects on man
Although people can generally protect themselves with repellents, there are occasional
days in the County of Athabasca when people are driven indoors along with their cattle to
escape savage attacks. Normally individuals of S. arcticum do not attack man but when
attacks occur, medical attention may be required to alleviate severe swelling and itching of
the affected limbs.
Black Fly
353
DESCRIPTIONS OF OUTBREAKS OF
5. ARCTICUM IN ALBERTA
Outbreaks adjacent to the North Saskatchewan River
A black fly outbreak in the Municipal District of Minburn No. 72 was investigated on
July 9 and 10, 1961 at the request of Mr. J. B. Gurba, Supervisor of Crop Protection and
Pest Control for the Alberta Department of Agriculture. The outbreak had actually occurred
during the third week in May but was of such short duration that by the time the black flies
had been identified as S. arcticum conditions had already returned to normal. Thus the
purpose of my visit in July was to determine the extent and severity of the outbreak by
interviewing livestock owners. The following were interviewed at a Municipal Council meet-
ing and others on their farms: Mr. C. Gamble, a local livestock feeder and Field Supervisor
for the Municipal District, Mr. Ed. McLaughlin, farming about 6 miles N. W. of Mannville,
Mr. A. W. Roland, about 5 miles S. of Minburn and Mr. G. Grabos, Vi mile S. of Innisfree.
They stated that black flies were pests of mammals, especially milking cows, every year for
a few days around the third week in May. Outbreaks were reported to have been especially
severe in dry years, the worst outbreaks having occurred in 1956 and 1961 when an area of
354
Fredeen
about 900 square miles centering on Mannville was affected (Fig. 1 ). During each of these
two outbreaks a few cattle were reported to have been killed and the udders of milk cows
were red with blood. Among others, Mr. McMillan farming east of Mannville lost cattle dur-
ing the 1956 outbreak.
Perhaps the most serious aspect of these two outbreaks was the requirement by many
people for medical attention as a result of complications following black fly bites. This has
also been a common feature of S. arcticum outbreaks in Saskatchewan. Dr. Hasinoff of the
Municipal Hospital at Minburn informed me that he had treated many people for black fly
bites in 1956. They had come from an area 30 to 40 miles in diameter surrounding Mann-
ville and Minburn. Black flies collected while attacking people were all S. arcticum.
Two men in particular had had such severe reactions in 1956 that they required long
medical attention and one of these, a Mr. S., was still unwell in 1961. The following symp-
toms, and the results of clinical tests in Edmonton, led Dr. Hasinoff to believe that Mr. S.
exhibited a distinct case of Arthus’ syndrome (Brown et al., 1938) or localized anaphylactic
reactions following sensitization by injections of black fly toxins. Dr. Hasinoff stated that
Mr. S. reported to him on June 10, 1956 in respiratory distress, itchy and with evidence of
black fly bites. Later he developed ulcers down to the bone on limbs that had not been
bitten. These were not due to infection but could only be ascribed to allergic reactions to
the black fly toxins. Mr. S. spent 5 months in the hospital before the ulcers finally healed.
Gudgel and Grauer (1954) and others have described severe reactions in humans that
necessitated medical treatment at the actual locations of black fly bites, but this may be the
first case where reactions have been observed in areas of the body remote from locations
where the bites occurred.
In 1962, May 22 to 25, another outbreak of S. arcticum occurred in the Mannville area
but this was of moderate intensity. By the time we received word of it and visited the area
on May 26 to 30 the outbreak had ended. However, specimens collected from barn windows
were all S. arcticum. On May 26, only a few adults could be collected around cattle and
these were a mixture of S. arcticum and S. venustum Say. The weather became cool and wet
about this time and individuals of these two species disappeared. Near Two Hills and
Myrnam a few specimens of Cnephia saskut chew ana S. and F. were collected flying around
cattle but none were actually seen on the cattle.
Since 1962 no further reports of black fly outbreaks have been received from this district.
On May 22, 1963 only a few S. venustum and S. vittatum Zett. were found around cattle.
These outbreaks of S. arcticum all occurred following northerly winds and thus the
extensive rapids on the North Saskatchewan River at several sites north of Willingdon and
Myrnam may have been the sources (Fig. 1 ). I have never examined these rapids for larvae
or pupae because of high water levels at the times I was in the area. However, on several
occasions I found abundant S. arcticum larvae in the North Saskatchewan River or its tribu-
taries above Edmonton (Table 1) and also at several sites in the province of Saskatchewan.
Individuals of this species have never been collected from the Vermilion or Battle Rivers
despite several thorough searches. On the other hand species such as S. venustum breed
in these smaller rivers but not in the Saskatchewan River. Experience has shown that indivi-
duals of S. venustum can be annoying to animals and man living within a half mile of their
Black Fly
355
breeding places but not far beyond that distance. Individuals of S. arcticum are known to
have a much longer flight range and thus when animals have been killed or damaged several
miles from the nearest potential breeding site, S. arcticum rather than S. venustum has
always been the culprit.
The possibility should not be overlooked that the Athabasca River rather than the
Saskatchewan may have been the source of the black flies in the outbreaks in the Minburn
area. This would have required flights of about 150 miles, but in one outbreak of S.
arcticum in Saskatchewan, livestock were killed by black flies that had been carried by the
wind more than 140 miles from their point of origin (Fredeen, 1958). Two separate sources
however are suggested by the differences in the seasons of the outbreaks, those at Minburn
occurring between late May and early June, and those at Athabasca between late May and
July and occasionally August or September.
In the North Saskatchewan above Edmonton, S. arcticum larvae and pupae are sometimes
abundant but severe outbreaks have never been reported from that area. A farmer near
Rocky Mountain House reported in 1949 that black flies were common and troublesome
throughout June, July and August every year. He had also observed swelhngs under the jaws
of horses during the black fly seasons and reported that a disease locally called “swamp
fever” used to kill many horses. Horses newly brought into the area were said to have been
particularly susceptible to swamp fever. The symptoms included swellings under the belly,
bleeding from the mouth and nostrils and rapid mortalities. These symptoms suggest black
fly damage but for the fact that fatalities in cattle did not occur according to the reporter.
Outbreaks adjacent to the Athabasca River
Residents in the County of Athabasca report that since the earliest years of settlement,
annual outbreaks of “sand flies” or black flies have occurred. They were believed to origin-
ate in the large swampy areas near the Athabasca and La Biche Rivers. Although they make
their first appearance during northerly winds in late May or in June and often persist into
September, the swarms doing the greatest damage are always exacted in late June or early
July.
Livestock has been killed apparently only in three years, 1955 (or 1956), 1963 and
1964. However, every year livestock productivity is affected to some degree. These out-
breaks were regarded as uncontrollable events until 1963, following establishment of a
County Agricultural Service Board. When an unusually severe outbreak occurred in 1963,
the County Agricultural Fieldman, Mr. H. Armfelt, recognized the problem as one that
required immediate attention. He obtained the assistance of Mr. J. B. Gurba and on July 12
they surveyed the outbreak area and obtained the following information (Gurba, 1963). The
“sand flies” had appeared suddenly that year after the first warm period in June. Arriving
on northerly winds they caused greatest damage in an area measuring about 20 miles north
and south by 5 miles east and west, centered on Grassland and Spruce Valley. Scattered
instances of attack occurred as far south as Boyle and Newbrook. Two bulls and four calves
were killed by the black flies about June 1 5 and many other animals became ill, some re-
quiring a veterinarian’s attention. Livestock and people new to the area were most severely
affected. Insecticide sprays and smudges gave some relief to milking cows but in general
356
Fredeen
milk and beef production declined noticeably during June and July. Black flies collected
from cattle on July 12 were all S. arcticum. As a result of this outbreak I was asked to help
locate breeding areas and recommend control measures.
In July of 1964 a careful examination of the Athabasca River between Whitecourt and
Smith proved the existence of numerous rapids well populated with S. arcticum larvae and
pupae. Unfortunately a sudden rise in the water level prevented examination of rapids below
Smith at that time.
Also in July of 1 964, a number of livestock owners in the eastern half of the County of
Athabasca were interviewed and their reports showed that another extensive outbreak had
occurred. Black flies had been first seen around cattle on May 30 after an all-day wind from
the north. However, June 10 was the first day that the cattle were noticeably irritated by
the black flies. On June 16 a new influx of black flies on a north wind forced man and live-
stock alike to remain indoors and killed at least one bull. This appearance of the first dam-
aging swarms of the year during the first warm weather in mid-June was said to be typical.
For about three weeks in 1964 these attacks continued to affect normal grazing and breed-
ing activities, milk production and weight gains.
Black flies collected alive from cattle and horses in many localities in the County on July
18 and 19, 1964 were 92% S. arcticum, 6%S. venustum and 2 %S. vittatum. Although these
black flies were moderately abundant around cattle, the latter were grazing normally in the
pastures. Collections of dead flies from the windows of a barn six miles northwest of Grass-
land, presumably trapped during the spring outbreak of 1964, contained 99% S. arcticum
and 1% S. venustum. Thus although S. arcticum seemed to be the main species involved in
the outbreaks of 1964, precise information as to the relative importance of it and other
local species throughout all spring and summer outbreaks was still lacking. This information
was obtained in two ways: by a widespread survey for the immature stages in streams and
rivers in and near the County in 1964, 1966 and 1967, and by collecting adults from wide-
spread attacking swarms throughout the entire outbreak seasons of 1966 and 1967.
The area in and around the County of Athabasca is traversed by many small streams, a
few small rivers such as the La Biche and Wandering Rivers, and one large river, the Atha-
basca (Fig. 2). Early in these investigations it was thought that S. venustum , especially from
the La Biche and Wandering Rivers, might have been at least partly responsible for the
severe outbreaks in nearby farmlands almost directly south of these rivers. The survey of
rivers and streams showed that S. venustum was actually widespread, but abundant in only
two streams, Pine Creek (May 31, 1967) and Wandering River (May 18 and June 7, 1966,
and May 31, 1967) (Table 4). Other species whose immature stages were also widespread
included S. verecundum S. and J. (a close relative of S. venustum although apparently non-
biting (Stone and Jamnback, 1955) ), S. vittatum and S. tuberosum (Lund.). Individuals of
S. vittatum are large and grey, commonly seen in the ears of livestock, but are not usually
considered to be serious pests. They were occasionally abundant in the La Biche River,
Calling River and Pine Creek. S. tuberosum, although widespread, was never abundant. S.
arcticum was abundant only in the Athabasca River. Insignificant numbers were found
breeding in Calling River and Deep Creek but these sources were too small to have contrib-
uted significantly to the outbreaks.
TABLE 4. Species of black fly larvae and pupae found in rivers and streams in Athabasca County and its environs, 1964 to 1967 inclusive
Black Fly
357
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May 18 and June 7, 1966, May 31, 1967
S. verecundum S. and J. Widespread, June and July; abundant in Bear Creek and Deer Creek, June 16, 1966
S. vittatum Zett. Widespread, May to October inclusive; abundant in La Biche River, July 19, 1964 and May 17,
1 966, Calling River, July 18,1 964 and Pine Creek, July 2 1 , 1 966
358
Fredeen
Altogether the immature stages of 14 species were discovered. Of these, S. arcticum,
because of the immensity of its breeding site, was considered to have the greatest potential
for developing large, damaging swarms.
Despite these extensive collections of the immature stages, decisive evidence as to the
actual species involved in the outbreaks was obtained only with the co-operation of 1 4 resi-
dents, situated in and near the County (Table 5, Fig. 2) who netted more than 46,000 black
flies from swarms attacking their livestock in 1966 and 1967 (Table 6). At some of the sites
black flies were netted every three days or so (oftener during outbreaks), throughout much
of the spring and summer flight periods; at other sites, collections were taken only during
prominent outbreaks.
In 1966, S. arcticum represented 92.2% of all black flies netted and in 1967, 89.6%. S.
venus turn appeared in moderate abundance on only one occasion, May 31, 1967, (site no.
14, Fig. 2) and S. vittatum on one occasion, September 17, 1967 (site no. 9, Fig. 2). At all
TABLE 5. Collectors and observers of black fly activity in and near the County of Atha-
basca in 1966 and 1967.
Years of
observations
Reference symbol
(See map, Fig. 2) 1966 1967
Direction of winds re-
quired to bring black flies
from major breeding sites
Name on the Athabasca River
A
B
C
D
E
F
G
H
I *
J *
K *
L*
M
N
* These sites are north of the La Biche River but, except for L, are located near the Wander-
ing River.
Black Fly
359
TABLE 6. Species of black flies collected from swarms near farm animals, in and near the
County of Athabasca*, Alberta, 1966 and 1967
1966 1967
* See Fig. 2 for locations of the collection sites.
**Some of the S. arcticum were collected from barn windows after detaching from livestock
that had carried them into the buildings.
other times only relatively small numbers representing species other than S. arcticum were
collected (Table 6). Thus it seems certain that the several severe and sometimes damaging
outbreaks in 1966 and 1967 were caused by individuals of S. arcticum , only.
Although the immense numbers of S. arcticum observed could only have had their origin
in a large river, i.e. the Athabasca, additional evidence as to the sources of these outbreaks
was obtained by relating the times of the outbreaks as reported by the co-operators, to
hourly wind data obtained from the Canada Department of Transport Meteorological Sta-
tion at the Lac La Biche Airport located 29 miles east and four south of Grassland P. O.
These data are listed in Tables 7 and 8. Almost invariably, each fresh invasion of black flies,
as indicated by a distinct increase in the severity of attack, was found to have been preceded
by winds blowing from some section of the Athabasca River (Fig. 2). On a few occasions,
however, the wind had been blowing from some other direction, or had been virtually calm.
Perhaps some of these anomalies were due to the fact that the wind and black fly data came
from sites separated by 30 miles or more. However, individuals of S. arcticum have been
observed to move upwind for short distances during outbreaks in Saskatchewan. In one
instance they seemingly moved 1.5 miles against a wind strong enough to raise dust from the
TABLE 7. Wind conditions prior to outbreaks of S. arcticum Mall, in and near the County of Athabasca, Alberta, in 1966
360
Fredeen
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Black Fly
361
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July 15 1900 B From cattle 112 0 (wind light, SE, NE)
TABLE 7. (continued)
362
Fredeen
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TABLE 7. (continued)
Black Fly
363
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**Wind data were obtained from the Canada Department of Transport, Meteorological Branch Station at Lac La Biche Airport, Alberta.
TABLE 8. Wind conditions prior to outbreaks of S. arcticum Mall, in and near the County of Athabasca, Alberta, in 1967
364
Fredeen
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June 22 0730 D Yard was full of black flies 385 102 NW 435
June 22 1600 D Flies came by the millions 262 110 NW 496
June 23 1200 E Many flies, mostly around noon 42 130 NW 514
Black Fly
365
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366
Fredeen
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July 3 0745 J Around cows in yard 248 25 S
Data from observers Number of hours that
the wind** had been
Time of report blowing from the Net wind vector**
and/or collection No. of S. Athabasca River
Black Fly
367
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July 1 1 0700 A From cattle in yard 62 0 (calm)
July 11 0730 E Lots of sand flies around cattle 404 61 WNW 440
July 11 1030 M From barn window 50 5 ENE 11
Data from observers Number of hours that
368
Fredeen
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July 17 0830 N Around cattle in yard 374 57 SE 164
July 18 0830 F Flies have been quite bad last few days 844 9 NW 27
July 18 0900 M Flies from barn window 59 0 (light N winds)
July 19 1815 A From cattle in yard 80 1 SSE 5
July 19 0800 E Many flying and feeding 192 6 N 5
Data from observers Number of hours that
the wind** had been
Time of report blowing from the Net wind vector**
and/or collection No. of S. Athabasca River
Black Fly
369
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** Wind data were obtained from the Canada Department of Transport, Meteorological Branch Station at Lac La Biche Airport, Alberta.
370
Fredeen
fields. The livestock they attacked was pastured near the edge of a wooded valley connected
to the Saskatchewan River Valley and the black flies presumably made most of their upwind
approach in the shelter of the valley walls and trees.
The annual outbreaks in the County of Athabasca are of unusual duration. Outbreaks at
Minburn and in the Province of Saskatchewan last for only a few days whereas at Athabasca
they often last for three or four weeks. For instance, in 1966 an outbreak began with
typical violent suddenness on July 2 and continued with slowly declining severity until
August 1 . Remissions occurred mainly on those days when the wind was blowing towards
the river rather than from it. An apparent second outbreak in 1966, lasting only two days,
occurred in late August (Table 7). (Similar autumn outbreaks have occurred in Saskatch-
ewan).
In 1967 the annual outbreak began suddenly on June 19 and continued with only a few
brief remissions until at least July 25. Again there was a distinctly separate autumn out-
break, this time on September 17 and 18, but the outbreak was sufficiently mild that it was
observed at only two sites.
Abdelnur (1968) recorded four apparent cycles of S. arcticum larval abundance in the
Athabasca River in 1965, with peaks in late May, early July, early August and mid-Sep-
tember. However, his data, based on direct counts of larvae, presumably obtained by wading
into shallow marginal areas of the river bed, would have been indirectly affected by daily
changes in the water level. For example, samples obtained from marginal waters when the
river level was stationary or falling could be expected to contain relatively large numbers of
larvae from permanent colonies whereas samples obtained from newly colonized areas dur-
ing periods of rising levels would contain relatively few larvae.
DISCUSSION, CONCLUSIONS AND FUTURE OUTLOOK
S. arcticum is widespread in Alberta and individuals typically breed in mountain-fed rivers
and streams. Although individuals of a few other species of black fly also breed occasionally
in these rivers, only those of S. arcticum are believed capable of developing in such large
numbers that damaging outbreaks can occur. The precise factors affecting abundance and
the subsequent development of outbreaks are not known.
Outbreaks in the Minburn area, presumably arising from the North Saskatchewan River
downriver from Edmonton are rare. Only two brief damaging outbreaks have been recorded,
one having occurred in late May of 1956 and the other in late May of 1961 . Perhaps records
of additional outbreaks of earlier years may eventually be discovered.
Outbreaks in the Athabasca area, arising from the Athabasca River, occur annually and
generally last for several weeks at a time. Beginning as early as mid-June, they have lasted
until about mid-September.
An abatement programme is presently being developed for the Athabasca area. Biological
abatement does not seem feasible with our current state of knowledge of the species; thus
chemical larviciding will be attempted. The development of a programme suitable for the
Athabasca River requires a large number of field trials. Quantitative assessments of the
Black Fly
371
effects on black fly larvae and other aquatic organisms are difficult in this river because of
the irregular and often large fluctuations in the water depth. The study should also include
the accumulation of information on the durability, distribution and effects of not only the
insecticide used but also of its break-down products. The major sources of black flies in the
river should be accurately determined to allow best use of chemical larvicides, especially any
with a short life. Possibly the entire river contributes to these outbreaks but in view of the
lengthy flight range known for this species in Saskatchewan (Fredeen, 1958), the major
sources may well be the extensive rapids downriver from Pelican Portage.
Another profitable area of research is the documentation of events associated with the
development of outbreaks, eventually to permit, if possible, prediction of the time and
severity of each impending outbreak. Such information would not only remove the elements
of suspense that occur annually in this area but would also be useful in planning livestock
management procedures and in planning the most economic utilization of larvicides.
ACKNOWLEDGMENTS
I am very much indebted to J. B. Gurba and L. K. Peterson of the Alberta Department of
Agriculture, Edmonton, and the many co-operators at Athabasca, for collecting thousands
of specimens during the 1966 and 1967 outbreaks at Athabasca, and for reporting many of
the details of these outbreaks. I also wish to thank co-operators in the Minburn area and
particularly Dr. Hasinoff for information about the Minburn outbreaks. I am very much
indebted to Gordon Glen of the Canada Agriculture Research Station, Saskatoon for his
assistance in sorting the specimens collected in 1 966 and 1 967 and for careful preparation
of the two maps illustrating this paper. I am also indebted to M. E. Taylor and L. Burgess of
the Canada Agriculture Research Station, Saskatoon, K. R. Depner of the Research Station,
Lethbridge, and L. K. Peterson for editing the manuscript.
REFERENCES
Abdelnur, O. M. 1968. The biology of some black flies (Diptera: Simuliidae) of Alberta.
Quaest. ent. 4: 113-174.
Brown, A., T. H. D. Griffitts, S. Erwin and L. Y. Dyrenforth. 1938. Arthus’s phenomenon
from mosquito bites. Southern Med. J. 31 : 6, 590-596.
Cameron, A. E. 1922. The morphology and biology of a Canadian cattle-infesting black fly,
Simulium simile Mall. Canada Dept. Agr. Tech. Bull. 5, n.s.
Canada Department of Energy, Mines and Resources. 1967. Surface water data, Alberta,
1965. Queen’s Printer, Ottawa (i-xii) 241 pp.
Curtis, L. C. 1954. Observations on a black fly pest of cattle in British Columbia (Diptera:
Simuliidae). Proc. ent. Soc. British Columbia 5 1 : 3-6.
Fredeen, F. J. H. 1958. Black flies (Diptera: Simuliidae) of the agricultural areas of Mani-
toba, Saskatchewan and Alberta. Proc. 10th Int. Congr. Ent. Montreal, 1956: 3, 819-823.
372
Fredeen
Fredeen, F. J. H. 1963. Oviposition in relation to the accumulation of bloodthirsty black
flies ( Simulium (Gnus) arcticum Mall. (Diptera) ) prior to a damaging outbreak. Nature
(London) 200: 4910, 1024.
Fredeen, F. J. H., J. G. Rempel and A. P. Arnason. 1951. Egg-laying habits, overwintering
stages, and life cycle of Simulium arcticum Mall. (Diptera: Simuliidae). Can. Ent. 83: 3,
Fredeen, F. J. H., A. P. Arnason and B. Berck. 1953. Adsorption of DDT on suspended
solids in river water and its role in black-fly control. Nature (London) 171:700.
Gudgel, E. F. and F. H. Grauer. 1954. Acute and chronic reactions to black fly bites
(, Simulium fly). A. M. A. Arch Derm. Syph. 70, 609-615.
Gurba, J. B. 1963. Personal communication.
Hearle, E. 1932. The black flies of British Columbia. Proc. ent. Soc. British Columbia. 29:
5-19.
Hutcheon, D. E. and V. S. Chivers-Wilson. 1953. The histaminic and anti-coagulant activity
of extracts of the black fly. Rev. Can. Biol. 12:1, 77-85.
Millar, J. L. and J. G. Rempel. 1944. Livestock losses in Saskatchewan due to black flies.
Can. J. Comp. Med. Vet. Sci. 8: 334-337.
Rempel, J. G. and A. P. Arnason. 1947. An account of three successive outbreaks of the
black fly, Simulium arcticum, a serious livestock pest in Saskatchewan. Sci. Agr. 27: 9,
428-445.
Rubzov, I. A. 1940. Fauna of the U. S. S. R. Diptera, Simuliidae. Acad. Sci. U. S. S. R.
Moscow 6: 6, 1-532.
Stone, A. and H. A. Jamnback. 1955. The black flies of New York State (Diptera: Simulii-
dae). N. Y. St. Mus. Bull. 349, 1-144.
Strickland, E. H. 1938. An annotated list of the Diptera (Flies) of Alberta. Can. J. Res., D,
16: 175-219.
Twinn, C. R., B. Hocking, W. C. McDuffie and H. F. Cross. 1948. A preliminary account of
the biting flies at Churchill, Manitoba. Can. J. Research (D) 26: 334-357.
73-76.