£ii — - %

V

Quaestiones

cntomologicae

A periodical record of entomological investigations,

published at the Department of Entomology, Uni-

versity of Alberta, Edmonton, Canada

VOLUME III

1967

CONTENTS

Guest editorial - The role of the anateur in entomology 1

Adisoemarto - The Asilidae (Diptera) of Alberta 3

Book review 91

Book review 93

Frank - A serological method used in the investigation of

the predators of the pupal stage of the winter moth,

Operophtera brumata (L. ) (Hydriomenidae) 95

Book review 106

Sarai - Effects of temperature and photoperiod on embryonic

diapause inNemobiusfasciatus (DeGeer) (Orthoptera,

Gryllidae) 107

Book review 136

Guest editorial -Fascinating taxonomy 137

Madge - A revision of the Genus Lebia Latreille in America north

of Mexico (Coleoptera, Carabidae) 139

Book review 243

Guest editorial - The religious faith of the scientist 245

Murdoch - The biology of the lilac leaf miner,

Fabr. (Lepidoptera : Gracillariidae) 247

Bueger - Sense organs of the labra of some blood-feeding Diptera . 283

Book review 291

INDEX

11

AbaXy 95, 100

paralletepipedus f9S ,99 yl02

Acacesia, 277

Acheta commodus ,19^

diapause in, 114, 119, 125, 131

Adisoemarto,S. , 3

Alberta, ecological regions, 4, 70

Alex,A.H. , 6, 66

Alexander, G. , 127, 134

Alexander, N. , 132, 134

Alexander, R.D. , 110, 133

Alexiopogon tevrioola, 12

Allan, J.A., 3, 66

Altica chalyhea, 141

ohliterata, 179

woodsi, 141

anatrepsis, 132

Andrewartha, H.G., 107, 133

Andrewes, H.E., 225

animal communities, 93

animal ecology, 93

Anisopogon, 27

Antheraea pernyi , 114

antibodies,

formation, 103

of Operophteva, 95

antigens,

preparation of, 96

antiserum,

absorption technique, 101

preparation of, 96, 103

production of, 97

Aphelogenia, 166

hilineata, 197

hivittata, 196

furcata, 190

guttula, 199

spraguei, 190

vittata, 189, 190, 194

Arctia caia, 127

Ash, see Fraxinus

Asilidae,

cannibalism, 6

distribution, 3

general description, 3

habitats of adults, 4

predators of, 7

seasonal succession, 7

subfamilies, 3

Asilidae (of Alberta)

distribution, 4

feeding habits, 5

subfamilies, 7

taxonomy, 3, 7

Asilinae, 7, 54

Asilus , 58

abdominalis , 23

aestuans , 56

aridalis , 3,6,59,63,81,90

auriannulatus , 59, 62, 90

caoopilogus, 55, 56

callidus , 5, 58, 60, 73,77,79,89

oumhipilosus , 3,6,7,59,62,90

oylindvicus , 53

delusus , 58, 59, 81, 89

evythoonemius , 5,6,59,61,81

gilvus , 52

gramalis , 3,5,7,59,64,81,90

maculatus , 55

maroooanum , 44

mesae , 5, 6, 7, 59, 90

nitidifaoies , 5, 59, 61, 82, 89

oooidentalis , 58, 59, 79, 89

paropus , 59, 60, 77, 81

rufioornis , 30

sabaudus , 19

snowi , 59, 61, 77

assassin flies, see Asilidae

Baotvia, 55

Bail, T., 269, 280

Baker, R.H. , 4, 66

Balfour -Browne, F. , 225

Ball, E.D., 110, 133

Bates, H.W. , 140, 225

Beck, S.D., 132, 134

Bigelow, R.S., 120, 134

Birch, L.G., 107, 133

birds, population studies, 91

Blackwelder, R.E., 138, 225

Blanton, F.S., 4, 66

Blatchley, W.S., 110, 134, 225

blowfly, 114

Bombomima, 44, 45

oolwnbioa, 5, 46, 88

fernaldi, 46, 47, 88

insignis , 5, 46, 48

partitor, 46, 88

postioata , 5, 46, 47, 88

Bombyx mori, 107, 132

Bonelli, F.A., 225

Bofkovec, A.B., 243

Boyden, A.A. , 103, 104

Bradley, J.G., 140, 225

Bristowe, W.S., 6, 66

Bromley, S.W. , 4, 66

Brooks, A.R., 3, 66

Brown, J.H. , 3, 67

Browning, T.O., 114, 134

Brulle, G.A., 225

Buerger, G., 283, 288

Ill

Burdick, H.C., 118, 134

Burgess, A., 291

cannibalism (in Asilidae) , 6

carabids, 95

Carabus bivittatus , 196

oyanooepha lus , 1 64

vittatus , 166, 189

Casey, T.L., 140, 225

CeraturguSf 25

Chaboussou, F., 141, 225

Chaudoir, M. , 140, 225, 226

chemos ter Hants, 243

Chevrolat, L. , 140, 226

Chortophaga viridi fas data y 127

Chrysolina varians , 141

Chrysopsy 276

nigripes y 283, 286

Church, N.S., 118, 134

ClavatoT y 29

sabulommy 29

Clethrionomy s glareolus y 102

Clubionay 111

CocdnelZa septempunotatay 101

Cole, F.R. , 4, 67

Cole, P., 114, 134

Comantellay 8, 9, 42

falleiy 42, 43, 77, 87

rotgeriy 42, 43

Coquillett, D.W. , 28, 67

Cragg, J.B., 114, 134

crickets, 107

Crowle, A.J., 95, 104

Csiki, E. , 139, 226

Culicoides vaviipennis y 283, 284, 2i

Curran, C.H., 3, 67

Curtis, J., 226

Cushman, R.A. , 141, 226

see dlsOy Isely, D.

Cymindis y 218

Cyrtopogoriy 4, 9, 30

albitavsis y 32, 36

albovarians y 5, 32, 36

auratus y 32, 77, 78, 86

aurifeXy 3, 32, 33

auvipilosus y 11

bimaoulay 5, 32, 34, 81

dasyllis y 5, 32, 38, 81

distinoti tarsus ,3,5,32,34,71,81

glavealis y 3, 32, 36

inversus y 3, 32, 36

lineotarsus y 31, 37, 77

me lanop leurus , 34

montanus y 32, 35, 86

nebula y 38

nigator y 31

Cyrtopogon , (cont . )

nugatory 32, 38

praepes , 32 , 34

sansoniy 5, 31, 37

willistoni y 5, 32, 33, 77

Danilyevsky, A.S., 107, 134

Dasyllis insignis y 48

pubescens y 49

vivoxy 51

Dasypogon aeacus y 23

argenteus , 9

oandidus y 10

diadema , 6

elegantulus y 9

fasdventris y 10

fimbriatus , 31

gelascens y 10

manioatus y 27

nigripennis y 25

piles ellus y 11

politus y 29

spatullatus y 24

trifasdatay 9

trifasciatus y 10

Dasypogoninae , 7, 8

genera, 8

Dasypogonini , 8

Daulopogony 11

terrioolay 12

Defalco, R. , 103, 104

Dejean, P. , 226

Dempster, J.P., 95, 105

Dendroica petechia y 276

Dendrolimus pini y 127

Deutziay 261

Dianohomena , 166

abdominalis y 198

aemulay 196

bilineatay 197

oonvictor y 198

devinotay 196

miranday 188

quadrivittata y 196

soapularis y 187

diapause, 107

Dickson, R.C., 107, 134

Diotyna annulipes y 277

Digitaria sanguinalis y 110

Dimmock, F., 264, 280

Diogmites angustipennis y 6

Diptera, see also Asilidae, 3

blood- feeding, 283

Dromius apioalis y 208

Echimuthus y 164

ecology, 93

, 283

IV

ecological methods, 136

Efferia, 56

candidus , 56

electrophoresis, 102

Elton, C.S., 93

embryogenesis, 114

entomologists, 1

entomology, amateur, 1

beginnings, 1

classification, 1

World Congresses, 1

Erannis , 102, 103

antigen, 100

auran ti avia , 9 9

defoliavia, 99

EvaXy 56

hioaudatus , 57

oanus , 57

costalis , 58

vufihavbis , 56

suboupveus y 5, 57

Euavmostus , 30

bimaoula, 30, 34

Eucyvtopogon , 8, 9, 38

albibavbis y 5, 39, 40, 87

oalcavatay 39, 41

comantis y 39, 40, 82

divevsipilosus y 39, 42, 82

inoompletus y 3,39,40,73,76,80

nebulOy 39, 42

spinigevay 39, 41

Euomymus y 261

Eupalamus y 30

alpestvis y 30

Evans, H.E., 106

Fabricius, J.C. , 140, 226

Feinberg, J.G., 105, 106

flea beetle, 141

Forbes, W.T. , 226, 255, 280

Formica neoclavay 276

Fourcroy, A.G., 226

Frank, J.H. , 95, 105

Freund, J. , 97, 105

Fvaxinus y 248, 261

amevicanay 251

divevsifo lia , 251

excelsior y 249, 250, 251, 281

pennsylvanica y 251

potamophila y 251

pubes cens y 251

rotundi folia y 250

Fulmek, L., 255, 280

Fulton, B.B., 110, 134

Galerucella cavicollis y 141

luteolay 141

gamma globulins, 98, 101

Gemminger, M. , 226

Geoffroy, E.L., 226

Gladstone, G.P., 98

Glossina morsitans y 284, 286, 290

palpalis y 283, 286

Gonypes , 53

Gooding, R.H. , 243

Gracillaria syringellay 247

adaptations, 263

dimensions, 256, 257

dispersal, 278

distribution, 249, 250

history, 249

host plants, 261

larval structure, 265

leaf rolling, 268

life cycle, 258

mines, 267

mortality, 275

populations, 270, 271

Gradwell, G.R., 95, 105

Graham , P . , 91

Grapholitha molestay 129

Gryllulus mitratus y 119

Gryllus pennsylvanicus y 119, 132

habitat systems, 93

Eaematobia irritans y 284, 287, 289

Haldeman , S . S . , 226

Halliburton, W.H. , 127, 134

Harvey, W.R. , 107, 134

Hatch, M.H. , 140, 226

Heald, F.D., 269, 280

Hebard, M. , 110, 134

Eeteropogony 9, 27

wilcoxi y 3,28,73,80,81,86

Hentz, N.M. , 226

Hering, E.M. , 261, 263, 266,280

Hine, J.S., 56, 67

Hogan, T.W. , 114, 135

Hogan’s method, 108

EolopogoHy 9, 25

albipilosay 26

albipilosus y 5,71,78,80,86

nigripilosay 3, 5, 26, 27, 81

seniculus y 26

Eomalops y 164

Horn, G.H., 140, 227

Hull, F.M. , 3, 67

Hutchings, C.B., 251

Eybomitra rupestris y 283,285,289,290

Hydr iomenidae , 9 5

Immunoelectrophoresis, 102

insect populations, 136

Isely, D. , 141, 226

V

Itopleotis quadrioingulata f

James, M.T. , 6, 68

Jobling, B. , 283, 288

Johnson, C.M. , 104, 105

Kaltenbach, J.H., 261, 280

Kogure, M. , 107, 135

Kozhanchikov, I.V. , 114, 135

labrum, 283

Lack, D. , 91

Lconprias j 164, 219

oyanellus , 177

oyanipennis , 176

limbioollis y 180

Laphria, 44, 48

aeatus , 5, 49, 50, 80

aimatis f 48, 52, 88

fulvithovaXy 45

gilva, 5, 48, 52, 88

index ^ 3, 5, 49, 51

5, 49, 51, 73, 78, 82, 88

postioata, 47

saokenif 88

sadales , 5, 48, 49, 80

, 3 , 5 , 49 , 50 , 72 , 78 , 80 , 87

vivaXy 49, 51

vultur, 88

xant/zippe, 48, 49, 76, 79,80,82,87

Laphriinae, 7, 43

de LaPorte, F. , 111

Lasiopogon, 4, 8, 11

aldriohiy 12, 18, 79, 80

oanusy 3, 12, 17

ainereUS yl2 ylS yll ylS yll ySO ySl)

hineiy 3, 5, 12, 18

primay 3,12,16,80,81,83

quadrivittatus ,5,6,7,11,14,76,78

ripicolay 12, 15, 77

tevricolay 5, 11, 12, 71, 83

trivittatusy 3, 5, 11, 13, 77,83

Latreille, P.A. , 140, 111

Leach, W.E., 111

leaf miners, 247

Lebia^

abditUy 201

abdomina lis , 198, 229

abruptUy 170

adolesoens y 178

aemulay 196

af finis y 180

connioolay 190

analiSy 184, 208, 230, 234, 239

anohoray 184

appendioulata y 184

arizonicay 175

ashenvillensis y 208

(cont . )

atrioeps jylSl) y 232, 241

atriventris y 153, 232

axillavis y 208

bavbavae, 176

bioinotay 111

bilineatay 197

bitaeniatay 171, 230

bivittatay 196, 229, 231, 235

bonelliiy 184

boreay 170

bvaoatay 178

bvunneay 208

brunnioollis y 218

biMieliae y 213

calliope y 111

callizonay 111

canonioay 203

oastigatay 178

characters of, 142

ohloropteray 218

cobaltinay 178

color of, 142

collavis y 208, 214

ooncinnay 165

Qong ungens , 190

conviotoTy 198

cyaneoy 177, 181

cyanellay 111

oyanipennis , 176

cynicOy 111

debiliceps y 190

deceptrixy 158, 232, 229, 230

depiotay 190

devinotOy 196

distribution, 144

divisoy 165, 230, 233

doubtful species, 218

duluthianOy 178

enormis y 154

erythrooephalay 208

esuridhs y 111

evolutOy 178

flaviventris y 218

flavolineata y 187, 218

flavovittata y 189

fluviatilis y 208

frigidOy 208

frontalis y 170

frugalis y 215

furcatay 189, 231

fusoatay 203, 208

grandisy 161, 229, 230, 232

guttulay 199

histricay 178

247,276 Lebiay

VI

Lehia (cont . ) ,

histvionioa , 193

mini, 215

inoitata, 178

insulata, 202, 229

key to species, 148

lecontei, 163, 229

lecta, 213

lobulata, 207, 218

ludovioiana, 215

maculicornis , 215

magica, 178

magistev, 161

maQUScula, 161

male genitalia, 143

mavginella, 208

melaena, 176

metuens, 199

miranda, 188, 235

moesta, 177

montana, 176

morphology, 142

nanulina, 155

nigrioapitata , 195

nigvipennis , 208

nigrosignata , 193

omata, 184, 208, 218, 231,236,239

papago, 178

peotita, 194

perallida, 206

perita, 182

phylogeny, 218

planifera, 178

pleuritioa, 173

primalis , 159, 232

prominens , 178

pulohella, 167, 168, 230, 233

pumila, 215, 229

puno tifera , 218

quadrivittata , 196

reperta, 208

rhodeana, 170

rhodopus , 215

rivularis , 161

ruficollis , 176

rufop leura , 172

soalpta, 186, 234

scapula, 183, 230, 234

scapularis , 166, 187,189,190,218

soutellata, 193

smaragdula, 177

solea, 187, 229, 230, 235

somonae, 190

subaf finis , 178

subdola, 157, 232

Lebia (cont . ) ,

subfigurata, 218

subgrandis , 160, 230, 232

sublimata, 218

subrugosa, 205

tahoensis , 167

tempeana, 190

tertearia, 215

testacea, 163, 164

tricoloT, 156, 232, 239

tvuoheensis , 178

tuckeri, 174

vermiculina, 178

virginica, 208

viridipennis ,170,229,233, 238

vividis, Yll , 180, 215, 229,230

vittata, 189, 190, 231, 235, 237

vivida, 167

websteri, 187

Lebida, 165

Lebis solea, 167

LeConte, J.L., 111

Lees, A.D. , 107, 135

Leng, C.W. , 227

Leone, C.A., 103, 105

Lepidoptera, 95

Leptinotavsa deoemlineata , 141

Leptogaster, 53

ap^S^^s,3,5,7,53,54,73,78,80,82

coloradensis , 3, 54

Leptogastrinae, 7, 53

Lestomyia , 9, 29

sabulonum, 5, 6, 7, 29,76,80,81

Lia femovata, 171

Libby, R.L., 103, 105

Ligustrum , 248, 261

ca lifornicum , 251

japonioum, 251

ovali folium, 251

vulgare ,251

lilac, see Syringa

Lindroth, C.H., 181, 111

Linnaeus, C., 1, 111

Lissoteles , 9

hermanni , 9

Listroderes obliquus , 131

Locustana pardalina, 132

LoxopezOy atrioeps, 155

atriventris , 153

enormis , 154

grandis, 161

magistev, 161

magusoula, 161

nanulina, 155

pimalis , 159

VI 1

Loxopeza (cont . ) ,

rivulariSf 161

testacea, 164

tricolor, 156

tuokeri, 176

Lucilia sericata, 114, 131

Lugger, 0., 110, 135

Lutshnik, V.N., 111

Maar, A., 254, 259, 261, 280

Madge, R.B., 139

mammals, bibliography, 137

experimental , 96

Martin, C.H., 3, 4, 68, 69

Masaki, S. , 118, 135

Matthee, J.J., 132, 135

Melander, A.L., 4, 68

Me lanop lus^ hivi ttatus^ 125

devastator, 127

mexioanus , 125

Melin, D. , 4, 68

Metabola, 166

rufopyga, 166

vivida, 167

Miorohemhix, 106

Miller, R.S., 93

Moss, E.H. , 3, 69

Motschoulsky, V., 140, 227

Myoohaoteriym hutyrieum, 97

McKelvey, S.D., 248, 281

tiebria, 101

brevicollis , 101

Negasilus, 65

belli, 5, 65, 77, 81

Nemobius,

age, 115

alternating temp, effect, 124

collection of, 108

diapause, 107

distribution, 110

ecology, 108

eggs, 126

diapausing, 120

post -diapause, 122

pre-diapause, 119

fasoiatus , 107

lab study, 115

life cycle, 110

low temperature effect, 127

nymphs, development, 116

effects of light on, 129

oviposition, 132

photoperiod, 115, 127

seasonal rhythm. 111

submergence of, 117

temperature sensitivity, 112

Nemobius (cont . ) ,

voltinism, 113

yezoensis , 127

Neopogon , 9

Nerax , 56

bioaudatus ,6 ,St> ,72 ,77 ,79 ,^1 ,S2 ,^9

oanus , 57, 82

Qostalis , 57, 58, 82

subcupreus , 57, 82

iHoocles , 8, 29

analis , 29

punotipennis , 30

utahensis , 30, 72, 73, 77, 78, 80

Nomadaoris septem fas data, 127

nomenclature, see also taxonomy

code of, 2

Norris, M.J., 127, 135

Nossall, G.J.V., 95, 105

nymphal development,

in Nemobius fasoiatus , 116

Nyssoninae, 106

Odynsky, W. , 3, 69

Oliver, A.G., 228

Omalomorpha, 164

Operophtera brumata , 95

antigen, 96, 100

collection of predators of, 96

Orthoptera, systematics of, 3

Ospriooerus , 8, 23

abdominalis , 23, 24, 71, 80

aeacides , 24

consanguineus , 23, 24, 80, 83

pumilus , 23, 24, 25

ventralis , 24

Ouchterlony concept, 104

Fa lamopogon , 31

alpestris , 31

Parasemia plantaginis , 127

Phalera bucephala, 131

Philammosius , 31

Philodromus aureolus , 277

Philonthus decorus , 95, 100, 101, 102

Phlebotomus papatasii , 132

Phormia regina, 100

photoperiod, 107

photronreflectometer, 103

Phyllyrea media, 261

Pinaoodera, 218

Podootria, 25

Poeoilostola, 163

Pogonosoma , 43 , 44

ridingsi , 7 9

strioklandi ,7) ,72 , 73,76,80,81

Polyoheloma, 163, 219

Populus nigra, 261

Vlll

precipitin reaction, 100, 102

precipitin test, 98

predator-prey studies, 95

predators, of Asilidae, 7

Pritchard, G. , 94, 136

privet, see Ligustrwn

Prootacanthella f 55

cacopi toga ,6,7,56,72,77,79,80,89

Promaohus , 54, 55

dimidiatus , 55, 79, 82

Proom, H. , 103, 105

proteins, precipitin reaction, 102

Pterostichus f 95

madidus , 100, 102

melanarius , 100, 102

Pussard, R. , 254, 258, 281

Putzeys, J.A.A.H., 228

Pygostolus , 29

Quedius lateralis ^ 101

Rakshpal, R. , 118, 135

Readio, P.A. , 107, 135

Reaumur, R.A.F. de, 249, 281

robber flies, see Asilidae

i?esa, 276

Roubaud, E., 132, 135

Salt, R.W. , 118, 134

sand wasps, 106

Sarai, D.S., 107

Say, T., 140, 228

Soamhus hispae , 247, 276

Schaeffer, C.F.A. , 228

Schwarz, E.A. , 228

SoleropogoYLf 19

negleotus f 22

piotioomis f 19

pwnilus , 25

Scudder, H.I., 283, 288

seasonal succession,

of Asilidae, 7

sense organs, 283

serological method, re:

winter moth predators, 95

Sich, A., 261, 281

silkworm, 114

Silvestri, F. , 140, 228

Simpson, G.G. , 139, 228

Simuliwn venustum, 283, 284, 289

vittaturrij 283, 284

Snodgrass, R.E., 228

Southwood , T . R . E . , 136

Spencer, K.A. , 2

Speirs, R.S., 95, 105

Sphecoidea, 106

Spilosoma menthastri ^ 131

Stager, R. , 261, 281

Steiner, A.L., 106

Stenopogon , 8, 19

oonsanguineus , 24

ee^ete,6,7,20,23,71,81

gratus , 20, 21, 84

inquinatus ,5,6,20,71,76,78,80,85

latipennis , 24

modestus f 21

morosus f 21

negleotus , 6, 7, 20, 22, 74, 85

ohsouriventris , 5, 20, 71, 73, 84

pumilus , 24

rufiharhis, 20, 84

univittatus , 21

Stenopogonini , 8

Stiohopogon ,8,9

argenteus, 9, 71

trifasoiatuSf 9, 10, 71,74,77,78

Stichopogonini, 8, 11

Stilopogon, 9

aequioinotus , 9

Stomoxys oaloitrans , 284, 287, 289

Strickland, E.H. , 3, 69

stridulation, 291

Strohecker, H.F., 110, 135

Strokov, V.V. , 249, 258, 281

Symphoricarpos , 261

Syringa, 2M , 261

amurensis y 281

ohinensis , 248

distribution, 248

emodi, 250

history, 248

gosikaea, 250

pekinensis f 250

persioa, 250

reflexa, 250

rothomagensis , 248

villosa, 250

vulgaris, 248, 250, 251

Sy sterna Naturae, 1

taxonomy, see also nomenclature, 2

of Asilidae, 7

general, 137

Telegoneura, 55

Te tragnatha , 277

Theobald, F.V., 258, 281

Thiodina, 111

Thomas, E.S., 110, 133

ticks (of Alberta) , 3

Tragardh, I., 254, 281

Trupanea, 55

Tuxen, S.L., 291

Varley, G.C., 95, 105

Vickery, V.R., 110, 135

voltinism, in Nemohius fasoiatus y 113

Von Gemet, G., 283, 288

Wadsworth, C. , 104, 105

Way, M.J., 94

Webb, R.A. , 98

West, A.S., 95, 105

Wieme, R.J., 95, 105

Wigglesworth, V.B., 245

Wilcox, J., 4, 69

Williams, C.A. , 102, 105

winter moth, see Operophtera hrumata

World Congresses, 1

World Weather Records, 113

Wytham Woods, 93, 95

Quaestiones

entomologicae

A periodical record of entomological investigations,

published at the Department of Entomology, Uni-

versity of Alberta, Edmonton, Canodo.

VOLUME III

NUMBER 1

JANUARY 1967

1

!

QUAESTIONES ENTOMOLOGICAE

A periodical record of entomological investigations, published at

the Department of Entomology, University of Alberta, Edmonton, Alberta.

Volume 3 Number 1 11 January 1967

CONTENTS

Guest Editorial 1

Adisoemarto - The Asilidae (Diptera) of Alberta 3

Book Review ,91

Guest Editorial - The Role of the Amateur in Entomology

Entomology has now become an essentially professional science.

Many younger entomologists - professionals in the true meaning of the

word and experienced in all the modern techniques of entomology, such

as genetics and statistics andusing complex tools like the electron micro-

scope - are possibly not fully conscious these days of the dominant role

the amateur has played in the past.

Progress in the science of entomology proceeded slowly from the

early beginnings in China, Greece and Rome, but already in the 17th and

early 18th centuries accurate observations and illustrations were being

recorded by such people as Maria Sibylla Merianin Holland and E. Albin

in England and John Ray made a significant advance with his classification

of insects published in 1710. The great leap forward, however, took

place from the middle of the 18th century following publication of the

first edition of the "Systema Naturae" by Linnaeus in 1738.

The great era of amateur entomology followed from the end of the

18th century and throughout the 19th with a flourishing of individual col-

lectors, local societies and journals. Vastnumbers of new species were

described and the first major systematic works were published. This

was largely the work of amateurs. It is true that great confusion has

been caused by the uncritical nature of much 19th century work but, as

any taxonomist knows, the problems of clas sification ar e not easy. Mis-

takes were inevitable. The important thing, however, was that the enor-

mous problem of attempting to name and classify all insects throughout

the world had begun. Without the enthusiasm, dedication and passionate

interest in insects shown by the amateur in the past, the tremendous ad-

vances in entomological knowledge and achievement in recent years would

not have been possible.

Just prior to and following the Second World War, the dominance

of the professional in world entomology became complete and a major

expansion in the organisation of entomology began which is still continuing

at the present time. The dramatic development in entomology during the

past 50 years is clearly reflected by the attendance at World Congres ses .

Both at the 1910 Congress in Brussels and the 1912 Congress in Oxford

there were less than 150 members, while at the second Congress held in

England in 1964 in London the numbers were approaching 2, 000. While

amateurs played an important part in the organisation of the early Con-

gresses, only a handful were members of the 1964 Congress and papers

were read by an even smaller number.

2

Spencer

In view of the overwhelming professionalisation of entomology to-

day, what contribution of importance can now be made by the amateur?

The life histories even of many of the commoner insects are not fully

known and the detailed work of rearing individual species is the sort of

work admirably suited to the limited time and means of younger amateur s .

Careful observation will reveal many unknown facts . The journals of

local societies will normally provide an outlet for publication of such

small scale studies . For the pure collector the preparation of local lists

either of all orders in a limited area or of one order in a wider field will

always provide information of value. The collection of any of the lesser

known orders or of particular biological groups such as galls or leaf

mines offers unlimited scope for exciting new discoveries.

The achievement of any original taxonomic work by amateurs be-

comes increasingly difficult. The multiplicity of species, the complexity

of the code of nomenclature and the proliferation of literature, now pub-

lished in numerous languages, necessitate years of patient study and

work before one can venture into original publication. Few amateurs

are prepared to devote their entire free time for years on end to ento-

mological study of this kind which alone will qualify them as taxonomists.

A further major difficulty for any taxonomist is the study of types.

Numerous species cannot be identified with certainty from the descrip-

tion alone and, in cases of doubt, personal examination of the holotype

will always become es sential. Confronted with this problem, the amateur

is not in a position to apply for a grant and fly off to examine the type in

question. He is dependent on the institution, wherever it may be, sending

the type to him as a loan. Unfortunately, many leading museums and

institutions refuse to lend types and here I would like to emphasise how

much this negative policy must be deprecated. The inability to study

types can represent a serious handicap to scientific work and not only

amateurs suffer in this way. There is even one well known European

museum which, in the past, has refused to allow a number of its types

even to be examined at all "owing to their great historical value". With

such a policy the scientific value of the types concerned becomes nil.

Surely the whole policy in relation to types must hinge on their value for

the advancement of scientific knowledge and any policy which frustrates

this must be wrong.

The early Chines e around 500 B.C. listed in the Erh-ya 53 species

of insects, while Aristotle recorded 47 species and Pliny in Rome 300

years later 61. Today the number of described species is approaching

the million mark, while almost certainly an equal number remain to be

discovered and described. Despite the immense progress of the past the

tasks still facing entomology are formidable and any contribution which

can be made by the dedicated amateur should be given every possible

encouragement by professionals. The enthusiasm of the one combined

with the experience and skills of the other will always prove a felicitous

combination which can only benefit the science to which we are all devoted.

Kenneth A. Spencer

19 Redington Road

London N. W. 3, England

Adisoemarto

3

THE ASILIDAE (DIPTERA) OE ALBERTA

SOENARTONO ADISOEMARTO

Entomology Research Institute Quaestiones entomologicae

K.W. Neatby Building, Carling Avenue „ fo<7

Central Experimental Farm

Ottawa, Ontario.

A taxonomic treatment for the adults of 85 species of Asilidae of Alberta and brief notes

on their ecological relationships and habitats are presented. To the 66 species listed by Strickland

(1938, 1946) the following eleven are added: Lasiopogon trivittatus Melander, L. hinei Cole and

Wilcox, L. canus Cole and Wilcox, Heteropogon wilcoxi James, Cyrtopogon aurifex Osten-Sacken,

C. inversus CtTrran, C. giaiealis Melander, haphria scorpio Me Atee, L. index A/cA^ee, Leptogaster

aridus Cole, and L. coloradensis James. Eight new species, Lasiopogon prima, Holopogon

nigripilosa, Cyrtopogon distincti tarsus, Eucyrtopogon incompletus, Pogonosoma stricklandi,

Asilus aridalis, A. gramalis, and A. cumbipilosus, are described.

The Asilidae^ also known as robber flies or assassin flies, are

predaceous insects, easily recognized by their morphological characters

as well as by their activities. All of these flies have an excavated front

and vertex to form a V-shaped depression. Both sexes are dichoptic.

Some characters show sexual dimorphism.

The family consists of about 5000 known species, distributed over

the six zoogeographic regions. Hull (1962) listed the number of species

occurring in each region. According to Martin (1965), the Nearctic

Region has the greatest number of species. There are five subfamilies:

Dasypogoninae, Leptogastr inae, Laphriinae, A silinae, and Megapodinae.

The last subfamily is confined to the Neotropical region. Fifteen of 18

genera described from the Eocene, the Oligocene, and the Miocene, are

still living (Hull 1962).

Alberta is a province where many differ ent zoogeographic elements

meet and play roles in the diversity of the living beings occurring there.

Allan (1943) has described the geology. Moss (1955) has provided the

description of the plant communities in Alberta, and Odynsky (1962) has

presented a map of soil zones of Alberta.

Few groups of insects or other arthropods of Alberta, have been

studied on a regional basis. The systematics of the acridoid Orthoptera

of southern Alberta, Saskatchewan, and Manitoba, have been presented

by Brooks (1958), and a study of the spotted fever and other Albertan ticks

has been made by Brown (1944). Lists of some groups of insects have

been published, including an annotated list of the Diptera of Alberta by

Strickland (1938, 1946). This study was mainly of taxonomy and geo-

graphical distribution, but some ecological notes, which may be useful

for further ecological study, are also presented. The A silidae of Alberta

have not been studied in detail, although a few new species have been

described from this province (Curran 1923). In Strickland's lists, 66

species of Asilidae were included, but eleven of these species probably

do not occur in Alberta. In this study, 19 species have been added to the

Alberta list, and of these eight are new species. The newly collected

4

Asilidae of Alberta

specimens are deposited in the collection of the University of Alberta,

and the holotypes and the allotypes of the new species are deposited in

the Canadian National Collection in Ottawa. The deposition of the speci-

mens examined is indicated by abbreviations between par entheses follow-

ing locality names. The abbreviations are:

AMNH American Museum of Natural History, New York City, N. Y.

CAS California Academy of Sciences, San Francisco, California.

CNC Canadian National Collection, Ottawa, Ontario.

DE Mr. D. Elliott, Calgary, Alberta.

LMK Mr. L. M. Kenakin, Edmonton, Alberta.

UA University of Alberta, Edmonton, Alberta.

USNM United States National Museum, Washington, D. C,

WSU Washington State University, Pullman, Washington.

Habitats of the Adults

Habitats of some species of the Asilidae have been described.

Melin (1923) studied the biology of the Asilidae of Sweden. James (1938)

discussed the habitat prefer ence of the Asilidae of Colorado, and Bromley

studied the habitats of the adults of Connecticut (1946) and of Florida

( 1950) . The following publications contain habitat data for certain groups:

Wilcox and Martin (1936) for Cyrfopo^on Loew; Melander (1923b) iov Lasiopogon

Loew; Baker (1939) for some species of robber flies from Coahuila,

Mexico; Blanton (1939); and Cole (1916).

The robber flies are sun-loving insects of dry open areas. Habitats

most commonly frequented are: dry fields, pastures, open bush country,

sandyareas, and edges of woods. According to Hull (1962), in desert or

semidesert country, these flies are attracted to small streams, and in

temperate regions, a few species occur in swampy areas and in deep

forest. Bromley (1946) stated that asilids were practically absent from

deep dark woods and swamps. For the state of Colorado, James (1938)

gave a list of five differ ent habitats in which he found asilids . The grass-

land habitat had the greatest number of species, 36, representing nine

genera. Bare areas and thickets were poor in Asilidae . Bromley (1946)

listed nine habitats for Connecticut. Seventeen genera containing 43

species were recorded from woodlands and bushy pastures . The species

of bushy pastures were similar to those of the woodlands, but more abun-

dant. Adults are collected consistently in Alberta in grasslands: pas-

tures, semi-arid short grass prairie, beach grassland (close to lake or

river), openings in the parkland forests; river banks and lake beach;

woodland paths; sandpits and sand dunes; coniferous forests; and bare

fields .

Distribution of Asilidae

Each ecological region of Alberta seems to have certain asilid

species. These species are more or less limited in their distribution

by the boundaries of these regions. The zonation of the province of

Alberta applied here is based on that of Moss (1955) and Brooks (1958).

Prairie

This region is by far the richest in asilid species. Of 37 species

Adisoemarto

5

recorded, 16 have not been found elsewhere in the province. Lasiopogon

terricola Johnson, L. quadrivittatus Jones, Stenopogon obscuriventris Loew,

Eucyrtopogon albibarbis Curran, and Asilus gramalis new species, have also been

found in the Parkland; Lasiopogon trivittatus Melander, Stenopogon inquinatus

Loew, and Asilus erythocnemius Hine, in the Boreal and the Bor eal-Cordil-

leran Transition; while Holopogon albipilosus Curran, Lestomyia sabulonumOsten-

Sacken, Erax subcupreus Schaeffer, Asilus mesae Tucker, Asilus gramalis new

species, Negasilus belli Curran, have also been found in the Subalpine and

the Montane regions . On the other hand, some species, Cyrtopogon willistoni

Curran, Bombomima columbica Walker, Laphria gilva L. , and Leptogaster aridus

Cole, may have been extending from the Subalpine and the Montane regions

into the Prairie region. Two species, Stenopogon inquinatus Loew and Cyrtopogon

bimacula Walker, are more or less ubiquitous in Alberta.

Boreal forest

There are 17 species recorded from the Boreal forest, but the

following species are limited to it: Holopogon nigripHosa new species, Bombomima

posticata Say, Laphria Scorpio McAtee, Laphria aeatus Walker, Laphria index

McAtee, and Asilus nitidifacies Hine . The species: Lasiopogon hinei Cole and

Wilcox, Bombomima insignis Banks, and Laphria janus McAtee, have extended

southward to the Parkland, and westward (except Bombomima insignis Banks)

to the Bor eal-Cordilleran Transition and Subalpine regions . Some other

species presentinthe Boreal region may have been the result of ' 'invasion"

from the Prairie, such as Lasiopogon trivittatus Melander and Asilus

erythocnemius Hine, or from the Subalpine or the Montane region, such as

Cyrtopogon dasyllis Williston, Laphria sedales Walker, and Asilus callidus

Williston. Cyrtopogon distinctitarsus new species is found in the Boreal forest

and in the Prairie.

Subalpine and Montane regions

The Subalpine region is the second richest in the species of Asilidae

in the province. Of 23 species recorded, only two are confined to this

region: Cyrtopogon sansoni Curran and Cyrtopogon albovarians Curran. The

remainder are elsewhere, mostly in the Montane region, with some

others in the Boreal and the Prairie regions. Another species found

here, Asilus erythocnemius , might have entered this zone from the Prairie

region.

Parkland and Boreal-Cordilleran

These regions are transitional. As one might have expected, asilids

found here are a mixture of species from two or more regions. In the

Parkland region, the species are mostly from the Prairie region, while

those in the Boreal-Cordilleran region, are mostly from the Subalpine

or the Montane regions. The Boreal species seem to have extended

equally to these two transitional regions.

Feeding Habits

Without exception, all species of the Asilidae are predaceous in

the adult stage. Food selection of this group, according to Hull (1962),

has been studied considerably, by Hobby and Poulton for the British

6

Asilidae of Alberta

Asilidae, Carerra for South American species, and in Notth America

by Bromley. However, according to James (1938), as far as food is

concerned, the Asilidae are indiscriminate. The food is variable, in-

cluding dragonflies, grasshoppers, Hemiptera, Hymenoptera, Coleoptera,

Lepidoptera, and Diptera. Spiders as food have been reported by Bristowe

(1924) and Bromley (1946). Cannibalism has been reported in Diogmites

angustipennis Loew by Alex (1936), and in Alberta, there is a cannibalistic

tendency in Stenopogon inquinatus Loew. Cannibalism in association with

courtship has been reported in Dasypogon diadema Fabricius by Poulton

(1906).

TABLE 1. Dates of first appearance of some species of the Asilidae

and numbers collected, from some localities in southern Alberta.

Adisoemarto

7

Predators

The asilids have enemies : spiders, wasps, birds, lizards, mantids

(Hull 1962), and in very rare cases, the larvae of Cicindela Linnaeus. A

species of red mite was found attached to the external parts of some

specimens of the species Lasiopogon dnereus Cole and L. trivittatus Melander.

Seasonal Succession of Species

Seasonal succession occurs in the adult Asilidae. Bromley (1934)

mentioned the occurrence of four distinct groups of the Asilidae, in

Brazos County, Texas, according to the period or time of appearance.

Data from a few Alberta localities are presented in Table 1. Although

no conclusion can be drawn, there is slight indication, that in the southern

parts of Alberta, the adults occur as two seasonal groups. The first

group appears in early June. Included here are Lasiopogon quadrivittatus

Jones, Lestomyia sabulonum Osten- Sacken, Asilus mesae Tucker, Asilus cumbipilosus

new species, Asilus aridalis new species, and Asilus gramalis new species.

The second group appears in early August, and includes Stenopogon coyote

Bromley, Stenopogon neglectus Bromley, Leptogaster aridus Cole, Proctocanthella

cacopiloga Hine, Nerax bicaudatus Hine, and Asilus erytbocnemius Hine.

TAXONOMIC TREATMENT

All of the subfamilies except the Megapodinae, which occurs only

in the Neotropical Region (Hull 1962), are representedin Alberta. Eighty

five species of 23 genera were recorded. The Dasypogoninae is the

largest subfamily: 11 genera with 49 species, followed by the Asilinae

with 19 species in eight genera. The Laphriinae is represented by 15

species of three genera, while Leptogastrinae has only two species of

Leptogaster Meigen.

KEY TO THE SUBFAMILIES OF ASILIDAE OF ALBERTA

1. Abdomen slender and cylindrical (fig. 102); second abdominal seg-

ment six times as long as first (fig. 103); wings with alula

greatly reduced or absent; hind femora club-shaped (fig. 85).

Leptogastrinae

Abdomen not slender, almost as broad as thorax (fig. 87); second

segment at most three times as long as first (fig. 104); alula

present; femora not club-shaped 2

2. Wings with marginal cell open (fig. 57) Dasypogoninae

Marginal cell closed (fig. 161) 3

3. Abdomen gradually tapering apically (fig. 104); second segment

three times as long as first; mediocubital crossvein of wings

absent, or M3 and Cu^ fused for a short distance at the place

of crossvein (fig. 165) Asilinae

Abdomen, up to sixth segment, parallel- sided, or broader at the

middle (fig. 101); second segment subequal to first; wings with

mediocubital crossvein present (fig. 161) Laphriinae

8

Asilidae of Alberta

Subfamily Dasypogonihae

Ten tribes comprise this subfamily, of which the Stichopogonini,

Stenopogonini, and Dasypogonini, are represented in Alberta. The

Stichopogonini is represented by Stichopogon Loew and Lasiopogon Loew;

Stenopogonini by S^enopo^on Loew, Ospriocerus Loew, H olo pogon L,oev^, Cyrtopogon

Loew, Eucyrtopogon Curran, and Heteropogon Loew; Dasypogonini by Comantella

Curran, Lestomyia Williston, and Nicocles Jaennicke.

Hull (1962) distinguished the Dasypogonini from the Stenopogonini

by the presence of a bent spine at apex of the front tibia, and placed

Comantella Curran in the former, Eucyrtopogon Curran in the latter. Based

on some other character s, these two genera should be placed in the same

group. Eucyrtopogon is more similar to Comantella Curran than to the rest

of the Stenopogonini.

Key to the Genera of Dasypogoninae of Alberta

1.

2.

3.

4.

6.

7.

Face bare, except on the oral margin; gibbosity not conspicuous

(fig. 3); ocellar bristles absent; dor socentrals absent (also

from Lasiopogon terricola Johnson) Stichopogon Loew

Face with hairs or bristles between oral margin and at least half-

way to antennal base (figs. 5, 10); ocellars and dor socentrals

present (except in Lasiopogon terricola Johnson) 2

Palpus one- segmented (fig. 40); vertex with posterior margin at

least twice as wide as front at antennal base (figs. 7, 9)

Lasiopogon Loew

Palpus two- segmented (fig. 42); vertex not widened posteriorly,

posterior margin at most one and half times as wide as front

at antennal base (figs. 15, 22) 3

Head (including eyes) higher than wide (fig. 14); front narrow, at

most as wide as long; upper occiput behind eyes strongly con-

vex ^ 4

Head wider than high (fig. 19); front wider than long; upper oc-

ciput behind eyes flat 5

Third antennal segment with promineht excision on inner side (figs.

123, 125) Ospriocerus Loew

Third antennal segment without such excision Stenopogon Loew

Wings with branches of third longitudinal vein slightly distad of

posterior crossvein (fig. 156) 6

Branches of third longitudinal vein clearly proximad to posterior

crossvein (fig. 158) 10

Dor socentrals (at least behind mesonotal suture) and scutellars

present; front forming an almost right angle with vertex. ... 7

Dor socentrals and scutellars absent; front almost horizontal,

slightly arched, not forming sharp angle with vertex 9

numerals absent Nicocles Jaennicke

numerals present 8

Metapleuron pilose or with bristles; third antennal s egment taper-

ing apically (fig. 130); no bent spine on apex of front tibia . . .

Adisoemarto

9

Heteropogon Loew

Metapleuron bare, third antennal segment dilated subapically (fig.

128); apex of front tibia with a bent spine ventrally

Lestomyia Willis ton

9. Hind basitarsus and hind tibia swollen distally (fig. 84); face al-

most flat (fig. 16); mesopleuron pilose Holopogon Loew

Hind basitarsus not swollen, hind tibia slender (fig. 79); gibbosity

rounded; mesopleuron with stiff hairs. Cyrtopogon Ljoew

10. Front tibia with curved spine at apex ventrally (fig. 77)

Comanfe//a Curran

Front tibia without such a spine Eucyrtopogon Curran

Genus Stichopogon Loew

Stichopogon Lioew 1847:499. Type species: Dasypogon elegantulus Wiedemainn.

Stilopogon Costa 1883 : 62. Type species: Stilopogon aequidnctus Costa.

Neopogon Bezzi 1910a : 147. Type species: Dasypogon trifasciata Say.

Lissoteles Bezzi 1910b : 177. Type species: Lissoteles hermanni Bezzi.

The genus Neopogon Bezzi was consider ed as a different genus from

Stichopogon Loew by Hull (1962) on the basis of the characteristics of the

chaetotaxy, the palpi, the vertex, and the body size. Curran (1934)

recognized only one genus under the name Neopogon Bezzi. Both Hull and

Curran treated Lissoteles Bezzi as a differ ent genus from Stichopogon Loew.

Hull further added three subgenera to the genus Stichopogon (s. s. ), namely

Dichropogon ezzi , Echinopogon Bezzi, and Cryptopogon White. Bromley (1951)

considered Neopogon Bezzi, Lissoteles Bezzi Echinopogon Bezzi, and Dichropogon

Bezzi, as synonyms of Stichopogon Loew.

Two species of this genus, argenteus Say and triiasciatus Say, are

known from Alberta.

Key to the Species of Stichopogon of Alberta

Uniformly silvery pollinose; mesonotal and abdominal pile long;

bristles on fir st abdominal segment weak, hardly distinguishable

from pile argenteus Say

Abdomen with black markings on dorsum of second, third, fifth,

and sixth segments; mesonotum with short suppressed hairs;

bristles on first abdominal segment strong and distinct

triiasciatus Say

Stichopogon argenteus Say

Dasypogon argenteus Ssiy 1823 : 51; 1869 : 65.

Stichopogon argenteus Back 1909 : 334.

This species is easily recognized by its uniform silvery color of

the pollen and the hairs. Antennae brownish black, bristles present on

lower sides of first two segments and upper side of second segment;

10

Asilidae of Alberta

face flat with slight elevation above epistoma (fig. 2); mystax of few

rows; palpus one-third as long as labium, pilose sub-basally; proboscis

with basal one-third of lower side silvery ^ollinos e, basal half pilose.

Thoracic pile long; scutellum with long hairs along posterior margin;

one presutural, onepostalar, and a row of metapleural bristles present.

Legs with pile anterolaterally on front coxa, laterally on middle coxa and

hind coxa, and on lower sides of femora; bristles present on tibiae and

tarsi; last tarsal segment, empodium, and pulvilli of equal length,

claws slightly longer than last tarsal segment. Wings clear, evenly

covered with micr otrichiae; mediocubital vein absent, and Cui fused

for a short distance. Abdomen elongate, pile longest on first two seg-

ments; ventral pile absent from first segment, shorter in females.

Slight variation occurs among the specimens, in the length of the first

segment of the style and the length of the fusion of M3 and Cu^.

The presence of this species in Alberta is doubted, but Strickland

(1938) included it in his list. It has been recorded from Manitoba to

Colorado, west to California, east to New York, south to Maryland; 22

specimens were examined.

Localities - MANITOBA: Onah (CNC). ONTARIO: Grand Bend (CNC). NEW YORK: Oak

Beach, Long Island (UA); Fire Island (AMNH); New York City (USNM). NEW JERSEY: Avalon

(USNM); Sea Side Park (USNM). ILLINOIS: Lake Forest (USNM). KANSAS: Medora (USNM).

CALIFORNIA: Los Angeles.

Stichopogon trifasciatus Say

Dasypogon trifasciatus Say 1823 : 51, 1869 : 64.

Dasypogon candidus Macquart 1846 : 67.

Dasypogon fasciventris Macquart 1850 : 69.

Dasypogon ge/ascens Walker I860 *. 277.

Stichopogon trifasciatus Williston 1886 : 289.

This species is easily distinguished from argenteus Say, by the char-

acters in the above key. Antennae with bristles on lower sides of first

two segments, and apical upper side of second; front and vertex golden

yellow pollinose; face pale yellow pollinose; mystax white, single row,

along upper mar gin of epistoma; palpus one -fifth as long as labium; pro-

boscis black, with silvery white pollen on basal half of lower side. Thor-

acic pile spar se, absent from mesonotum; the latter provided with semi -

appressed short black hairs; one presutural, one postalar, a row of six

metapleural bristles present. Legs with sparse pile; bristles present

onapicesof femora, on tibiae, and on tarsi. Wings clear, evenly cover ed

with micr otrichiae; veins brown; mediocubital cr os svein present, short.

Abdomen silvery white pollinose, with triangular black markings on

second, third, fifth, and sixth segments, with apices facing forward

(fig. 87); pile very sparse, short, but longer on side of first segment.

The antennae and the mediocubital crossvein vary slightly.

This species seems to prefer bar e areas, including exposed rocks,

wind blown areas, and stream sides (James 1938), open beach, sand

plains, restricted sandy or gravelly areas (Bromley 1946), and also

pastures or bare fields (including unpaved roads) near streams. It is

widely distributed in the United States and Canada; 63 specimens were

examined.

Adisoemarto

11

Localities - ALBERTA: Edgerton; Medicine Hat (UA).

Other localities - MANITOBA: Aweme; Onah (CNC). CALIFORNIA: San Diego Co. (CNC).

ARIZONA: Madera Canyon, St. Rita (U.A). NEW MEXICO: Silver City (UA), TEXAS: Brazos;

Madison; Frio; Be.xar; Travis; Burleson. WYOMING: New Castle, Weston Co. (AMNH) ,

NEBRASK-^: Broken Bow (AMNH). IOWA: Iowa City (AMNH); Ames (AMNH). ONTARIO:

Point Pelee (UA CNC); Orilla (UA); Grand Bend (CNC). QUEBEC: Hull (CNC). NEW YORK:

Long Island (AMNH).

Genus Lasiopogon Loew

Lasiopogon Loew 1847 : 508. Type species: Dasypogon pilosellus Loew.

Daulopogon Loew 1874 : 377.

This genus and Stichopogon Loew are grouped in the tribe Sticho-

pogonini (Hull 1962). Species of both genera have a complete prosternum

(fig. 66), wide front and vertex (fig. 9), but they are distinguished by

the shape of the gibbosity and the mystax (fig. 8).

Antenna with abundant strong hairs on lower sides of first two seg-

ments and apical upper side of second; third segment with apical style

(figs. 118-121); ocellar bristles present; palpus one-segmented; left

and right car dostipites separate, held together by membrane; upper half

of occiput usually with bristles, at least present behind orbital margin

as continuation of verticals; bristle-like hairs present on fr ontovertex,

except in Melander and terricola Johnson. Bristles on thorax

mostly onmesonotum; dor socentrals absent from terricola Johnson; Meta-

pleuron always with a vertical row of bristles; short or longer bristle-

like hair s on humeri; posterior mar gin of mesopleur on with long bristle-

like hairs, except in trivittatus Melander and terricola Johnson; pale pile on

upper posterior corner of sternopleur on. Bristle-like hairs present

anteriorly on front coxa, laterally on middle coxa, and sparsely on hind

pair; femoral bristles if present, subapical; tibial bristles arranged in

five rows, nine to 11 subapical bristles also present; tarsal bristles

arranged in circles. Wings hyaline, evenly cover ed with micr otrichiae;

marginal cell open, two submarginal cells always present, open; four

posterior cells always open; anal cell always closed; anterior cr os svein

always before middle of discal cell. Sides of first abdominal segment

usuallywith bristles and pale pile; male genitalia rotated 180°, bristles

present on hypandrium; ovipositor with acanthophorite s and spines,

valves of eighth sternum prominent (figs. 106, 107).

Nine species of this genus are present or listed as occurring in

Alberta. The record of one of them, ripicola Melander, is doubted,

while prima is described as a new species.

Key to the Species of Lasiopogon Loew of Alberta

1. Mystax entirely white 2

Mystax entirely black or mixed black and white 5

2. Dor socentrals and scutellars absent terricola Johnson

Dor socentrals and scutellars present 3

3. Scutellars white quadrivittatus Jones

Scutellars black 4

4. Two scutellars (fig. 67); mesonotum with few setulae; meta-

pleural bristles vdiite trivittatus Melander

12

Asilidae of Alberta

5.

6.

7.

8.

9.

10.

Scutellar bristles numerous (fig. 68); mesonotum with more or

less numerous long hairs ripicola Melander

Mystax mixed white and black prima new species

Mystax entirely black 6

Apical abdominal bands absent; pollen if present, not forming

definite bands hinei Cole and Wilcox

Pollinose apical bands present and definite 7

Abdominal bands golden yellow, less than one-fourth of correspon-

ding abdominal segments, the rest of segment more or less

shining black .canus Cole and Wilcox

Abdominal bands greyish, wider than one-fourth of corresponding

segments, the rest of segment mostly brown 8

Male 9

F emale 10

Genitalia with superior forceps (surstyli) broad, length less than

twice the width; hypandrial bristles convergent

aldrichi Melander

Superior forceps with length four times the apical width (fig. 180);

hypandrial bristles more or less parallel cinereus Cole

Ovipositor not entirely black, valve of eighth sternum orange . . .

aldrichi Melander

Ovipositor entirely black cinereus Cole

Lasiopogon terricola Johnson

Daulopogon terricola Johnson 1900 : 326.

Lasiopogon terricola Back 1919 : 300-301.

Alexiopogon terricola Curran 1934 : 183.

Curran (1934) separated this species from the rest of Lasiopogon

Loew, and erected a new genus for it, Alexiopogon . However, the following

characters of this species show that it belongs in Lasiopogon Loew: the

shape of the gibbosity and the mystax, the size of the front and vertex,

the presence of the ocellar and the occipital bristles, the presence of the

long sparse pile on the vertex, the presence of the short hairs on the

humerous, the mouthparts, the male genitalia, and the ovipositor . Front

and vertex widened posteriorly, golden yellow pollinose, with a pair of

parallel grooves, convergent toward neck (fig. 7); bristles on head pale

yellowish; face oale yellowish pollinose. Thorax golden yellow pollinose,

paler on pleura; one or two presuturals, one or two intraalars, one or

two postalar s, black; metapleural bristles pale yellowish; dor socentrals

and scutellar s absent; mesonotum with a pair of brownish vittae and short

semi-appressed pale yellow hairs. Legs brownish; coxae black, pol-

linose; femora black dorsally; tibiae with pale yellowish hairs and pale

andblackbristles; tarsi with blackbristles; claws reddish brown ba sally,

black apically; empodium black; pulvilli yellow. Wings slightly longer

than abdomen; veins brownish. Abdomen shiny black, reddish brown

apically; very short, sparse, appressed pale hairs present; side of

first segment with weak bristles or bristle-like hairs; male genitalia

reddish, hypandrial bristles pale yellow, convergent; ovipositor reddish

Adisoemarto

13

brown.

This species is found on the low damp ground (Johnson 1900; Cole

and Wilcox 1938) as well as on dry sand bars or bare sand dunes. It

ranges from Alberta to Mas sachusetts, south to Virginia; 69 specimens

were examined.

Localities - ALBERTA; Fabyan (UA); Wainwright (UA); Provost (UA); Manyberries-

Orion (UA); Writing- on-Stone Provincial Park (UA); Lethbridge, OldmanRiver (UA); Medicine

Hat.

Other localities - NORTH DAKOTA: Mott (CNC), INDIANA: Bare Sand, Lafayette. OHIQ

Pine Creek, Hocking Co. VIRGINIA: Great Falls. MARYLAND: Plummer ' s I sland; Beltsville.

NEW JERSEY: Clementon; Lahaway, Ocean Co.; Riverton; Wenonah. MASSACHUSETTS:

Amherst; Chicopee.

Lasiopogon trivittatus Mel and er

Lasiopogon trivittatus Melander 1923b : 144-145.

Males of this species are described for the first time. Vertex

golden yellow pollinose; short stiff hairs present as a row of three to

four between grooves and orbital margin, and two pairs in front of ocel-

lar plate; ocellars black; occiput yellowish grey pollinose, occipital

bristles on upper margin and transversely behind vertex; mystax pale

yellow; antennae black, hairs on lower and apical upper sides of first

two segments black; proboscis black, pile on lower basal half white;

beard white. Prothorax golden yellow pollinose, paler toward ventral

sides; pile on pronotum, on episternum, and on epimeron, pale yellow-

ish; mesonotum grey to yellowish pollinose; dor socentral and acrostichal

vittae present, complete; presutural, intraalar, and postalar bristles

always single; presutural dor socentrals usually two, rarely one; post-

humerals sometimes present; postsutural dor socentrals always two;

setulae present on mesonotum in front of suture; scutellars black, two,

sometimes with two black and few white setulae; metapleurals pale yel-

lowish. Legs grey pollinose; coxal hairs pale yellowish; lower sides

of femora with pale bristles and pile; tibial and tarsal bristles black,

absent from ventral surfaces; claws brown, tips black; empodium black.

Wings hyaline, evenly covered with micr otrichiae; anterior crossvein

at basal one-third or half of the length of discal cell; anal cell closed.

Abdomen brownish pollinose basally, posterior one-third to half greyish

pollinose, extending forward on lateral margins, sides of first segment

with bristles and sparse pile; appressed setulae on all abdominal seg-

ments; male genitalia (figs. 175-179), black, yellowish grey pollinose;

hypandrial bristles convergent. Number and color of bristles and setulae,

and total length (6. 0 - 9. 0 mm) vary. In very rare cases, one or two

black bristles are present among white mystax.

This species is abundant along river banks, often re sting on rocks .

Red mites were found attached to the ventral side of the neck and behind

the hind coxae (membraneous parts) of a female specimen from Luscar,

Alberta (UA). Another species, Lasiopogon dnereus Cole, collected from

the same locality, was also found to have the same species of mites

associated with it. Lasiopogon trivittatus Melander in some localities, is

associated with L. dnereus Cole and L. quadrivittatus Jones. This species

has been recorded from Montana and Alberta; 148 specimens were ex-

14

Asilidae of Alberta

amined.

Localities - ALBERTA: Flatbush, Pembina River (UA); Edmonton, Emily Murphy Park

(UA), Beverly Municipal Dump (UA and LMK); Luscar, McLeod River (UA); Red Deer River,

Red Deer (UA and LMK); Drumheller (UA); Dinosaur Park (UA); Nordegg, North Saskatchewan

River Valley (UA and LMK); Crowsnest Forest, Dutch Creek (UA); Banff, Eisenhower lookout

(CNC).

Other localities - MONTANA: Gold Creek.

Lasiopogon quadrivittatus Jones

Lasiopogon quadrivittatus Jones 1907 : 278.

Among the species of Lasiopogon Loew occurring in Alberta, this is

the most easily recognized, for the bristles are all pale. In general

appearance it is similar to ripicola Melander, but the latter has black

scutellar bristles. Face and lower occiput gr ey pollinose; front vertex,

and upper occiput yellowish pollinosej bristle s and hair s pale yellowish.

Thorax golden yellow pollinose; dorsocentral vittae rusty brown, with

golden orange lining; acrostichal vittagrey or golden yellow; space be-

tween dorsocentral and acrostichal vittae brownish, giving appearance

of four vittae; six dor socentrals, two to three before suture; post-

humerals present or absent; presuturals two to three; intraalars two;

postalar s two; scutellars six; mesopleural and sternopleural pile white;

metapleural bristles five to eight. Legs light yellowish pollinose; middle

and hind pairs less pilose; hind femora with a row of bristles on anterior

sides; claws reddish brown basally, black apically; empodium black.

Wings hyaline, vein brownish; fourth posterior cell open, narrower or

wider than the first; anal cell closed. Abdomen grey pollinose; bristles

and pile pale yellowish; a pair of basal semicircular brownmarkings on

each, except fir st segment;, male genitalia black, golden yellow pollinose,

hairs and bristles pale yellowish; hypandrial bristles convergent; ovi-

positor black, yellowish pilose. Total length 7.0- 10. 0 mm inmales,

and 8.0- 11. 5 mm in females; number of metapleural and mesonotal

bristles exhibit variation.

In southern Alberta this species is common in late spring, but it

appears later in the northern parts of the province. It has been found

associated with Lasiopogon terricola Johnson, L. trivittatus Melander, L. cartus

Cole and Wilcox, Eucyrtopogon albibarbis Curran, and Asilus aridalis n. sp.

This species inhabits several different habitats: bare paths, along river

banks, and sand dunes near river . It ranges from Alberta and Wyoming,

east to Nebraska; 134 specimens were examined.

Localities - ALBERTA: Edmonton, Beverly Municipal Dump (UA and LMK), Country Club

(LMK), Emily Murphy Park (UA), White Mud Park (UA); Fabyan, Campsite (UA); Bind loss (UA);

Empress (UA); Sandy Point Bf-idge (UA); Army Expt. Sta. (UA); Medicine Hat (UA; CNC);

Seven Persons (UA); Burdett (UA); Pendant d'Oreille (UA); Writing- on-Stone Provincial Park

(UA); Milk River (CNC); Lethbridge (UA; CNC); Taber (UA) ; Dinosaur Park (UA) ; Drumheller

(UA); Calgary (CNC).

Other localities - MONTANA: "Montana, C.U.". WYOMING. nArASKA: Halsey War

Bonnet Canyon; Bad Lands; Mouth of Monroe Canyon. NORTH DAKOTA: Bismarck.

Adisoemarto

15

Lasiopogon ripicola Melander

Lasiopogon ripicola Melander 1923b : 143-144.

This species is similar to Lasiopogon quadrivittatus Jones, but is dis-

tinguished by the black color of the scutellar bristles; the male genitalia

are also different.

The presence of this species in Alberta is doubted, but it was in-

cluded by Strickland (1946) in his list. It ranges from Washington and

Idaho to California; seven specimens were examined.

Localities - WASHINGTON: Wayawai (CNC) ; Pasco (USNM); Cashmere. IDAHO: Lewiston.

OREGON: The Dalles.

Lasiopogon cinereus Cole

Lasiopogon cinereus Cole 1919 I 229.

This species is distinguished from the others by the following char-

acters: the black mystax, the wide grey bands on the abdominal posterior

sides, the shape of the superior forceps of the male genitalia (tapering

apically), and the entirely black ovipositor . Face grey pollinose, mystax

as long as antenna; front and vertex yellowish tinged; frontal and ver-

tical hairs weak; brownish transverse band across lateral ocelli; an-

tennae black, fir st two segments with black hairs. Thorax grey pollinose;

prothorax yellowish pilose; mesonotal hairs black, long; dorsocentral

bristles weak, two before suture, three to four behind suture; two to

three presuturals; posthumerals present or absent; humeri yellowish

tinged, black hairs present; mesopleuron with hairs on front half of

upper margin and posterior upper corner, pale yellow pile present on

posterior corner of sternopleur on; metapleural bristles black, mixed

with white pile; scutellum yellowish grey pollinose, bristles black. Legs

average for the genus, with long pale pile on lower sides of femora;

bristles black. Wings hyaline, slightly infuscated; halteres brownish.

Basal three-quarters of abdominal segments rusty brown pollinos^

apical one-fourth grey pollinose; long pale yellowish pile present on

lateral sides of first four of male and first two of female abdominal seg-

ments; last four segments of males and last five segments of females

with black setulae; bristles present on sides of first segment; venter

long yellowish pilose; male genitalia (figs. 180-184) black, superior

forceps yellowish grey pollinose, blackhaired; hypandrial bristles black,

convergent; ovipositor black, sparsely yellowish pilose, spines black.

This species has been found associated with Lasiopogon trivittatus

Melander. The adults are active, flying from rock to rock in the river,

or along river banks. It ranges from Alberta to California, east to Utah

and Colorado; 51 specimens were examined.

Localities - ALBERTA: Nordegg, North Saskatchewan River (LMK); Luscar, McLeod

River (UA); Red Deer (UA); Cr owsne st F or est , Wilkins on Creek (U A) , Dutch Creek (UA) ; Banff

(CNC); Frank (CNC); Waterton (CNC); Blakiston Brook, Waterton Park (UA).

Other localities - WASHINGTON: Blewett; Buckley; Cle Elum; Gaynor; Goldendale;

Kalama River; Lake Cushman, Mason Co. ; Mt. Rainier, Ipaut Creek Camp, Old White River

Entrance; Naches; Rainier National Forest, Indian Flat Camp, Lodgepole Camp; Satus Creek;

Virden; Walla Walla (CNC). OREGON: Mehama (AMNH); Hood River; Joseph; Lebanon;

16

Asilidae of Alberta

Wallowa Lake. CALIFORNIA; Tuolumne Meadows , Yosemite Park. MONTANA: edge of Mus sel-

shell River, Winnecook. WYOLIING: near Lander; Thumb Station, Yellowstone National Park.

UTAH; Uinta Mountains; Duchesne Mountain; Sheep Creek, Duchesne Co. COLORADO: Rock-

wood (USNAl).

Lasiopogon prima new species

This species is readily distinguished from the rest of the Albertan

species by the color of the mystax, which is mixed black and white. The

genitalia are also diagnostic of the species; the superior forceps (sur styli

of Cole) are provided with disc-like projections on the inner sides (fig.

187).

Male. Face greyish yellowpollinose; lower side of mystax white,

upper side black; front and vertex dull greyish yellow pollinose; fronto-

orbital hairs two rows; hairs in front of ocellar plate black, abundant;

ocellar bristles black; upper half of occiput dull greyish yellowpollinose,

lower half greyish pollinose; antennae black (fig. 119); hairs on first

two segments black; style half as long as third s egment; proboscis black,

palpus black, one-eighth as long as labium. Prothorax brownish grey

pollinose, white pilose; mesonotum greyish pollinose, dor socentral vittae

brown, acr ostichal vitta faint, ended at mesonotal suture, lateral meso-

notal margins brownish; hairs and bristles black; four left and three

right presuturals, one left and two right intraalars, one pair postalar,

six pair s presutural dor socentrals, three left and unidentified right post-

sutural dor soc 3ntrals ; scutellum black, greyish pollinose, eight bristles

black, mixed with black hairs; mesopleuron yellowish grey pollinose,

paler on lower side, black hairs on upper posterior corner; upper pos-

terior corner of sternopleur on whitish pilose; a row of eight metapleural

bristles black. Legs black, average for Lasiopogon Loew; pile on coxae,

on femora, and on tibiae white; left front femur with two, right front

femur with four bristles on dor soposterior surface, two bristles on mid-

dle pair, a row of six on left and five on right anterior side of hind fem-

ora; front and hind tibiae with three rows of four bristles on dorsal sur-

faces, middle pair with four rows; tibiae with nine to twelve apical bris-

tles; tarsal bristles arranged in circle subapically; claws brown basally,

black apically; pulvilli tawny, empodium black, as long as pulvilli.

Wings hyaline, evenly cover ed with microtrichiae, veins brown; anterior

crossvein at middle of discal cell; fourth posterior cell open, as wide

as first, fifth three times as wide as fourth; anal cell closed at margin

(fig. 148). Abdomen shining black, yellowish grey pollinose on apices,

extending forwa-»-'i at sides and middle, leaving a pair of black spots on

each segment; yellowish white pile on lateral sides of first four seg-

ments, semiappr es sed on the rest, venter grey pollinose, white pilose;

male genitalia black, hypandrial bristles black, convergent, superior

forceps broad basally, tapering apically, with disc-like projection on

ventral inner side (fig. 187).

Females. Except for the number and position of the bristles, fe-

males of this species are similar to the males; bristles on first ab-

dominal segment mixed with black; last four segments with setulae on

lateral sides; ovipositor black.

This species varies individually in the number of bristles, es-

Adisoemarto

17

pecially those on the‘‘mesonotum, the position of the anterior crossvein,

and the width of the fourth posterior cell. Total length is from 7.0 - 9.0

mm. The habitat is the same as that of the other species of Lasiupogon

Loew.

Holotype: male, Nordegg, North Saskatchewan River Valley,

Alberta, 28-V-1963 (Adisoemarto, Freitag, Ball, collectors); deposited

in CNC.

Paratypes: one male, three females, same data; male, female.

North Saskatchewan River, near Rocky Mountain House, Alberta, 29- V-

1963, same collectors; one male, three females. Garth, Alberta, same

collectors; male, Brazeau Dam, Lodgepole, Alberta, 9-VII-1964 (L.

M. Kenakin) . All these localities are on the eastern slopes of the Rocky

Mountains, in the vicinity of the North Saskatchewan River. Except the

last specimen, kept in LMK collection, the paratypes are deposited in

UA collection.

The name prima has been chosen, because this species was the first

asilid collected in 1963, on an expedition to the Rocky Mountains.

Lasiopogon canus Cole and Wilcox

Lasiopogon canus Cole and Wilcox 1938 : 32-34.

According to Cole and Wilcox (1938), this species in general ap-

pearance resembles European rather than North American members of

the genus . It is distinguished from the other species by the black mystax

and the narrow golden yellow abdominal bands. Face, front, and vertex

golden yellow pollinose; hair s and bristles black; hair s in front of ocellar

plate weak; antennae and hairs on first two segments black; occiput

golden yellow pollinose, paler toward chin, bristles black; beard yel-

lowish white. Thorax golden yellow pollinos e; dor socentral vittae brown,

not reaching humeri, acrostichal vitta faint, incomplete; bristles and

hairs entirely black. Legs golden yellow pollinose; coxae greyish pol-

linose; pile entirely golden yellow. Wings hyaline, brownish, covered

with microtrichiae; fourth posterior cell varies from closed to widely

open; anal cell closed with stalk. Abdomen black, slightly goldenyellow

pollinose, pile yellow; in some specimens, last three segments with

blcLck setulae on lateral sides; male genitalia shining black, superior

forceps broad basally, tapering apically, bristles black, conv'ergent on

hypandrium; ovipositor with orange valves, but comparatively shorter

and broader than those of aldrichi Melander. The bristles on the sides of

the first abdominal segment vary in number, from six to eight, and are

all black, or black mixed with white.

The species is found on barepaths or gravelly river banks. A few

specimens, which were probably just emerged, have the male genitalia

not completely inverted or still in uninverted situation. These specimens

are kept in UA. It is known from Alaska and Alberta; 32 specimens

were examined.

J-.ocali ti es - A LBER TA ; R ocky Mountain House, North Saskatchewan R iver (UA) ; Edmonton,

Whitemud Park (UA), Emily Murphy Park (UA), Country Club (LMK).

Other localities - ALASKA; Savonoski, Naknek Lake; Healy; Fairbanks.

18

Asilidae of Alberta

Lasiopogon hinei Cole and Wilcox

Lasiopogon hinei Cole and Wilcox 1938 : 51-53.

This species is recognized by the obscure abdominal pollen^, not

arranged as apical bands, and also by the long and dense pile on the ab-

domen of the males. Face, front, and vertex yellow pollinose; frontal

and vertical hairs long, black, abundant, continued to occiput; occiput

greyish pollinose; proboscis black, pile yellowish, palpus black, yellow-

ish pilose. Prothorax pale yellowish grey to golden yellow pollinose,

pile yellowish; mesonotum golden yellow pollinose, dorsocentral vittae

brownish to velvety black with golden yellow lining on inner sides, acros-

tichal vitta paler, greyish or golden yellow, incomplete, space between

dorsocentral and acrostichal vittae brownish; dorsocentral bristles

weak, varying from five to eight (two to three presutural, and three to

five postsutural) ; posthumerals present or absent; two to three pre-

suturals; two to three intraalars; two pairs of postalars; usually eight

scutellars, sometimes hardly distinguishable from hairs; metapleural

bristles eight to nine in a row, black, mixed with pale yellowish pile.

Legs golden yellow pollinose; numerous long black bristles present on

apical halves of femora and tibiae; claws brownish basally, black apically.

Wings hyaline, slightly infuscated; fourth posterior cell as wide or half

as wide as first; anal cell always closed, with stalk. Ground color of

abdomen black, pollen yellowish grey; long yellowish pile present on

first; anal cell always closed, with stalk. Ground color of abdomen

black, pollen yellowish grey; long yellowish pile present on first four

abdominal segments of males or first three of females, the rest seg-

ments with brownish black hairs; bristles on first abdominal segment

pale yellowish, on male genitalia black; hypandrium orange, bristles

convergent; ovipositor black, spines black, valves brownish orange.

The number of bristles on the mesonotum varies.

The adults of this species, in Alberta, have been found along bare

paths near streams or rivers with grasses or bushes next to them. It

is known from Alaska and Alberta; 14 specimens were examined.

Localities - ALBERTA; Flatbush, Pembina River (UA); Edmonton, R ainbow Valley (UA),

Whitemud Park (UA); Rocky Mountain House, North Saskatchewan River (UA).

Other localities - ALASK-V; Katmai.

Lasiopogon aldrichi Melander

Lasiopogon a/cfric/?/ Melander 1923b : 139-140.

The females of this species and of canus Cole and Wilcox have

orange valves of the ovipositor, but they are distinguished from one

another by shape. The males of this species ar e recognized by the shape

of the superior forceps. Slight variation occur s in the shape of the third

antennal segment (figs. 120, 121). The number of mesonotal bristles

varies . The fourth posterior cell varies from completely closed to widely

open. A female specimen from Drumheller (UA), Alberta, is slightly

different from the other specimens with respect to the shape of the ovi-

positor (fig. 107). This specimen might belong to Lasiopogon pacificus Cole

and Wilcox.

This species ranges from British Columbia and Alberta to Califor-

Adisoemarto

19

nia, east to Utah and Colorado; 48 specimens were examined.

Localities - ALBERTA: Banff (UA and CNC); Drumheller (UA).

Other localities - BRITISH COLUMBIA: Robson (CNC). OREGON: Mt. Hood (USHM);

Anthony Lake; Blue Mountains, Tollgate; Fish Lake, Steins Mts.; Haines; Strawberry Mt. ,

Grant Co. ; Sumpter; Wallowa Lake, Aneroid Lake Trail. WASHINGTON: Blue Mts. ; Signal

Peak; White Rock Springs, Steven Pass, Cascade Mts.; Mt. Spokane (USHM). IDAHO: Moscow

Mt. (CNC); Long Valley, Alpha. WYOMING. COLORADO: La Veta Pass. UTAH: Beaver

Creek. CALIFORNIA: Samoa (USNM).

Genus Stenopogon Loew

Stenopogon Loew 1847 : 483. Type species: Asilus sabaudus F. 1794.

Scleropogon Loew 1866 : 26. Type species: Scleropogon picticornis Loew 1866.

This genus contains robust species. Curran (1934) and Hull (1962)

considered this genus { sensu stricto) different from Scleropogon Loew, on the

basis of the absence of pile or hair s from the metapleur on. The definition

was thought by Bromley (1937) to be trivial; he (1951) treated Scleropogon

Loew as a synonym of Stenopogon Loew. Back (1909) was the first to con-

sider these two groups as congeneric.

Head slightly higher than wide; face, front, and vertex narrow;

gibbosity of two types: in ” inquinafus group" gibbosity very prominent,

starting close to antennal base (figs. 12, 13), in "coyote, group", gibbosity

starting farther away (fig. 11), third antennal segment tapers apically,

without obvious apical excavation (fig. 126); style tapers apically. Pro-

thorax with bristles, pile present among bristles and on anterior corner

of sternopleuron and posterior one-third of sternopleuron; bristles or

hairs present on, or absent from, metapleuron; mesonotal bristles more

abundant on posterior half. Legs pilose; front femora with bristles on

apices, middle pair with a row of bristles on anterior sides, hind pair

with two rows on anterior sides; tibiae with three or four rows of bris-

tles; tarsi with bristles subapically. Wings hyaline, axillary cell and

alula fuscous or smokey; second and third veins slightly recurved; an-

terior crossvein at, or slightly before, middle of discal cell; fourth

posterior cell open or closed; anal cell closed or narrowly open (figs.

150, 151). Abdomen more or less cylindrical, elongate, tapering api-

cally; male genital organ not inverted (fig. 93); ovipositor withacantho-

phorites and spines (figs. 89-92).

According to the definition of the genus Ospriocerus Loew by Martin

(pers. comm.), consanguineus Loew and pumilus Coquillett belong to that

genus, not to Stenopogon Loew.

There are five species in Alberta: obscmiventris Loew, rufibarbis

Bromley, inquinatus Loew, coyote Bromley, and neglectus Bromley, but

Strickland (1938 and 1946) included also gratus Loew in his lists.

Key to the Species oi Stenopogon Loew of Alberta

1. Metapleuron with hairs, or with weak or strong bristles 2

Metapleuron without hairs or bristles, at most tomentose or pol-

linose 3

Wings with first and fourth, posterior cells open; abdomen black-

2.

20

Asilidae of Alberta

3.

4.

5.

6 .

ish or less pollinose; first antennal segment blackish

neglectus Bromley

First posterior cell narrow at tip or sometimes closed with stalk

(figs. 150, 151); fourth posterior cell closed with stalk; ab-

domen greenish grey pollinose; first antennal segment brownish

orange coyote Bromley

Abdominal dorsum uniformly black 4

Abdominal dorsum reddish brown, black only on sides 6

Humeri orange brown inquinatus Loew

Humeri black 5

Evenly greyish pollinose species; pile and bristles yellow

obscuriventris Loew

Darker, bright orange pollinose; pile and bristles bright orange.

rufiharbis Bromley

Humeri orange-brown, covered withgreyish pollen, inquinatus Lioew

Humeri black, covered with yellowish orange pollen. . gratus Loew

Stenopogon obscuriventris Loew

Stenopogon obscuriventris hioew 1872 : 30.

This species is easily recognized by the uniform greyish pollen

and yellow bristles and pile. Back (1909) treated this species as con-

specific with californiae Walker. Antennal segments unicolored; style

orange brown apically; palpal segments equal (fig. 42); gibbosity almost

touching antennal base (fig. 12). Thorax unicolored; prothorax with

bristles only on pronotum; presutural dor socentrals absent; humerals

absent; dorsocentral vittae blackish brown; pile on anterior corner and

posterior half of sternopleur on long. Coxae and basal three-fourths of

femora black, the rest yellowish; claws brownish ba sally, black apically.

Wings hyaline, veins brownish; in males, axillary cell and alula tinged

silvery white, less obviously so in females; all posterior cells open;

anal cell opennarrowly or almost closed. Abdomen unicolored; pile on

first three segments long; male genitalia (figs. 194-198) and ovipositor

orange brown. Number of bristles varies. Sexual dimorphismis shown

only by the white infuscation on the wings of the males.

This species ranges from Alberta andColorado, west to California;

23 specimens were examined.

Localities - ALBERTA; Czar (UA); Medictne Hat (CNC); Lethbridge (CNC).

Other localities - SASKATCHEWAN: Pike Lake; Great Sand Hills, west of Swift Current.

OREGON; Summer Lake; Chewaucan R., near Paisley. IDAHO: Victor (AMNH); Giveout

(AMNH); Mt. Pelier (AMNH). WYOMING: Jackson (AMNH); Rawlins (AMNH); Green River

(AMNH); Medicine Bow (AMNH); Carbon (AMNH); Rock Spring (AMNH); Centenial (AMNH).

UTAH: Promontory Point (USNM); Huntsville (USNM); Logan Canyon (USNM). COLORADO:

Animas (AMNH); Monte Vista (AMNH); Ouray (AMNH); Jefferson (AMNH); Blanca (USNM).

ARIAONA: Kaibab Forest, Grand Canyon. CALIFORNIA: Mone Lake (AMNH); Mariposa Co.

(A.MNH); Mount Diablo (AMNH); Mt. Hamilton (AMNH).

Stenopogon rufibarbis Bromley

Stenopogon rufibarbis Bromley 1931 : 431.

Adisoemarto

21

This species is very similar to obscuriventris Loew. The male geni-

talia (figs. 199-203) and the ovipositors of these two species are very

slightly different from one another . The two species may be distinguished

by the different color of pollen, pile, and bristles.

This species ranges from British Columbia to Arizona, and east

to Utah. Strickland (1938) included this species in his list, but the record

was based on misidentified specimens of Stenopogon obscvriventris Loew. It

probably does not occur in Alberta; 31 specimens were examined.

Localities - BRITISH COLUMBIA: Osoyoos; Anderson Lake; Seton Lake; Oliver.

WASHINGTON. OREGON: Cherry Creek, Klamath Lake; Alberta Lake. IDAHO: Giveout

(AMNH). CALIFORNIA: Keddi Plumas Co. (AiVINH) ; Sierra Nevada (AMNH) ; Coleville (AMNH) ;

Philo Mendocino (AMNH); Mt. Hamilton (AMNH); Feather River (AMNH); Butte Co. (AMNH);

Cedar ville (AMNH) ; CTioPlumas Co. (AMNH); Lassen Co.; San Antonio, Ontario; LosAngeles;

Pasadena; Lake of W'oods; Echo Portals, Eldorado Co. UTAH: St. George (AMNH) . ARIZONA:

Lacobs I.ake.

Stenopogon grains Loew

Stenopogon grains Ljoew 1872 : 31.

Stenopogon nnivittatns ~LiOew 1874 : 358.

This species is similar to californiae Walker, but can be distinguished

by the mesonotal vestiture and the color of the pile and bristles. The

bristles and pile are orange, darker than those of californiae Loew, and

the mesonotum is provided with longer dorsocentral black hairs. The

male genitalia are also differ ent in the shape of the hypandrium and of the

superior forceps (figs. 204-217).

This species is known from California only, but Strickland (1938)

included it in his list; two specimens were examined.

Localities - CALIFORNIA: San Francisco (USNM).

Stenopogon inqninatns Loew

Stenopogon inqninatns ]_,oeV\/ 1866 : 47.

Stenopogon modestns Loew 1866 : 46.

Stenopogon morosns Loew 1874 : 356.

This species is distinguished from grains Loew, by the reddish

brown humeri. There are two forms: one with reddish brown abdomen,

the other with black abdomen.

Brown form: front and vertex greyish yellow pollinose; gibbosity

very prominent (fig. 13); antennae brownish or reddish black; proboscis

and palpi black. Thoracic ground color black, humeri reddish brown;

pollen greyish yellow; prothorax pilose, pronotum and episternum with

bristles; dorsocentral vittae brownish black; presutural dorsocentral

bristles absent; metapleuron bare; scutellum reddish brown with black

posterior edge. Coxae and dor sal sides of femora black, the rest reddish

brown; basal one third of claws reddish brown, the rest black. Wings

hyaline, s emi - infuscated; posterior cells open; anterior crossvein

slightly before, or at the middle of discal cell. Abdomen reddish brown

on the middle, black on lateral sides; venter black; pile long on sides

of first two segments, shorter and sparser on the following segments;

22

Asilidae of Alberta

ventral pile long; male genitalia orange brown with black hairs; apical

end of eighth segment of females with lateral pits submarginally (fig.

91); acanthophorite orange brown, spines black.

Black form: this differ s only in the coloration. Trochanter s black,

femora black with reddish brown apices; femoral bristles black; ab-

domen black, eighth segment in both sexes reddish brown with apical

black band; male genitalia the same as those in brown form; acantho-

phorites reddish brown, spines black.

In addition to these forms, there is also intermediate form, with

broad lateral sides of abdomen black and narrow middle part reddish

brown.

In Colorado, this species inhabits wheat grassland (James 1938).

In Alberta, it has been collected in various habitats, such as in grassland

of long grass, in semi-arid prairie grassland, on gravelly river banks,

in sand pits, at the edge of, or in the openings in the coniferous forests.

It has been found associated with Asilus callidus Williston. This species

has been recorded from British Columbia eastward to Minnesota, and

south to Arizona; 143 specimens were examined.

Localities - ALBERTA: Peace River (UA); Lac la Biche (UA and LMK); Opal-Coronado

(UA and LMK); Celestine Lake, Jasper National Park; Jasper (CNC); Nordegg, North Saskat-

chewan Valley (UA); Seebe (DE); Banff (UA and CNC); Gorge Creek (UA); Redrock Canyon,

Waterton Lakes Park (UA); Calgary (UA); Lethbridge (CNC); Bow River (CNC); Orion (UA);

Medicine Hat (UA); Steveville- Wardlow (UA); Consort (UA).

Other localities - BRITISH COLUMBIA: Vernon; Nicola Valley; Lillooet (CNC); Aspen

Groove (CNC); Seton Lake (CNC). SASKATCHEWAN; Pike Lake (CNC). MANITOBA: Aweme

(CNC). MINNESOTA: R ed R iver of the North. IDAHO: Victor (AMNH); Giveout (AMNH) ; Mt.

Pelior (AMNH). WYOMING: Jackson (AMNH); Rawlins (AMNH); R ock Springs (AMNH) . NEB-

RASKA: Glen, Sioux Co. ; Spring View Bridge, Point Co. ; West Point. COLORADO: Walsenburg

(AMNH); Monte Vista (AMNH); Alamosa (AMNH); Cochetopa National Forest (AMNH). UTAH;

Hatch (AMNH). ARIZONA: N. Rim Grand Canyon (AMNH); Oracle (AMNH). CALIFORNIA:

Benton (AMNH); Clio Plumas Co. (AMNH).

Stenopogon neglectus Bromley

Stenopogon neglectus Bromley 1931 : 430.

Sc/eropo^on neg/eefus Hull 1962 : 126.

This species is readily recognized by the presence of hairs on the

metapleuron, Stenopogon coyote Bromley has also hairs on the metapleur on,

but the two species are readily distinguished by the differ ence in the wing

venation. The male genitalia are also different (figs. 213-217). In

neglectus Bromley, the superior forceps and the gonopods vary from red-

dish brown to black.

The habitats of this species are mainly pastures, wheat grass of

the grassland (James 1938), long grass prairie, and semi-arid short

grass prairie. It ranges from Alberta to Arizona; 19 specimens were

examined.

Localities - ALBERTA: Medicine Hat (UA); Comrey, Milk River Valley (UA).

Other localities - OREGON: Castle. IDAHO: Lewiston. WYOMING: Lander; Jackson

(AMNH); Carbon Co. (AMNH). COLORADO: Creeds. UTAH: Ac. SL. Dsrt. NEVADA: Fallon

(AMNH). ARIZONA: White Mts. (AMNH).

Adisoemarto

23

Stenopogon coyote Bromley

Stenopogon -coyote Bromley 1931 : 42 9.

This species is similar to neglectus Bromley in having the meta-

pleural hairs, but it is easily distinguished by the wing venation. The

first posterior cell is always narrower apically, and the fourth posterior

cell is always closed with long stalk (figs. 150, 151). The habitat is

similar to that of neglectus Bromley. It ranges from Alberta to Arizona

and New Mexico; 47 specimens were examined.

Localities - ALBERTA: Drumheller (UA and CNC); Steveville- Wardlow (UA); Dinosaur

Trail, Dinosaur Provincial Park (UA); Lake Newell, Kinbrook Island Park (UA); Brooks (CNC);

Medicine Hat (UA); Orion (UA); Writing-on-Stone Park (UA); Conirey, Milk R iver Valley (UA);

Lethbridge (CNC).

Other localities - WYOMING: Lander; Lusk. SOUTH DAKOTA: Custer (USNM) ; Piedmont,

Nowlin Co. (USNM). COLORADO: Walsenburg (CNC); Salida; Poncha Spring; Colorado City.

ARIZONA.

Genus Ospriocerus Loew

Ospriocerus Loew 1866 : 29. Type species: Asilus abdominalis S^y 1823.

This genus is known only from the New World. It is very similar

to Stenopogon Loew, but readily distinguished by the third segment of the

antenna, which has a pit or excavation on the apical lower side (figs.

123, 125). There are three types of style in this genus (Martin pers.

comm.): the hidden type with a spine inside ( Ospriocerus abdominalis Say) ;

the short type with apical pit and spine inside ( Ospriocerus latipennis Loew);

and the Mexican type. The second type of style is like that of Neoscleropogon

Malloch, as described and illustrated by Hull (1962). Most of the char-

acters are like those of Stenopogon Loew; gibbosity not prominent (fig. 10);

metapleuron with hair s; wings broad, fourth posterior cell always closed.

In Alberta, there are two species, Ospriocerus abdominalis Say and

O. consanguineus Loew, but another species, pumilus Coquillett was also in-

cluded by Strickland (1938), probably on the basis of misidentified speci-

mens; two male specimens of Stenopogon coyote Bromley were labelled as

Stenopogon pumilus Coquillett by Curran and Strickland.

Key to the Species of Ospriocerus Loew of Alberta

First antennal segment four times as long as second; style hidden

(fig. 124) abdominalis Say

First antennal segment at most twice as long as second; style

short, with apical pit (figs. 122, 123)

consanguineus Loew and pumilus Coquillett’’'

Ospriocerus abdominalis Say

Asilus abdominalis Ssiy 1823 : 375.

Dasypogon aeacus Wiedemann 1823 : 390.

’•'These two species are hardly distinguishable; they are possibly conspecific.

24

Asilidae of Alberta

Dasypogon spatullatus'Bell3iTdi 1861 : 82.

Ospriocerus ae&cides Loew 1866 : 51.

Ospriocerus abdominalis Coquillett 1898 : 37.

Ospriocerus ventralis Coquillett 1898 : 37.

This species is easily distinguished from the other two by the color

of the abdomen and the wings . Coquillett (1898) distinguished ventralis from

abdominalis Say on the basis of the color of the venter of abdomen, orange

in the former and black in the latter. Head, thorax, legs, and all bris-

tles, and pile black; antennae black, style cryptic; second segment of

palpus spindle-shaped. Wings broad, purplish, infuscated. Abdomen

mostly orange, with first and basal half of second segment black, in

some females lateral mar gins of each segment black; eighth segment of

females black, sixth and seventh segments of some females black; venter

black, orange, or'black and orange; male genitalia and female acantho-

phorites black.

This species has been recorded from the Northwest Territories to

Arizona and Texas, east to Pennsylvania; 24 specimens were examined.

Localities - ALBERTA: Medicine Hat (UA); Chin, prairie coulee (UA).

Other localities - NORTHWEST TERRITORIES. BRITISH COLUMBIA: Oliver (CNC).

SASKATCHEWAN; Roche Percee (CNC). NORTH DAKOTA: Beach (CNC). WYOMING: Carbon

Co. (AMNH); Jackson (AMNH). UTAH: Stockton (CNC); Howel, Dolemite (CNC); Moab, Grand

Co. COLORADO: Mesa Verde (AMNH): Pagosa Spring (AMNH) : Palisade (AMNH) ; Fort Collins;

Colorado Springs ; Spaniard Peak. NEBRASKA: Sioux Co. KANSAS: Golden City. OKLAHOMA:

Optima (AMNH) : W'ichita National Forest (CNC). TEXAS: Travis Co. (AMNH); Austin (AMNH);

Round Mts. NEW MEXICO: Cortez; White's City, Eddy Co. ARIZONA; Carr Canyon, Huachuca

Mts., Cochise Co. ; ’Wilcox (AMNH); Tucson (AMNH). CALIFORNIA. IDAHO: Snake Co.

MONTANA: Lombard. WASHINGTON: Squaw Creek.

Ospriocerus consanguineus Loew

Stenopogon consanguineus Loew 1866 : 48.

Stenopogon latipennisl^oevi/ 1866 : 49.

Ospriocerus consanguineus Martin, pers. comm.

Specimens of this species are easily distinguished from abdominalis

Say by the size of the antennae, coloration, and wing venation. Abdominal

segments greyish pollinose; pile yellowish, longer on lateral sides of

first segment; male genitalia and ovipositor orange brown; gonopods of

male genitalia with hair lamellae subapically (fig. 190),

James (1938) recorded this species from moist sedge meadows,

arid mixed and bunch grassland, and tall weed wasteland, where natural

vegetation has been disturbed. It ranges from Alberta to Manitoba, and

south to Texas; 16 specimens were examined.

Localities - ALBERTA; Medicine Hat (UA).

Other localities - MANITOBA: Onah (CNC). SOUTH DAKOTA: Sioux Co. WYOMING:

Douglas (AMNH)., NEBRASKA: Pierre; Chandron; Agate (CNC). COLORADO: Boulder (AMNH) ;

La Junta (AMNH); Regnier; Wray; Rocky Ford; Roggen; Denver. NEW MEXICO: San Jon

(AMNH). OKLAHOMA: Greer Co.; Chickasha. TEXAS: Dallas.

Ospriocerus pumilus Coquillet

Stenopogon pumilus Coquillett 1904 : 33.

Adisoemarto

25

Scleropogon pumilus Hull 1962 : 126.

Ospriocerus pumilus Martin, pers. comm.

This species is strikingly similar to consapguineus Loewand may be

conspecific. The male genitalia of the two are not different.

This species is known from Texas and Kansas. Strickland (1938)

included it in his list, but this was probably based on misidentified speci-

mens; five specimens were examined.

Localities - K.-XNSAS: Clarke Co.; Ellis Co. (USNM). TEXAS: Brownsville (USNM); Spur

(USXM); Hidalgo Co. (USNM).

Genus Holopogon Loew

HolopopogonLioew 1847:473. Type species: Dasypogon nigripennis Meigen 1820.

Podoctria Megerle (Ms) in Meigen 1820 : 279. N omen nudum .

Ceraturgus Rondani, not Wiedemann 1856 : 156.

The species of this genus are small, 4. 5 - 9. 0 mm, mostly black

with long curly pile. In the United States and Canada, 17 species have

been described (Martin 1959). They are grouped into two subgenera:

three in Dasyholopogon Martin, and the rest in the subgenus Holopogon Loew.

The species of the subgenus Holopogon Loeware very similar to one another;

the male genitalia are non-diagnostic, and most of the remaining char-

acters are relative and variable. Because of this, Martin grouped the

species into four species complexes: seniculus complex, acropennis complex,

phaenotus-oriens complex, and guttula complex. Further, he stated that the

taxonomic status of these complexes is not certain. They may be indeed

more than one species, two or more subspecies, or each may be a single

highly variable species.

Head broad and short (figs. 16, 17); face broad and flat; front

slightly narrower at antennal base, with depression in front of ocellar

plate; the latter elevated, more or less rounded; front with lateral pro-

tuberance; antennae black, first two segments equal, third elongate,

tapering apically, style with two micros egments (fig. 137), the first very

small; palpi two segmented; face, front, and vertex pilose and pollinose.

Thoracic ground color black; pleura white pollinose; presutural meso-

notum, except dorsocentral vittae and posterior inner quarter, white

pollinose, the r est of mesonotum and scutellum black; pile on prothorax,

mesopleuron, anterior and posterior corners of sternopleuron, meta-

pleuron, and mesonotum and scutellum, long, sometimes shorter on

mesonotum; lower slope of metanotum golden yellow tomentose; bristles

weak, hardly distinguishable from pile. Legs black; coxae greyish pol-

linose, with long pile on anterior sides of front, and lateral sides of

middle and hind pairs; femora stout, with pile; hind tibiae club-shaped,

ventral sides of front and hind tibiae golden yellow tomentose; ventral

sides of tarsi golden yellow tomentose, hind basitarsi swollen; claws

curved, empodium short. Wings hyaline, alula small; venation varies

slightly within the species; fifth vein slightly curved anteriorad; bran-

ching of third vein at or slightly beyond the tip of discal cell; marginal,

submarginal, and posterior cells open; anal cell closed, with or without

stalk, in some others open (Hull 1962). Abdomen pilose laterally, more

26

Asilidae of Alberta

or les s shining dor sally; venter pilose; bristles absent, or undetectable;

male genitalia short, reddish, partly rotated (90°); gonopods with arms

and spine-like process (fig. 220); clasper also spine-like; ovipositor

reddish, acanthophorites with four to five pair s of spines . The coloration

of the pile shows sexual dimorphism.

Three species are present in Alberta: albipilosa Curran, seniculus

Loew, and nignpi/osa new species . All three species belong to the subgenus

Holopogon Lioew.

Key to the Species of Holopogon Loew of Alberta

1. Wipg veins yellow seniculus Loew

Wing veins brown 2

2. Pile on mesonotum and scutellum white albipilosa Curran

Pile on posterior mesonotum and scutellum black

• nigripilosa new species

Holopogon albipilosa Curran

Holopogon albipilosusCuTTSin 1923 : 207.

This species shows sexual dimorphism in the coloration of the

pile. The pile on the front and the vertex is black in the males, white in

the females; the mystax is black with few white hairs in the males,

white with one or two black hairs in the females; the antennal hairs are

black in the males, white in the females; the rest of the pile is brownish

in the males and white in the females.

This species ranges from British Columbia to Manitoba, south to

Nevada and Wyoming; holotype, allotype, and 19 additional specimens

were examined.

Localities - ALBERTA; Wainwright (UA); Drumheller (UA) ; Medicine Hat (UA and CNC);

Orion (UA); Lethbridge (CNC); Oldman River, Lethbridge (CNC); Picture Butte (UA).

Other localities - BRITISH COLUMBIA: Vernon (type locality : CNC); Chilcotin (CNC).

SASKATCHEWAN: Saskatoon (CNC); Saskatchewan Landing (CNC). MANITOBA. IDAHO;

Montpelier (AMNH) . WYOMING; Carbon Co. (AMNH); Green River (AMNH); Jackson (AMNH);

near Lander (AMNH). NEVADA.

Holopogon seniculus Loew

Holopogon seniculus LiOew 1862 : 62.

This species is readily distinguished from the others by the yellow

wing veins. The pile is long, white in the males and yellowish in the

females.

This species is known from Alberta and Saskatchewan, south to

Colorado, and west to Nevada; 10 specimens were examined.

Localities - ALBERTA: Scandia (CNC); Medicine Hat (CNC); Lethbridge (CNC).

Other localities - SASKATCHEWAN: Saskatoon (CNC). WYOMING. NEBRASKA; Chandron

(WSU). COLORADO; Lamar (AMNH). NEVADA.

Adi soemarto

27

Holopogon nigripilosa new species

This species is easily distinguished from albipilosa Curran and

seniculus Loew by the color of the mystax, which is black in females, and

the black pile on the posterior mesonotum and on the scutellum. It is

described from three female specimens. Length: 8. 0 mm.

Female. Face, front and vertex, pale golden yellowish pollinose;

pile on vertex, front, and ocellar triangle golden yellow, mixed with

black on frontal protuberance; mystax black, pale golden yellowish pile

present along lateral margins of face; antennae black, with black hairs

on first two segments; occiput black, lower half golden yellowish pol-

linose, bristles and hairs on upper part black; pile on lower half of

occiput, on proboscis and palpi, and beard, white. Prothorax yellowish

white pollinose and pilose, middle pronotum brownish tomentose; dorso-

central vittae brown; humeri, anterior lateral margins of mesonotum,

white pollinose, the rest of mesonotum brownish tomentose; white pile

present on anterior one fourth of mesonotum, anterior lateral margins,

to sutures, the rest of mesonotal pile black; scutellum brownish tomen-

tose, black pilose; mesopleuron brownish pollinose, paler on anterior

half, white pilose; metanotum brownish pollinose. Coxae greyish brown

pollinose, white pilose; femora, and front and middle tibiae white pilose,

hind tibiae with black hairs; bristles on tibiae and tarsi black; claws,

basal half reddish brown, black apically. Wings hyaline, micr otrichiae

brownish, veins brown; venation of average Holopogon Loew (fig. 152),

anal cell closed with stalk. Abdomen shining black with lateral sides of

first two segments yellowish brown pollinose; pile white, longer on

sides of first two segments, shorter on succeeding s egments , very short

and sparse on dorsum; ventral pile long, white; acanthophorites black,

with four black spines.

This species is called nigripilosa , because of the black mesonotal

and scutellar pile, which distinguishes this species from the other two

species from Alberta.

This species was collected from a glade in a conifer ous for est, with

short grass and herbs.

Holotype: female, Opal-Coronado, Alberta, 5. VII. 1963 (L, Kenakin

and S. Adisoemarto); deposited in CNC.

Paratypes: same data; deposited in UA.

Genus Heteropogon Loew

Heferopogon Ljoew 1847 : 488. Type species: Dasypogon manicatus IsAeigenlSZO .

Anisopogon Loew 1874 : 377.

The name Anisopogon Loew was used as a substitute for Heteropogon

Loew, the latter name having been used for a plant (Back 1919). How-

ever, Anisopogon Loew was used by Hull (1962) for the second subgenus of

Heteropogon Loew.

Head wide and flat or short; face and occiput pilose; pile similar

to "plume"; first two antennal segments equal, third segment tapering

apically, one and half times as long as first two segments together (fig.

28

Asilidae of Alberta

150); style two- segmented, the first segment small. Thorax with more

or less rounded mesonotum; anterior mesonotum and mesopleuron pilose;

humerals, presuturals, intraalar s, dor socentrals, postalars, andscutel-

lars present. Legs slender; coxae pilose; bristles present on anterior

sides of femora, several rows on tibiae, subapically on tarsi; basitarsi

long, at least twice as long as second segment; ventral sides of tarsi

setulate. Wings hyaline, partly smokey or diffusedly maculate (fig. 153);

venation normal, all posterior cells open, anal cell open very narrowly

apically or closed, alula present. Basal abdomen as broad as thorax,

tapering apically to one third basal width; male genitalia shiny dor sally,

more or less pointing downward (figs. 222-225).

Coquillett (1893a) and Wilcox (1941) gave synopses of the species

of Heteropogon Loew of North America north of Mexico. A single species,

Heteropogon wilcoxi James, is known from Alberta.

Heteropogon wilcoxi James

Heteropogon wilcoxi James 1934 : 84.

Mystax, frontal and vertical pile, and beard, white; four ocellar s

white; antennae black, one bristle on apical lower side of second anten-

nal segment white; occiput black, white pilose, bristles white; palpi

two- segmented, subequal, first segment excavated laterally (fig. 45).

Thorax greyish yellow pollinose; prothoracic pile long; mesonotal pile

present marginally, dor socentrally and acr ostichally; long pile also

present on anterior and posterior corners of sternopleur on, on meso-

pleuron, on metapleuron, and on upper center of hypopleuron. Coxae

and femora black, tibiae and tar si yellow to or ange br own; coxae yellow-

ish pollinose with long white pile; femora with pile on ventral sides,

bristles on basal ventral and apical posterior sides of front, and anterior

sides and apices of middle and hind pairs; ventral sides of front and

hind tibiae, and tarsi, golden yellow tomentose; claws strong, curved,

black; empodium short, brown. Wings slightly longer than abdomen,

veins brown; anal cell open narrowly; branching of third vein above tip

of discal cell; anterior crossvein behind the middle of discal cell (fig.

153).

Abdomen yellowish grey pollinose; white pile present on lateral

margins, shorter on posterior segments; dorsum covered with short,

sparse pile; ventre white pilose; last three segments of female shining

black, acanthophorites black, bearing five pairs of black spines; male

genitalia shining brownish orange (figs. 222-225).

This species ranges from Alberta to Arizona; seven specimens

were examined.

Localities - ALBERTA: Lethbridge (UA and CNC).

Other localities - WYOMING. COLORADO: Model, Hochne; Mesa de Maya, Tobe; Sprin-

ger. ARIZONA: Holbrook. ILLINOIS: Joliet.

Adisoemarto

29

Genus Lestomyia Willi ston

Lestomyia Williston 1884 : 19. Type species: Clavator sabulonum Osten-

Sacken 1877.

Clavator Osten-Sacken not Philippi 1877 : 391.

In appearance, these flies resemble Lasiopogon Loew, but are dis-

tinct in the antennae and some other characters, such as the vertex, the

front, the gibbosity, and the presence of a bent tibial spur on front tibiae.

Male genitalia are rotated about 90°; hypandrium subtriangular ; aedeagus

long; superior forceps more or less like those of Heteropogon Loew (fig. 58).

Face broad, gibbosity not too prominent; front and vertex convex

marginally, ocellar plate elevated, rounded; first two antennal s egments

subequal, third swollen apically, style single segmented, truncate, hollow

on tip (fig. 128); palpi two- segmented. Thorax with strong bristles,

markedly pollinose, less pilose. Legs slender; pile short, appressed;

bristles stout, mostly on tibiae and tarsi; claws long; empodium two

thirds as long as claws, sharp. Wings hyaline, all posterior cells open,

anal cell open narrowly; branching of third vein above or beyond tip of

discal cell; anterior crossvein slightly beyond middle of discal cell;

alula well developed (fig. 154). Abdomen elongate, pile short and semi-

appressed, longer on first segment; bristles present on sides of first

segment. Seven species are included in this genus, all Nearctic in dis-

tribution. In Alberta this genus is represented by one species, Lestomyia

sabulonum Osten-Sacken.

Lestomyia sabulonum Osten-Sacken

C/avator sa6u/onum Osten-Sacken 1877 : 392.

Lestomyia sabulonum Williston 1884 : 20.

This species is yellowish grey pollinose; all bristles are white.

Size 7.0- 11. 0 mm in males, 8. 0 - 12. 0 mm in females. There is no

sexual dimorphism. The number of bristles varies individually; the

ocellar bristles three to four pairs; metapleural bristles in a row of

four to six; humerals three to four; post-humerals none to two; intra-

alars two to three; dor socentrals eight to ten; scutellars three to four

pairs .

This species lives in mainly dry fields, with short grass and cacti,

near to, or far from, water.

This species is known from British Columbia and Alberta, south

to California, east to Wyoming; 35 specimens were examined.

Localities - ALBERTA; Burdett (UA); Medicine Hat (UA); Comrey, Milk River Valley

(UA); Writing-on-Stone Provincial Park (UA); Little Bow Park, McGregor Lake (UA).

Other localities - BRITISH COLUMBIA: Oliver (CNC). CALIFORNIA: Claremont (CNC).

WYOMING: Rawlins (AMNH) .

Genus Nicocles Jaennicke

A^jcoc/es Jaennicke 1867:355. Type species: Nicocles analis Jaennicke 1867 .

Pygostolus Loew 1866 : not Haliday 1833. Type species: Dasypogon politus Say

1823.

30

Asilidae of Alberta

This group includes flies with rather flat abdomens. The head is

similar to Heteropogon Loew and Lestomyia Willi ston, but the shapes of the

antennae (fig. 129) and the mystax (figs. 20-22) readily distinguish the

two groups. The humeral bristles are absent from Nicocles Jaennicke.

Face flat, bristles present along epistomal margin; first two an-

tennal segments subequal, third segment tapering apically, bristles

present on lower side of second segment; vertex and front broad, semi-

parallel (fig. 22); proboscis short; palpi two- segmented, subequal (fig.

44). Thorax with bristles on posterior half of mesonotum; humerals

absent; presuturals present; metapleur on with bristle-like hair s . Legs

slender; bristles present on middle femora, on tibiae, and on tarsi;

front and hind basitarsi twice as long as second tarsal segments (figs.

7 5, 76). Wings longer than abdomen, maculate in some species; discal

cell elongate; third vein branch above or beyond tip of discal cell; an-

terior crossvein at apical two-thirds of discal cell; all posterior cells

open except anal cell narrowly open at tip or closed; alula not well deve-

loped. Abdomen shiny and rather flat (figs. 94-96). In males: seventh

segment concealed under broader sixth segment; male genitalia small,

notrotated, concealed under sixth abdominal s egment. Infemales: eighth

segment concealed inside seventh segment; acanthophorites with five

pairs of spines.

This genus is represented in the Neotropical R egion by one species,

and in the Nearctic Region by 14 species. One species, Nicocles utahensis

Banks, occurs in Alberta.

Nicocles utahensis Banks

Nicocles utahensis 'Ba.nks 1920 : 66-67.

Nicocles punctipennis 'bAelandev 1923c : 217-219.

This species is easily recognized by the shiny black abdomen and

incomplete silvery white marking on the fifth segment of males and fe-

males. The silvery markings differ between the sexes. In the males,

the marking on the fifth abdominal segment is incomplete, interrupted

medially, broader laterally, and on the sixth segment, the marking is

entire (fig. 94). In the females, the markings are present on the last

three segments, broad on the lateral margins, tapering, and separated

by a small gap medially (fig. 96).

This species ranges from British Columbia and Alberta, south to

Oregon and Utah.

Localities - ALBERTA: Medicine Hat (CNC).

Other localities - BRITISH COLUMBIA: Robson (CNC).

Genus Cyrtopogon Loew

Cyrtopogon Lioew 1847 : 516. Type species: Asilus ruficornis F. 1794.

Euarmosf us Walker 1851 : 102. Type species: Euarmostus bimacula Walker

1851.

Eupa/amus Jaennicke 1867 : 86. Type species; Eupalamus alpestris Jaennicke

Adisoemarto

31

1867. Preoccupied in Hymenoptera, Wesmael 1844, andinCole-

optera, Schmidt- Goebel 1846.

Palamopogon Bezzi 1927 : 61. Type species: Palamopogon alpestris Jaennicke

1867.

Philammosius Rondani 1856 : 156. Type species: Dasypogon fimbriatus Meigen

1820.

Wilcox and Martin (1936) included 68 species in this genus. The

species were arranged in 21 groups and five "single" species: falto

Walker, laphriformis Curran, lyratus Osten-Sacken, alleni Back, and tenuis

Bromley.

This genus seems to be the most successful group in North America

north of Mexico; so far it has not been reported from Mexico (Wilcox

and Martin 1936). Twenty three species have been reported from the

Palaearctic, two from the Ethiopian, and three from the Oriental Region

(Hull 1962). There are 14 species known from Alberta.

There are many characters for the identification of the species,

depending on the group, such as shape, color, and ornamentation of the

tarsi; ornamentation of abdomen; markings on the wings; shape of the

mystax; the scutellum; the metapleura; the legs; the claws; the gib-

bosity and width of the face; and the antennae (Wilcox and Martin 1936).

Back (1909) noticed that some species were aberrant forms of the

genus; these were placed in different genera; Eucyrtopogon by Curran

(1923), Metapogon by Coquillett ( 1904) ; Nannocyrtopogon byWilcoxand Martin

(1936).

Most species live inareas near or within coniferous forests. Other

known habitats are: sand near willows along running water, and open

desert. Limited data on the phenology and mating behaviour of some of

the species were presented by Wilcox and Martin (1936). Melin (1923)

provided information on the biology of the Palearctic species, Cyrtopogon

lateralis Fallen.

See Wilcox and Martin (1936) for the description of the genus.

Key to the Species of Cyrtopogon Loew of Alberta

1. Last segment of front tarsus elongate, as long as three proceeding

segments together, flattened (fig. 81); fir st abdominal segment

with a posterior pollinose fascia . Hneotarsus Curran

Fore tarsus with subequal segments; first abdominal segment with-

out a posterior pollinose fascia 2

2. Hind tibiae entirely black 3

Hind tibiae entirely or partly reddish or orange brown 6

3. Mystax entirely black; tibial pile short 4

Mystax with white or yellow pile; tibial pile long, black or mixed

with white 5

4. Tibial pile in both sexes black; hairs of male genitalia black. . ..

nigator Osten-Sacken

Tibial pile white; hairs of male genitalia white .... sansoni Curran

5. Silvery hairs on segments 1 to 5 of male front tar sus not noticeably

longer apically; first two abdominal segments with yellow hairs;

32

Asilidae of Alberta

hind femora yellowish haired praepes Williston

Silvery hairs on segments 2 to 5 of male front tarsus longer api-

cally; more than two basal abdominal segments with pale yellow

hairs; hind femora with black hairs . . wilUstoni Curran

6. Abdomen with dense, erect, light colored pile, covering at least

the dorsum of abdominal segments 2 and 3. . . . c/asy//is Williston

Pile of abdomen not as above 7

7. Hind tibia with long white pile montanus Loew

Hind tibia without such long pile 8

8. Metapleural bristles entirely black 9

Metapleural bristles mixed with orange, or entirely orange or

pale yellow 10

9. Third antennal segment orange; tarsal segments mostly black. .

aurifex Osten-Sacken

Third antennal segment black; last tarsal segment black, the re-

maining reddish brown 6imacu/a Walker

10. Scutellum silvery pollinose; hind tibia black; metapleural bristles

entirely orange or pale yellow nugator Osten-Sacken

Scutellum not or hardly pollinose; hind tibia partly or entirely

orange, reddish, or yellow; metapleural bristles mixed orange

and black 11

11. Antennae entirely black 12

Third antennal segment orange 14

12. Basal one third of hind tibia black, the remaining orange or red-

dish brown 13

At least basal half of hind tibia orange, tibial apex black 14

13. Anterior tibia black; tibial pile long inversus Curran

Anterior tibia orange brown; tibial pile practically absent

albovarians Curran

14. Abdominal bands interrupted medially . . . .distinctitarsus new species

Abdominal bands complete, orange 15

15. Male 16

F emale 17

16. Front tibia and tarsus with white fringe of hairs, as long as dia-

meter of segments; pile on face orange; 2-4 abdominal seg-

ments with orange pile across the segments auratusCole

Front tibia and tar sus without fringe of hairs; bristles or hairs on

face black mixed with white ox yellow; pile of abdomen only on

lateral sides, absent from fourth segment . . . glarealis Melander

17. Pile on third abdominal segment as long as that on second

auratus Cole

Pile on third abdominal segment shorter than that on second ....

glarealis Melander

Cyrtopogon auratus Cole

Cyrtopogon auratus Cole 1919 • 230.

Cyrtopogon albitar sis Curran 1922 : 278-279.

Cyrtopogon albitarsis Curran 1924 : 279.

Adisoemarto

33

This species belongs to the aurifex group, in which the male ab-

dominal segments 2-4 are provided with dense fulvous pile across the

segments .

Males. Face withmane-like orange pile; front tibia and front tar sus

with fringe of white hairs on outer sides (fig. 78); fifth to seventh ab-

dominal segments short, tectiform; male genitalia black, form as in

figs. 226-229.

Females. Mane on gibbosity spar ser; fringe of white hairs absent

from front tibia and front tarsus; abdominal segments not tectiform,

yellow pile on first three segments shorter than that of male, still shorter

on fourth, and almost absent from fifth.

A female specimen from Yellowstone Park, Cascades Y.R., 22.

Vii. 1923 (A. L. Melander), was chosen as allotype of albitarsis Curran,

1922, but determined by G. Stuart Walley, 1932, as not the allotype.

This species rangds from Alberta to Oregon, southeast to Colorado;

holotype and 14 additional specimens were examined.

Localities - ALBER TA: Banff (type locality; CNC); Banff, Lake Minnewanka, Davil's Gap

Trail (UA); Waterton (AMNH); Gorge Creek (UA).

Other localities - WASHINGTON': Mt, Spokane. OREGON: Strawberry Mtn. , Grant Co.

(CNC): Wallowa Lake. IDAHO: Long Valley, Alpha (UA). WYOMING: Yellowstone National

Park, Medison Junction (AMNH); Yellowstone N. P. , Cascades Y.R. (CNC); Sylvan Pass,

Yellowstone Park. COLORADO: Malta (AMNH). UTAH: Uintah Mts.

Cyrtopogon aurifex Osten-Sacken

Cyrtopogon aurifex Osten-Sacken 1877 : 301-302.

This species is similar to auratus Cole, but the two are distinguished

by the color of the metapleural bristles: entirely black in aurifex Osten-

Sacken, mixed with orange in auratus Cole.

This species ranges from Alberta and British Columbia, south to

California; two specimens were examined.

Localities - ALBERTA: Seebe, Kananaskis Forest (DE).

Other localities - BRITISH COLUMBIA: Vancouver Island. WASHINGTON: Mt. Adams,

Clearwater; Mt. Adams, Signal Peak. OREGON: Mary's Peak; Crater Lake. CALIFORNIA:

Weber Lake, Sierra Nevada; Gold Lake, Sierra Co.

Cyrtopogon willistoni Curran

Cyrtopogon willistoni Curran 1922 : 277-278.

This species belongs to the calHpedilus group, in which the last two

segments of the middle tarsus of the males are provided with a disc of

black hairs (fig. 80). The males of this group are more or less easily

separated from one another by the shape of the silvery hairs on the front

tarsi, but the females are hardly distinguishable.

This species ranges from Alberta and British Columbia, south to

California and Colorado; 32 specimens were examined.

Localities - ALBERTA: Banff (UA and CNC); Calgary (UA and DE); Mountain View (CNC);

Twin Butte (CNC); Waterton Lakes Park (CNC and UA).

Other localities - BRITISH COLUMBIA: Chilcotin (AMNH); Aspen Grove (AMNH) . Minnie

Lake; Nicola, Oliver. WASHINGTON: Signal Peak (AMNH); Blue Mts., Godman Springs ; Col-

ville; Mt. Adams; Mt. Spokane; Tampico; Yakima. OREGON: Fish Lake; Steins Mts.,

34

Asilidae of Alberta

Harnery Co. ; Ontario; Strawberry Mt. , Grant Co. IDAHO: Dong Valley, Alpha. MONTANA:

GallatinCo.; Madison Co. ; Bozeman. WYOMING: Mammoth Hot Springs , Y ellowstone National

Park (AMNH); Grand Teton NatT Park; COLORADO: Elbert (AMNH); Electra Lake (AMNH);

Ouray (AMNH); South Fork (AMNH). UTAH; Roosevelt Creek, R aft R iver M ts . ; Zion NatT

Park. CALIFORNIA; Coleville, Mono Co. (AMNH); Sacramento.

Cyrtopogon praepes Williston

Cyrtopogon praepes Williston 1884 : 12,

This species is similar to wiUistoni Curran. The males are dis-

tinguished by the presence of the silvery hairs on the first tarsal segment

of the front tarsus, and the females are distinguished by the yellowish

hairs on the hind femora.

Strickland (1938) included this species in his list, but I do not be-

lieve it occurs in Alberta. This species ranges from British Columbia

to California and Nevada; four specimens were examined.

Localities - BRITISH COLUMBIA: Vaseaux (CNC); Penticton (CNC); Robson (USNM).

WASHINGTON: Olympia; Roy. OREGON. IDAHO. NEVADA: Elko (USNM). CALIFORNIA:

San Francisco; Santa Cruz; Santa Rosa.

Cyrtopogon bimacula Walker

Euarmostus bimacula Walker 1851 : 102.

Cyrtopogon melanopleurus Loew 1866 I 61.

Cyrtopogon bimacula Loew 1874 : 365.

This species is easily recognized by the wings of the males: macu-

lated at the apex and the tip of the anal cell (fig. 155); in the females,

there is a tendency to light infuscation on the wings of the same pattern

as in the males; both sexes have largely yellowish white pile and black

metapleural bristles.

This species is transcontinental in the North, ranging from the

Northwest Territories to New Mexico; 71 specimens were examined.

Localities - ALBERTA: High Level (UA); Flatbush, Pembina River (UA); Lac la Biche,

Owl River (UA); Chipewyan (CNC); Opal (UA); Sandy Lake (UA); Beaverlodge (UA); Nordegg;

Columbia Icefield (UA); Calgary (UA); Wilkinson Creek, Bow River Forest (UA); Lethbridge

(UA); Morrin (CNC); Medicine Hat (CNC); Elkwater (CNC); Cypress Hills (UA).

Other localities - NORTHWEST TERRITORIES: McKenzie Delta, Reindeer Depot (CNC).

BRITISH COLUMBIA: Steelhead; Lorna. WASHINGTON: Mt. Rainier, Sunrise, Paradise;

Mt. Baker. OREGON: Aneroud Lake Buie Mts.; Horst Mts., Lane Co.; Frog Meadows, Lane

Co. IDAHO. MONTANA: Skalkadho Pass, Ravalli Co. WYOMING: Yellowstone NatT Park.

COLORADO: Camp Creek R. Station; Aspen; South Peak; Ward. NEW MEXICO: Las Vegas

Mts. SASKATCHEWAN: Dandrum (CNC); Saskatoon (CNC); St. Victor (CNC). MANITOBA:

Douglas (CNC) . ONTARIO: Sand Lake (CNC); Sadbury. QUEBEC; Megantis (CNC); Seven Isles

(CNC). NOVA SCOTIA: Truo. NEW HAMPSHIRE; Breton Woods; Mt. Washington; White

Mountains .

Cyrtopogon distinctitarsus new species

This species resembles bimacula Walker to some extent, but is

distinguished by the color pattern of the legs and the color of the meta-

Adisoemarto

35

pleural bristles.

Female. Face, front, and vertex, golden yellow pollinose; mystax

black, mixed with golden yellow pile on center of gibbosity; hairs on

front, vertex, occiput, first two antennal segments, and second palpal

segment, black; beard, pile on first palpal segment and on lower side

of proboscis, white; antennae black; gibbosity prominent near antennal

base (fig. 18). Thorax golden yellow pollinose; pleura without shiny

bare area; pollinose color pattern similar to bimacula Walker; pile on

propleur on white, onpronotum, metanotum, and scutellum, black; meta-

pleural haris orange yellow. Legs bicolored; basal halves of tibiae,

basal three-fourths of basitarsi, basal halves of tarsal segments 2-3,

orange brown; the remainder of legs black; basal half of claws orange,

apex black; empodium very short, orange; pulvilli broad; pile on coxae

yellowish, on lower sides of femora white, on upper sides black, short,

appressed, longer on apices of hind pair, on tibiae, black, short, sparse;

bristles of tibiae and tarsi black. Wings hyaline, with micr otrichiae,

brownish maculate on the following: anterior crossvein, base of discal

cell, anterior branch of cubitus and mediocubital crossvein, apex of

discal cell and branching of third vein (fig. 156); anterior crossvein at

basal one fourth of discal cell; halteres orange. Abdomen black, more

or less similar to bimacula Walker; pile yellowish; spines reddish brown.

This species has beennamed distinctitarsus , becausethe color pattern

of the tarsi is quite distinct from the remaining species of Cyrtopogdn from

Alberta.

Holotype: Female, Opal, Alberta, 5. VII. 1963 (L. Kenakin and

S. Adisoemarto); deposited in CNC .

Paratypes: Female, Lac la Biche, sand dunes, N. E. shore, Al-

berta, 2-4. VII. 1964 (L. M. Kenakin and S. Adisoemarto); female ,

Lethbridge, Alberta, 24. VI. I960 (D. Larson); deposited in UA.

Cyrtopogon montanus Loew

Cyrtopogon montanus Loew 1874 : 362.

This species is easily recognized by the color of the mystax, and

the long pile on the abdominal segments and the legs, mostly black in

the males and white in the females. Upper middle part of mystax white,

the remaining black; frontal, vertical, and upper occipital pile black;

beard white; antennae black, third segment slightly orange, pile on first

two segments white. Propleural pile white, pile on the remainder of

thorax black. Legs mostly black, hind tibiae and hind tarsi reddish

brown; pile on coxae, lower basal femora, and dorsal sides of hind

tibiae, white, the remaining pile of legs black. Abdominal pile of male

bicolored, on posterior corners of each segment white, the remaining

black; male genitalia (figs. 230-233) black, with black hairs; abdominal

pile of females entirely white.

This species is found in central western North America, from

British Columbia, south to California, and eastto New Mexico; holotype

(CNC) and eight additional specimens were examined.

36

Asilidae of Alberta

Localities - ALBERTA: Banff (type locality; CNC),

Other localities - BRITISH COLUMBIA: Seton Lake (UA); Vernon; Departure Bay; Gold

Stream; Lillooet; Oliver. WASHINGTON: Cle Elum; Mt. Adams, Signal Peak, West Klickitat;

Mt, Rainier, Sunrise, White River; Olympia; White Rock Spring, Steven Pass, Cascade Moun-

tains. OREGON: Anthony Lake; Canby; Fox; Hood River; Marys Peak; McKenzie Pass; Mt.

Hood; La Grande; North Powder, IDAHO: Lake Waha; Long Valley, Alpha; Mosco Mt. ;

Potlach. UTAH: Ogden. CALIFORNIA: Towle (AMNH); Emigrant Gap (AMNH); Gold Lake

(AMNH); Sierra Nevada; Fallen Leaf Lake; Lake Tahoe; San Bernardino Mts. ; Mt. St. Alens;

Sequoia Nat'l Park, Welverton; Truckee; Yosemite Valley. COLORADO: Boulder; Gold Hill;

Longs Peak Inn; Ward; Poncha Pass. NEW MEXICO.

Cyrtopogon albovarians Curran

Cyrtopogon albitarsis Curran 1923 : 134-135, not Curran 1922 : 278-

279.

Cyrtopogon albovarians Curran 1924 : 279-280.

The first specimen was first described as the allotype of albitarsis

Curran 1922 (= amatus Cole), but then Curran (1924) realized that the

specimen belongs to a different species, and described it as albovarians .

It differs from amatus Cole (= a/6harsis Curran) in the following char-

acters: antennae entirely black; upper one third of mystax white, the

remainder black; pile on posterior mesonotum white; pile on meso-

pleuron longer, white on lower side, black on upper side; metapleural

pile white; metanotal pile black; pile on sides of third and fourth ab-

dominal segments mixed with black.

This species is known only from Alberta; holotype (CNC) and one

additonal specimen were examined.

Localities - ALBERTA; Banff (type locality; CNC); Wabamun (UA),

Cyrtopogon inversus Curran

Cyrtopogon inversus Curran 1923 : 172-173.

This species is similar to albovarians Curran, but can be distinguished

by the black anterior tibiae and the long tibial pile.

This species ranges from Alberta and British Columbia, south to

Oregon, and east to Colorado; two specimens were examined.

Localities - ALBERTA: Seebe, Kananaskis Forest (DE).

Other localities - BRITISH COLUMBIA: Aspen Grove; Darcy; Nicola; Chilcotin; Hadley;

Kamloops; Lillooet, Seton Lake. WASHINGTON: Signal Peak, R anger Station; Virden. OREGON:

Eagle Ridge, Klamath Lake. WYOMING: Yellowstone Nat'l Park. COLORADO: Longs Peak.

Cyrtopogon glarealis Melander

Cyrtopogon glarealis Melander 1923a : 113-114.

This species belongs to the pulcher group, but is distinguished from

pulcher Back by the color of the pile on the second abdominal segment of

the males (black in glarealis Melander, orange in pulcher Back), and by the

color of the metapleural hair s in the females (largely black in the former,

orange in the latter). Males of this group are easily recognized by the

form of the abdominal segments, which are gradually compressed later-

Adisoemarto

37

ally toward the posterior end, and by the orange third antennal segment.

This species ranges from Alberta and British Columbia, south to

California and Wyoming; three specimens were examined.

Localities - ALHLR I A: Kootenay Plains, 116'Z5'W 52'^7'N (LMK).

Other localities - BRITISH COLUMBIA: Salmon Lake, Nicola District. WASHINGTON:

Spokane; Wolf Fork, Touchet River, Blue Mts. OREGON: Wallowa Lake, Aneroid Lake Trail.

IDAHO: Gold Hill, Latan flo. ; Moscow Mts. ; Long Valley, Alpha (UA). MONTANA: Big Hole

Battle Field, I'^eavcrhead Co.; Gallatin Co. WYOMING: Madison Junction, Yellowstone Nat'l

Park; Dunroven Pass. CALIFORNIA: Angora Peak, I'ahoe.

Cyrtopogon lineotarsus Curran

Cyrtopogon lineotarsus Curran 1923 : 187-188.

This species is a member of the leptotarsus group, in which the last

tarsal segment of the front tarsus is elongate (fig. 81). In the males,

the gibbosity is almost triangular from the anterior aspect; it reaches

the antennal base and is very prominent. A female specimen from Glacier

Park, Montana, 5. VIII. 1925 (G.A. Mail) was doubtfully identified by

Wilcox (1935) as Cyrtopogon lineotarsus Curran, and also described as the

female of lineotarsus Curran by Wilcox and Martin (1936), although they

were doubtful, because this specimen differed in the color of the mystax

from the type specimen; the mystax is entirely white in the female, and

some other differences are also obvious: the thoracic, abdominal, coxal,

and femoral pile, is entirely white. According to Wilcox and Martin

(1936), lineotarsus Curran could be the same species as predator Curran,

based on a comparison of the specimens of both sexes with the types of

both species.

This species ranges from Alberta to Montana; holotype (CNC) and

two additional specimens were examined.

Loralitifs - Ai. BERTA: Banff (type locality; CNC); Kananaskis Valley, Pocaterra Creek

(CA.S).

Other locality - MONTANA: Glacier Park.

Cyrtopogon sansoni Curran

Cyrtopogon sansoni Curran 1923 : 138-139.

This species belongs to the nugator group, in which the scutellum

is flat, largely pollinose on the center, shining on the edge. The hypo-

pleural hairs are entirely white, and the abdominal bands are interrupted

medially. Face broader than long; gibbosity golden brownish pollinose,

more or less rounded; front, vertex, and occiput greyish pollinose,

black pilose; mystax black; beard white. Meson^otal pile and scutellar

hairs brownish black; mesonotal bristles black.

This species is known from Alberta only; holotype (CNC) and allo-

type (CNC) were examined.

LocalitU's - ALBLR'l'A: Banff (type locality; CNC).

38

Asilidae of Alberta

Cyrtopogon nugator Osten-Sacken

Cyrtopogon nugator Osten-Sacken 1887 : 307.

This species is strikingly similar to sansoni Curran and hardly

distinguished from it. See Wilcox and Martin (1936) for the diagnostic

characters.

This species ranges from British Columbia, south to Arizona and

New Mexico. Strickland (1938) included this species in his list, but I do

not believe it occurs in Alberta; six specimens were examined.

Locahtu's - F’)RrilSi[ COLUMIilA: Saanich; Vernon; Agassiz. WASHINGTON; Signal

Peak; Rainier Nat'l loresl; Sumner (UA). OREGON; Mt. Hood. IDAHO; Lake Wala. CALI-

FORNIA: Weber Lake, Sierra CIo. (CNC); Grass Lake, Tahoe; Tioga Road; Yosemite.

COLORADO: Aspen (AMNH). NEW MPiXICQ: Cloudcroft. ARIZONA: Santa Catalina Mts.

(AMNH and ESNM).

Cyrtopogon dasyllis Williston

Cyrtopogon dasyllis Williston 1893 : 66.

Males have maculate wings, but the pattern is different from that

of bimacula Walker: specimens of dasyllis Williston have one black macula

on the apex of the wing, and a narrow one around the second cubital vein

(fig, 157); the abdomen is provided with yellowish long pile on the entire

first four abdominal segments, black on the remaining; hypandrium of

the male is provided with a pair of spine-like structures (fig. 236). The

wings of the females are not distinctly maculate, but are slightly infus-

cated in the place of the maculae.

This species ranges from Alaska and the Northwest Territories,

south to Oregon, and Colorado; 16 specimens were examined.

Localities - A LIH:IR TA : Banff (UA, CNC, andAMNH); Lake Louise (CNC); Jasper (CNC).

Other localities - AIASKA; Skagway (AMNH). NORTHWEST TERRITORIES: Cameron

Bay, Great Bear Lake (CNC). YUKON TER R ITOR lES: Whitchor s e (CNC ) . BR I TISH COLUMBIA:

Davie Lake (CNC); Robson (CNC); Shaswap (CNC); Juktakamin (CNC); Kaslo. WASHINGTON:

Mt. Rainier, Sunrise; Mt. Rainier, Paradise Inn; Mt. Rainier, White River Camp; Randle.

OREGON; Strawberry Mt., Grant Co. IDAHO. COLORADO; Deer Mt.

Genus Eucyrtopogon Curran

Eucyrtopogon Curran 1923 : 95. Type species Cyrtopogon nebulo Osten-Sacken

1877.

This genus is confined to the Nearctic Region, and includes 11

species (Hull 1962). In Alberta, Eucyrtopogon is represented by seven

species, one of them, incompletus , is new.

Face, front, and vertex, broad, narrower at antennal base; front

less slanting than Cyrtopogon Loew; third antennal segment elongate,

tapering apically (fig. 131); antennal style half as long as third segment;

fir st two antennal segments oval, subequal; proboscis short, stout (fig.

61); palpi two-segmented, long (fig. 43). Thorax slightly elongate;

mesonotum pilose; dorsocentral bristles weak; metapleural hairs always

present. Legs slender; femora and tibiae subequal, the former thicker;

bristles short, strong on tibiae and tarsi; pile present on tibiae; claws

Adisoemarto

39

strong, long. Wings hyaline, maculate in certain areas (figs. 158, 159);

one and a half times as long as abdomen; microtrichiae usually present,

but absent from albibarbis Curran. Abdomen semiparallel, curved dor-

sally, about one and a hcdf times as long as thorax; posterior corners of

segment 1 to 6 always pilose, seventh segment in some species bare;

pile longer on sides, short and appressed on dorsum; male genitalia

black, concealed, almost constant in form within the genus (figs. 238-

242); acanthophorites with four to six pairs of spines . Sexual dimorphism

very slight, present in some species on the pattern of pile and hairs,

pattern of costal setulae, and pollen of the abdomen.

1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

12.

Key to the Species of Eucyrtopogon Curran of Alberta

Male 2

F emale 7

Wing clear, microtrichiae absent a/6i6ar6is Curran

Wing with microtrichiae 3

Middle line of thorax with very conspicuous mane-like white and

black hairs * comantis Curran

No condensation of pile to form a mane 4

Wings with a double row of setulae, black, curved 5

Costal setulae pale brownish or orange, not curved 6

From side aspect mystax with ends of hairs pale yellowish or white

diversipilosus Curran

Mystax with ends of hairs brownish nebulo Osten-Sacken

Costal setulae end between tip of subcosta and first longitudinal

vein incompletus n. sp.

Costal setulae complete, reaching wing tip

calcarata Curran or spinigera Curran

Wings without microtrichiae albibarbis Cur r dm

Microtrichiae present 8

Seventh abdominal segment without sericeous pollen

ne6u/o Osten-Sacken

Seventh abdominal segment with sericeous pollen 9

Mesonotum with acrostichal white mane anteriorly 10

Mesonotum without such mane 11

Front tibia and all tarsi with prominent white mane-like pile ....

comantis Curran

Front tibia and tarsi with less conspicuous mane-like pile

incompletus new species

Sericeous pollen on sixth abdominal segment extending broadly

almost to base of segment calcarata Curran

Pollen not extending over two-thirds the distance to base, or only

very narrowly so on sides 12

Acanth.ophorite with four pairs of spines diversipilosus Curran

Acanthophorite with five or six pairs of spines . . . .spinigera Curran

40

Asilidae of Alberta

Eucyrtopogon comantis Curran

Eucyrtopogon comantis Cur T3in 1923 : 116--117.

This species is recognized by the presence of the mane-like white

pile on the acrostichal area of the mesonotum; this vestiture is also

present in albibarbis Curran, but these two species are easily distinguished

by the presence or absence of micr otrichiae of the wings. Sides of face

with white mane; each bristle of mystax bicolored, brownish black bas-

ally, yellowish white apically; ocellar and antennal bristles brownish

yellow. Legs black; coxal pile yellowish; femoral pile long, white,

appressed on dorsal sides; pile on front and middle tibiae white, short,

appressed, mane-like, half aslongas tibia diameter , continued to dorsal

sides of tarsi, less conspicuous on hind pair; pile in females longer;

claws black. Posterior corners of fourth, fifth, and sixth abdominal

segments with bristle-like brownish hairs, absent from females; venter

white pilose; male genitalia black; acanthophorite with five pairs of

spines .

This species is known from British Columbia and Alberta, south

to Colorado; holotype (CNC) and nine additional specimens were exam-

ined.

Localities - ALBERTA: Fabyan, Campsite (UA); Medicine Hat (UA and AMNH); Calgary

(CNC); Magrath (CNC).

Other localities - BRITISH COLUMBIA: Chilcotin (type locality; CNC); Vernon (CNC);

Departure Bay (CNC). WYOMING. COLORADO: Maez Creek, Huerfano Co. (USNM); Wet

Mts., Huerfano Co. (USNM).

Eucyrtopogon albibarbis Curran

Eucyrtopogon albibarbis Curran 1923 : 117.

This species is similar to comantis Curran, but is readily distin-

guished by the absence of villi or micr otrichiae from the wings. Upper

two thirds of face white pilose; each bristle of mystax bicolored, black

basally, white apically. Mesonotum is provided with acrostichal mane.

Dorsal faces of front tibia and of all tar si with appressed mane-like white

pile, of equal size in both sexes.

This species is known from Alberta and Saskatchewan; holotype

(CNC) and 16 additional specimens were examined.

Localities - ALBERTA: Fabyan, Campsite (UA); Medicine Hat (CNC); Calgary (UA).

Other localities - SASKATCHEWAN: Saskatoon (CNC); Moose Jaw (type locality; CNC);

Regina (USNM).

Eucyrtopogon incompletus new species

This species resembles comantis Curran and albibarbis Curran, but

differs in the following respects: the male has curved costal setulae on

the wing, which end a considerable distance before the wing tip; the

wing tip; the front tibia of the females is without obvious white mane.

This species is distinguished from comantis by the following char-

acters: mesonotal pile less abundant, acrostichal mane less obvious;

costal setulae not complete, ending between subcosta and first longi-

Adi soemarto

41

tudinal vein; costa from this point to apex bare; front tibia of female

without obvious white mane. From albibarbis Curran, it is distinguished

by the following characters; micr otrichiae present on wings of both sexes;

costal setulae present.

Male. Face white pollinose; mystax black, mixed with white pile

on upper and lateral margins; front, vertex, and occiput brownish pol-

linose; hairs on front, vertex, and occiput brownish; six ocellar bris-

tles black half basally, white apically; first two antennal segments sub-

equal, black, white pilose; third antennal segment missing; beard and

pile on proboscis and palpi, white. Thorax more or less similar to

comanUs Curran, but acrostichal pile shorter. Legs similar to comantis

Curran. Wings with micr otrichiae, costal setulae end between subcosta

and first longitudinal vein; shape and color of costal setulae similar to

those of comantis Curran.. Abdomen similar to comantis Curran.

Female. Pile on front tibia normal, not produced into long, mane-

like pile as in comantis Curran; the remainder similar to comantis Curran.

The name incompletus has been chosen for this species on the basis

of the shape of the incomplete costal setulae.

Holotype; Male, Cypress Hills, Alberta, 26. V. 1964 (S.

Adisoemarto) ; deposited in CNC.

Paratypes: Two females, same locality, 24. V. 1964 (S.

Adisoemarto); deposited in UA.

Eucyrtopogon calcarata Curran

Eucyrtopogon calcarata Curran 1923 : 119.

The males of this species are readily recognized by the costal

setulae, which are longer than the diameter of the costa, and by the

presence of a conical tubercle on the anterior apex of the hind coxa. The

sixth abdominal segment of the females is provided with broad s ericeous

pollen on the sides.

This species is known from British Columbia and Alberta; holo-

type (CNC) and 12 additional specimens were examined.

Localities - ALBERTA: Waterton Lakes (CNC); Cowley (UA); Coleman (CNC); Banff

(type locality; CNC); Jasper (UA); Edmonton (UA).

Other localities - BRITISH COLUMBIA: Robson (CNC); Cranbrook (CNC); Princeton

(CNC).

Eucyrtopogon spinigera Curran

Eucyrtopogon spinigera Curran 1923 : 117-118.

The males are very similar to calcarata Curran, but the females

are distinguished by the size of the pollinose marking on the sixth ab-

dominal segment, being less than two thirds of the length of the seg-

ment.

This species is known from the Northwest Territories to Alberta

and British Columbia; holotype (CNC) and 10 additional specimens were

examined.

Localities - ALBERTA: Calgary (CNC); Cowley (CNC); Medicine Hat (UA).

42

Asilidae of Alberta

Other localities - BRITISH COLUMBIA; Victoria (type locality; CNC); Pass Creek (CNC) ;

Copper Mtn. (CNC).

Eucyrtopogon diversi pilosis Curran

Eucyrtopogon diversipilosis CuTT3.n 1923 : 118.

This species is similar to spinigera Curran. The males are dis-

tinguished by differences in costal setulae: diversipilosis Curran, they

areblackand curved; in spinigera Curran, they are orange and not curved.

This species is known from British Columbia and Alberta; holo-

type (CNC) and four additional specimens were examined.

Localities - ALBERTA: Banff (CNC); Coleman (CNC).

Other localities - BRITISH COLUMBIA: Chilcotin (type locality; CNC); Lavington (CNC);

Wilmer (CNC).

Eucyrtopogon nebulo Osten-Sacken

Cyrtopogon nebulo Osten-Sacken 1877 : 309.

Eucyrtopogon nebulo Curran 1923 : 120-121.

This species is similar to diversipilosis Curran, especially the males,

but the females are readily distinguished by the absence of the pollen

from the seventh abdominal segment.

This species ranges from British Columbia and Alberta, south to

California, Wyoming, and Utah; six specimens were examined.

Localities - ALBERTA: Waterton (CNC).

Other localities - BRITISH COLUMBIA: Royal Oak (CNC); Duncan (CNC); Trinity Valley

(CNC); Cranbrook (AMNH). IDAHO: Moscow (USNM). WYOMING: Jackson's Lake (AMNH).

UTAH: Logan Canyon (USNM); Logan Peak (USNM). CALIFORNIA.

Genus Comantella Curran

Comantella Curran 1923 : 93. Type species: Cophura fallei Back 1909.

This genus is similar to Eucyrtopogon Curran, but is distinguished

by the presence of a curved spur on the apex of the front tibia. The male

genitalia show similarity in general appearance to those of Eucyrtopogon

Curran (figs. 243-247). Four species are included in this genus, all

found in the Nearctic Region. In Alberta, two species are known. The

species of this genus are very similar to one another. James (1937) has

presented a key to the species of this genus.

Key to the Species of Comantella Curran of Alberta

Thoracic mane on a clearly defined black vitta rotgeri James

Medial vitta of thorax at most poorly defined fallei Back

Adisoemarto

43

Comantella fallei Back

Cophura tallei Back 1919 • 378-379.

Comantella maculosa Curran 1923 : 93-94.

Comantella fallei Cut T Sin 1923 : 311-312.

This species is strikingly similar to rotgeri James. For the diag-

nostic characters, see James (1937). This species has been recorded

from Medicine Hat, Alberta, in late winter, and early and mid fall. It

ranges from Alberta southeast to Colorado and Nebraska; 12 specimens

were examined.

Localities - ALBERTA: Medicine Hat (UA and CNC).

Other localities - WYOMING. COLORADO: Denver (CNC). NEBRASKA: Crawford (CNC) .

Comantella rotgeri James

Comantella rotgeri James 1937 : 61.

This species is distinguished from fallei Back on the following

characters: medial vitta of thorax definitely demarcated, mystax coarser,

not white tipped, pale hairs and bristles deeper yellow, pale pile coarser,

less dense, ventral pile coarser, more extensively black.

This species ranges from Alberta to New Mexico; two specimens

were examined.

Localities - ALBERTA: Medicine Hat (USNM).

Other localities - COLORADO: Rio Seco, Costilla Co. NEW MEXICO.

Subfamily Laphriinae

This subfamily is represented by three genera, which belong to

two tribes. They live either in coniferous or parkland forests. Laphria

Meigen and Bombomima Enderlein are difficult to distinguish, but there is

a tendency in Bombomima Enderlein towards a more rounded abdomen.

There is also a difference in the shape of the pseudoclasper s of these

two groups: in Laphria Meigen, they are relatively simple; in Bombomima

Enderlein, they are forked (figs. 266, 270, 273, 276). Before the genus

Bombomima Enderlein was erected, its species were treated under DasylHs

Loew. Banks stated that DasylHs Loew (s.l. : DasylHs Loew and Bombomima

Enderlein) was an offshoot of the genus Laphria Meigen. Hull (1962) placed

these genera in different tribes.

Key to the Genera of Laphriinae of Alberta

1. Proboscis on apical half compressed dor socentr ally (fig. 62); third

antennal segment dilated, as long as first two segments together

(fig. 132); wings with first submarginal cell divided into two

(fig. 160) Po^onosoma R ondani

Proboscis compressed laterally; third antennal segment slender

(fig. 133), longer than first two segments together; first sub-

marginal not divided (fig. 161) 2

2. Abdomen robust, more or less rounded, densely pilose; pile on

44

Asilidae of Alberta

mesonotum covers ground color; pseudoclasper forked (fig.

266) Bombomima Enderlein

Abdomen parallel- sided, less pilose; mesonotal ground color,

not entirely covered by pile; pseudoclasper simple (fig. 251

pci) Laphria Meigen

Genus Pogonosoma Rondani

Pogonosoma Rondani 1856 : 160. Type species: Asilus maroccanum F. 1794.

In theNearctic Region, this genus has only three species. Cresson

(1920) treated melanoptera Wiedemann as conspecific with dorsata Say, but

Hull (1962) treated them as two different species. The third species is

ridingsi Cresson.

Face thick, with slight slit under antennal base; gibbosity rounded,

starting at about middle of face (fig. 23); vertex deeply excavated (fig.

24); ocellar plate with one or two pairs of bristles; bristles or, bristle-

like hairs present on orbital margin of front (opposite to antennal base,

fig. 24); first antennal segment stout, second shorter and smaller in

diameter, third narrow at base, dilated and oval apically, bristles present

on apical lower side of first segment (fig. 132); proboscis compressed

dor socentrally, pointed apically; palpi two- segmented, second segment

flattened, thin, and scoop-like (fig. 47). Thorax opaque, thinly pollinose;

hairs present on prothorax, posterior half of mesopleuron, upper half

of pteropleuron, and scanty on mesonotum, semierect; bristle-like hair s

present on metapleuron; dor socentral bristles absent. Femora slightly

thicker subapically; tibiae slightly curved, provided with hairs, hind

pairs with bristles; bristles present on tar si; second to fourth segments

of tar sus heart-shaped; claws strongly curved apically; empodium long.

Wings longer than abdomen (fig. 160), evenly covered with microtrichiae,

sometimes infuscated along veins; marginal cell closed with long stalk;

first submarginal cell divided by crossvein; first posterior cell open

or closed, sometimes with stalk; fourth posterior cell closed with stalk;

anterior crossvein at basal one-third of discalcell; alula well developed.

Abdomen semiparallel in males, slightly wider subapically in females;

first five segments with two to four bristles on middle of each side; pile

short, longer on venter, subappressed on dorsum.

Strickland (1938) recorded one species, ridingsi Cresson, from

Alberta.

Key to the Species of Pogonosoma Rondani of Alberta

Beard on lower orbital margin black; coxal pile mixed black and

white; front femoral and tibial hair s entirely black; metapleural

hairs in female black; abdominal hairs in female entirely black

stricklavdi new species

Beard entirely white; coxal pile entirely white; front femoral and

tibial hairs mixed black and white; metapleural hair s infemale

white; abdominal hairs infemale white on first three segments

Adisoemarto

45

ridingsi C res son

Pogonosoma ridingsi Cresson

Pogonosuma ridingsi Cresson 1920 : 214-215.

This species is similar to dorsata Say (Cresson 1920), but is dis-

tinguished from the latter mainly by the difference of the color of the

pile, hairs, and bristles.

This species ranges from British Columbia southeast to California

and Texas. Strickland (1938) included this species in his list. So far,

I have seen only a female specimen of Pogonosoma from Alberta, which is

stricklandi new species; holotype (USNM) and 10 additional specimens were

examined.

Localities - BRITISH COLUMBIA; Robson (CNC and USNM): Copper Mtn. (CNC); Kamloops

(CNC); Departure Bay (CNC); Victoria (CNC). CALIFORNIA: Plumas Co. COLORADO;

Florissant (type locality; USNM). TEXAS: Waco (USNM).

Pogonosoma stricklandi new species

This species is easily distinguished from ridingsi Cresson by the

presence of black pile on the lower orbital mar gin and the entirely black

pile or hairs on the front and the middle legs. The female differs from

those of ridingsi Cresson in the color of the metapleural hairs, entirely

black, and th§ abdominal pile, also entirely black.

Female. Pile, hairs, and bristles on face white, few black bristles

on middle of gibbosity; beard white, mixed with black on maxillae; oc-

cipital pile white on lower half, black on upper half and on orbital margin;

frontal, vertical, and ocellar pile black; ocellar bristles black; antennal

first two segments orange, bristles black, pile white on lower sides,

black on upper sides. Thorax white pollinose; pile mostly black, white

on proepimeron, and on anterior and posterior corners of mesopleuron;

metapleural hairs black; all bristles black. Legs black, coxae white

pollinose; pile, hairs, and bristles black, sparse white pile on front

coxae, few white hairs on subapical dorsal side of hind femur. Wings

covered with brown microtrichiae; halter es black. Abdominal pile,

hairs, and bristles entirely black.

This species is named in honor of the late Dr. E. H. Strickland.

Holotype: Female, Waterton, Alberta, 12. VII. 1923 (E.H.

Strickland); deposited in CNC.

Genus Bombomima Enderlein

Bombomima Enderlein 1914 : 253. Type species: Laphria fulvi thorax Fabricius

1805.

This genus resembles Laphria Meigen. The distinguishing character s

are, so far, not satisfying. The females of Bombomima usually have a

broad abdomen, but the abdomen of the males is slender as in Laphria

Meigen. Another character which may be used for distinguishing these

46

Asilidac- of Alberta

two groups is the shape of the pseucloelaspers. Psc'udoelasper s of 13

species of Bombomima and 11 species of Laphria have Ijeen examined. In

Laphria sackeni Wilcox, the ps eudoc la sper is not as in the other species of

Laphria Meigcn, but rather pr ovided with a "tooth", although not as complex

as those of Bombomima Enderlcin (figs. Z66, Z70, Z73, Z76).

This genus is known only from the Nearctic Region. In Alberta,

five species have been recorded, most of them were from the Montane

or Subalpine region.

Key to the Species of Bombomima Ended ein of Alberta

1. Third and fourth, sometimes also fifth, abdominal segments with

vivid orange-red hairs posteriorly; the succeeding segments

with yellow hairs fernaldi Back

Abdominal hairs unicolored, yellow Z

Z. Humerus, and usually also pronotum and upper occiput with black

hairs 3

Humerus, pronotum, and upper occiput with pale yellow hairs . . .4

3. Two apical abdominal segments entirely yellow haired

columbica Walker

Three apical abdominal segments yellow haired partitor Banks

4. Posterior half of mesonotum vivid orange-red haired, contrasting

with yellow hairs of anterior half .insignia Banks

Mesonotum uniformly with pale yellow hairs posticata Say

Bombomima columbica Walker

Laphria columbica Walker 1868 : 338.

Bombomima columbica Hull 196Z : 3Z5.

This species is similar to partitor Banks, but is distinguished by

the absence of yellow pile from the fourth abdominal segment. The male

genitalia are similar (figs. Z64-Z7 0).

This species ranges from British Columbia and Alberta, south to

California; 14 specimens were examined.

Localities - ALBLiR T^'V: Grinishaw (U.A).

Other localities - BRITISH COLUMBIA; Robson (CXC); Sugar Lake(CNC); A gas siz (CNC) ;

Victoria (CNC). WASHINGTON: I-.llensburg (AMNH). OREGON, CALIFORNIA.

Bombomima partitor Banks

Dasyllis partitor Banks 1917 : 54.

Bombomima partitor Hull 196Z : 3Z5.

This species is easily recognized by the pattern of the pile. The

pile is unicolored yellow; on the anterior half of the mesonotum it is

erect, and from dorsal view the ground color of the mesonotum is visible ;

on the posterior half of the mesonotum, the pile is decumbent, and con-

ceals the ground color. Pile is present on the last five abdominal seg-

ments .

Adisoemarto

47

This species is known from British Columbia and Alberta, south

to Idaho and Oregon; 36 specimens were examined.

Localities - ALBERTA; Banff (CNC); Banff, Lake Minnewanka, Campsite (UA); Seebe

(CNC); Nordegg, North Saskatchewan River Valley (UA).

Other localities - BRITISH COLUMBIA: Robson (CNC); Copper Mtn. (CNC); Chilcotin

(CNC); Sugar Lake (CNC); Seton, Lillooet (CNC); Uclucet (CNC); Nicola (CNC); Douglas Lake

(CNC); Shuswap Falls (CNO; .'^t. St. James (CNC); Ft. Steele (CNC); Westwold (CNC); Chase

(CNC); Vernon (CNC); Keremeos (CNC) ; Kelowna (CNC); Penticton (CNC) ; Summer land (CNC) .

IDAHO: Victor. WASHINGTON. OREGON.

B ombomima fernald'i Back

Dasyllis fernaldi Back 1904 : 2 90.

B ombomima femaldi Bromley 1929 I 160.

This species is similar to columbica Walker and partitor Banks, but

is distinguished by the color pattern of the pile and by the male genitalia

(figs. 271, 273). The pile on the presternum varies from entirely black

to mixed black and yellow; on the mesopleuron, the pile is of three dif-

ferent patterns, entirely black, entirely yellow, or mixed black and yellow.

This species ranges from Alberta and British Columbia, south to

Arizona and New Mexico; 127 specimens were examined.

Localities - ALBERTA; Jasper (CNC); Waterton (CNC and UA).

Other localities - BRITISH COLUMBIA; Robson (CNC); R evelstoke Mtn. (CNC); Lillooet

(CNC); Victoria (CNC); Departure Bay (CNC). WASHINGTON. OREGON. IDAHO; Moscow

Mts. (AMNH). WYOMING: Yellowstone (AMNH): Jackson (AMNH). UTAH. COLORADO:

Summit Road (AMNH); Aspen (AMNH); Ouray (AMNH); Electra Lake (CNC); Pingree Park

(AMNH). NEW MEXICO: Santa Fe Canyon (AMNH). ARIZONA: Flaggstaff (AMNH); Grand

Canyon (AMNH); San Francisco (AMNH). MONTANA; Glacier Park (UA).

Bombomima posticata Say

Laphria posticata Say 1824 : 374.

Bombomima posticata Bromley 1929 ’ 160.

This species is recognized by the uniformly yellow pilose meso-

notum. The mesonotal pile is mor e or less decumbent on the mesonotum,

erect only on the acrostichal line and transverse suture, from dorsal

view it appears as an inverted-T black marking. Bromley (1929) des-

cribed two varieties, brunnea and scufe/Zaris, which were treated as differ ent

species by Hull (1962) . The specimens found in Alberta belong to scutellaris

Bromley, buthere they are treated as poshcaZa Say, becaus e the differ ence

between these two forms is slight: scutellar bristles are blackin posticata

Say, and yellow in scutellaris Bromley.

This species ranges from Alberta and the Northwest Territories,

east to New Brunswick, and south to New York and Connecticut; holo-

type of scutellaris Bromley (CNC) and 26 additional specimens were ex-

amined.

Localities - ALBERTA; Lesser Slave Lake (UA); Cross Lake (UA); Nordegg (CNC).

Other localities - NORTHWEST TERRITORIES. MANITOBA: Victoria Beach (CNC);

Sandilands (CNC). ONTARIO: Lake Nipigon (CNC); Sadbury (CNC); Ottawa (CNC); Guelph

(CNC); Orilla (CNC); Sand Lake (CNC). QUEBEC: Aylmer (CNC); Fairy Lake (CNC). NEW

BRUNSWICK: St. Leonard (CNC); Nerepis (CNC). MAINE: Great Pond (USNM). NEW

HAMPSHIRE; Franconia (AMNH). MASSACHUSETTS; Amherst (USNM) ; Boston (AMNH) . NEW

YORK: North Elba (AMNH). CONNECTICUT; Avon (AMNH).

48

Asilidae of Alberta

Bombomima insignis Banks

Dasyllis insignia Banks 1917 : 54.

Bombomima insignis Hull 1962 : 325,

This species is similar to posticata Say, but is distinguished by the

presence of orange pile on the posterior half of the mesonotum. The

male genitalia are in general similar (figs. 274-276).

This species ranges from Alberta to Nova Scotia, south to Min-

nesota; 21 specimens were examined.

Localities - ALBERTA: Bilby (UA and CNC); Chipewyan (CNC).

Other localities - SASKATCHEWAN: Attons Lake (CNC); Weskesin Lake (CNC). MANI-

TOBA: Aweme (CNC); Teulon (CNC). MINNESOTA: Duluth (AMNH). ONTARIO: Lake Nipigon

(CNC). QUEBEC: Norway Bay (CNC). NOVA SCOTIA: Kentville (CNC),

Genus Laphria IVI eigen

Laphria Meigen 1803 : 27 0.

In this genus the abdomen tends to have parallel sides in both sexes,

but in janus McAtee, the abdomen of the females broadens slightly, as

in the species of Bombomima Enderlein. The pseudoclasper in the male

genitalia of janus McAtee, has a projection (fig. 261). This same form

of pseudoclasper is found in Laphria vultur Osten-Sacken and L. sackeni

Wilcox (figs. 262-263). The mesonotal and abdominal pile varies from

very sparse and short, as in felis Osten-Sacken and xanf/tippe Williston, or

appressed, as in aimatis McAtee and gilva L. , to erect, as in janus McAtee.

There are nine species known from Alberta, Most were collected

near coniferous forest.

Key to the Species of Laphria Meigen of Alberta

1. Dorsum of abdomen usually without pile, but if pile is present it

is spar se and very appressed; third antennal segment cylindrical

(fig. 133) 2

Abdominal pile erect, at least on sides; third antennal Segment

dilated subapically (fig. 137) • 3

2. A.bdomen entirely black; femora entirely orange; third antennal

segment five times as long as second (fig. 136) . . sac/a/es Walker

Abdomen black anteriorly, orange posteriorly; at least anterior

femora entirely black; third antennal segment seven times as

long as second (fig. 133) xanfft/ppe Willis ton

3. Ground color of abdominal dorsum entirely black 5

Abdominal dorsum with triangular orange markings 4

4. Three abdominal segments with orange markings, sixth segment

black gilva Linnaeus

Four abdominal segments with orange marking .a/maD's McAtee

5. Beard and pile on coxae white; bristles on face mainly black,

mixed with yellow pile in mane 7

Beard and pile on coxae yellow or orange; bristles on face mainly

yellow or orange 6

Adisoemarto

49

6. Bristles on face orange and black; humeral hair s orange or yellow

vivax Willis ton

Bristles on face orange-yellow; humeral hairs black in males,

mixed with orange in females janus McAtee

7. Pile and mane on face yellow or orange; abdominal pile concolorous

orange-yellow scorpio McAtee

Pile and mane on face white; pile on face white; pile on first ab-

dominal segment white or paler than the rest of abdominal pile

8

8. Metapleural hairs dark brown or black; all or few of scutellar

bristles black aetus McAtee

Metapleural hairs white; scutellar bristles yellowish

index McAtee

Laphria Xanthippe Willis ton

Laphria Xanthippe Williston 1884 : 31-32.

This species is easily recognized by the almost bare abdomen and

the reddish brown hind femora. There is slight sexual dimorphism in

this species. The reddish color of the abdomen and of the hind femur is

broader in the males. The beard is white in the males and black in the

females. The facial mane is entirely white in the males, mixed with

black in the females.

This species ranges from British Columbia and Alberta, south to

Oregon, and east to Colorado; 26 specimens were examined.

Localities - ALBERTA: Banff (CNC); Banff, Lake Minnewanka, Campsite (UA); Bow

River Forest, Wilkinson Creek (UA); Water'ton Lakes (CNC).

Other localities - BRITISH COLUMBIA: R evelstoke Mtn. (CNC) ; Fort Steele (CNC) ; Hedley,

Nickel Plate (CNC); Jesmond (CNC). OREGON: Mt. Hood (USNM). WYOMING: Yellowstone

Nat'l Park (AMNH). COLORADO: Electra Lake (AMNH).

Laphria sadales Walker

Laphria sadales Walker 1849 : 378-379.

Dasy//rs puiescens Williston 1884 : 32.

Laphria sadales McAtee 1918 : 161.

This species is similar to Xanthippe Williston, but is easily distin-

guished by the entirely black abdomen and reddish legs, except for the

black coxae. Sexual dimorphism is very slight. The abdomen of the

males is paler posteriorly, with golden yellow dorsal pile. The pile on

the abdomen of the females is entirely black.

This species ranges from Alberta to California and Colorado, east

to New Hampshire and Connecticut. The western and eastern populations

are probably connected by geographically intermediate populations in the

Boreal forests; 22 specimens were examined.

Localities - ALBERTA: Assineau River, near Lesser Slave Lake (UA); Banff (CNC);

Clymont (UA); Waterton (UA).

Other localities - BRITISH COLUMBIA: Robson (CNC); Trinity Valley (CNC); Nicolum

River; HopeMts.; Kaslo. WASHINGTON: MoscowMts. (USNM); Mt. R ainier (USNM) ; Electron

(USNM); Olympia; Pullman. OREGON: Strawberry Mt. (AMNH); Marys Peak (AMNH); Mt.

50

Asilidae of Alberta

Hood. IDAHO; Long Valley, Alpha (UA). WYOMING: Yellowstone (AMNH). CALIFORNIA.

Towle (AMNH); Fieldbrook; Humboldt Co. COLORADO; Chatanqua (USNM). ONTARIO: (CNC).

QUEBEC: Laniel (CNC). VERMONT: Rutland; Chittenden. NEW ITAMPSHIRE: Franconia

(AMNH); White Mts.; Mt. Washington (AMNH), MASSACHUSETTS: Southbridge (USNM).

CONNECTICUT: Avon (AMNH). NEW YORK; Axton.

Laphria scorpio McAtee

Lap/iria Scorpio Me A tee 1918 : 163-164.

This species can be distinguished from the two proceeding species

by the presence of erect pile on the abdomen, and from the other species

of the genus by the black bristles on the face and the concolorous ab-

dominal pile. Facial protrusion near antennal base not too obvious (fig.

25); a row of black bristles present on each facial submargin; facial

mane and pile orange; beard white; hairs on first palpal segment whit^

on second black; third antennal segment blade- shaped, without groove

(fig. 137). Thorax black; mesonotum and scutellum shiny, the rest

yellowish white pollinose; pile mostly black, white on propleuron and

anterior corner of sternopleuron, and golden yellow, appressed, on

mesonotum and scutellum; metapleuralhair s yellow mixed with few black.

Legs black; pile on coxae, on loser sides of femora, and on front and

middle tibiae of female, white; hajrs and bristles black; tomentum on

tarsi and frontal tibiae, brownish; claws black, empodium brownish

orange. Wings cover ed with brownish microtrichiae, darker along veins,

halter es yellow. Abdomen black; male sixth segment elevated medio-

posteriorly, with a pair of stumpy projections, seventh also with a median

projection, which more or less fits into a space between projections on

sixth segment (figs. 99, 100); pile short, orange, appressed on dorsum;

venter orange pilose; male genitalia black (figs. 243-258). In Alberta,

this species was collected on the edge of coniferous forest.

This species has beenrecorded mostly from eastern central North

America, from New Hampshire to Virginia, Ontario and Alberta; holo-

type and 19 additional specimens were examined.

Localities - ALBERTA; Kinuso, near Lesser Slave Lake (UA).

Other localities - ONTARIO: Trenton (CNC); Lake Nipigon (CNC), QUEBEC: Laniel

(CNC). VERMONT: Camel's Hump. NEW HAMPSHIRE: White Mts. (type locality; USNM and

AMNH); Mt. Washington (AMNH). NEW YORK: North Elba (AMNH); Chateaugay. PENNSYL-

VANIA: Springboro (USNM). VIRGINIA: Skyland (USNM).

Laphria aeatus Walker

Laphria aeatus Walker 1849 : 381.

This species resembles scorpio McAtee, but the color of the facial

mane and pile immediately distinguishes it from the latter. Other diag-

nostic characters are as follows: the third antennal segment has a nar-

row apical slit (fig. 138), and the abdominal pile is white on the first

segment, and yellow on the remainder . This species is found near coni-

ferous forests in northern Alberta.

This species ranges from Vermont and Ontario to Alberta; five

specimens were examined.

Localities - ALBERTA; Assineau River, near Lesser Slave Lake (UA).

Adisoemarto

51

Other localities - ONTARIO: Lake Nigipon (CNC).. VERMONT: Laurel Lake (USNM).

Laphria index McAtee

Laphria index McAtee 1918 : 164.

This species is similar to aeatus Walker, but is distinguished by

the longer mystax, the pattern of the mesonotal pile, triangular from

dorsal view, and entirely yellow scutellar bristles.

The species scorpio McAtee, index McAtee, and aeatus Walker, have

one character in common: the tubercles on the ends of the sixth and the

seventh abdominal segments. In Alberta, this species was collected

from the same habitat as that of LapAria aea^us Walker .

This species is known from eastern central North America and

Alberta; holotype (USNM) and 29 additional specimens were examined.

Localities - ALBERTA: Assineau River, near Lesser Slave Lake (UA).

Other localities - MANITOBA: Aweme (CNC). ONTARIO: Lake Nipigon (CNC); Lake

Abitibi (CNC); Point Pelee (CNC); Guelph (CNC); Jordan (CNC); Orilla (CNC); Bobcaygeon

(CNC); Ottawa (CNC). QUEBEC: Aylmer (CNC); Chelsea (CNC); Montreal (CNC); Hemming-

ford (CNC); Wakefield (QNC); Quoey Hill (CNC). NEW YORK: Nepara Park Yonkers, Flushing

(USNM); New York. CONNECTICUT: Avon (USNM). NEW JERSEY: Ramsey (USNM); Fort

Lee. PENNSYLVANIA: Harrisburg (type locality; USNM); Linglestown; Stoverdale. VIRGINIA:

Dead Run.

Laphria janus McAtee

Laphria janus h/ic Ate e 1918 : 153-154,

This species is readily distinguished from the other species of

Laphria Meigen of Alberta, by the bright orange color of the abdominal

pile, and the yellow mesonotal pile. The abdomen of the female is rather

rounded (fig. 101), and is more or less similar to that of Bombomima

Enderlein. The male genitalia are also similar to those of Bombomima

Enderlein, but the fork of the pseudoclasper is not very strong (fig, 261).

Most of the specimens from Alberta were collected near coniferous

forest.

This species is known in eastern and western central North

America. In the west it is distributed from Alberta to Washington, east

to Colorado, and in the east it is recorded from Maine to Michigan;

holotype (USNM) and 23 additional specimens were examined.

Localities - ALBERTA: High Level (UA); Kinuso (UA); Assineau River, near Lesser

Slave Lake (UA); Cross Lake (UA); Bilby (UA); Edmonton (UA); Nordegg, North Saskatchewan

River Valley (UA); Gorge Creek (UA); Banff (CNC).

Other localities - BRITISH COLUMBIA: Kaslo. WASHINGTON: Brodie. WYOMING.

COLORADO: Creede; Tolland. ONTARIO: Heyden; Sault St. Marie. MICHIGAN: Isle Royal;

Dickinson Co. NE W HAMPSHIR E: Mt. Washington (type locality; USNM); Mt. White; Ottolengui.

NEW YORK. MAINE.

Laphria vivax Williston

Dasyllis vivax Williston 1884 : 30.

Laphria vivax McAtee 1918 : 156.

This species is recognized by the pattern of the pile. The pile is

52

Asilidae of Alberta

yellow; on the abdomen it is decumbent on the posterior margins and

from a dorsal aspect it is visible only on the lateral and posterior mar-

gins.

This species ranges from Alberta and British Columbia, south to

Washington, and east to Colorado; 10 specimens were examined.

Localities - ALBERTA: Banff (UA); Banff, Sulfur Mt. (CNC).

Other localities - BRITISH COLUMBIA: Robson(CNC); Chilcotin (CNC) ; Copper Mountain

(CNC); Kaslo (CNC). WASHINGTON. IDAHO: Moscow (USNM). COLORADO: Marshall Pass

(USNM); Summit Co.

Laphria aimatis McAtee

Lap/jria aimahs McAtee 1918 : 160-161,

This species is easily recognized by the presence of orange-yellow

markings on some of the abdominal segments.

This species ranges from Alberta and British Columbia south to

California and Colorado; holotype and 21 additional specimens were ex-

amined.

Localities - ALBERTA: Brule Lake, near Jasper (USNM).

Other localities - BRITISH COLUMBIA: Robson (CNC); Midday Valley (CNC); Merritt

(CNC); Vernon (CNC); Oliver (CNC). IDAHO: Moscow Mts. (AMNH); Krasel (USNM)

CALIFORNIA: Baron (type locality; USNM); Midway (AMNH); Carrville, Trinity Co. (AMNH);

Shasta (AMNH); Edwards; Sierra Nevada; Placerville. COLORADO: El Paso (USNM); Lead-

ville (AMNH).

Laphria gilva Linnaeus

Asilus gilvus Linnaeus 1858 : 605,

Laphria gilva McAtee 1918 : 155-156.

This species is similar to aimatis McAtee, but is distinguished by

a difference in the number of orange markings on the dorsum of the ab-

domen: there are three on Laphria gilva L. , and one each on the third,

fourth, and fifth segments; the sixth segment is entirely black. The

male genitalia are also different from those of aimatis McAtee: in gilva

L. , each of the superior forceps is provided vAth two lamellate appen-

dages, while in aimatis McAtee, each forceps has orte lamellate appen-

dage (figs. 259, 260).

This species occurs in eastern as well as western central North

America, and is also knownfrom Europe; 65 specimens were examined.

Localities - ALBERTA: Medicine Hat (UA); Whitla (CNC); Lethbridge (CNC); Castle

Mountain (CNC); Banff (UA and CNC).

Other localities - NORTHWEST TERRITORIES: Cameron Bay, Great Bear Lake (CNG).

BRITISH COLUMBIA: Robson (CNC); Co'^per Mt. (CNC); Tuktakamin (CNC); Vavenby (CNC).

WASHINGTON. OREGON. MONTANA: Lame. WYOMING: Lander (AMNH). COLORADO:

Empire; Estes Park. ARIZONA; St. Catalina Mts. (AMNH). ONTARIO: Sudbury (CNC);

Ottawa (CNC); Thor Lake (CNC); Fort Williams (AMNH); Macbeth (AMNH); Sault St. Marie;

Whitefish Point. QUEBEC: Fort Cologne (CNC); Cascapedia (CNC); Trinity Bay (CNC) ;

Abbotsford (CNC); Laniel(CNC). NEW BRUNSWICK: Bathurst (CNC); NOVA SCOTIA; Baddeck

(CNC). MASSACHUSETTS: Tyingsboro; Blanchard; Dedham; Beverly; Burgess. MICHIGAN:

Alpena; Dickinson.

Adisoemar to

53

Subfamily Leptogastrinae

In 1909 Back recorded only one genus of this subfamily in North

America north of Mexico. He presented a synopsis of 15 species . Later,

additional genera were erected; Tipulogaster By Cockerell (1913), Psylonyx

by Aldrich (1923), Beameromyia and Apachekolos by Martin (1957).

In Alberta, the subfamily is represented by one genus with two

species, Leptogaster aridus Cole and L. coloradensis James.

Genus Leptogaster Meigen

Leptogaster Meigen 1803 : 269. Type species: Asilus cylindricus De Geer

1776.

Gonypes Latreille 1805 : 309. Type species: Asilus cylindricus De Geer

1776.

This genus is recognized by the following combination of charac-

teristics: the wings are without bands or spots, with five posterior cells,

the legs are slender, but the femora are somewhat club-shaped (fig. 85),

and the pulvilli are absent.

Head silvery white tomentose; face narrow, epistoma broader;

frontnarrow, wider toward vertex; mystax present along epistomal mar-

gin; first antennal segment small, second wider, with short hairs on

apical lower and upper sides, third elongate and attenuate, style long,

slender, spine present (fig. 139); palpi one-segmented, shining, clavate,

borne on a tubercle (fig. 46); occiput convex on lateral sides, few hairs

present on lower side, few short bristles on upper side behind vertex.

Thorax white tomentose; mesonotum convex, slightly protruding anter-

iorly (fig. 64); mesonotal vittae present, not reaching hind margin; two

bristles present above wing base. Legs shining, pale orange; coxae

white pollinose; hind femora swollen distally; tibiae slender, hind pair

gradually thicker apically (fig. 85); basitarsi as long as second tarsal

segments; claws long, empodium half as long as claws; pulvilli absent.

Wings shorter than abdomen, covered with micr otrichiae; alula absent;

all peripheral cells open; third branch of media and anterior branch of

cubitus fused for a considerably long distance (fig. 163); second branch

of cubitus and second anal vein almost parallel; halteres brownish with

long stalk. Abdomen slender , elongate (fig. 102); second segment much

longer than first; posterior segments wider; posterior margin of first

with one or more pairs of bristles; superior forceps of male genitalia

with ventral excavation; ovipositor short.

Key to the Species of Leptogaster Meigen of Alberta*

Occipital bristles black; superior forceps of male genitalia with-

out spine-like projection apically aridus Cole

Occipital bristles white, pale, or tinged with color; superior for-

ceps of male genitalia with spine-like projection apically ... .

’"From Martin (1957).

54

Asilidae of Alberta

coloradensis James

Leptogaster aridus Cole

Leptogaster aridus Cole 1919 • 229.

According to Martin (1957), this species is easily recognized by

the characteristics of the male genitalia. A female specimen was col-

lected from Writing- on-Stone Provincial Park, an almost arid area,

where the vegetation was short grass and cacti. This species was also

found in association with Nerax bicaudatus Hine.

This species is known from Alberta, and according to Martin (1957),

is found along the Pacific coast and some localities in California; 10

sDecimens were examined.

Localities - ALBERTA: Writing-on-Stone Provincial Park (UA).

Other localities - WASHINGTON: Yelm (UA). OREGON: Mt. Hood. CALIFORNIA: Big

Bear Lake, Hannah Flats (AMNH); Idyllwild, San Jacinto Mts. (AMNH); Strawberry, Tuolumne

Co. ; Snowline Camp, Eldorado Co. ; Yosemite Nat'l Park; Whitney Portal, Inyo Co. ; Tanbark

Flat, Los Angeles Co. ; Glendale.

Leptogaster coloradensis James

Leptogaster coloradensis Ja.mes 1937 : 14.

Variation in the pollen color was described by Martin (1957).

This species ranges from Alberta to Kansas; one specimen was

examined.

Localities - ALBERTA: Lethbridge (CNC).

Other localities - WYOMING. SOUTH DAKOTA: Cedar Canyon; Cottonwood; Buffalo;

Highmore; Presho; Kennebec; Desmet; Gettysburg. COLORADO: Boulder (type locality);

Berthoud Pass. KANSAS: Ellis Co. ; Sheridan Co.

Subfamily Asilinae

This subfamily is a highly specialized and complex group. Special-

ists have devoted much effort to defining the genera. The members of

this group inhabit various habitats, such as open grassland, sandy beach,

and near coniferous forest.

Key to the Genera of the Asilinae of Alberta

1, Wing with three submarginal cells (fig. 164) Promachus Lioew

Wing with two submarginal cells 2

2. Antennal style longer than third antennal segment (fig. 141) .... 3

Antennal style as long as or shorter than third segment (fig. 143)

4

3. Facial gibbosity very prominent, bulging on top (fig. 27); antennal

style, including spine, at least twice as long as third segment

(fig. 141); scutellum at base less than twice its length (fig. 70);

male genitalia longer than high (fig. 280); ovipositor almost

Adisoemarto

55

three times as long as seventh abdominal segment (fig. 114) ..

Nerax Hull

Facial gibbosity almost flat on top (fig. 26); antennal style one and

a quarter as long as third antennal segment (fig. 140); scutellum

at base twice as long as its length (fig. 69); male genitalia

higher than long (fig. 277); ovipositor less than twice as long

as seventh abdominal segment (fig. 113); ninth sternum provided

with spines (fig. 113) Proctacanthella Br omley

4. Scutellum with bristles (fig. 71, 72) Asilvs complex

Scutellum without bristles (fig. 73) Negasilus Curran

Genus Protnachus Loew

Promachus Loew 1848 : 390. Type species: Asilus maculatus F. 1775

Trupanea Macquart 1838 : 91. Type species: Asilus maculatus F. 1775.

Preoccupied by Schrank 1803, Diptera.

Telejoneura Rondani 1864 : 48. Unnecessary change of name.

Bactria Megerle (Ms) in Meigen 1820 : 307. Nomen nudum.

Promachus can be easily recognized by the character of the wing

venation: three submarginal cells, with the radial crossvein near the

middle of the first (fig. 164).

There is one species, dimidiatus Curran, known in Alberta.

Promachus dimidiatus Curran

Promachus dimidiatus Curran 1927 : 87-88.

According to Curran (1927) this species can be easily confused

with bastardi Macquart, but is distinguished by the absence of black hairs

from the first abdominal segment of the male. By comparing two females

and one male of bastardi Macquart with six pairs of dimidiatus Curran, the

following characters, which are more or less constant, have been found

useful for distinguishing these two species: in dimidiatus Curran, the

metanotal hair s and the hair s on the third abdominal segment, are entirely

white, while in bastardi Macquart, the metanotal hairs are mostly black,

and the hairs on the third abdominal segment are mixed black and white.

Both species have black bristles on the first abdominal segment.

This species ranges from Alberta to Manitoba, south to New Mexico;

holotype (CNC) and 18 additional specimens were examined.

Localities - ALBERTA: Orion (UA and CNC); Milk River (CNC); Dunes (CNC).

Other localities - MANITOBA: Aweme (type locality; CNC), Onah (CNC). COLORADO:

Master, Plainview (USNM). NEW MEXICO: Arroyo, Pecos R iver (USNM) .

Genus Proctacanthella Bromley

Proctacanthella Bromley 1934 : 96. Type species: Asilus cacopilogus Hine

1909.

This group was separated from Asilus by Bromley (1934) on the

56

Asilidae of Alberta

basis of the absence of hairs from the metanotal slopes, and by the cy-

lindrical ovipositor, which is provided with a circlet of spines.

There are five species of this genus, all Nearctic, and in Alberta,

this genus is represented by one species, cacopiloga Hine.

Proctacanthella cacopiloga Hine

Asilus cacopilogus Hine 1909 I 165-166.

Proctacanthella cacopiloga Bromley 1934 : 96.

Hine ( 1909)placed this species and /eucopo^on Williston in the Rhadiurgus

group of the genus Asilus L. Curran (1924) was the first to realize that

Asilus cacopilogus Hine was different from the other species of Asilus L. ,

and suggested it belonged to Erax Scopoli. The males of this species are

easily recognized by the shape of the male genitalia (figs. 277-279), but

the females are hard to distinguish from one another.

This species ranges from Alberta to T exas, and east to New Jersey;

32 specimens were examined.

Localities - ALBERTA: Medicine Hat (UA); Orion (UA); Writing-on-Stone Provincial

Park (UA).

Other localities - MANITOBA: Aweme (CNC). WYOMING: Lance Creek (AMNH). NEB-

RASKA: Mitchell (CNC); Fromont. COLORADO: White R ock (AMNH) ; Wray (AMNH); La Junta

(AMNH); Pueblo (AMNH); Olney (AMNH); Fort Collins (AMNH). KANSAS: Clark Co.

OKLAHOMA: Admore. TEXAS: Forestburg (AMNH); Rosser. ILLINOIS; Havana. INDIANA:

Mineral Springs (CNC). NEW JERSEY: Anglesea.

Genus Nerax Hull

?Verax Hull 1962 : 476. Type species: Asilus aestuans L. 1767.

Erax Macquart 1838. Type species: Erax rufibarbis Macquart 1838.

Efferia Coquillett 1893, Type species: Efferia candidus Coquillett 1893.

This group is readily recognized by the form of the male genitalia

(fig. 280) and the ovipositor (fig. 114), the shape of the third antennal

segment and the style (fig. 141), and the wing venation (figs. 165-168).

This genus is confined to the New World. In Alberta, TVerax is re-

presented by four species.

Key to the Species of Nerax Hull of Alberta

1. Third vein of wing branched before tip of discal cell (figs. 165,

166) 2

Third vein branched beyond tip of discal cell (figs. 167, 168). . . .3

2. Acrostichal line with long hairs and bristles; last two segments

of abdomen of male silvery pollinose, hairs sparse, few in

number; abdomen of female yellowish pollinose

bicaudatus Hine

Acrostichal line without hairs and bristles, but with short setulae;

all abdominal segments of male silvery pollinose, hairs num-

erous, long, except for a longitudinally-directed bare’ line at

Adisoemarto

57

middle; abdomen of female silvery white pollinose

canus Hine

3. Tibiae black subcupreus Schaeffer

Tibiae orange-brown costalis Williston

Nerax bicaudatus Hine

Erax bicaudatus Hine 1919 • 138.

Nerax bicaudatus Hull 1962 : 478,

This species is recognized by the wing venation, the pollen pattern

of the males and the females, and the presence of comparatively long

hairs on the acr ostichal line of the mesonotum. The color of the bristles

varies from entirely white to entirely black in almost every arrangement.

In the males, this. variation occur s in the ocellar bristles, frontal hairs,

few bristles of the mystax, upper occipital bristles, presutural dorso-

central bristles, postalar bristles, mesopleural bristles, and scutellar

bristles. In the females, the variation is less obvious. This species

inhabits arid grassland, pastures, and short grass areas with cacti. It

has been found associated with Stenopogon neglectus Bromley a.nd Leptogaster

aridus Cole.

This species ranges from British Columbia to Manitoba, and south

to Texas; 65 specimens were examined.

Localities - ALBERTA: Drumheller (UA); Wardlow (UA); Medicine Hat (UA and CNC);

Burdett (UA); Manyberries (UA); Orion (UA and CNC); Comrey, Milk River Valley (UA);

Writing-on-Stone Provincial Park (UA); Lethbridge (UA and CNC),

Other localities - BRITISH COLUMBIA: Summerland (UA), MANITOBA: Aweme (CNC),

COLORADO, TEXAS: Amarillo; Plainview; Hereford; Coyote Lake, Bailey Co,

Nerax canus Hine

Erax canus Hine 1916 : 22.

Nerax canus Hull 1962 : 478.

This species is similar to bicaudatus in the wing venation (figs. 165,

166), but is distinguished by other characters: the acrostichal bristle-

like hairs are absent, the abdomen is evenly silvery white pollinose in

both sexes, and the ocellar bristles are two in number (six in bicaudatus

Hine).

This species ranges from British Columbia to California. I have

not seen Alberta specimens, but Strickland (1946) included this species

in his list; 14 specimens were examined.

Localities - BRITISH COLUMBIA: Kamloops (CNC); Seton Lake (CNC); Nicola (CNC);

Lone Pine (CNC) ? Vernon (CNC); Oliver (CNC), CALIFORNIA: CrescentCo, (USNM); Mariposa

(USNM); Westgabd Pass Plateau (USNM); Ivfidway (USNM); Sierra Nevada (USNM); Antioch

(USNM),

Nerax subcupreus Schaeffer

Erax subcupreus Schaeffer 1916 : 66.

58

Asilidae of Alberta

Nerax subcupreus Hull 1962 : 478.

This species belongs to the carinatus group (Hine 1919), in which

the acrostichal mane is present, and the costal and the subcostal veins

of the wings of the males are slightly bent (fig. 167). This species ranges

from Alberta to Arizona, and from Colorado westward to California;

holotype (USNM) and nine additional specimens were examined.

Localities - ALBERTA: Medicine Hat (UA and CNC).

Other localities - IDAHO: Victor (AMNH). WYOMING: Stewart R. Sta. (AMNH).

COLORADO: Alamosa (AMNH); Electra Lake (AMNH). ARIZONA: Prescott (type locality;

(USNM). CALIFORNIA: Essex (AMNH).

Nerax costalis Willi ston

Erax costalis Williston 1885 : 64,

Williston (1885) included this species in his key, but did not describe

it, Aldrich (Hine 1919) and Hull (1962) did not recognize this species,

but it is accepted by some other authors. It is similar to shbcupreus

Schaeffer, but readily distinguished by its orange-brown tibiae.

This species ranges from Alberta and Saskatchewan, south to

Colorado; seven specimens were examined.

Localities - ALBER TA: Medicine Hat (UA) ; Lethbridge (CNC) ; Oldman R iver, Lethbridge

(CNC).

Other localities -SASKATCHEWAN: Rockglen (CNC). MONTANA. WYOMING. COLORADO.

The Asilus Complex

This complex probably includes several related genera, but no

attempt is made to separate them. All of the species involved in this

discussion are treated under the name Asilus Linnaeus.

There are 12 species of this complex present in Alberta.

Key to the Species of the Asilus Complex of Alberta

1. Apical margins of abdominal segments provided with distinct bris-

tles 2

Apical margins of abdominal segments without bristles 9

2. Femora black, at most only with apical reddish brown or yellow

bands 3

Femora reddish brown or yellow on posterior sides

delusus Tucker

3. Posterior margin of eighth sternum of abdomen of the male extended

into a subtriangular lobe, provided with tuft of'hairs; ovipositor

three to four times as long as sixth and seventh segments to-

gether (figs. 115, 116) occidentalis Hine

Posterior margin of eighth sternum of abdomen of male not exten-

ded; ovipositor at most twice as long as sixth and seventh seg-

ments together 4

4. Four scutellar bristles ca//idus Williston

Adisoemarto

59

Two scutellajr bristles, sometimes with additional small ones ... 5

5. White bristles on poster o-ventral side of front tibia 6

Black bristles on poster o-ventral side of front tibia 8

6. Antennal style two-thirds as long as third antennal segment

erythocnemius Hine

Antennal style less than half as long as third antennal segment. .

7

7. Male genitalia and eighth sternum of abdomen of female orange;

penis as in figure 297; black hairs on sternum of abdomen of

female mesae Tucker

Male genitalia and eighth sternum of abdomen of female dark brown;

penis as in figure 300; white hairs on eighth sternum of ab-

domen of female cumbipilosus new species

8. Ventral side of front femur with rather stout black bristles

paropus Walker

Ventral side of front femur with pale long hairs and pile

snowi Hine

9. Metanotal slope without pile or hairs auriannulatus Hine

Metanotal slope with pile or hairs 10

10. Posterior sides of femora reddish brown aridalis new species

Femora uniformly black 11

11. Mystax and mesonotal pile black nitidifacies Hine

Mystax and mesonotal pile white gramalis new species

Asilus delusus Tucker

Asilus delusus Tucker 1917 : 92.

This species is readily recognized by the color of the femora:

black on the anterior sides, orange on the posterior sides. Another

species, aridalis n. sp. , has the same color pattern of the legs, but the

abdomenis without bristles on the posterior sides. Antennal style slen-

der, as long as third segment (fig. 142). Upper side of mystax black,

lower side white; bristles on mesonotum and legs mostly black; two

black scutellar bristles present; superior forceps of male genitalia

twice as long as gonopods (fig. 282).

This species ranges from Alberta southward to Arizona, and from

Utah eastward to Kansas; seven specimens were examined.

Localities - ALBERTA: Medicine Hat (UA and CNC); Lethbridge (UA).

Other localities - MONTANA. UTAH: Glacier Lake (USNM). COLORADO. KANSAS.

ARIZONA: Santa Rita Mts. (USNM); Chiricahua Mts. (USNM).

Asilus occidentalis Hine

Asilus occidentalis Hine 1919 : 147-148.

This species is readily recognized by the presence of a lobe-like

extension on the posterior margin of the eighth sternum of the abdomen

of the male (figs. 115, 116), and the ovipositor is three to four times as

long as the sixth and seventh abdominal segments together.

This species ranges from British Columbia to California, and the

60

Asilidae of Alberta

presence of this species in Alberta is doubted. Strickland (1938) might

have based his record on a misidentified specimen of Asilus callidus

Williston; six specimens were examined.

Localities - BRITISH COLUMBIA: Royal Oak (CNC); Aspen Grove (CNC); Nicola Lake,

Merritt (UA); Keremeos (UA); Victoria (CNC), OREGON: Antelope Mt. , Harney Co. (UA).

CALIFORNIA. NEVADA.

Asilus paropus Walker

Asilus paropus Walker 1849 I 455.

This species is similar to callidus Williston and erythocnemius Hine.

It is distinguished from callidus Williston by the number of the bristles

present on the scutellar mar gin (figs. 71, 72), and from erythocnemius Hine

by the size of the antennal style (figs. 143, 144). This species also re-

sembles snowi Hine, but is distinguished by the presence of stout bristles

on the ventral side of the front femur, instead of hairs and pile (figs.

82, 83).

This species ranges from Alberta to New Mexico, and eastward

to New Hampshire and Connecticut; 64 specimens were examined.

Localities - ALBERTA: Bilby (UA); Golden Spike (UA); Devon (UA); Cypress Hills (UA);

Calgary (CNC).

Other localities - SASKATCHEWAN: Saskatoon (CNC); Skipton (CNC). MANITOBA:

Teulon (CNC); Melita(CNC). ONTARIO; Ottawa (CNC). QUEBEC: Hull (CNC); Aylmer (CNC).

NOVA SCOTIA: DigbyCo. (CNC). NORTH DAKOTA. WYOMING. UTAH. CALIFORNIA. NEW

MEXICO.

Asilus callidus Williston

Asilus callidus Williston 1893 : 75.

This species can be easily mistaken for occidentalis Hine, but can be

distinguished by the absence of a lobe-like projection from the eighth

sternum of the abdomen of the male; the female has a comparatively

short ovipositor. The male genitalia of the two species are slightly dif-

ferent from one another (figs. 283-286). This species inhabits open

spaces near or within coniferous forests. In Alberta, this species has

been found in numbers together with Stenopogon inquinatus Loew. It ranges

from Bi-itish Columbia to Massachusetts, southward to California and

Utah. In Alberta it is not found in the prairies. It is probably a boreal

and subalpine species; 169 specimens were examined.

Localities - ALBERTA: Lac la Biche (UA); Opal (UA); Tawatinaw (UA); Bilby (UA);

Golden Spike (UA); Jasper, Lake Celestine (UA); Gorge Creek (UA); Flat Creek (UA); Banff

(CNC); Banff, Lake Minnewanka, Davil's Gap Trail (UA); Nordegg (UA); Seebe (DE); Cowley

(CNC); Coleman, Lake Island (CNC) ; Waterton Lakes (CNC) .

Other localities - BRITISH COLUMBIA. WASHINGTON. OREGON. CALIFORNIA: Colville,

R ock Creek (CNC) ; Coalinga (CNC); Carson Pass (CNC); Lone Pine (CNC). NEVADA: Ormsby

Co, (USNM). UTAH: Zion Nat'l Park (USNM). MASSACHUSETTS: Springfield (USNM) ,

Adisoemarto

61

Asilus erythocnemius Hine

Asilus erythocnefhius Hine 1919 : 163.

This species resembles callidus Williston, but it is paler, yellowish

pollinose, and it has only two scutellar bristles. It is distinguished

from other species by the length of the antennal style (fig, 144). The

pollinose excresence of a number of specimens is bright golden yellow

instead of yellowish, and the mesonotal vittae are clearly defined blackish

brown. All other characters are the same. This species is found in

open grassland or in the open spaces within the parkland or coniferous

forests. It ranges from British Columbia to Wyoming, and east to Mas-

sachusetts and Maryland; 126 specimens were examined.

Localities - ALBERTA: Peace River (UA); Lac la Biche (UA); Edmonton (UA and LMK);

Consort (UA); Rosebud (UA); Cassils (UA); Oyen (UA); Medicine Hat (UA); Orion (UA); Many-

berries (UA); Burdett (UA); Lake Newell, Kinbrook Island Provincial Park (UA); Scandia (UA);

Cypress Hills (UA); Lethbridge (UA); Seebe (UA); Waterton (UA).

Other localities - BRITISH COLUMBIA: Trinity Valley (CNC); Prince George (CNC).

QUEBEC: Natashqua (CNC). MASSACHUSETTS. MARYLAND. MONTANA. WYOMING.

Asilus snowi Hine

Asilus snowi Hine 1919 l 160,

This species is similar to paropus Walker, but the ventral side of

the front femur is provided with hairs and pile only, and is without bris-

tles.

This species occur s mainly in eastern central North America, from

South Dakota to Kansas, eastward to Nova Scotia, and south to Virginia.

The presence of this species in Alberta is doubted, but Strickland (1938)

listed it, possibly on the basis of misidentified specimens; 13 specimens

were examined.

Localities - ONTARIO: Ottawa (UA). QUEBEC: Hemmingford (CNC). NOVA SCOTIA:

Truro (CNC). CONNECTICUT: Cornwall; Hamden; Stratford; Marlborough; Wallington;

Stamford. DISTRICT OF COLUMBIA: Little Falls (USNM), VIRGINIA: Great Falls (USNM) .

ILLINOIS: Fort Sheridan (UA).

Asilus nitidifacies Hine

Asilus nitidifacies nine 1919 : 165,

This species is recognized by the black mesonotal pile, and also

by the shape of the superior forceps of the male genitalia (fig. 288).

The wings are evenly covered with microtrichiae, and brownish mar-

kings are absent from the middle of the subcostal cell (fig. 169). This

species is mostly found on the edges or in the open spaces of coniferous

forests. It ranges from British Columbia to Oregon, eastward to

Quebec; 19 specimens were examined.

Localities - ALBERTA: Cross Lake Trail (UA): Flatbush (UA); Opal (UA) ; Banff (UA);

Flat Creek (UA); Moraine Lake (CNC).

Other localities - BRITISH COLUMBIA: Terrace (USNM); Ainsworth (USNM). OREGON:

Mt. Hood (USNM), ONTARIO: Larder Lake (CNC). QUEBEC: Seven Isles (CNC) .

62

Asilidae of Alberta

Asilus auriannulatus Hine

Asilus auriannulatus Hine 1906 : 29.

This species is easily recognized by the color pattern of the legs.

The legs are mostly orange-yellow, but the coxae are black, the front

and the middle femora are black on the anterior basal two-thirds. The

male genitalia are easily distinguished from those of the other species

(figs. 291-294).

This species ranges from Alberta, British Columbia, south to

California; 21 specimens were examined.

Localities - ALBERTA: Seebe (DE); Banff (UA and CNC); Gorge Creek (UA); Waterton

(CNC).

Other localities - BRITISH COLUMBIA: Fernie (CNC) ; Mara (CNC) . OREGON: Prineville,

Hood River (USNM), CALIFORNIA. WYOMING.

Asilus mesae Tucker

Asilus mesae Tucker 1907 : 92.

This species is easily recognized by the orange color of the male

genitalia and of the eighth sternum of the abdomen of the females. This

species is similar to erythocnemius Hine, but can be distinguished by the

size of the antennal style: less than half of the third antennal segment

in mesae Tucker, and more than two-thirds in the latter. It is also

similar to cumbipilosus new species. For details, see under cumbipilosus ,

In Alberta, this species inhabits the grasslands of the paririe.

This species ranges from British Columbia and Alberta, southward

to Colorado; 23 specimens were examined.

Localities - ALBERTA: Drumheller (UA) ; Lake Newell, Kinbrook Island Provincial Park

(UA); Medicine Hat (UA); Manyberries (UA); Taber (CNC).

Other localities - BRITISH COLUMBIA; Oliver (CNC). IDAHO: Bear Lake (CNC); Paris

(CNC); Mt. Pelier (CNC). WYOMING: Green R iver (CNC) ; R ock Springs (CNC) ; R awlins (CNC) ;

Carbon Co. (CNC). UTAH: Benson(CNC); Snowville (CNC). COLORADO: Pagosa Spring (CNC);

(CNC); Regnier (CNC); White Rock (CNC); Walsenburg (CNC); Grand Junction (CNC); Animas

(CNC). KANSAS.

Asilus cumbipilosus new species

This species is markedly similar to mesae Tucker, and is distin-

guished by differences in the male genitalia and the ovipositor. The

other characters are more or less similar in both species. These two

species occur in the same habitats. Other characters, except male

genitalia and ovipositor, similar to those of mesae Tucker; male genitalia

dark brown; penis with short arms, as long as penis sheath (fig. 300);

ovipositor dark brown, eighth sternum of abdomen of female with white

hairs.

The name cumbipilosus hSiS been chosen, because of the appressed

hairs present on the legs. This species is an inhabitant of short grass

prairie.

Holotype: Male, Etzikom Coulee, Alberta, 3 . VIII . 1963 (J. &C.D.

Sharplin and S. Adisoemarto); deposited in CNC.

Adisoemarto

63

Paratypes: three females, same data as for holotype; one female.

Welling, Alberta, 19. VII. 1922 (H. L. Seamans); one male Medicine Hat,

Alberta, 8. VII. 1932 (F.S. Carr); one female, Orion, Alberta, 9. VII,

1950 (E.H. Strickland); two n;iales, two females, Kinbrook Island Pro-

vincial Park, Lake Newell, Alberta, 10. VI. 1964 (S. Adisoemarto); all

are deposited in UA.

Asilus aridalis new species

This species is more or less easily recognized by the greyish pol-

linose body, and distinguished from mesae Tucker and cumbipilosus new

species by the absence of the bristles from the posterior sides of the

abdominal segments and the color pattern of the femora.

Male. Head yellowish - grey pollinose; gibbosity from about the

middle of face; mystax white with few black hairs on top; antennae black,

first segment one and one half times as long as second, third segment

attenuate apically, as long as first two together; style about two-thirds

as longas third antennal segment (fig. 145); frontal hairs white; ocellar

hairs black; occipital bristles mostly white, mixed with a few black on

upper side; beard and pile on lower side of proboscis white; palpi black,

long, one- segmented, white haired; proboscis black. Thorax yellowish

grey pollinose; pile white; hairs on postsutural dorsocentral area, on

scutellum, and on metanotal slope, white; mesonotal setulae white, black

onacrostichal line; mesonotal bristles black, two pr esuturals, two intra-

alars, onepostalar, and five dor socentrals; two scutellars black; meta-

pleural bristles white. Legs with coxae greyish pollinose; pileand hairs

on front and middle coxae white; anterior sides of femora black, pos-

terior sides reddish yellow, provided with appr ess ed, short, white setu-

lae, and long white hairs also present on ventral sides; tibiae orange-

yellow, black on tips, with appressed, short, white setulae and golden

tomentum on antero- ventral sides of front pair and on posterior sides of

hind pair; tarsi orange-yellow, black haired, tomentum present as con-

tinuation from tibiae; claws black; empodia as long as claws, black;

femoral bristles white, tibial bristles mostly black; tarsal bristles

entirely black. Wings hyaline, microtrichiae light brownish; halteres

greyish yellow, black tinged. Abdomen greyish, pollinose, brownish

markings on mid-dorsum of each segment; hairs white, on first seg-

ment erect, appressed on the remaining; white bristles present on sides

of first segment; male genitalia reddish brown (figs. 301-303).

Female. Most characters, similar to those of male, different in

the followings: mystax mostly black, white bristles only on epistomal

margin; hairs on first two antennal segments black, few white on dor-

sal sides; frontal hairs black; mesonotal setulae entirely black; bris-

tles on legs mostly black; dorsal abdominal hairs black; ovipositor

black, twice as long as seventh abdominal segment, black haired.

This species is called aridalis , because the specimens live in the

arid areas. The habitat of this species is similar to that of cumbipilosus

new species.

Holotype: Male, Dinosaur Park, Steveville, Alberta, 9. VI. 1964

64

Asilidae of Alberta

{S. Adisoemarto) ; deposited in CNC.

Allotype: Female, same data as for holotype; deposited in CNC.

Paratypes: one male, two females, same data as for holotype

(UA); male, female, Kinbrook Provincial Park, Lake Newell, Alberta,

10. VI. 1964 (S. Adisoemarto) (UA); two males, female, Scandia, Al-

berta, 20. VI. 1956 (E.E. Sterns) (CNC); female. Medicine Hat, Alberta,

16. VII. 1956 (E.E. Sterns) (CNC); male, Lethbridge, Alberta, 4. VII.

1923 (H. E. Grey) (CNC); female, Lethbridge, 6. VII. 1956 (E.E. Sterns)

(CNC); male, female, Lethbridge, 7. VII. 1956 (E.E. Sterns) (CNC).

Asilus gramalis new species

This species is similar to mesae Tucker and cumbipilosus new species,

but is readily distinguished by the absence of the bristles from the pos-

terior sides of the abdominal segments. It is distinguished from aridalis

new species by the golden pollinose body and entirely black femora.

Total length ranges from 9. 0 mm to 13. 0 mm.

Male. Face, front, and vertex golden yellow pollinose; gibbosity

from about the middle of face; mystax mostly white, three black bris-

tles present on top and few on lower corners; antennae black, first two

segments black haired, third segment tapers apically, one and a half

times as long as first two together; style one-third as long as third an-

tennal segment (fig. 146); frontal hairs white; ocellar hairs black; oc-

cipital bristles entirely white; beard white, pile on lower side of pro-

boscis white; palpi black, long, one- segmented, black haired; proboscis

black. Thorax golden yellow pollinose, pile, hairs, and bristles mostly

white; hairs on mesonotum not setula-like, more or less erect; black

hairs present on space between humeri and dor socentral area; mesonotal

postalars, and six dor socentrals (eight on right side), mostly black; two

scutellars black; metapleural bristles white. Coxae golden yellow pol-

linose, anterior pairs with white pile and bristles; femora black, red-

dish brown streaks present on ventr o-posterior sides, with appressed,

short, white setulae, and a row of white bristles on ventral sides; tibiae

reddish brown, black on tips, with appressed, short white setulae, and

golden tomentum on antero- ventral sides of front pair, and on posterior

sides of hind pair, bristles white; tarsi reddish brown, white haired,

tomentum present as continuation from tibiae, bristles mixed black and

white; claws black, empodia orange-yellow. Wings hyaline, micro -

trichiae brownish; halter es slightly brownish yellow. Abdomen greyish

yellow pollinose; hairs white, appressed; bristles white, present only

on sides of fir st segment; male genitalia reddish brown; superior forceps

one and a half times as long as gonopods (figs. 304-306); hairs white.

Female. Similar to male; ovipositor black, one and a half times

as long as seventh segment, black haired.

This species has been named gramalis , because the specimens live

in grass fields. The habitat of this species is similar to that of aridalis

and cumbipilosus.

Holotype: Male, Scandia, Bow River, Alberta, 10. VI. 1964 (S.

Adisoemarto) in copula with allotype; deposited in CNC.

Adisoemarto

65

Genus Negasilus Curran

Negasilus Curran 1934 : 184.

This genus is distinguished from Asilus complex by the absence of

the scutellar bristles. The other character s are not different from those

of the Asilus complex.

The genus is monotypic.

Negasilus belli Curran

Negasilus belli Cur ra.n 1934 : 184.

This species is similar to Asilus cumbipilosus new species, but is

easily distinguished by the absence of the scutellar bristles and the bris-

tles on the posterior margins of the abdominal segments. The genitalia

are also different. A female specimen from Lethbridge, Alberta differs

slightly from the others in the following respects: occipital bristles

bleck, frontal hairs black, body with golden yellow pollen, brighter than

in the other specimens; kept in CNC.

This species ranges from Alberta to Colorado, and west to Califor-

nia; 25 specimens were examined.

Localities - ALBERTA; Consort (CNC); Claresholm (CNC); Bow Island (CNC): Cypress

Hills (UA); Scandia (CNC); Taber (CNC); Lethbridge (CNC).

Other localities - SASKATCHEWAN: Assiniboia (CNC). WYOMING: Laramie (USNM).

COLORADO: Creede (USNM). UTAH: Laketown (USNM); Manila (USNM). NEVADA: Fallon

(AMNH). CALIFORNIA: Cedarville (USNM); Lake Mono Co. (AMNH).

ACKNOWLEDGEMENTS

I would like to express my sincere gratitude to Dr. G. E. Ball, who

has guided this study, for his helpful suggestions and his valuable aid in

the preparation of this manuscript. I would also like to thank Dr. C. H.

Martin for his valuable suggestions and help in identification. I wish to

offer my sincere appreciation to the following entomologists and insti-

tutions, from which I have received loaned material: Dr. P. H. Arnaud,

Jr., California Academy of Sciences; Mr. Don Elliott, For est Entomology

and Pathology Laboratory, Calgary, Alberta; Dr. M. T. James, Wash-

ington State University; Mr. L.M. Kenakin, Department of Entomology,

University of Alberta; Dr. G. Steyskal, United States National Museum,

Washington, D.C.; Dr. J. R. Vockeroth, Canada Department of Agri-

culture, Ottawa, Ontario; Dr. W. W. Wirth, United States National

Museum; Dr. P. Wygodzinsky, American Museum of Natural History,

New York, N. Y. I express gratitude to the External Aid Office, Ottawa,

for support in this project through scholarship. Finally, I would like to

extend my gratitude to Mr . RobinE. Leech, with whom lengthy discus sions

were held, and from whom some ideas, directly or indirectly, have

been derived and applied herein.

66

Asilidae of Alberta

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Meigen, J. W. 1820. Systematische Beschreibung der bekannten euro-

paischen zweifliigeligen Insekten. Achen.Vol. 2 : 239-345, pi. 20, 21.

Melander, A. L. 1923a. The genus Cyrfopo^on (Diptera;Asilidae). Psyche,

Camb. 30 : 102-119.

Melander , A . L. 1923b. The genus Lasiopogon (Diptera; Asilidae) . Psyche^

Camb. 30 : 135-145.

Melander, A. L. 1923c. Studies in Asilidae (Diptera) . Psyche Camb

30 : 207-219.

Melin, D. 1923. Contribution to the knowledge of the biology, meta-

morphosis and distribution of the Swedish Asilids, in relation to

the whole family of Asilids. Academical Dissertation. Almqvist

Adisoemar to

69

& Wiksells Boktryckeri AB. Uppsala.

Moss, E. H. 1955. The vegetation of Alberta . Bot, Rev. 21 : 493-567.

Odynsky, W. 1962. Soil zones of Alberta, as published by Alberta soil

surveys. Map. Revised. Department of Extension, University of

Alberta, Edmonton, Alberta.

Osten- Sacken, C.R. Baron von. 1877. Western Diptera : Descriptions

of new genera and species of Diptera from the region west of the

Mississippi and especially from California. Bull. U. S. Geol.

Geogr. Sur. Terr. 3 : 189-354.

Poulton, E. B. 1906. Predaceous insects and their prey. Part 1. Trans,

ent. Soc. Lond. 1906 : 323-409.

Rondani, A.C. 1856. Dipter ologiae italicae prodromus 1 : 1-126.

Rondani, A.C. 1864. Dipterorum species et genera aliqua exotica revisa

et annotata, novis nonnullis descriptis. Arch. Zool. Anat. Fisiol.

3 : 1-99, 1 pi.

Say, T. 1823. Description of dipterous insects of the United States. J.

Acad. nat. Sci. Philadelphia 3 : 9-54, 7 3-104,

Say, T. 1824. In W. H. Keating: Narrative of an expedition to the sour ce

of the St. Peter's River, Lake Winnepeek, Lake of the Woods 1823

under the command of Major Long. (Say : 268-378).

Strickland, E. H. 1938. An annotated list of the Diptera (flies ) of Alberta.

Canad. J. Res. D 16 : 175-219.

Strickland, E. H. 1946. An annotated list of the Diptera (flies) of Alberta.

Additions and corrections. Canad. J. Res. D 24 : 157-173,

Tucker, E. S. 1907. Some results of desultory collecting of insects in

Kansas and Colorado. Kansas Univ. Sci. Bull. 4 : 51-112.

Walker, F, 1849. List of specimens of dipter ous insects in the collection

of the British Museum. Part 2 : 231-484.

Walker, F. 1851. Characters of undescribed insects in the collection

of William Wilson Saunders, Esq. Ins. saundersiana 1 : 76 - 156

2 pi.

Walker, F. 1851. Characters of undescribed insects in the collection

of W. W. Saunders. Trans, ent. Soc. Lond. ser. 2, 5, part 2:

268-334.

Walker, F. 1868. In J. K. Lord: The naturalist in Vancouver Island

and British Columbia.

Wiedemann, C.R.W. 1828. Aus ser eur op’aische z •./ eifliigelige Insekten,

als Fortsetzung des Meigenschen Werkes. 1 : 1-608, 7 pi.

Wilcox, J. 1941. New Heteropogon with a key to the species (Diptera,

Asilidae). Bull. Brooklyn ent. Soc. 36 : 50-56.

Wilcox, J. and C. H. Martin. 1936. A review of the genus Cyrtopogon

Loew in North America (Diptera-Asilidae) . Ent. Amer. 16 : 1-85,

5 pi.

Williston, S. W. 1884. On the North American Asilidae (Dasypogoninae,

Laphriinae), with a new genus of Syrphidae. Trans. Amer. ent.

Soc. 11 : 1-35, 2 pi.

Williston, S. W. 1885. On the North American Asilidae (Part II) . Trans.

Amer. ent. Soc. 12 : 53-76.

Williston, S. W. 1886. Dipter ological notes and descriptions. Trans.

Amer. ent. Soc. 13 : 287-307.

70

Asilidae of Alberta

Williston, S. W. 1893. Newor little known Diptera. Kans. Univ. Quart.

2 : 59-78.

Fig. 1; map of Alberta showing ecological regions (after Moss 1955 and Brooks 1958) and col-

lecting sites . 1, Prairie; 2, Boreal Forest; 3, Subalpine and Montane; 4, Parkland and Boreal-

Cordilleran.

Adisoemarto

71

Figs. 2-19 head; 2 Stichopogon argenteus Say, 3-4 S. trifasciatus Say, 5-7 Lasiopogon terricola Johnson,

8-9 h. cinereus Cole, 10 Ospriocerus abdominalis Say, 11 Stenopogon coyo(e Bromley, 12 S. obscmiventris

Loew, 13-15 S. inquinatus Loew, 16-17 Holopogon albipilosa Curran, 18-19 Cyrtopogon distinctitarsus new

species .

72

Asilidae of Alberta

Figs. Z0-Z8 head; ZO-ZZ Nicocles utahensis Melander, Z3-Z4 Pogonosoma stricklandi new

Z5 Laphria Scorpio McAtee, Z6 Proctacanthella cacopiloga Hine, Z7-Z8 A'erax bicaudatus Hine.

species,

Figs. Z9-37 mouthparts; Asilus calHdus Williston, 29 labrum epipharynx, ventral aspect, 30

cross-section of proboscis, 21-33 hypopharynx, 34 maxillary blade, 35 maxillae and labium,

ventral aspect, 36 palpus, 37 labium, dorsal aspect, 38 apical part of labium, Laphria janus McAtee,

39-40 cardostipites and palpi, 41 Leptogaster aridus Cole, lateral aspect.

47

Figs. 42-47 palpus; 42 Stenopogon obscvriventris Loew, 43 Eucyrtogpogon incompletus new species, 44

Nicocles ute/iensjs Melander , 45 Heteropogon wilcoxi James, 46 Leptogaster aridus Cole, 47 Pogonosoma stricklandi

new species.

Figs, 48-53 Asilus callidus Williston, 48-50 head, 51 thorax, lateral aspect, 42 same, dorsal

aspect, 53 prothorax, anterior aspect, 54 Stenopogon neglectus Bromley, prothorax, 55 same, dorsal

aspect, 56 Stichopogon trifasciatus Say, thorax, lateral aspect; Ba, basalare; cx, coxa; e, eye

em, epimeron; es, episternum; gb, gibbosity; gn, gena; h, humerus; hp, hypopleuron; ht,

haltere; msn, mesonotum; msp, mesopleuron; mtn, metanotum; mtp, metapleuron; pc,

posterior callus; pn, pronotum; pp, pteropleuron; ps, presternum; pst, prosternum; ptn,

postnotum; s, suture; Sa, subalare; sc, scutellum; sp, spiracle; spl, sternopleuron.

Adisoemarto

75

Fig, 57 Lasiopogon cinereus Cole, wing; A, anal vein; Ac, anal cell; Ax, axillary cell; B, basal

cell; Be, basal costal cell; C, costal vein; Cc, costal cell; Cu, cubital vein; D, discal cell;

hv, humeral vein; M, medial vein; m, medial crossvein; Me, marginal cell; m-cu, medio-

cubital crossvein; P, posterior cell; R, radial vein; r-m, anterior crossvein; Sc, subcostal

vein; See, subcostal cell; Sm, submarginal cell.

76

Asilidae of Alberta

Figs. 58-59 male genitalia; 58 Lestomyia sabulonum Osten Sacken; A, lateral aspect; B, dorsal

aspect; C, ventral aspect; D, aedeagus; E, proctiger; 59 Lasiopogon quadrivittatus Jones; ae,

aedeagus; cl, clasper; gp, gonopod; ha, hypandrium; hpr, hypandrial process; pgr, proctiger;

ss, superior forceps.

Figs. 60-63 proboscis; 60 Stenopogon inquinatus Loew, 61 Eucyrtopogon incompletus new species, 6Z

Pogonosoma stricklandi new species, 63 Laphria Xanthippe Williston; 64 Leptogaster aridus , thorax, lateral

aspect.

Adisoemarto

77

Figs. 65-66 prothorax, anterior aspect; 65 Stichopogon trifasdatus Say, 66 Lasiopogon cinereus Cole;

67-73 scutellum; 67 Lasiopogon trivittatus Melander, 68 L. ripicola Melander, 69 Proctacanthella cacopiloga

Hine, 70 N erax bicaudatus Hine, 71 Asilus paropus Walker, 72 dsi7us ca///dus Williston, 73 Negasilus belli

Curran; 74-88 leg; 74 Lasiopogon quadrivittatus Jones, 75 Nicocles utahensis Melander, front tibia and

tarsus, 76 same, hind tibia and tarsus, 77 Comantella fallei Back, front tibia and tar sus , 78 Cyrtopogon

auratus Cole, front tibia and tar sus, 79 Cyrtopogon auripilosus Wilcox and Mar tin, hind leg, 80 Cyrtopogon

willistoni Curran, middle tarsus, 81 Cyrtopogon //neo^arsus Curran, front tibia and tarsus, 82 Asilus

paropus Walker , 83 Asilus snowi Hine, front femur.

78

Asilidae of Alberta

Fig, 84 Holopogon albipilosus Curran, hind leg; 85 Leptogaster aridus Cole, same; 86 L. aridus Cole, last

tarsal segment; 87-105 abdomen; 87 Stichopogon fn/asc/a^us Say, ?, 88 Lasiopogon quadrhdttatus Jones,

$, 89-91 Stenopogon inquinatus Loew, eighth segment and ovipositor, 92 same, acanthophorite., 93

same, cf. 94-95 Nicocles utahensis Melander, d, 96 same, ?, 97 Cyrtopogon auratus Cole, d, dorsal, 98

same, lateral, 99 Laphria scorpio McAtee, d, dorsal, 100 same, lateral, 101, L. janus McAtee, ?,

102-103 Leptogaster aridus Cole, ?, 104-105 Asilus callidus Williston, ?,

Adisoemarto

79

114

Figs. 106-117 ovipositor; 106 Lasiopogon aldrichi Melander, specimen from Grant

107 same, specimen from Drumheller, Alberta, 108-119 Laphria Xanthippe Williston,

ridingsi Cresson, 112 Promachus c/lm/c//a^us Curran, 113 Proctacanthella cacopiloga Hine, 114

Hine, 115-116 Asilus ocddentalis Hine, d, 117 Asilus callidus Williston.

Co. , Oregon,

111 Pogonosoma

Nerax bicaudatus

8Q_ Asilidae of Alberta

Figs, 1 18- 140 antenna; 118 Lasiopogon c/nereus Cole, 119 L. prima newspecies, 120 L. a/cfnc/ii Melander,

specimen from Grant Co., Oregon, 121 same, specimen from Drumheller, Alberta, 122-123

Ospriocerus consanguineus Loew, 124-125 O. abdominal is Say, 126 Stenopogon inquinatus Loew, 127 Holopogon

a/6/p/7osus Cur ran, 128 Lestomyia sabulonum OstenSacken, 129 Nicocles utahensis Melander, 130 Heteropogon

wilcoxi James, 131 Eucyrtopogon mcompletus new species, 132 Pogonosoma stricklandi new species, 133- 134

Laphria xan^ft/ppe Willi ston, 1 3 5 same , fir s t two s egments , dor sal, 1 3 6 L. sedales Walker, 137 L. scorpio

McAtee, 138 L. aeatus Walker, 139 Leptogaster aridus Cole, 140 Proctacanthel la cacopiloga Hine ,

Adisoemarto

81

Figs, 141—147 anteiMlia.J 141 Messm bi^andatms Himej 14Z A“®’nisiBS Xiickcr, 143 Asilms pairopviis Walker,

144 Asilem etymocmernikm Hiee, 145 A. anehillis mew species, 146 A. §rammlis mew species, 147 Wsgasihms

betm Cuirram.

Figs, 148-157 wing; 148 Lasiopogon prima new species, 149 same, cf. 3rd longitudinal vein, 150

stenopogon coyote Bromley, 151 same, cf. first posterior cell, 152 Holopogon nigripilosa new species,

153 Heteropogon wilcoxi James, 154 Lestomyia sabulonum Osten Sacken, 155 Cyrtopogon bimacula Walker, ?,

156 C. distinctitarsus new species, ?, 157 C. dasyllis Williston, ?.

82

Asilidae of Alberta

Figs. 158-169 wing; 158 Eucyrtopogon comantis Curran, 159 E. diveTsipilosus Curran, 160 Pogonosoma

stricklandi new species, 161 Laphria Xanthippe Williston, 162 L. janus McAtee, 163 Leptogaster aridus Cole ,

164 Promachus dimidiatus Curran, 165 Nerax bicaudatus Hine, 166 N. canus Hine, 167 N. su6cupreus Schaeffer ,

168 N. costalis Williston, 169 Asilus nitidifacies Hine.

Adisoemarto

83

Figs. 170-193 male genitalia; 170-174 Lasiopogon terricola Johnson, 175-179 L. trivittatus Melander,

180 - 184 L. cinereus Cole, 185 - 189 L. prima new species 190 - 193 Ospriocerus consanguineus Loew.

84

Asilidae of Alberta

Figs, 194-207 male genitalia; 194-198 Stenopogon obscuriventris Loew, 199-203 S. rvfibarbis Bromley,

204-207 S; gratus Loew,

Adisoemar to

85

Figs. Z08-Z17 ijiale genitalia; Z08-Z1Z Stenopogon inquinatus 'Loew, Z13-Z17 S. neglectus Bromley,

86

Asilidae of Alberta

Figs. 218-Z37 male genitalia; 218-221 Wo/opogon a/6jpiZosa Curran, ZZZ-ZZ5 Heteropogon wilcoxi Ja.mes ,

226-229 Cyrtopogon auratus Cole, 230-233 C. montanus Williston, 234-237 C. dasyllis Williston.

Adisoemarto

87

Figs. 238-258 male genitalia; 238-242 Eucyrtopogon albibarbis Curran, 243-247 Comantella fallei Back,

248-252 Laphria Xanthippe Williston, 253-258 L. Scorpio McAtee; cl, clasper; pci, pseudoclasper .

88

Asilidae of Alberta

Figs. 259 Laphria aimatis McAtee, gonopod, 260 L. gilva Linnaeus, same, 261-263 ps eudoc lasper ,

261 L. janus McAtee, 262 L. vultur Osten Sacken, 263 L. sackeni Wilcox; 264-276 male genitalia;

264-267 Bombomima partitoT Banks, 268-270 fi. co/um6/ca_ Walker , 271-273 6. lernaldi Back, 274-276

B. posticata Say.

Adisoemarto

89

Figs. 277-Z90 male genitalia; 277-279 Proctacanthella cacopiloga Hine, 280-281 Nerax bicaudatus Hine,

282 Asilus delusvs Tucker, 283-284 A. occidentalis Hine, 284 gonopod and clasper, inner side, 285-

287 A. callidus Williston, 288-290 A. nitidifacies Hine.

90

Asilidae of Alberta

Figs. Z91-306 male genitalia; Z91-294 Asilus auriannulatus Hine, Z95-Z97 A. mesae Tucker, Z98-300

A. cumbipilosus new species, 301 - 303 A. aridalis new species, 304 - 306 A. gramalis new species.

91

Book Review

LACK, DAVID, 1966. Population studies on birds, v + 341 pp. 1 pi.,

31 figs. + 29 line drawings. Clarendon Press, Oxford. Price - 63 shil-

lings.

This represents a sequel to the author's 1954 work, "The Natural

Regulation of Animal Populations", now out of print. Thirteen longterm

studies of birds, with one exception all over four years duration, are

considered in detail and supported by eleven minor studies. Chapters

were sent to the principal authors concerned for criticism.

The main body of the work sets out these studies in detail and con-

clusions on bird populations are drawnfrom them. In the appendix, Lack

summarises chapter by chapter his 1954 work and points out various

errors and ideas which have not stood the test of time. He then goes on

to summarise his ideas on the natural regulation of animal populations

and to examine in some detail the theories of Andrewartha and Birch

(1954) and Wynne-Edwards (1962) and to explain why, in his opinion,

these do not fit the facts.

Lack's basic ideas, that animal populations are regulated by den-

sity dependent factors which have arisenasa result of natural selection,

are unchanged, and this book provides further evidence for these in res-

pect of birds. He admits that there is, at present, no conclusive evidence

for this, but points out that natural selection itself had to wait some 70

years for a satisfactory mathematical formulation (Fisher 1930) and until

1956 for a field proof (Kettlewell 1956).

Entomologists will find the evidence that birds have little influence

on insect populations in the breeding season but considerable effect on

overwintering ones, interesting.

Unlike most recent books on animal populations there are no for-

mulae representing hypothetical mathematical models, a point which

should please the less mathematically minded.

There are few errors, but Gough Island (p. 268) is in the South

Atlantic, not Pacific.

This book represents an important contribution to the literature on

animal populations and should be read by all interested in this field. All

ecologists should read the appendix.

The book is clearly printed on good quality paper and well bound.

Attractive line drawings by R obert Gillmor illustrate the principal species

referred to and some techniques.

Peter Graham

Re-printed from: Hooke^ R.^ 1665. Miarographia or some physiologioal descriptions

of minute bodies made by magnifying glasses idith observations and enquiries there-

on. Jo Martyn and Ja Allestry , London xxxiv + 246 pp.

Quaest

iones

entomologicae

A periodical record of entomological investigations,

published at the Department of Entomology, Uni-

versity of Alberta, Edmonton, Canoda.

VOLUME III

NUMBER 2

APRIL 1967

QUAESTIONES ENTOMOLOGICAE

A periodical record of entomological investigations, published at

the Department of Entomology, University of Alberta, Edmonton, Alberta.

Volume 3 Number 2 3 April 1967

CONTENT S

Book review 93

Frank - A serological method used in the investigation of

the predators of the pupal stage of the winter moth,

Operophtera brumata (L. ) (Hydr iomenidae) 95

Book review 106

Sarai - Effects of temperature and photoperiod on embryonic

diapause in Nemobius fasdatus (DeGeer) (Orthoptera,

Gryllidae) 107

Book review 136

Book Review

ELTON, CHARLES S. 1966. The pattern of animal communities, xx

+ 412 pp. Methuen, London. Price - 90 shillings.

Much of the controversy in ecology has stemmed from the practice

of generalising too far from inadequate field data. As O. W. Richards

has said (A. Rev. Ent. 6 : 147, 1961), "disagreement probably arises

from the absence of sufficient evidence of the right kind". It is welcorhe,

therefore, to see two books (see p. 136), from the same publisher, which

although rather different in outlook and purpose are concerned basically

with providing 'evidence of the right kind'.

Charles Elton, the founder of 20th century animal ecology, has for

the last 20 years been leading an 'ecological survey' of Wytham Woods,

Oxford University's field site in Berkshire, England. This book is an

attempt to communicate some of the results from this project and to

analyse the pattern that exists there. Consequently, the results are

strictly applicable only to this area, although the methods and major

generalizations should be useful in studying other areas, at least those

with similar climate and physiography. The basis for the survey is the

classification of habitats devised by Elton and R.S. Miller (J. Ecol. 42 :

460-496, 1954), which is modified only slightly in this book. This clas-

sification, depending on Habitat Systems , divided laterally into F ormation

Types and stratified into Vertical Layers, with a small series of Qual-

ifiers, will be well known to ecologists. After five chapters which des-

cribe the methods of ecological survey and the history and geography of

Wytham Hill, there are twelve chapter s each dealing with the communities

associated with certain subdivisions of the habitat classification. The

emphasis is on terrestrial communities, although one chapter deals with

the small water bodies of woodland, and is predominantly concerned with

Wytham, although there is one chapter on the terrestrial maritime zone.

94

A chapter on dispersal and invaders and one on the whole pattern com-

plete the main body of the book.

The well known pyramid of numbers is now matched by an inverse

pyramid of habitats. Herbivorous animals tend to show greater special-

ization of feeding habits than do the carnivores or the scavengers, which

have a greater habitat range. The result is the gradual dissolving of the

primary community pattern, and the realization that every community

unit is partly interlocked with others. This 'girder system', it is sug-

gested, has a stabilizing influence on communities, for even simple com-

munities can exist in a stable state. The distinction is made, however,

between instability and vulnerability. Relatively simple communities

may be stable but at the same time vulnerable to invasions from other

areas. Complexity, then, at theintra-and inter-community level, would

seem to carry with it some stabilizing property.

One might ask, what has been achieved that might be considered a

basic advance in ecological theory during the 20 years of this ecological

survey, or even in the 40 years since the publication of Elton's first

book on ecology? Unfortunately the answer seems to support those who

consider that this approach to ecology has, for the present at least,

reached a plateau of usefulness. A tremendous amount of information

has been collected and this is perhaps an advantage of the ecological sur-

vey approach, in that the collection of data is not biased by preconceived

ideas on what is or is not important. But the sorting and analysis of this

data must present enormous problems, especially by the hand- sorting

method used at the Bureau of Animal Populations and to someone with

less experience and insight than Dr. Elton. In order even to confirm the

suggestions that have beenmade, the ecological survey may well have to

turn to themethodsof those who "have embarked on various quantitative

investigations", but who appear not to be held in very high esteem by the

author of this book.

Ecology as a socially useful science - as work for entomologists,

conservationists, wildlife management, and fisheries - will find little

guidance from ecological survey. The methods are quite at odds with

the urgency of practical problems. Even the generalizations from the

Wytham Survey are of little help. This book finishes with one page on

the regulation of numbers in populations and two and one half pages on

conservation. The plea, put forward in Elton' s previous book. The Ecology

of Invasions (1958), for preserving in as rich a form as possible all the

communities that may, be interspersed among croplands is repeated in

these last two pages. While the charm of diversification is obvious, the

solution of pest problems ("invaders of unfamiliar kinds ") does not neces-

sarily follow, as pointed out by M. J. Way (J. appl. Ecol. 3(suppl) : 29-32,

1966).

On the other hand, this book is a joy to read - it is an important work

of English literature which is informative and inspirational. It is 'scien-

tific natural history', andas such should recommend its elf to bothlaymen

interested in natural history and to professional biologists, especially

those who need reminding that animals live outside! An important part

of the book is the 88 beautiful black and white photographs which will be

particularly appealing to those acquainted with the British countryside.

Gordon Pritchard

A SEROLOGICAL METHOD USED IN THE INVESTIGATION OF THE PREDATORS

OF THE PUPAL STAGE OF THE WINTER MOTH, operophtera brvmata

(L.) (HYDRIOMENIDAE)

J.H. FRANK Quaestiones entomologicae

Hope Department of Entomology 3 : 95 — 105 1967

University Museum, Oxford*

A method is described for producing antibodies to winter moth pupae antigens in experi-

mental rabbits. This method proved useful in estimating the extent of feeding by certain predatory

beetles on winter moth pupae when more direct methods were impracticable.

The populations of oak-defoliating Lepidoptera in a study area at

Wytham Wood, Berkshire, England have been under investigation for a

number of years by Varley and Gradwell (I960, 1963a, 1963b). One of

the most abundant of these species is the winter moth, which is in the

pupal stage under ground between the end of May and the end of November.

These pupae suffer a high mortality. Varley and Gradwell (1963a) sug-

gested that the principal causes of this mortality might be moles, mice,

and predatory beetles, particularly species of the carabid genera

Pterostichus and Abax and the staphylinid Philonthus decorus {Gr . ) , Laboratory

feeding tests (Frank 1967) showed that several species of Pterostichus j

Abax parallelepipedus (Pill, and Mitt. ) and Philonthus decorus wer e able to penet-

rate thecocoonand integument of the winter moth pupae and eat the con-

tents. In order to evaluate the number of pupae taken by each species of

predator a serological technique has beenused, based on that of Dempster

(I960) but with several differences.

This is not an exhaustive study of serological methods applied to

ecology, but indicates techniques which may be of use to those wishing

to study predator-prey relationships, and is a sequel to West (1950).

Crowle (1961) and Wieme (1965) are useful reference works.

Successive injections of a foreign protein into a mammalian blood

stream lead to the formation of antibody as described by Nossall (1964)

and Speirs (1964). These antibodies may be highly specific to individual

antigens. Usually a given antigen induces maximal antibody formation

when it is pure. If immunisation is done with a mixture of several anti-

gens it is less likely that an antibody will be formed against any given

antigen.

Experimental animals injected with winter moth proteins might

therefore be expected to produce antibodies specific to the proteins pre-

sent. Winter moth proteins derived from predator guts, if unaltered

by enzymatic action, should give an immune reaction with the antibody.

A white precipitate, suitable precautions having been taken, would be

proof of this reaction.

* Present address: Department of Entomology, University of Alberta, Edmonton, Alberta.

Serological Method

96

PREPARATION OF MATERIALS

The account given here relates the steps used in the preparation of

an antiserum in experimental rabbits to winter moth pupal antigen. The

extraction of predator gut contents and the testing of these against the

prepared antiserum are recounted. After the results of this experiment,

I have explained the reasons for some of the processes and mentioned

some of the alternative methods available.

Collection of Predator Meals

Overnight catches from pitfall traps in the study area were made

twice a week in the summer and autumn of 1964 and 1965. Possiblepre-

dators were removed from the traps and killed with ethyl acetate vapour .

Immediately upon returnto the laboratory the crops of these possible

predators were removed and smeared onto filter papers. The filter

papers were labelled with date, species and sex of the animals, and

stored in a desiccator over phosphorus pentoxide. It is stated that such

smears have been stored for over 2 years without deterioration (Hall

etal. 1953).

The Experimental Mammals

Three adult male rabbits were obtained as experimental animals,

and a series of injections given. Mature animals are usually better anti-

body producers than young ones. Prolonged immunisation should be

avoided because of increase in the chance of antibody formation against

minor impurities, and because of the risk of production of sera with less

specificity.

Preparation of Antigen

Winter moth larvae were allowed to pupate in peat in rearing cages

in the laboratory. Within a few weeks of pupation, healthy pupae were

removed from their cocoons and weighed.

One g of pupae at a time was crushed by pestle and mortar in 25 ml

of 0. 9% sodium chloride solution. The contents of the mortar were

washed into a flat-bottomed flask, and a few drops of M / 1000 potassium

cyanide were added to precipitate melanins (Dempster I960). The flask

was left for 24 hr in a refrigerator at 5 C. The contents of the flaskwere

then divided equally between 2 centrifuge tubes and centrifuged for 1 hr

at 2500 rpm, and approximately 1400 g. The clear liquid was then passed

through a Seitz EK sterilising filter pad and collected in a thick-walled

flask with a side arm to which a filter pump was fitted. The total volume

of saline used in crushing the pupae and washing the mortar, flat-bot-

tomed flask, and centrifuge tubes was 50 ml.

Prior to filtration the filter -pad holder, thick-walled flask, and a

freeze-drying flask had been sterilised by autoclaving. The liquid in

the thick-walled flask was freeze-dried using liquid air. The freeze-

dried material was stored in a sterile, stoppered tube in a refrigerator

until needed. When sufficient antigen had been freeze-dried it was pooled

in order to minimise variation and re- stored.

Care in the preparation of the antigen is neces sary to avoid denatur-

F rank

97

ing the proteins present. Normal methods of drying may cause a change

in the protein structure. Sterile equipment is necessary to prevent bac-

terial attack and accidental introduction of foreign protein.

Production of Antiserum

Approximately 0. 2 g of the freeze-dried antigen was reconstituted

with 10 ml of distilled water, and the pH was buffered at 6. 8 by the ad-

dition of disodium hydrogen phosphate and dihydrogen sodium phosphate.

To the solution was added an equal volume of a mixture of the emulsify-

ing agent Arlacel 'A' and a light oil, Bayol 'F‘, in the ratio 3 : 17. The

liquids were thoroughly emulsified by drawing into and expelling from a

fine glass pipette. The emulsion was divided into three portions, as

accurately as pos sible, and injected subcutaneously into the experimental

animals at four sites: over each scapula and over each flank. All the

equipment was autoclaved before use, including the hypodermic syringes

and '20 gauge' needles.

Anintervalof a week was left before the second series of injections,

whena bacto-adjuvant was used. This antigen had the same composition

as that previously injected but contained, in addition, a suspension of

killed, dried Mycobacterium butyricum , This enhances the immune response

(Freund et al. 1948), A third and a fourth series of injections was given

at intervals of one week and using incomplete adjuvant, i, e, without

Mycobacterium butyricum .

After a further week rabbit no, 1 was starved fora day in an attempt

to eliminate some lipid material from the blood stream, and was then

bled from the ear. The reason for attempting to get rid of dissolved

lipids in the blood is that these might separate on standing and thus ob-

scure the reaction, A small cut was made in the posterior edge of the

ear under sterile conditions and 5 ml of blood were run into a 'Lusteroid'

plastic centrifuge tube. Lusteroid is a non-wettable commercial plastic

to which blood will not readily adher e. The blood was allowed to clot at

room temperature for one hour, and then the serum was centrifuged for

one hour at 2500 rpm to precipitate fibrin and blood corpuscles. The

clear serum was then decanted into a stoppered plastic tube.

The following day all three rabbits were given a fifth series of in-

jections. On the sixth day after the injections all the rabbits were starved

for 24 hours, and on the seventh day all three were bled from the ear.

The serum was treated as previously, decanted into labelled plastic

tubes and stored in a refrigerator.

Further injections were givenat intervals of 6 weeks to keep up the

level of immunity, Exsanguination was done every 3 weeks because it

was found that the serum tended to deteriorate after being removed from

and replaced in the refrigerator several times.

METHODS OF TEST

Preliminary Test

A very small volume of the clear serum was allowed to run up short

lengths of glass capillary tubing of 1 mm internal diameter and about 3

98

Serological Method

cm long, and the lower ends of the tubes were pushed into plasticine. A

small quantity of freeze-dried antigen was reconstituted with distilled

water, taken up in a micropipette, and layered carefully on top of the

serum in the capillary tubes. The tops of the tubes were then sealed

with plasticine and the tubes were allowed to stand at room temperature.

In less than 10 min a white precipitate had formed at the serum antigen

interface. This was taken to indicate that the rabbit had formed anti-

bodies against the winter moth antigen.

The Precipitin Test

Gamma globulins

Various methods have been used by different workers to test the

immune reaction between antigen and antibody and some of these are

mentioned in the discussion. Basically, the dried predator gut smears

are reconstituted with distilled water and reacted with either whole anti-

serum or a gamma globulin extract of the serum. All immunoglobulins

are antibodies, but gamma globulins constitute more than 90% of immuno

globulins, at least in human serum (Wieme 1965), Because several

antibodies are likely to be produced to the several components of the

water soluble protein from winter moth pupae, it is most probable that

some of these will be gamma globulins. By usinga solution of the gam-

ma globulins alone from the immune serum, a clearer liquid is produced

in which immune reactions are less likely to be obscured.

Method used

The method used in the following experiments was modelled on that

used by R.A. Webb and G, P, Gladstone at the Sir William Dunn School

of Pathology, Oxford. An agar of the following composition was made

up: ‘Bacto Agar' 3,0 g, sodium chloride 1.7 g, barbitone 1.48 g, gly-

cine 4. 5 g, distilled water 180 ml. The chemicals were weighed out in-

to a large beaker, and the distilled water was added and stirred well.

Difficulty was experienced in getting the barbitone to dissolve, so the

beaker and contents were autoclaved for about 10 min, which proved ef-

fective in dissolving the barbitone. Twenty ml of a 10“"^ M solution of

merthiolate (thiomersal) were added as an antibiotic, and the agar was

then filtered through sterile cotton wool and stored in sterile screw-

topped jars kept in an oven at 50 C. It was found that the agar tended to

deteriorate if kept for more than 10 days.

An apparatus was made up consisting of a levelling table onto which

was fitted a perspex tray. This tray held eight glas s microscope slides

side by side, and eight holes were punched through the tray, one under

the position of each slide and slightly towards one end. The sides of the

tray were of such height that when the microscope slides were in position

1 mm of the rim was exposed, and thus a layer of agar of depth 1 mm

could be poured over the slides.

Two rosettes, each containing a central and six peripheral well-

cutters were fixed into another piece of perspex which was of such con-

struction that it would run along the longitudinal rims of the tray, and

could be pressed down over each slide to cut wells in the agar. The

peripheral cutters of each rosette were each at a distance of 7,5 mm

F rank

99

from the central one, and each cut a circular well of diameter 2 mm.

The agar remaining in the wells was sucked out by a glas s pipette attached

to a filter pump.

In operation the agar is poured over 8 slides of standard thickness,

allowed to solidify, and then the wells are cut and their centres sucked

out. Antibody is put into the central well of each rosette, and antigen

into some or all of the outer wells by means of glass micropipettes. The

tray of slides is allowed to stand in a water - saturated atmosphere for

the precipitates to develop. After this, cuts are made carefully through

the agar at the edges of each slide, and the slides are removed from the

tray. The slides and agar layers on them are placed inpetri dishes con-

taining 0. 9% sodium chloride solution for 24 hr. Staining is done for

about 30 sec in a 5% solution of amido black in 60% alcohol, and the stain

is differentiated for about 24 hr in a 2% solution of acetic acid in distilled

water. The slides are then washed in tap water to remove the acid. A

photographic record of the precipitate lines may be made at this stage,

and a visual assessment of the strength and position of the lines is made.

The agar is now dried in situ on the microscope slides in an oven at

50 C. Alternatively the layer of agar may be separated from the micro-

scope slide and dried on a lantern slide for direct projection, or dried

on a stiff, glossy card for filing. During the staining process the slides

must be handled carefully in order not to loosen or damage the agar.

There is some danger of splitting of the agar during the drying process.

Incorporation of glycerol into the acetic acid solutionused for destaining

will prevent this but makes the agar somewhat sticky. Some very faint

precipitin lines are liable to disappear during the drying process.

Reconstitution of predator meals and testing

Strips of filter paper carrying the predator meals were cut out and

put into 2 ml centrifuge tubes, and 1 ml of 0. 9% sodium chloride was

added. The tubes were allowed to stand for several hours at room tem-

perature and then placed in a refrigerator overnight to allow solution of

the material. Before testing they were allowed to stand at room tem-

perature for 30 min.

The peripheral wells of each rosette were filled with antigen by glass

micropipettes and the central wells were filled with antiserum by the

same method. The micropipettes and tubes were sterilised for re-use

by boiling in dilute hydrochloric acid and washing in distilled water.

Control tests

Reconstituted antigens as well as fresh material from winter moth

pupae reacted with the prepared antiserum at full strength. In addition,

fresh material from pupae of the species Erannis aurantiaria (Esper) and

E. defoliaria (Clerck) reacted with the prepared antiserum, but much less

strongly. Antiserum from rabbit 3 gave much the strongest reactions.

It was then neces sary to ascertain whether the antis erum would react

with protein recovered from the guts of predators, and to discover how

long it took for the enzymatic action in the (predator) guts to denature

the protein so that it would not give the precipitin response.

In order to test this, a number of Abax parallelepipedus which had been

100

Serological Method

starved for a week, were confined in plastic containers, each with a

winter moth pupa. Feeding was observed, and when the greater part of

the pupa had been destroyed, the remains of the pupa were removed from

the container and a note was made of the date and time. Two specimens

of Ahax were killed every 24 hr after feeding, and their guts were re-

moved, smeared onto filter paper, dried and stored. For testing, after

allowing time for the soluble gut contents to dissolve, drops of this liquid

were tested against winter moth pupal antiserum. It was found that the

guts of those Abax which had been killed 24 hr after feeding gave the

strongest precipitin reaction, and the reaction diminished rapidly with

time, so that it was only just discernible for those individuals killed 96

hr after feeding.

Specimens of Pterostichus melanarius (111,) gave results identical to those

obtained from Abax specimens and it was assumed that specimens of

Pterostichus madidus (F. ) did not differ in this respect.

Specimens of both Abax and Philonthus decorus fed on larvae and pupae

of Phormia regina (Meigen) gave no precipitin reaction with winter moth

pupal antiserum, showing both the dissimilarity of this dipterous and

winter moth protein, and that it is not merely the presence of food in the

predator gut which causes the reaction, but the presence of winter moth

protein which is of importance.

The precipitin reaction of winter moth antiserum with winter moth

antigen, the latter either fresh or freeze-dried and reconstituted, gave

2 strong lines of precipitate, and occasionally a third, weak one, prob-

ably indicating that antibody had been formed against more than one pro-

tein, This formation of more than one line may have been an artifact

(Crowle 1961) due to change in temperature during the reaction or to

high antibody concentration but, since it occurred many times during the

course of the experiments the former explanation seems likely. The

quantity of antibody formed against any one of these proteins would pro-

bably be less than if a single pure protein had been injected, but there

were obvious difficulties in attempting to separate the various winter

moth proteins and it was unnecessary to do this for the purposes of this

study. In actual tests of predator gut contents, of course all lines were

found to be weaker than when using pure antigen, it was often found that

there was one strong line and a second barely discernible.

The antigen prepared from pupae of Erannis species gave only one

line which was in the same position as the strongest line formed by win-

ter moth antigen, but weaker than this, when tested against winter moth

pupal antiserum. The apparent similarity of these proteins shown by the

similar position of the precipitin lines seems to stress the affinity of the

genera Operophtera and Erannis, It should be possible to distinguish rep-

resentatives of the two genera by the difference in pattern of the lines

in the precipitin reaction if their antigens were concentrated enough,

but the antigens recovered from predator guts were dilute, A large

amount of Erannis protein taken from a predator gut could be confused

with a much smaller amount of winter moth protein from a similar source.

In this experiment it was necessary to consider that a small proportion

of the positive reactions was due to genera related to Operophtera , The

smallness of this proportion was partly because of the great numerical

F rank

101

superiority of winter moth larvae over those of related genera in 1964

and 1965 at Wytham, and partly because Operophtcra antigens gave much

stronger reactions with Operophtcra antiserum than did those of related

genera. These latter reactions, in many cases, were probably not de-

tected.

A fact emer ging is that Philonthus decorus ^ Quedius lateralis (F. ) and Nebria

brcvicollis (F. ) fed on adult winter moth in November 1964, and subs equently

tested against winter moth pupal antiserum did not give any indication

of a precipitin reaction. Similarly, an extract of adult winter moth pre-

pared in December 1965 did not react with pupal antiserum, whilst fresh

pupal antigen prepared on dates up to the middle of November continued

to give a strong reaction. It must be concluded that there is a rapid

change in protein structure just prior to adult emergence, or that the

proteins to which antibodies are formed lie in the pupal integument. The

former seems to be the more likely because the antiserum was prepared

against the water - soluble proteins of the winter moth pupa, and the pro-

teins of the pupal wall must be highly insoluble.

The criticism that antibody may be formed against plant material

in the gut of the winter moth larva, and present in the pupa, and which

may also be present in the guts of suspected predators, thus giving an

immune response, can be dismissed. The winter moth larva evacuates

its gut before pupation (Gradwelb per sonal communication). If this were

not the case then various plant materials would have to be tested against

the pupal antiserum.

It was found on testing the predator gut contents with undiluted anti-

serum, that other components of the serum in addition to gamma glob-

ulins tended to be deposited around the antibody well, and these to some

extent obscured the precipitate lines. Because of this, a solution of

gamma globulins was prepared by adding a saturated solution of ammo-

nium sulphate to whole serum (giving a concentration of about 50%) to

precipitate the gamma globulins, which were then centrifuged down and

redissolved in 0. 9% sodium chloride. In all experiments whole serum

was put into the central well of one rosette of each slide, and gamma

globulins into the central well of the other rosette. This would indicate

whether there were any antibodies present which were not gamma glob-

ulins and it also helped in orientating the slide after the staining proces s.

A plan of the distribution of the reagents was drawnat the beginning

of each test, and the precipitin lines were drawn on this when they be-

came apparent. I found it preferable to use only three of the outer wells

of each rosette for the precipitin lines to be at their clearest.

Absorption techniques

An absorption technique was used by Dempster (I960) to make his

antiserum more specific to the protein of his prey species. He found

that the precipitin reaction of his prey species, Phytodecta olivacea Forst.

was about 400 times as strong as the reaction to the Phytodecta antiserum

of Coccinella septempunctata L. , which also occurred in his study area. He

therefor e added just sufficient Coccinella material to precipitate the anti-

bodies to it, and then used the serum for his precipitin reactions. A

slight excess of Coccinella would greatly reduce the s ensitivity of the ser-

102

Serological Method

um to Phytodecta .

This method would have presented problems in the present experi-

ment because of the several related genera pre sent. Adequate sensitivity

was obtained without it. After precipitation from the serum of antibody

material common to the winter moth and to the various species of Erannis

and other related genera present in the study area, the antibodies re-

maining in the serum would probably produce very weak reactions with

winter moth pupal antigens . There would be practical difficulties in per-

forming accurately a series of absorption reactions. The population of

Erannis in the Study area was much smaller than that of the winter moth

so that its effects could be largely ignored.

RESULTS

Of 122 Abax parallelepipedus taken during the time the winter moth was

in the pupal state during 1964 and 1965, 21 gave a positive precipitin

reaction, indicating that they had fed on winter moth pupae not more than

4 days before they were captur ed. Six of 226 Philonthus decorus taken under

similar conditions had fed not more than 3 days before they were cap-

tured, The period after which winter moth pupal remains were no longer

detectable in Philonthus decorus was found to be 3 days, a shorter time than

for the carabid species tested. This is probably correlated with the

fact that the food of Philonthus decorus is ingested in liquid form and is more

easily digested. Three of 18 Pterostichus madidus and 1 of 5 Pterostichus

melanarius had fed within 4 days on winter moth pupae. No other species

of beetle and no beetle larvae gave positive results, but a male and a

female of the vole Clethrionomys glareolus Schr. taken in May 1965 were both

shown to have fed on winter moth. Usingthese figures and the population

sizes of the beetles it was possible to estimate the number of winter moth

pupae taken in a unit area by each species of predator, the number of

winter moth pupae per unit area having already been estimated (Frank

1967).

DISCUSSION

The winter moth protein giving the precipitin reaction seems to be

a complex of several proteins because, in well-defined reactions, three

distinct lines of precipitate could be seen. There are techniques for the

separation of proteins in such systems. Moving boundary electrophoresis

and zone electrophoresis can be used only when fairly large quantities

of the reagents are available. They allow proteins to be distinguished

by the rate at which these migrate towards an electrode in a conducting

solution.

The method of immunoelectr ophor esis (Williams I960) separates

the precipitin lines of the various proteins present. An agar plate is

used with a central, longitudinal trough filled with antibody. The antigen

is placed in two wells, one on either side of the trough, and the various

proteins in the antigen diffus e in a longitudinal direction because of elec-

F rank

103

trodes placed at either end of the plate. The antibody diffuses outwards

towards the antigen, and precipitin arcs are formed in various positions

on the plate depending upon the proteins present. By comparing the posi-

tions of the arcs, it is possible to determine whether two antigens are of

the same composition. This method would be of value in comparing the

proteins present in closely-related lepidopterous pupae, such as those

of Erannis and Operophtera ^ in an initial study. It would be too time con-

suming to use in routine sampling such as was done by simple diffusion.

In the preparation of the antiserum the incorporation of the antigen

in a water-in-oil emulsion prepared from paraffin oil (e. g, Bayol 'F')

and Arlacel (mannide monooleate) enhances and sustains antibody for-

mation and the alteration of sensitisation may occur concomitantly, but

theyappear to be distinct (Freund et al. 1948). The immune response is

not potentiated when vegetable oil is substituted for paraffin oil, nor when

the antigen is administered in an oil-in-water emulsion,

Dempster ( I960) used 0, 4% potas sium alum to precipitate the soluble

proteins so that they should diffuse more slowly into the blood stream

of the rabbit and thus sustain antibody formation, a method described by

Proom (1943).

The factors in promoting and sustaining the immune response are

probably prolonged absorption as well as the protection of antigens against

destruction and elimination, favourable cellular reactions about the anti-

genic depots, and the production of multiple foci of antibody formation

in lymph nodes (Freund et al. 1948).

The test method used by Dempster ( I960) was to dr aw a small volume

of the smear extract into a capillary tube, followed by an equal volume

of the antiserum. Because of density differ ences an interface is produced

where a white precipitate of antibody combined with antigen forms and

is viewed by indirect light against a black background. Clarity of the

liquids is essential for satisfactory results. The method has several

disadvantages. The precipitate formed is not permanent, and for recor-

ding purposes it is necessary to photograph thecapillary tubes and their

contents. The layering of the antisera must be done with extreme care

to obtain sharp interfacial division. Jarring of the tubes should be a-

voided; very fine precipitates easily become diffuse and may be lost due

to carelessness.

Oudin used a similar method, but coated the interior of the tubes

with a layer of agar (Williams I960). A layer of precipitate forms in

the agar and is less ephemeral thanin the first method described. When

several antigen - antibody systems are present, several layers of pre-

cipitate form at different levels in the tube depending upon the rates at

which the antigens diffuse into the agar. The drawbacks are the difficul-

ties inherent in the layering of the liquids and in the placing of the agar

coating, and also in obtaining a permanent record.

Leone (1947) followed Boyden and Defalco (1943) in using a more

elaborate method which has some of the drawbacks of the method used

by Dempster (I960), but gives a quantitative measurement. He used the

Libby photr onr eflectometer to measure the turbidity of the precipitin

reaction (Libby 1938). Here it is necessary to dilute the antisera by an

appropriate factor because of the weakness of the insect antigens. A

104

Serological Method

disadvantage was that the exact protein concentration had to be known in

all antigen solutions in order for the reaction titres to be comparable.

The technique developed by Ouchterlony (Williams I960) allows anti-

gen and antibody to diffus e into an agar-filled petri dish which originally

contains neither reagent. Small wells are cut in the agar and a few drops

of the antigen and antibody solutions are placed in these. The antigen

and antibody diffuse outwards towards each other at rates in proportion

to their concentration and diffusion coefficients. Where the antigen en-

counters its specific antibody a line or precipitate forms. The lines for

different antigens are distinctly separated because of differences in dif-

fusion rate. This clear separation of lines makes it possible to distin-

guish more reactions by this technique than with the Oudin tubes.

Various modifications of this basic Ouchterlony concept have been

used. Wadsworth (1957) devised a micro-immunodiffusion technique ,

which was further modified by Crowle (1958), based on theuseof a tem-

plate for cutting wells in the agar. Feinberg (1964) fur ther modified this

by floating a pattern-perforated disc of thin, rigid, transparent plastic

onto molten agar, and then allowing the agar to solidify in contact with

the disc. The disc is not subsequently removed from the agar, and the

reagents are applied at the apertures of the disc where agar is exposed.

The difference between this method and that of Wadsworth and Crowle is

that no holes are cut in the agar, and the disc does not need to be of such

thickness as to supply wells to contain the reagents. Antigens and anti-

bodies lie on the exposed agar and will notunder-run the disc because of

the effective seal formed.

Another method uses a similar technique to that of Wadsworth and

Crowle, but with cellulose acetate instead of agar. The cellulose ace-

tate strips used can be handled with less risk of damage during the stain-

ing process than can agar (Johnson era/. 1964).

ACKNOWLEDGEMENTS

1 am indebted to Professor G.C. Varley for granting me facilities

for carrying out the study of which this was part. I thank Professor R.

A. Webb and Dr. G. P. Gladstone for their advice, and Dr. S. L. Sutton,

University of Leeds, for his collaboration. My gratitude is also due to

Mr. Peter Ford, Oxford University, for help in the inoculation of the

experimental rabbits, and to the National Environment Research Council

for financial support.

REFERENCES

Boyden, A. A, and DeFalco, R. 1943. Report on the use of the photron-

r eflectometer in serological comparisons. Physiol. Zool, 16 : 229.

Corcoran, A. C. (ed. ). 1952. Methods in medical resear ch. Year Book

Publishers, Chicago. Vol. 5.

Crowle, A. J. 1958. A simplified micro double-diffusion agar precipitin

technique. J. Lab. din. Med. 52 : 784.

F rank

105

Crowle, A. J. 1961. Immunodiffusion. Academic Press. 333 pp.

Dempster, J. P. I960. A quantitative study of the predators on the eggs

and larvae of the broom beetle, Phytodecta olivacea (Forster) using the

precipitin test. J. Anim. Ecol. 29 : 149-167.

Feinberg, J. G. 1964. A new device for immunopr ecipitation in agar

gels. Nature, Fond. 201(4919) : 631.

Frank, J, H. 1967. The insect predators of the pupal stage of the winter

moth. J. Anim. Ecol. (in press).

Freund, J, et al. 1948. Antibody formation and sensitisation with the

aid of adjuvants. J. Immun. 60 : 383.

Hall, R.R., Downe, A.E.R., Maclellan, C.R. and West, A.S. 1953.

Evaluation of insect predator-prey relationships by precipitin test

studies. Mosquito News 13 : 199-204.

Johnson, C. M. , Westwood, J.C.N. , and Beaulieu, M. 1964, A con-

tinuous-flow double diffusion technique with cellulose acetate. Na-

ture, Bond. 204(4965) : 1321-1322.

Leone, C.A. 1947, A serological study of some Orthoptera. Ann. ent,

Soc. Am. 40 : 417-433.

Libby, R . L. 1938. The photronreflectometer - an instrument for the

measurement of turbid systems, J. Immun. 34 : 71.

Nossall, G. J. V. 1964. How cells make antibodies. Scient. Am. 211 :

106-115.

Proom, H. 1943, The preparation of precipitating sera for the identi-

fication of animal species, J. Path, Bact, 55 : 419-426.

Speirs, R.S. 1964. How cells attack antigens . Scient. Am. 210 : 58-64.

Varley, G. G, and Gradwell, G.R. I960. Key factors in population stu-

dies, J, Anim. Ecol. 29 : 399-401.

Varley, G.C. and Gradwell, G.R. 1963a, Predatory insects as density

dependent mortality factor s . Proc. 16th Int. Congr. Zool. Washing-

ton 1 : 240.

Varley, G. G. and Gradwell, G.R. 1963b. The interpretation of insect

population changes. Proc. Geylon Assn, Adv, Sci. 1962, 18 : 142-

156.

Wadsworth, G. 1957. A slide microtechnique for the analysis of im-

mune precipitates in gel. Int. Archs. Allergy appl, Immun, 10 :

355.

West, A.S. 1950, The precipitin test as an entomological tool. Can,

Ent. 82 : 241-244.

Wieme, R. J. 1965. Agar gel electrophoresis. Elsevier, Amsterdam/

London/New York. 425 pp.

Williams, C.A. I960. Immunoelectrophoresis. Scient. Am. 202 : 130,

106

Book Review

EVANS, H. E, 1966. The comparative ethologyand evolution of the sand

wasps. 526 pp. Harvard University Press.

H. E. Evans is well known for his studies of various solitary wasps,

especially "sand wasps" (Nys soninae) . We find in this latter group a

particularly high proportion of very original, or specialized, or evolved

traits of behavior (some of which even suggest a comparison with social

or sub-social forms). We can mention for example: night clustering;

oviposition in an empty cell (instead of on a prey); special behavior re-

lated to the larva such as provisioning it day after day (instead of "mass

provisioning", prior to hatching of the larva); cell cleaning, etc. ; the

range of prey selection tends also to be wider than in many other groups

of Sphecids; some Microbembix accept all kinds of insects (occasionally

other arthropods, such as spiders, etc.) and are scavengers.

Thus the study of this group is particularly interesting perse and

also with respect to the evolution of some features in the entire group

of Sphecoidea (and perhaps even in some social or sub-social forms).

H. E. Evans has gather ed, discussed, compared, and evaluated car e-

fully the different elements of information concerning this group and at

times even other groups, when useful; he emphasizes also the need for

additional information and experimentation. The integration of these

various elements leads to a final general synthesis, and also to appealing

working hypotheses. The conclusions are established on a wide base in-

cluding for example morphology of adults, larvae and cocoons, fossils,

phylogenetic and zoogeographical considerations, and especially ecolo-

gical and ethological features . The illustrations and especially the photo-

graphs, are excellent and attractive.

This very good book is particularly interesting and useful for students

of the behavior of insects and the puzzling complexity of this at the level

of the higher Hymenoptera.

Andre L. Steiner

Department of Zoology

University of Alberta

Sarai

107

EFFECTS OF TEMPERATURE AND PHOTOPERIOD ON EMBRYONIC DIAPAUSE

IN NEMOBIUS FASCIATUS (DE GEER) (ORTHOPTERA, GRYLLIDAE)

DARSHAN SINGH SARAI Quaestiones entomologicae

Department of Agriculture 3 : 107 — 135 1967

Courthouse, Kelowna, B.C.

The effects of temperature and photoperiod on invoking and terminating embryonic

diapause of Nemobius fasciatus were studied in relation to the adaptation of this cricket to

the long winter of a continental climate at 53 40’ N.

N. fasciatus ;'s univoltine in Alberta. Eggs undergo facultative diapause at the

termination of anatrepsis, overwinter, and hatch in early July of the succeeding year. There

are six nymphal instars; adults appear early in August and survive until frosts occur in late

October. Diapause appears to be induced mainly by the wide diurnal temperature range in late

summer. Middle aged females deposit higher percentages of diapausing eggs than younger or

older females. A high incubation temperature (29 C) suppresses diapause. Diapause is terminatejd

by exposure of pre-diapausing or diapausing eggs to temperatures of -15 C before incubation.

Heavy mortality occurs, however, in freshly laid eggs when they are exposed to 0 C. A temper-

ature of 5 C halts development of post-diapause eggs but this is resumed within 24 hours at

an incubation temperature of 29 C. Exposure of any stage to constant or varying photoperiod

does not apparently influence the incidence of diapause.

A single generation per year and resistance to low temperatur e^ at

least in the resting stages, are commonly found in insects which inhabit

high latitudes. In these regions climatic variations which are responsible

for these adaptations are very wide. In the resting stage, developmental

proces ses and metabolism are very much retarded. This phenomenon of

physiological rest or diapause, is followed by an active period of the life

cycle at the onset of favourable conditions . Danilyevsky (1965) attributes

the univoltine life cycle to the short duration of favourable conditions.

Where favourable conditions continue longer, insects are mostly multi-

voltine. Diapause in univoltine insects is mostly obligatory and in mul-

tivoltine species facultative (Andrewartha and Birch 1954, Lees 1955,

Danilyevsky 1965). In the former, diapause is commonly invoked by in-

ternal physiological processes regardless of the external conditions, but

the life cycle as a whole remains under the influence of temperature which

determines the breaking of diapause and also the duration of development

of the active stages. In multivoltine insects facultative diapause is mainly

controlled by external factors, such as photoperiod, temperature, food,

and humidity (Lees 1955, Danilyevsky 1965).

It was once believed (Readio 1931, Cousin 1932) that diapause was

invoked by unfavourable conditions in most insects. But the principal

stimulus for the inception of diapause is photoperiod, although temper-

ature, water, and diet may be involved (Harvey 1962). Although photo-

period has no direct effect on the development of insects, it may, through

the neurosecretory system, induce diapause immediately prior to un-

favourable conditions. The discovery of the dependence of the inception

of diapause on photoperiod was mainly the work of Kogure (1933) who

studied bivoltine races of Bombyxmori L. in Japan. Recent works include

those of Danilyevsky in 1948, 1949a and 1951, (cited in Danilyevsky 1965),

Dickson (1949), Lees (1955), and Corbet (1956),

108

Diapause in Nemobius

The present study was undertaken to determine how Nemobius fasciatus

(DeGeer), the striped ground cricket, is adapted to inhabit one of the

coldest locations in its distribution. As this adaptation is mainly through

embryonic diapause, this study is concerned with the effect of environ-

mental factors on induction and termination of diapause. This species is

omnivorous, as are most crickets, feeding on decaying organic matter

and leaves of grasses. Diet was discarded as a possible environmental

factor affecting diapause because crickets are general feeders . Similarly

water was eliminated, because eggs remain in moist soil from the time

of oviposition to hatching. T emperatur e and photoperiod were, therefore,

studied in various combinations.

Crickets were collected in 1964 in hummocky pastures near Atim

Creek 21 miles west of Edmonton city centre (53°40‘N, 113°50‘W). In

1965, another meadow two and a half miles from Atim Creek was used

because the previous field was flooded. Crickets break their metathoracic

legs easily if netted and to collect them uninjured the mouth of a wide-

mouthed fruit jar was put in front of adults and last instar nymphs which

were induced to jump into it. I sometimes collected 80 crickets in an

hour. It was easy to locate males by their sound. Each male was com-

monly found with one or two females. Young nymphs were collected in

the field with an aspirator. Crickets brought from the field were kept

in the laboratory in battery jars with rabbit pellets for food (Ghouri and

McFarlane 1958), inverted bottles of water plugged with absorbent cotton

for water supply, and strips of paper towels to increase the surface area

for walking and for hiding. About 2 0 to 3 0 crickets in sex ratio of one

were kept in each jar. Fresh moist soil in plastic petri dishes was pro-

vided for oviposition each 24 hours. Eggs were sieved out underwater in

a tray, damaged and small ones were discarded, others were put in plas-

tic petri dishes on a filter paper covering a thick pad of wet absorbent

cotton. These petri dishes were kept covered except for a few minutes

every day for observation. Eggs are highly susceptible to desiccation.

The cotton remained moist for about three weeks at 29 and 24 C and for

two months at 0, 5, and 15 C. A few drops of tap water were added when

needed. Dishes were marked on the cover and on the under surface with

the number of the experiment, the number of eggs, and the date of ovi-

position.

Eggs were observed after 24 hours for hatching. Newly hatched

nymphs were removed from the dish with an aspirator and counted as

they were transferred to a fruit jar. Egg shells were removed and

counted to check this.

Eggs for laboratory experiments were deposited by the crickets which

were collected in the field mainly in the first half of September, when

ovipositing adults were abundant. Eggs laid within 24 hours of the col-

lection of the crickets were used. Unless otherwise stated the experi-

ments of 1964 were repeated under similar conditions in 1965, on com-

parable dates.

Hogan's method (1959) for the Australian Acheta commodus Walker was

used to determine the stage of embryonic development at which diapause

occurs. The eggs were soaked in water in a watch glass for half an hour .

The water was then replaced by a mixture of two parts of glacial acetic

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109

acid, two parts of chloroform, and one part of absolute alcohol, for 25

minutes at 34 C. Eggs were then transferred to a mixture of one part of

glycerol and one part of 70 per cent alcohol. The embryo was then clearly

observed under the microscope without cutting sections or dissecting

eggs.

The percentage hatching of eggs shown in the results includes only

those which hatched within 13 to 15 days at 2 9 C or within 2 0 to 22 days

at 24 C, for non-diapausing or post-diapausing eggs. Throughout this

study these were the periods required for development without diapause

at these incubation temperatures. Diapause in many eggs results in a

prolonged hatching period. A typical curve of percentage hatching of

total eggs versus incubation days at 29 C is illustrated in fig. 1. Unless

otherwise indicated the limits of variation of controlled temperatures

throughout this work were less than ± 1 C.

Fig. 1. Frequency distribution of hatching of eggs of N. fasciatus at an

incubation temperature of 29 C. N = 250.

The crickets from the vicinity of Edmonton were very kindly deter-

mined by Dr. R . D. Alexander to whom eight male and female specimens

with notes on habitat and a tape recording of the song were sent for this

purpose.

110

Diapause in Nemobius

DEVELOPMENT IN THE FIELD

N. fasciatus is recorded in wet localities over most of eastern North

America including Newfoundland and Florida. By putting together various

records of its distribution (Lugger 1897, Blatchley 1920, Fulton 1931,

Hebard 1930, 1936, Strohecker 1937, Ball et al. 1942, Alexander and

Thomas 1959j Vickery 1963) it is found that this species is distributed

from southern Canada to northern Mexico and east of the great basin in

the United States. Characteristically, it inhabits marsh borders and

other poorly drained grassy situations such as stream banks, meadows

and pastures. It is commonly associated with crab grass ( Digitaria sanguinalis

Scop. ) .

In the northern United States this cricket overwinters in the egg,

matures in July, and continues to sing until the last individuals are killed

by winter. In Ohio, the earliest recorded singing date is July 8 and the

latest November 14. In the University of Michigan Museum there are adult

specimens from central and northern localities of Michigan, the earliest

of which was collected on July 17, the latest on November 16. There are

also adult specimens which were taken in Florida every month of the

year (Alexander and Thomas 1959). N. fasciatus starts maturing before

the middle of June in the Piedmont area and coastal plains of North

Carolina and probably there are two to three generations per year at this

latitude (Fulton 1931). Near Edmonton in 1964 and 1965 the song of this

cricket was not heard before the first week of August or after the last

week of October,

Seasonal History at Edmonton

The seasonal history of N. fasciatus was studied inmeadows near Atim

Creek from the end of April to the end of October in 1964 and 1965. In

1964 eggs hatched from July 8 to 15, adults were first seen on July 30,

but there was no singing although the day was warm (22.7 C) and singing

starts a day or so after the last molt. A week later, the buzzy chirps

were very noticeable, Oviposition had started by August 10, andreached

a maximum between the last week in August and the middle of September

(fig. 2). There was heavy oviposition on warm and particularly on sunny

days, especially when this fine weather continued for two or more days.

Below 15 C there was little or no oviposition even in September. Fifth

and 6th instar nymphs were seen for the last time on October 10. Heavy

frosts in the third week of October killed most of the crickets and they

were heard last on October 24, a partly sunny day with afternoon temper-

ature 11.6 C.

In 1965 there were 7.48 inches of rainfall in June compared to 1. 04

inches in 1964 and 3, 15 inches normal for this month. Meadows near

Atim Creek were flooded from June 21 to August 10 and muddy until the

middle of September; plant growth reappeared by the end of the month.

Neither crab grass nor crickets were present. To test the viability of

eggs deposited here in 1964, soil samples were taken in October 1965

from sites where oviposition had been observed. No eggs sorted from

the mud by sieving under water hatched. Seasonal life history was ob-

served in the unflooded edge of a spruce grove to the north and in another

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111

meadow about 2, 5 miles away which dried early. It was similar to that

of 1964; singing was fir st heard on August 10, some adults were present

but 5th and 6th instar nymphs predominated. On September 24-25 0. 9

inches of snow fell and stayed fora day and a half. This snow and nightly

frosts from September 22 to 28 killed most of the crickets. Singing was

last heard on October 13, a sunny day with afternoon temperature 12.2

C. The adult population was maximum by the end of August in each year.

Fig. 2. Number of eggs deposited by N. fasciatus per week per female

caged outside, 1964.

Seasonal incidence of Diapause

Almost synchronous hatching of N. fasciatus in July, and overwintering

of the egg stage suggest a regulating mechanism by which the seasonal

rhythm of development is adapted to this climate. This mechanism could

be the changes in photoperiod or daily temperature during late summer.

An experiment was conducted to determine whether the changing photo-

period or temperature influenced the females to lay diapausing eggs.

Methods and materials

Adults were collected weekly from August 7 to October 10, 1964 and

kept in battery jars in screen cages of two cubic feet volume outside

under natural conditions of photoperiod and temperature. The locations

of the cages were selected to avoid artificial light and shadows from buil-

dings or trees . Wet soil in plastic petri dishes was supplied for oviposition

and renewed every 24 hours. Throughout the experiment eggs were sieved

out in water and incubated at 29 C inpetri dishes. In all 4,644 eggs were

112

Diapause in Nemobius

incubated at this temperature.

Three other batches of five hundred and fifty eggs each were incu-

bated at 29 C, 22 C, and 20 ±1.5 C respectively. Lots of about 100 were

started one week apart from August 16 to September 20. Temperatures

in the cages were recorded with thermographs. Data for Edmonton photo-

periods for the complete year were obtained from the Dominion Public

Weather Office, Edmonton.

Results

The percentage hatches (within 15 days) of eggs deposited during

succeeding weeks of the summer of 1964 and incubated at 29 Care shown

in Table 1. These results show no obvious correlation between the inci-

dence of diapause and any weather factor, except the difference between

day and night temperatures or amplitude of temperature variation. There

was over 50 per cent diapause whenever the amplitude of temperature

difference was greater than 16 C. Either the eggs within the female are

directly sensitive to changes in environmental temperature, or they are

influenced indirectly through some physiological mechanism in the female

that is affected by fluctuating temperatures. It is apparent from Table 1

too that in the fir St halves of August and October the number of diapausing

eggs is less thanin September. Thus a second factor contributing to the

percentage of diapausing eggs could be the age of females.

TABLE 1. The percentage hatch of eggs deposited in successive weeks

by N. fasciatus under natural conditions of temperature and photoperiod

and incubated at 29 C, 1964.

In the batches at different incubation temperatures the overall per-

centage hatch in 15 days at 29 C was 35.6. At 22 C no hatching occurred

within 25 days, 3. 5 per cent hatched in 60 days and 4. 3 per cent in 90

days. At 20 C there was only 3 per cent hatch in 90 days. Thus all the

eggs incubated at 22 C or 20 C diapaused. It can be concluded that con-

tinuous higher incubation temperatures suppress diapause. No significant

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113

differences due to starting dates were found.

Discussion

Differences in the dates of the first and last singing in Ohio^^ Michi-

gan, and Edmontonare probably due to the differencein temperatures at

these places. North Car olina is warmer than Ohio and Michigan, and two

to three generations per year are possible (Fulton 1931). In Florida the

temperature remains fairly high throughout the year and the variation in

the mean monthly temperatures for the years 1941-50 (World Weather

R ecords 1959) j is only between 11. 4 and 27. 7 C, which allows this cricket

to breed throughout the year . Fig. 3 shows a comparison of meanmonthly

temperatures at four localities with the type of life cycle at these places.

It appears that the number of generations per year at different latitudes

is in fact determined by the period of high temperature available.

Fig. 3. Meanmonthly temperatures (C) at different localities in relation

to voltinism in N. fasciatus. a Jacksonville, Florida - Multivoltine; A

Cincinatti, Ohio - Probably univoltine; • Detroit, Michigan - Probably

univoltine; x Edmonton, Alberta - Univoltine.

114

Diapause in Nemobius

Since the mean maximum temperature in Edmonton during August,

September, and October rarely goes above 22. 5 C there is no hatching of

eggs deposited earlier in the season which is in accordance with the

laboratory results (4. 3 per cent hatch in 90 days at 22 C). A few nymphs

which were seen in the late summer probably hatched late from the pre-

vious yearns eggs or were exceptionally slow in developing. This point

was cleared up during the study of the life history in the laboratory where

some nymphs developed exceptionally slowly and some died before reach-

ing maturity.

Gragg and Cole (1952) observed that there were veryhigh fluctuations

in the number of non-diapausing larvae in the blowfly, Lucilia sericata Meig.

throughout the summer months. They stated (p. 603) that there was no

obvious correlation between the fluctuations and any one weather factor.

The same appears to be true of N. fasdatus j so far as photoperiod, number

of eggs laid, or hours of bright sunshine are concerned. Fluctuations in

the number of diapausing eggs laid during succeeding summer months

were apparently due to the range of temperatures experienced by the fe-

males during the time of oviposition. Crickets which were exposed to a

range of temperatures favourable for embryogenesis (15. 5 to 27. 7 C),

or favourable for breaking diapause (0. 6 to 15. 5 C) laid less diapausing

eggs. A large number of diapausing eggs were laid by crickets exposed

to widely fluctuating temperatures (below 10 to above 20 C).

Many articles have been published which deal with the effect of sea-

sonal fluctuations in temperatures on the induction and termination of

diapause. It is generally accepted that diapause termination in most in-

sects needs low temperatures whereas embryogenesis in post-diapause

stages requires high temperatures to proceed. Ther e has been, however,

very little work done on the effects of daily fluctuations of temperature

on diapause. Kozhanchikov (1949) associated diapause in the Chinese

oak silkworm Antheraea pernyi Guer. with the effects of daily fluctuations

in temperature on the larvae. He found that all of the larvae transformed

into non-diapausing pupae when the daily temperature varied from 19 to

22 C at a mean rearing temperature of 20 C. However, with a range of

18 to 25 C at the same mean temperature 86 per cent of the larvae dia-

paused, and all the larvae diapaused when the daily fluctuations in tem-

perature were from 18 to 32 C, 15 to 30 C or 15 to 32 C. At a slightly

lower rearing temperature (17 C) all larvae underwent diapause when the

daily temperatures varied from 15. 2 to 25. 0 C, 14. 0 to 25. 0 C and 14. 0

to 30.0 C. Therefore, in Antheraea pernyi Guer. both the mean rearing

temperature and the range of fluctuation are important in invoking dia-

pause.

Browning (1952b) observed in Acheta commodus Walk. (= Gryllulus commodus )

that as the incubation temperature is raised, an increasing percentage of

the eggs develops without diapause. His observations are confirmed by

Hogan (1960a) and also apply to N. fasciatus .

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PRELIMINARY LABORATORY STUDY OF DEVELOPMENT

Effects of Photoperiod and Age on Diapause

Since the previous experiment did not show any relationship between

photoperiod and the incidence of diapause, another experiment was con-

ducted to verify those results by changing the photoperiod in the laboratory

while other conditions such as food, humidity, and temperature were

controlled. Another reason for this experiment was to determine whether

diapausing eggs are deposited in response to changing photoperiod ex-

perienced by adults or nymphs.

Methods and materials

Last instar nymphs of both sexes were collected on July 29, 1964

and kept in battery jars in four lots of 20. These jars were placed in a

growth chamber provided with fluorescent and incandescent lights oper-

ated by time switches, and a humidity control. Two groups of these were

exposed to changing photoperiod and the other two served as controls kept

in constant darkness. The photoperiod corresponding to September 1 in

Edmonton was fir st used in order to break the continuity of the Julyphoto-

period experienced in nature by the nymphs. A light intensity of 500 foot-

candles was recorded on the surface of the shelf holding the battery jars.

The photoperiod was changed to that of September 6 after five days and

similar changes continued until the photoperiod was that of October 28,

Temperature was maintained in the chamber at 28. 9 C for 12 hours per

day and at 1. 5 C for the remaining 12 hours. The relative humidity was

maintained at 68 percent. The control battery jar s were cover ed secure-

ly with a light proof cover made of plastic which was black on the inside

and white on the outside. The crickets matured from July 3 0 to August

3 and eggs were laid mainly from August 10 to September 23, They were

collected every day and incubated at 29 C.

Results

As shown in fig. 4 only those eggs which were deposited by crickets

when under 12 hr 36 min or 9 hr 49 min photoperiod per day, showed a

high percentage hatch (45.9 and 43. 3 per cent, respectively) whereas

only 15,6 to 29.2 per cent of the rest of the eggs hatched. When these

results are compared with those of field adults laying eggs under the

same photoperiods, there is no similarity. Changing photoperiod has

no apparent relation to the incidence of diapause. Furthermore, the per-

centage hatch of eggs laid by the illuminated crickets (23.2) is signifi-

cantly higher at the 1 per cent level than that of eggs laid by the control

ones (12. 9). This could result from the absorption of radiant heat by the

illuminated crickets. In the control groups the difference in percentage

hatch of eggs deposited by females of different ages was not significant

evenat the 5 per cent level. This suggests that light is necessary to the

full expression of the tendency of middle-aged females to lay diapausing

eggs.

100

15

25

20

SEPT

10

60-

\

\

30

12 i1 io 9

DAILY PHOTOPERIOD IN HOURS

Fig. 4. The percentage of diapausing eggs laid on succeeding dates by

crickets kept in continuous darkness (above#— -•) and under decreasing

photoperiod (belowO 0)at 28. 9 C for 12 hours and 1. 5 C for remaining

12 hours per day. Incubation temperature 29 C.

Nymphal Development

There is no information concerning the number or duration of nym-

phal instars in N. fasciatus under field conditions or in the laboratory,

Nymphal development was studied in the laboratory to permit recognition

of stages collected from the field and allow experiments in which different

instars were given photoperiod treatments.

Methods and materials

Eggs were collected from females kept in the laboratory. They were

maintained at 5 C for 3 0 days to break diapause and then incubated at

29 C or 24 C. Nymphs emerging from eggs on any one day were divided

into groups of either ten or two which were kept separately in rearing

jars. Forty-five jars were kept at each of the incubation temperatures.

Twenty of these contained 10 nymphs each, and the remaining 25 two

nymphs each. The jars were covered with fine insect screening and the

crickets were individually observed for moulting by removing paper strips

and vials every 24 hours from the first instar to the adult stage. For

the first three nymphal instar s, the light colour of newly moulted nymphs,

body size, and width of the head capsule between the eyes were the cri-

teria used to determine new moultings. Exuviae were of little value as

they were usually consumed by the nymphs. Nymphs older than the fourth

instar were examined for the length of wing pads and ovipositor. Measure-

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merits were made with an ocular micrometer.

Results

There are six nymphal instars. The mean duration of the nymphal

instars and the mean width of the head capsule between the compound

eyes are set out in Table 2. The first three nymphal instars are almost

of the same duration at each of the two temperatures, whereas, the fourth,

fifth and sixth instar s differ considerably. There is a difference of about

0, 1 mm in the width of the head capsule between each of the first three

instars. Only injured and dying nymphs were attacked and eaten by

healthy ones. No differences were observed between the two sizes of

groups .

TABLE 2. The duration of nymphal instars of fasciatus in days and the

width of the head in mm. Means ± standard deviations.

From field observation and these experiments this species is uni-

voltine in Edmonton. Non-diapausing eggs cannot hatch the same year,

firstly because cumulative temperature is insufficient and secondly be-

cause the big differences between day and night temperatures induce

diapause.

Submergence in Water

It was observed previously that two to three days exposure to muddy

soil was not detrimental to eggs, while 51 days was. The effect of sub-

mergence of eggs in water in the laboratory was, therefore, studied.

Methods and materials

Eggs deposited on September 27 and 28 by females collected from

Atim Cr eekmeadow in 1964 were used. Four hundred and fifty eggs were

kept at 5 C from the end of October 1964 to July 5, 1965 when the fol-

lowing experiment was started. Five groups each of 75 eggs were treated

by submerging them under tap water in bottles for 5, 10, 15, 20, and 25

days, A remaining group of 75 eggs was incubated in the normal manner

to serve as a control. Each bottle was provided with one inch of wet soil.

Eggs were placed a few millimeters below the surface of the soil. The

bottles were then filled with tap water gently without disturbing the eggs

and kept at 29 C. After treatment eggs were sieved out and incubated at

118

Diapause in Nemobius

29 C.

Results

Table 3 shows the percentage mortality and the minimum incubation

period after immersion. Development seems to continue at about its

normal rate at 29 C under water for eight to nine days and then cease.

There is a significant correlation at the one per cent probability level

between duration of submergence and percentage mortality. Submer-

gence of eggs in water is clearly detrimental after 5 days and about three-

quarters of the eggs in the field will be killed by 25 days of flooding.

Barber and Dicke (1939) have shown that pupae of Heliothis armigera Hubner

are killed by water; moreover, mortality increases with a rise of tem-

perature.

TABLE 3. The lethal effect of submergence in water for different periods

on eggs of N. fasciatus .

r (correlation coefficient) between period

of submergence and percentage mortality

5% probability level

1% probability level

p< 0. 01; p< 0.05)

0.

0. 88

0. 95

EFFECTS OF LOW TEMPERATURES IN THE LABORATORY

ON DIAPAUSE

While both temperature and photoperiod may invoke or terminate

diapause, Lees (1955, p. 53) stated that temperature is by far the most

important environmental agency controlling the termination of diapause.

There are many examples of insects from several orders in which dia-

pause is broken by exposure to low temperatures. It has been shown in

a number of species of Orthoptera (Parker 1930, Burdick 1937,

Andrewartha 1943, Church and Salt 1952, Browning 1952a, b, Hogan

1960a, b, Rakshpal 1962a, b, Masaki 1962) that exposure of diapausing

eggs to low temperatures for an optimum time leads to ready development

when they are incubated at an appropriately higher temperature.

The effects of continuous low temperatures for different periods and

of alternating low and high temperatures on diapausing and post-diapaus-

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119

ing eggs were investigated. In discus sing the results of these experiments

I have arbitrarily described hatching within three days of first hatch of

less than 30 per cent of eggs as indicating full diapause and above 80 per

cent as indicating no diapause. Hatching of between 3 0 and 80 per cent

within 3 days is described as partial breakage of diapause.

Effects of Low Temperature on Pre-diapause Eggs

Eggs laid in August and September undergo a longer period of high

temperatures before being chilled than those laid in October, It has

been shown in some orthopteran species (Parker 1930, Church and Salt

1952, Browning 1952a, b) that diapause is broken in eggs which are ex-

posed to low temperature before the diapause stage is reached. But until

recently the effect of low temperatures on eggs of different ages has

scarcely been explored, except for some work by Hogan (1960b) on Acheta

commodus Walk,, Rakshpal (1962b) on Gryllus pennsylvanicus Burm. andMasaki

(1962) on the Emma field cricket, GrylMus mitratus (Burm. ).

An experiment was carried out primarily to determine the relation-

ship between low temperatures and the breaking of diapause in eggs of

various ages. It was also possible to see if low temperatures had any

detrimental effect on these eggs. Rakshpal (1962b) observed very high

mortality when one day old eggs of Gryllus pennsylvanicus Burm, were ex-

posed to low temperature.

Methods and materials

About 80 crickets were collected from the field and kept in the lab-

oratory in four rearing jars. Fourteen hundred eggs laid within 24 hours

were collected on each of two successive days. Each group of fourteen

hundred eggs was divided into lots of 50, Two replicates each of 50 eggs

were moved after keeping at 22 C for 0, 1, 2, 3, 4, 5, 6, 7, 8, 9, 10,

11, 16, and 21 days (referred toas the initial incubation period) to either

0 C or 5 C for 30 days. Thereafter these eggs were incubated at 29 C.

Zero days initial incubation meant that eggs which were laid within 24

hours were exposed directly to low temperatures, whereas one day initial

incubation meant that eggs laid within 24 hour s were kept at 22 C for one

further day, and so on.

Results

Table 4 shows higher mortality (33 per cent) of eggs which were

given zero days initial incubation and exposed to 0 C than of those which

were kept at 5 C (10 percent). Mortality was similar for the rest of the

eggs. Eggs incubated initially for zero to three days did not take up

water during their exposure to low temperatures, but did so eventually

during final incubation at 29 C within seven days. Only some eggs which

were given an initial incubation of four to six days completed water up-

take before being exposed to lowtemperatures. After a seven day initial

incubation period all eggs were fully swollen with water . Hatching of eggs

without initial incubation and for both low temperature treatments started

on the 13th day and continued up to the 18th day of incubation. Maximum

hatching occurred, however, on the 14th day of incubation. All the hat-

ching of eggs which were incubated initially for 2 to 11 days, occurred

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Diapause in Nemobius

within a range of two to three days; for the longer initial incubation,

however, this range was extended to a period of six to seven days. The

period to maximum hatching decreased from 14 to 10 days as the initial

incubation increased from zero to seven days. It seems from this dif-

ference of four days in these incubation periods that eggs without low

temperature treatment developed at 22 C in seven days to the stage they

had reached at 29 C in four days. Thus the total incubation period at

29 C for all the post-diapausing eggs is almost always 13 to 15 days.

After more than seven days initial incubation maximum hatching took

place on the 10th day (sometimes on the 9th day) of final incubation. Eggs

which were incubated initially for 16 and 21 days showed a somewhat

lower total percentage hatch than those given 1 to 11 days initial incub-

ation. They also showed a wide range of hatching (six to seven days) and

hence a higher mean incubation period. This suggests that eggs which

were kept at 22 C for longer periods before chillingmight have under gone

diapausemore strongly than eggs which were exposed to low temperature

earlier. They, therefore, might require longer low temperature treat-

ment to break diapause. Analysis of variance showed that the variation

in per cent hatch among treatments is significant at the one per cent pro-

bability level due to the interaction of low temperatures and initial in-

cubation periods . Variationin percent hatch among treatments of initial

incubation periods, however, is significant only at the five per cent level.

Effects of Low Temperature on Diapausing Eggs

N. fasciatus is found in places where winter temperatures differ wide-

ly, it follows that this cricket possesses some mechanism by which dia-

pause is terminated by locally available low temperatures. Lees (1955)

found that diapause termination in insects from warmer localities does

not require such low temperatures as in insects inhabiting colder environ-

ments. Danilyevsky (1965) and Masaki (1961) consider that genetical

differentiation for terminating diapause by local low temperatures is

often involved in insects with a seasonal rhythm of development adapted

to different climatic areas. Bigelow (I960, 1962) reported variations

in adaptability to local low temperatures in local populations of field

crickets .

I was interested, therefore, in determining the range and periods of

low temperatures effective in breaking diapause.

Methods and materials

Eight hundred eggs, which had been laid over a 24 hour period during

September 1964 were divided into 16 groups of 50. All groups were in-

cubated initially at 22 C for 10 days to allow them to reach the diapause

stage, so that mortality would be lessened. Four groups each were then

kept at the following temperatures: 0 C, 5 C, IOC, and 15 C. After 15,

30, 45 and 60 days one group of eggs from each room was incubated at

29 c. The same procedure was repeated in 1965 under similar conditions.

Results

The percentage hatching in 11 days (the normal incubation period for

post-diapause eggs incubated previously for 10 days) is plotted against

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121

time of exposure to low temperatures in fig. 5. All four temperatures

broke diapause, but forty-five day exposures were needed before this

approached completeness. Frequency distributions of the percentage

hatching of eggs kept for 15, 30, 45, and 60 days at 5 C (fig. 6) show that

the range of hatching period decreases with increase in time of low tem-

perature treatment. It was found by analysis of variance that the highly

significant variation in hatching resulted from different durations of low

temperature treatment, whereas, variations due to different low tem-

peratures and due to the interaction of low temperatures with durations

are not significant.

TABLE 4. The effect of exposure to low temperature on mortality and

termination of diapause in eggs of N. fasciatus kept at 22 C for different

periods after being deposited. Incubation temperature was 29 C.

1

Some eggs hatched after very long periods.

*F' value for different initial incubation periods =

5% probability level =

1% probability level =

3. 13

2. 60

3. 96

'F’ value for interaction of low temperatures and

initial incubation periods

5% probability level

1% probability level

4^49,;o;c

2, 13

2. 93

122

Diapause in Nemobius

Fig. 5. The percentage hatch of diapause eggs of N. fasciatus after ex-

posure to different low temperatures for different periods, prior to

incubation at 29 C.

Effects of Low Temperature on Post-diapause Eggs

Post-diapause eggs under natural conditions sometimes experience

low temperatures during development in early summer. Rakshpal (1962b)

has shown that exposure of post-diapause eggs of Gryllus pennsylvanicus to

low temperature causes some mortality. The following experiment was

conducted primarily to determine whether low temperature causes any

mortality in post-diapause eggs, and secondly to determine how long it

takes for eggs which recover from this chilling to develop at incubation

temperature.

Methods and materials

Six hundred eggs were counted from eggs which had been deposited

within 24 hours. Two days later another similar sample of eggs was ob-

tained for replication. Each of these samples was divided into 12 groups

of 50. All the eggs were left at 22 C for 10 days and then transferred to

5 C for 3 0 days to terminate diapause, then brought to 29 C. Three groups

(from each replicate) were then incubated continuously at 29 C to serve

as controls, whereas, the other nine were moved to 5 C (second chilling)

three at a time, after 4, 6, and 8 days of incubation (referred to as first

post-diapause incubation). From each of these three groups one was

brought to 29 C for final incubation after 5, 10, and 15 days of second

chilling.

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123

Fig. 6. Frequency distribution of hatching of eggs of N- fasciatus exposed

to 5 C for 15, 30, 45, and 60 days and then incubated at 29 C.

Results

The percentage hatch at 29 C in all the treatments varied from 80 -

96. Hatch was greater in eggs which were given six days first post-dia-

pause incubation and five days second exposure to low temperature. The

control eggs started hatching on the ninth day of incubation and this was

largely complete by the 11th day (normal for eggs exposed to low tem-

perature after 10 days of pre-diapause incubation at 22 C), with maximum

hatching on the 10th day. Eggs which were incubated at 29 C for first

post-diapause incubation for 4, 6, and 8 days (regardless of duration of

second chilling) started hatching in their final incubation after 5, 3, and

1 days with maximum hatch on the 6th, 4th, and 2nd day, respectively.

124

Diapause in Nemobius

It is clear that all the groups of eggs have the same number of days of

total incubation period after the termination of diapause regardles s both

of the time at which they were exposed to low temperature for the second

time and of the duration of that exposure. Thus, once diapause is ter-

minated development follows a simple time-temperature pattern. Ex-

posure to low temperatures during development decreases the rate of

development only. It was also observed that after the second chilling

eggs recovered and resumed their normal development within 24 hours

at 29 C. Statistically there is no significant variation in percentage

hatch among all the treatments (including the control).

Effects of Alternating High and Low Temperatures

The objectives of this experiment were to determine whether the

diapause-inducing influence of widely fluctuating daily temperature ex-

perienced by the females as shown in the field, can be attributed to the

direct effect of these temperature fluctuations on the eggs.

Methods and materials

The high and low temperatures used in the experiment were 29 C

and 5 C, respectively. Five hundred eggs deposited on September 13

and 500 laid on September 16 were used. Each sample was divided into

five groups of 100 eggs. Four such groups from each replicate were

exposed to 2, 4, 8, and 12 day cycles of high and low temperatures over

a total period of 48 days. Each cycle was formed of two equal durations,

one at a low and the other at a high temperature. The two day cycle of

alternating high and low temperature treatment would mean one day at

high and one day at low temperature alternately for 48 days. The re-

maining fifth group of eggs was exposed continuously to low temperature

for 24 days to serve as a control. Twenty-four days was the time period

chosen because, for all the treatments the total number of days spent by

the eggs at low temperature was 24. After treatments eggs were incu-

bated continuously at 29 C.

Results

Table 5 shows that the per cent hatch at high temperature during the

treatments increased from 0 to 70 with the increase of 2 to 12 days in the

duration of cycle. Some hatching occurred during the treatments because

eggs remained at high temperature for a total of more than 15 days, the

normal period for incubation. Furthermore, eggs which were subjected

to two or four day cycles of high and low temperatures showed a very

low percentage hatch when incubated continuously at the completion of

treatments, that is, they were mostly in diapause. Cycles longer than

four days are not as effective in inducing diapause. An analysis of var-

iance revealed that variation in percentage hatch among different treat-

ments was highly significant.

Discussion

Diapause in N. fasdatus is broken by low temperature even in freshly

laid eggs. Lees (1955) mentioned that in many insects, particularly

those of Orthoptera, exposure to low temperature should synchronize

Sarai

125

with some definite stage in the morphological or physiological develop-

ment of the embryo to break diapause. He called this stage the period

of sensitivity. In N. fasciatus this period extends from zero to seven days

at 22 C. The stages of embryogenesis in this period are all the stages

up to the end of anatrepsis, which were observed by studying 40 eggs

after every 24 hours from a sample of 1, 000 kept at 22 C. It was found

from these observations that eggs developed readily without exposure to

low temperature up to the completion of anatrepsis (which comes mostly

after seven days of incubation at 22 C), after which development was

blocked in most of the eggs. Rakshpal (1962b) observed a similar period

of sensitivity in Gryllus pennsylvanicus Burm.

TABLE 5. The mean percentage hatching of eggs of N. fasciatus during and

at the end of alternating 29 C and 5 C temperature treatments for 48 days.

‘F’ value for treatments = 62. 96’!'*!'

5% probability level = 9. 28

1% probability level = 29. 46

It has been noted by Parker (1930) in Melanoplus mexicanus (Sauss.),

Church and Salt (1952) in Melanoplus bivittatus {Sa.y) ^ and Browning (1952a,

b) in Acheta commodus Walk., that eggs which have not reached the diapause

stage but which were exposed to an adequate low temperature for an

adequate period and then incubated, hatched without diapause. Browning

(1952b) found that Acheta commodus is unusual in this respect, because in

most species undergoing diapause low temperature is most effective in

terminating diapause after the insects have entered diapause.

Hogan (1960a) criticized the observations of Browning from his own

experiments on Acheta commodus but confirmed them later (1960b) by stat-

ing that exposure of pre-diapause eggs to a temperature of about 13 C

so weakens the tendency of eggs to enter diapause that it is readily aver-

ted when they are transferred to a suitable incubation temperature (p,

528). In fact, in his experiments from which he criticized Browning a

suitable incubation temperature was not provided for the eggs after low

temperature treatment.

Only a small proportion of the pre-diapause eggs o£ Gryllus pennsylvanicus

Burm. were rendered free from diapause by low temperature (5 - 7 C)

and there was such a high mortality that the eggs which were incubated

for one day only at 22 - 23 C before exposure to low temperature were

126

Diapause in Nemobius

completely killed by the latter (Rakshpal 1962b).

The observation that eggs of N. fasciatus have presumably entered

diapause more strongly when kept for longer periods at high temperature

before low temperature treatment agrees to some extent with that of

Browning (1952a) in Acheta commodus Walk, He has shown that a preliminary

high temperature treatment for more than two days caused the eggs of

Acheta commodus Walk, to enter diapause more firmly than if kept for a

little or no time at high temperature before low temperature treatment.

He believed that diapause in eggs given a lengthy initial incubation at

high temperatures was more intense and so required a longer period for

its breaking. Moroga (1951, cited in Lees 1955) has also observed in

Bombyx mori L. that if eggs were given an initial incubation period ranging

from 3 to 80 days, before 40 days of chilling at 5 C, the percentage of

non- diapau sing eggs dropped from 84 to 8.

Masaki (1962) exposed eggs of the Emma field cricket, Gryllulus mitratus

(Burm, ) to a high temperature (30 C) for different periods and then kept

them at a low temperature (20 C) to determine the effect of duration of

high temperature on diapause intensity. He found that eggs were most

responsive to the diapause -intensifying action by 9 or 14 day high tem-

perature treatment when they were in the late pre-diapause and early

diapause stages. His findings, however, were based on the mean incu-

bation period at 20 C after high temperature treatment. These mean

incubation periods ranged from 108,5 to 132.4 days, which shows that

these eggs were still diapausing.

Alternating low and high temperatures apparently cause eggs of

N. fasciatus to undergo diapause and this effect increases as the duration

of the cycle length decreases to 48 hours. This characteristic could be

of high survival value by enabling the eggs laid in the late summer to

undergo diapause because of low night temperatures and high day tem-

peratures. This diapause is eventually broken by the continuous cold of

winter. Rakshpal (1962a) has shown that changing high and low temper-

atures for 56 days, in eggs of Gryllus pennsylvanicus Burm, has the same

effect on breaking diapause as 60 days continuous 6 to 7 C exposure. In

other words, 52 days low temperature treatment (the total number of

days for which eggs remained at low temperature during the treatment)

in changing high and low temperatures have the same effect as 60 days

continuous chilling.

In N. fasciatus the total number of incubation days at high temperature

required in post-diapause eggs remains the same regardless of the time

when these eggs were subjected to low temperature again during their

incubationand the duration of that period of chilling. Thus post-diapause

eggs in early summer continue to develop whenever temperatures are

favourable. Low temperatures retard or prevent development and eggs

resume their normal development shortly after the termination of low

temperature periods. Rakshpal (1962b) has shown somewhat similar

results in Gryllus pennsylvanicus Burm, but post-diapause eggs of this species

took three days to recover from the second chilling.

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127

COMBINED EFFECTS OF TEMPERATURE AND

PHOTOPERIOD ON DIAPAUSE

There are few papers on the effects of photoperiod on diapause in

the Orthoptera. Masaki (1963) studied the effect of photoperiod on the

development of Nemobius yezoensis Shiraki, a univoltine Japanese cricket

overwintering as late instar nymphs. He found that nymphal diapause is

terminated by long photoperiod (16 hours per day), but is maintained for

a long time by a short one (12 hours). Other important work is that of

Norris (1959, 1965) on the red locust, Nomadacris septemfasdata (Serv, ),

Woodrow (1964) on Melanoplus devastator Scudder, and Halliburton and

Alexander (1964) on Chortophaga viridifasdata (DeGeer),

This study was undertaken with the hope of determining whether the

univoltine life history of N. fasdatus at Edmonton is due to a short summer

or is regulated by photoperiod causing the production of diapausing eggs.

Experiments were conducted to study the effect of photoperiod on the

breaking of diapause in eggs during low temperature treatment, and on

the various post-embryonic stages by recording the incidence of diapause

in the eggs laid by them. The criteria for diapausing and non-diapausing

eggs are the same as in the previous sections.

Influence of PhotoperiocS on Diapause during Chilling

Gayspitz (1953, cited in Danilyevsky 1965) found that diapause in the

larvae of the pine moth, Dendrolimus pini L. (Lepidoptera; Lasiocampidae)

under conditions of 12 hours daily light and a temperature of 20 C lasts

about a month, whereas at the same temperature with continuous illum-

ination activity begins after 17 days. Danilyevsky (1965) also mentioned

a similar but stronger effect of photoperiod on the diapausing larvae of

Arctia caia L. and Parasemia plantaginis L. (Lepidoptera: Arctiidae).

N. fasdatus deposits eggs in the soil at a depth ranging from three to

sevenmillimeters which could bean adaptation counteracting a diapause

inhibiting effect of light.

Preliminary experiment

Five hundred eggs laid within 24 hours in September 1964 were

divided into five groups of 100. Another sample of 500 eggs was used

in 1965 for replication. All these eggs were left at 22 C in the dark for

10 days to allow them to reach the diapause stage. Thereafter one group

from each sample was kept under 8, 12, 16, and 24 hours daily light at

5 C in light-proof compartments for 30 days. The fifth group was kept

in total darkness to serve as a control. The light was provided by a 16

inch 15 w fluorescent "cool white" lamp operated by a time switch. The

light intensity was 85 - 90 foot-candles on the shelf holding the petri

dishes containing the eggs. All the eggs were incubated at 29 C after

treatment.

Results - The eggs in all the treatments were virtually free from

diapause. Although the difference in percentage hatch among the treat-

ments was not significant, eight hours daily photoperiod showed slightly

greater hatch (93 per cent) than the rest of the photoperiods (average

128

Diapause in Nemobius

87, 5 per cent). Thus it appears that low temperature alone is sufficient

to break diapause in this species.

The Effect of Incubation Temperatures

It was mentioned previously that the percentage hatch of eggs of

N. fasciatus is greater at higher incubation temperatures than at lower

ones, when they are incubated without any low temperature treatment to

break diapause. Moreover, eggs remain at low temperatures for six

months at Edmonton. The effect of very long exposure of these eggs to

low temperatures and their incubation at various temperatures to deter-

mine the degree to which diapause is terminated was therefore studied.

Secondly, as pointed out in the results of the previous experiment, the

effect of eight hours daily photoperiod during chilling needed further

clarification. This experiment was designed to determine the effects of

the above mentioned factors and their interactions.

Methods and materials _ In this experiment two replicates each of 2,475

eggs were used. For this purpose, 300 crickets were collected on Sep-

tember 10, 1964 and kept in 10 battery jars in the laboratory. Eggs for

one replicate were collected from those deposited on September 11,

These eggs were divided into 75 groups of 33. They were kept in the

dark at 22 C for 10 days and thereafter moved to 5 C. At this temperature

15 such groups were subjected to each of 0, 8, 12, 16, and 24 hours

daily light. Three groups were removed each time from every sample

of 15 groups after 6, 12, 24, 48, and 96 days. One group from each of

these three was incubated at 29 C, 24 C, and 20 C. This experiment was

repeated on the same dates in 1965 and under the same conditions to re-

plicate for analysis of variance. Unfortunately the temperature in the

cabinet set for 20 C started fluctuating so widely when the experiment

was already in progress during both years that the data for 20 C had to

be excluded from the statistical analysis. An analysis of variance for a

split plot design was used because eggs were subjected to different in-

cubation temperatures in different cabinets.

Those eggs which hatched within 16 days of incubation at 24 C were

taken as non-diapausing since 14 to 16 days is the average incubation

period at 24 C for post-diapause eggs which have been subjected to low

temperature after 10 days of an initial incubation at 22 C temperature.

Results - Table 6 shows that the per cent hatch for different photo-

periods at each of the two incubation temperatures for the same durations

of chilling are not significantly differ ent. Secondly, hatch increases with

the increase in duration of low temperature treatment at both the incu-

bation temperatures. At 29 C however, there is rapid increase in per-

centage hatch up to 24 days of chilling when eggs are virtually free of

diapause. For chilling durations longer than 24 days the percentage hatch

at this temperature does notvary much. On the other hand incubation at

24 C shows very little breaking of diapause by treatments up to 24 days

at low temperature. Diapause is terminated at an incubation temperature

of 24 C to a great extent when eggs had been chilled-for 48 days. Ninety-

six days chilling enabled almost all the eggs to develop readily at this

Sarai

129

temperature. This means that the lower the incubation temperature the

longer the low temperature treatment required to allow the eggs to develop

readily.

TABLE 6. The mean percentage hatch of eggs after exposure to different

photoperiods at 5 C for different periods and then incubated at 24 or 29 C.

The observations from the previous experiment that photoperiod

during chilling does not affect termination of diapause are confirmed from

this experiment, because statistically there is no significant difference

among hatching of eggs given different photoperiods . The only significant

variations in hatching among the different treatments are those which are

due to different durations at low temperature, and due to different in-

cubation temperatures. Both these variations are highly significant. It

was found that development in eggs proceeded normallyat 20 ± 3 C when

chilled for 96 days. It is possible, therefore, that eggs after such a long

chilling are capable of developing promptly at lower incubation temper-

atures. This confirms that no influence of photoperiod on diapause or

development can beshownand that eggs are rendered free from diapause

by low temperature alone. Secondly, eggs need longer exposure to low

temperature for termination of diapause if they are to develop subsequen-

tly at lower incubation temperatures.

The Influence of Light on Nymphs

There is much evidence showing the influence of photoperiod oncer-

tain stages in the life history of different insects affecting diapause either

in the succeeding stages or in the next generation. Dickson (1949) ob-

served in GraphoUtka molesta (Busck. ) in southern California that a substan-

tial proportion of the larvae entered diapause in the early autumn when

temperatures were still favourable for development. He found thatdia-

130

Diapause in Nemobius

pause in these larvae had already been determined by the photoperiod

experienced during the early larval instars.

Danilyevsky (1948, cited in Danilyevsky 1965) found that if Acronycta

ruwicis L. (Lepidoptera: Noctuidae) is exposed during the larval feeding

period to a short day-length (6 - 15 hours) virtually every individual en-

ters diapause in the pupal stage, whereas the incidence of diapause fell

to zero when the larvae experienced 17 hours daily photoperiod.

Kogure (1933) showed in Bombyxmori L. that light and temperature

acting on eggs and early larval instars induced a particular pattern of

diapause which appearedin the eggs laid by the adults arising from these

stages.

In all of the previous experiments on the influence of photoperiod on

diapause in A^. fasciatus only the egg and adult stages were considered. An

experiment was planned to determine whether photoperiod acting on the

nymphs has any effect on the proportion of diapausing eggs deposited by

the females arising from these nymphs.

Methods and materials

Newly hatched nymphs for this experiment came from eggs which

were deposited in September 1965 held after diapause at 24 C. They

were divided into 85 groups of ten, within 24 hours after hatching. Each

group was raised to maturity on rabbit pellets in a rearing jar. Five

such jars were exposed to 12 hours or 16 hours daily light in two growth

chambers for each of the nymphal instars and the adult stage. The light

intensity on the surface holding the jar s was approximately 500 foot-can-

dles and the temperature was kept at 24 C. Since the mean durations of

the six instars at 24 C are 5. 8, 5. 6, 4. 7, 6. 3, 8. 1 and 9. 8 days, res-

pectively, the first instar was exposed for six days only, the second for

six days only and so on to either 12 or 16 hours daily light. Five jars

were subjected to each of 0, 12, and 16 hours daily light for all the post

embryonic stages, to serve as controls. All the jar s were observed once

a week to change the water vial, jars on a dark regime under a 40 watt

red light. High mortality during handling resulted from this technique.

After reaching the adult stage all the five samples of each treatment

were transferred to a battery jar for convenience in getting eggs. For

oviposition wet soil was provided in petri dishes in the usual manner on

every alternate day and sometimes after every fourth day to minimize

the exposure of adults to red light. Eggs thus obtained were sieved in

water and incubated directly at 29 C.

Results

Table 7 shows the sex ratio in each treatment and the percentage

hatch of the eggs at 29 C laid by the adults resulting from each treat-

ment. It is apparent that neither of the photoperiods influences the laying

of diapausing eggs since the percentage hatch of the eggs laid by the con-

trol crickets reared in darkness throughout the post-embryonic part of

the life history is not much different from the other treatments. Only a

small proportion of the 50 nymphs in each treatment reached maturity,

which decreased the precision of the results.

Sarai

131

TABLE 7. The influence of photoperiod on different post-embryonic

stages expressed as the percentage hatching of eggs laid by the resulting

females. Incubation temperature was 29 C.

Control crickets reared in total darkness yielded 7 males and 5 females

and 25% of their eggs hatched.

Discussion

Marcovitch (1923, cited in Danilyevsky 1965) working with aphids

was probably the first to discover the effect of photoperiod on the life

cycle of an insect. It is only during the last few years that the subject

of photoperiodism has attracted close attention. In most of the insects

worked on so far , diapause has been found to be influenced by photoperiod.

But Danilyevsky and Gayspitz (1948, cited in Lees 1955) found that dia-

pause in P/ja/era 6ucep/ia/a (L, ) and Spilosoma menthastri Esp, (Lepidoptera) was

unaffected by photoperiod. Dickson (1949) also observed no effect of

photoperiod on adult diapause in Listroderes obliquus Klug. (Coleoptera) when

larvae were grown under conditions of 9, 15, and 24 hours daily light.

He also found Lucilia sericata Meig. (Diptera) independent of photoperiod for

its diapause. Egg diapause in N. fasciatus also is unaffected by photoperiod

in any stage.

Browning (1952b) observed that inalso Acheta commodus Walk, that the

incubation temperature influenced the termination of diapause after low

temperature treatment. Exposure for 30 days to 10,3 C gave 84 per

cent hatch at 29. 9 C, whereas following the same low temperature treat-

ment only 64 per cent hatched at 26. 5 C and none at 20. 9 C,

The observation that N. fasciatus is neutral to photoperiod, however,

should not be taken as valid for this species in general, because only the

Edmonton strain is involved. For the generalization of these observa-

tions, experiments with strains from other parts of its range will be

required.

132

Diapause in Nemobius

GENERAL DISCUSSION

N. fasciatus, like many other Orthoptera, undergoes diapause at the

close of anatrepsis. Wide variations were observed in the proportion

of diapausing eggs among eggs laid during succeeding summer weeks in

association with wide fluctuations in the daily range of temperature. The

diapause-inducing effect of alternating high and low temperatures on the

eggs in the laboratory suggests that this association is causal, and that

the eggs during thepre-and post- oviposition period are affected directly

by external temperatures. To resume development diapause eggs of

N. fasciatus require to be subjected to temperatures below IOC for periods

up to 96 days followed by temperatures above 20 C. The induction of

diapause by fluctuating temperatures enables most eggs to enter diapause

by the end of summer. Untimely hatching of eggs in the same summer

is thus greatly reduced, an adaptation of high survival value. Lees (1955,

p. 29) states that as a general rule high temperatures tend to avert dia-

pause while low temperatures favour arrest of development. This holds

good in N. fasciatus in which a high percentage hatch would not normally

occur in the latitude of Edmonton because temperatures in late summer

rarely rise above 25 C. None of these processes show evidence of being

affected by photoperiod or changes in photoperiod.

A second probable factor contributing to variations in the proportion

of diapausing eggs, is the age of the adult female at the time of ovi-

position. It has been observed in Bombyx mori L. (Kogure 1933), p/j/e6ofomus

papatasii Scop. (R oubaud 1935), Locustana pardalina Walk. (Matthee 1951),

Gryllus pennsylvanicus Burm. (Rakshpal 1962a) that the physiological condition

of the female at the time of oviposition affects the presence or absence

of diapause in the next generation. The incidence of diapause in the eggs

seems to follow a normal curve of frequency distribution through adult

life, but this is partially suppressed when the insects are kept in con-

tinuous darkness.

The only difference observed between diapausing and post-diapausing

eggs of N. fasciatus was that the latter were clearer in their consistency.

It is a general belief that during diapause some physiological changes

occur which ultimately result in the resumption of active development.

The difference in the consistency of diapausing and non-diapausing eggs

in N. fasciatus is probably due to some such physiological changes in the

egg during low temperature treatment. Zolotarev (1947 and 1950, cited

in Danilyevsky 1965) calls these changes the diapause processes.

Andrewartha (1952) introduced the term "diapause development" for the

same thing. His term has since been used by other authors (Browning

1952a, b, Lees 1955, Masaki 1962, Beck and Alexander 1964a, b).

Danilyevsky prefers the term "reactivation" for the same phenomenon,

becausehe thinks that the term development is commonly linked with pro-

gressive growth and differentiation. Probably in these physiological

changes different processes are involved in different insects (Harvey

1962).

It seems that N. fasciatus does not need an adaptive mechanism such

as a very intense obligatory diapause or diapause decided by the photo-

period to survive in this part of the world. Diapause is facultative, be-

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133

cause a certain percentage of the eggs always hatch providing the in-

cubation temperature is above 26 C. Changes in temperature are the

main influence determining the presence or absence of diapause in the

eggs. Under natural conditions, however, univoltinism is maintained,

firstly mainly by the induction of diapause through alternations of quite

low temperatures at nights with fairly high temperatures during days

for most of the oviposition period, and secondly by temperatures in Sep-

tember and October too low for the completion of development before

winter. Thus even those eggs which need only a short period of low tem-

perature for breaking diapause do not develop until June and July of the

following summer when temperatures are high enough for a long enough

time to complete development. By that time diapause will be broken in

all the eggs. There is, therefore, almost synchronous hatching in the

first half of July in nature.

ACKNOWLEDGEMENTS

1 wish to thank most sincerely Dr. W. G. Evans, my supervisor, for

his many valuable suggestions and for his readiness to discuss any prob-

lem arising during the course of this study; Dr. G. E. Ball for his help

in many ways, 1 wish to thank Dr. D. A. Boag, member of my commit-

tee, for many useful suggestions.

1 am also grateful to Miss J.G. Shore, and Miss A, Zalums for their

cooperation and help, and Mr. H. Welling, supervisor of greenhouses,

for providing space in the low temperature rooms.

I wish also to express my gratitude to Dr. B. Hocking forgiving me

the opportunity to work in Canada and to the University of Alberta for an

intersession bursary during the summer of 1965, enabling this work to

be carried on without interruption.

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Andrewartha, H. G. 1952, Diapause in relation to ecology of insects.

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Diapause in Nemobius

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developmental time of the embryos of the grasshopper, Melanoplus

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Church, N.S. and R. W. Salt. 1952, Some effect of temperature on

development and diapause in eggs of Melanoplus bivittatus , Can. J. Zool,

30 : 173-184.

Corbet, P.S. 1956. Environmental factor s influencing the induction and

termination of diapause in the emperor dragonfly, Anax imperator

(Odonata: Aeshnidae). J. exp, Biol, 33 : 1-14.

Cousin, G. 1932, Etude experimentale de la diapause des insectes.

Bull, biol, Fr. Belg. , Paris Suppl. 15 : 1-341,

Cragg, J. B. and P. Cole. 1952. Diapause in Lucilia sericata , J. exp,

Biol. 29 : 600-604.

Danilevskii (= Danilyevsky), A, S, 1965, Photoperiodism and seasonal

development of insects. Oliver and Boyd, Edinburgh and London,

ix + 283 pp,

Dickson, R, C. 1949. Factors governing the induction of diapause in the

oriental fruit moth, Ann. ent. Soc, Amer, 42 : 511-537.

Fulton, B. B, 1931. A study of the genus Nemobius (Orthoptera : Gryllidae),

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Harvey, W. R . 1962. Metabolic aspects of insect diapause. A. Rev,

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Hebard, Morgan. 1930. Orthoptera of Alberta, Proc. Acad, nat, Sci,

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Hebard, Morgan, 1936. Orthoptera of North Dakota. N. Dak. agric.

Sarai

135

Exp, SubstnRep, 284.

Hogan, T. W. 1959. A rapid method of examining diapause of eggs of

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Kogure, M. 1933. The infleunce of light and temperature on certain

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Lees, A.D. 1955, The physiology of diapause in arthropods. Cam-

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16-26.

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Norris, M. J. 1959. The influence of day length on imaginal diapause

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appl. 2 : 154-168.

Norris, M. J. 1965. The influence of constant and changing photoperiods

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J. Insect Physiol. 11 : 1105-1119.

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136

Book Review

SOUTHWOOD, T.R.E. 1966. Ecological methods, xviii + 391 pp. Methuen, Lon-

don. Price - 75 shillings.

Southwood, in his preface, points out the distinction between 'methods for ecol-

ogists' and 'ecological methods ' , which are "those concerning ... the measurement,

description and analysis of both the population and the community". This means

that measurement of environmental factors is not covered by this book, but this is

well documented in other compilations. "Emphasis is placed on those (methods) most

relevant to work on insects and other non-micr oscopic invertebrates of terrestrial

and aquatic environments, but it is believed that the principle s and general techniques

will be found of value in studies on vertebrates and marine animals".

The section on 'Materials and Methods' in a biological paper can often be the

most important part of the paper, not only because conclusions so obviously depend

on methods, but also it is here that the way to solve some other problem may be

found. (Editors who shy away from purely methodological papers or who demand

cuts from this section might take note.) This widely scatter ed literature has not be-

fore been brought together into one place and this is hardly surprising in view of the

magnitude of the problem. Dr. Southwood has, however, succeeded in doing this

with, as the book is subtitled, particular reference to insect populations. The work

of 1400 authors has been examined and this presumably means even more individual

papers, complete citations for which are given at the end of each chapter.

After 5 pages of introduction, there are chapters on: the sampling programme

and the measurement and description of dispersion (51 pages); absolute population

estimates using marking techniques (42 pages); absolute population estimates by

sampling air, plants, plant products and vertebrate hosts (34 pages); absolute esti-

mates by sampling soil and litter (25 pages); absolute estimates by sampling fresh-

water habitats (16 pages); relative methods of population measurement (55 pages);

estimates based on products and effects of insects (11 pages); methods for the es-

timation of natality, mortality, and dispersal (37 pages); the construction, descrip-

tion and analysis of age-specific life-tables (34 pages); age-grouping of insects and

time-specific life-tables (11 pages); experimental component analysis (4 pages); the

measurement of associationbetween species and the description of a fauna (28 pages) ;

the estimation of productivity and the construction of an energy budget (19 pages).

There are 101 clear figures and 24 tables showing equipment and explaining processes

of analysis. In many instances, the whole process from collection of data through

final mathematical analysis is given, making it unneces sar y to go to the original

literature.

It is very difficult to find fault with this book. It will be essential to anyone in

anyway involved in research on insect populations, particularly during the planning

of a new project. Economic entomology must benefit. Teachers of ecology, too,

will find the book extremely useful. The author expresses the hope that more pre-

cise studies and more critical analyses will be attempted as a result of this book.

If this should happen then ecology will advance and the ecological armchair may be-

come a more comfortable place to inhabit.

Gordon Pritchard

Department of Entomology

University of Alberta

Quaestiones

entomologicae

A periodical record of entomological investigations,

published ot the Department of Entomology, Uni-

versity of Alberta, Edmonton, Conada.

VOLUME III

NUMBERS

JULY 1967

QUAESTIONES ENTOMOLOGICAE

A periodical record of entomological investigations, published at

the Department of Entomology, University of Alberta, Edmonton, Alberta.

Volume 3 Number 3 31 July 1967

CONTENTS

Guest editorial

Madge - A revision of the Genus Lebia Latreille in America north

of Mexico (Coleoptera, Carabidae)

Book review

137

. 139

^43

Guest Editorial - Fascinating Taxonomy

It is often stated that insects form at least 75% of the kinds of

animals. It is less often noted that this class is one of the most diverse

in the animal kingdom, with several of its 30-or-so orders being some-

times thought of as more diverse than the entire phylum Vertebrata.

From the taxonomic viewpoint, it is seldom remarked that while the

vertebrates are in a highly classified state, with problems of taxonomy

and nomenclature in the background, insects are at the stage where there

is great taxonomic activity, much monographic work, and innumerable

problems of nomenclature of all possible kinds.

These conditions justify no one in thinking that the insects are

more important zoologically than the vertebrates, but they do make it

obvious that the study of insect taxonomy today is likely to be much more

complex than the current sort of work on vertebrate taxonomy. To many

of us, this means more varied and more interesting.

A recent very extensive bibliography of mammals (Walker 1964)

shows among the thousands of items only a very few monographs , virtually

no major catalogs (except of type specimens), and few substantial faunal

studies in the last thirty or forty years. There are a tremendous num-

ber of studies of single species, usually of some particular aspect of

that species or its name. In fact, all the aspects of taxonomy together

are virtually buried under an avalanche of more practical studies, oc-

casioned by man's direct interest in these large animals as game, in

zoos, and under conservation. This is the natural result of the fact that

the species taxonomy reached a high state nearly a hundred years ago,

with many monographic and faunal treatments at that time.

Although the formal taxonomic study of insects began at the same

time as that of mammals, it progressed much more slowly, probably

mostly because of the vastly larger number of kinds and the much greater

difficulty in accumulating the necessary specimens . Large faunal studies

are still being produced, as witness the Insects of Hawaii by E. C.

Zimmerman, Monographs of tribes, families, and even orders are not

uncommon in today's literature, even with the problems of obtaining

publication, and generic reviews are legion.

138

In this situation, insect taxonomists are encountering all the pos-

sible problems of taxonomy, including many never faced by some of the

vertebrate taxonomists. These latter may be the result of availability

of thousands of specimens of some species; or of genera including hun-

dreds or even thousands of species; or of the organisms being too small

in size to be studied without special techniques; or especially of the two-

hundred-year history of the names, over which span a variety of inter-

pretations have been made and superseded, often by a larger number of

reviewers .

In addition to all this, one major factor alone contrasts the pre-

sent-day taxonomy of insects with that of mammals. This is the exis-

tence of many still undescribed species and genera of insects. Although

there have recently been predictions that new species will soon taper

off, there is as yet no clear indication of this, and the undescribed species

now to be found inmuseums will keep taxonomists busy for years even if

collecting turns up no more new ones.

Monographs, catalogs, and faunal studies never were so much

needed in mammalogy as in entomology, simply because a mammalogist

can be reasonably competent over the entire range of three to four thou-

sand species. Many insect families contain more species than this, re-

quiring a much higher degree of taxonomic specialization. And further-

more, it is probably much more difficult to distinguish 1000 species of

one genus than 4000 species that are clearly distributed among a thousand

genera.

For these reasons the taxonomy of insects is today far more di-

verse than that of mammals; probably far more demanding in discern-

ment and discrimination; still wide open for major contributions of a

variety of sorts; much more generally adaptable to statistical analysis

of the variation of its species; less affected by the practical interests of

man; and for all these reasons more fascinating.

R.E. Blackwelder

Department of Zoology

Southern Illinois University

Carbondale, Illinois

Madge

139

A REVISION OF THE GENUS LEBIA LATREILLE IN AMERICA NORTH OF

MEXICO (COLEOPTERA, CARABIDAE)

RONALD BRADLEY MADGE

British Museum (Natural History)

S. Kensington, London S.W. 7

Quaestiones Entomologicae

3: 139-242 1967

Within the genus Lebia in America north of Mexico four subgenera and 47 species are

recognized as valid. The genus is defined in a broad sense to include several New World groups

recognized as distinct genera by some workers. This concept of Lebia is supported with mor-

phological and limited biological evidence. It is also shown that in this sense Lebia encompasses

many exotic groups recognized as distinct genera at the present time.

A key to the sub genera and species is given. Each subgenus and species is described and

synonyms are listed. The distribution of each species is presented by locality records and for

those species with extensive ranges distribution maps are given. Structures important in identifi-

cation, especially the endophallic armature of the male genitalia, are illustrated.

The subgenus Loxopeza includes eight species of which three, deceptrix, subdola, and

subgrandis, are described as new. Five names are reduced to synonymy.

The sz/bgenws Polycheloma is described as new. The name of its single species, lecontei,

is a replacement for an invalid homonym.

The subgenus Lebia includes 37 species of which four, nigricapitata, abdita, insulata,

and perpallida are described as new. Forty-nine species group names are relegated to synonymy.

Phylogenetic relationships are postulated for these subgenera and species.

The genus Lebia is a group of ground beetles almost world wide in

distribution, comprising several hundred species. The species are

usually colorful and range from about 2. 5 to 14 mm in length. Although

the majority of species occur in the tropical regions a large number occur

in temperate areas. Very few extend into the far north. The adults are

predaceous and the larvae, as far as is known, are parasitoids on the

immature stages of chrysomelid beetles. However, even though they

may be beneficial, nothing is known about the biology of the vast majority

of the species.

This study deals with the taxonomy of the adults of Lebia occur-

ring in America north of Mexico. The concept of this group of species

as constituting a single genus is upheld, thus agreeing with most North

American students of Lebia but opposing the view of many worker s in other

parts of the world. The various populations of North American Lebia are

reevaluated in the light of the currently accepted concept of species as

outlined by Simpson (1961). Forty-seven species are recognized, seven

of which are described as new. This compares with 94 species listed

from the area under study by Csiki (1932) in Coleopterorum Catalogus.

These 47 species are arranged in four subgenera, one of which is new.

A phylogeny of the species within these subgenera is presented.

140

Revision of Lebia

HISTORICAL SUMMARY

The concept of the genus Lebia has undergone considerable modi-

fication since its description by Latreille in 1802. Although segregates

from Lebia were recognized at an early date ( Lamprias Bonelli 1809,

Echimuthus Leach 1815 (= Lamprias ) these were almost completely ignored

in the nomenclature of North American Lebia, A few American Lebia were

placed by Motschoulsky under the generic names Lamprias and Lia. It was

not until Chaudoir (1870-71) produced his Monographie des Lebiidesthat

the classification of our species was seriously changed. Of the many

genera into which Le6/a was split by Chaudoir, four [Lebia Latreille,

Loxopeza Chaudoir, Aphelogenia Chaudoir and Dianchomena Chaudoir) were re-

cognized as occurring in America north of Mexico. Later, Lamprias , as

understood by Chaudoir, was added to this list when Horn (1882) recog-

nized-Lebia divisa cis a member of that group. Previously Chaudoir (1870-

71) and Horn (1872) were uncertain of the generic affinities of divisa,

Metabola Chaudoir was found to occur here also when Bates described

Metabola vivida from Sonora and Arizona. Horn (1872) recognized Chaudoir ' s

genera as valid but later (1882) thought it best to regard them as sub-

genera. Since then the opinion of Horn has been followed by some wor-

kers while others have recognized some of the segregates of Chaudoir

as being generically valid. Thus Casey (1920) recognized Lebia, Loxopeza^

and Dianchomena but not Aphelogenia', Bradley (1930) recognized Lebia and

Dianchomena and not the others.

The first two species of Lebia in America north of Mexico to be

described were Fabricius 1776 and bAhiaiws Fabricius 1798, both

under the generic name Carabus . Say (1825) described four species of

Lebia although one of these [ornata ) was known to Melsheimer under the

na-m e quadrinotatus , a nomen nudum . Following Say North American species

of Lebia were described by many workers, most notably by Dejean,

LeConte, Chaudoir, Horn, Bates, and Casey. All the known species of

Lebia in the United States east of the Rocky Mountains were listed by

LeConte (1848). He recognized 23 species. In 1872, the year after

Chaudoir had finished his monograph of the known species of the world,

Horn gave a revision of the American species. Horn's key was the last

dealing with all the known American species . Although most species re-

cognized in this study were described from the area north of Mexico, a

number of species with predominantly southern distributions were ori-

ginally described from Mexico by Chevrolat or Bates. In the twentieth

century the taxonomy of Lebia was s eriously complicated whenT. L. Casey

described many new species, most of which have turned out to be syno-

nyms, and recognized as distinct many forms previously regarded as

having no taxonomic validity. The last species to be described in our

area was Malaena Hatch, 1953.

BIOLOGY

Information is available on the life cycle and immature stages of

very few species of Lebia. Silvestri (1904) described the life cycle of

Madge

141

Lebia scapularis Fourcroyin Europe and Chabous sou ( 1939) that of the North

American Lebia grandis Hentz. Lindroth (1954) described larvae of the

European Lebia chi or ocephala Hoffman. These three species in their laA'val

stages attack the pupating larvae and pupae of chrysomelid beetles . Pos-

sibly this is a habit of all species of Lebia . Accompanying this mode of

feeding there has been a striking hypermetamorphosis developed. The

following brief account of the life cycle is based upon that of grandis with

differences from the other two species noted where these are known.

Eggs are laid singly in the soil and, being covered with a sticky

secretion of the accessory glands, are camouflaged by the particles of

dirt sticking to them. The soil must be moist as the eggs are quite sus-

ceptible to desiccation. At 250 C the eggs hatch in 11 to 12 days.

Chabous sou found that in one case two pair s of grand/s produced 2600 larvae

even though one of the females died prematurely.

When the larvae hatch out they look like typical carabid larvae

about 3 to 4 mm long in grandis, with well developed legs, mouth parts,

and body sclerites. In both grandis and chlorocephala the tergal sclerites

are entire while in scapularis they are divided. This first active stage

then seeks out in the soil the pupating larvae and pupae of its host (for

grandis _ Leptinotarsa decemlineata , for chlorocephala - Chry solina varians and for

scapularis - Galerucella luteola ) . After feeding on the host chrysomelid the

first instar larva becomes greatly distended and has only weak powers

of locomotion. In nature it would thus probably feed only on a single

larva or pupa although in laboratory studies Lindroth was able to feed

the larva of chlorocephala as many as four host larvae or pupae. In grandis

and chlorocephala the first instar then molts to the second instar. In

scapularis the first instar larva before molting spins a cocoon from silk

secreted by the malpighian tubules. The second instar larva differs in

appearance from the first in that the appendages are all reduced and the

body sclerites are lacking. The second instar larva does not feed. It

molts to fourth pupal stage in grandis and chlorocephala , but in scapularis the

second instar larva molts into a form termed the "prepupa" by Silvestri.

This "prepupa" has the pupal characteristics but an abdomen of 10 seg-

ments. The "prepupa" then molts to the true pupa. In grandis develop-

ment from eclosion to emergence of the adult requires 15 to 20 days at

25 C.

The adult beetles are nocturnal and predaceous . Adults of scapularis

pass the winter at the base of plants and in the spring seek out and feed

upon the eggs and larvae of their host chrysomelid. After becoming

sexually mature they lay their eggs. Adults from this first generation

emerge in mid July and from these a second generation develops, the

adults of which hibernate.

In addition to grandis several other species of North American Lebia

as adults have been found to feed on chrysomelids although nothing is

known about the larvae. Cushmanand Isely (1916) found that in confine-

ment individuals oi Lebia fuscata ornata by them) would readily attack

callow adults and pupae of the cherry leaf beetle Galerucella cavicollis (LeC).

Isely (1920) found that adults of L. viridis fed upon eggs, larvae and pupae

of the grape vine flea beetles Altica chalybea 111. and A. woodsi Isely. Also,

Isely found that adult L. ornata "fed upon pupae and prepupae of the flea

beetles in confinement".

142

Revision of Lebia

TAXONOMIC CHARACTERS OF ADULTS

Color

Color and especially color pattern are very important in the iden-

tification and classification of the species of Lebia and are used both to

unite species into major groups and to separate some of the closely re-

lated species. There are three groups of colors found in Lebia: pale

colors (usually some shade of yellow or orange), metallic colors (usually

blue or green), and dark colors (usually black or brownish). The inter-

mediate condition between dark and pale is termed infuscated. In the

descriptions color is described by these terms (pale, dark, infuscated,

or metallic) with the actual color often noted in parentheses as well. By

using this scheme of nomenclature it is not necessary to describe the

variation within a color group. Metallic colors are readily recognized

as being such and dark and pale colors, in any one species, are usually

quite distinct.

External Morphology

In Lebia there are few external morphological characters of much

use for taxonomic purposes . Usually those available are difficult to in-

terpret and are often applicable to only a few species. However, these

are used in the identification of the species whenever possible.

In the descriptions the term “mouth parts" refers not only to the

mandibles, maxillae, and the labium but also to the labrum and to the

gula. The epilobes of the mentumare triangular flaps on the mesal side

of the lateral lobes of the mentum (fig. 1). They are said to be present

or absent. However, Horn (1884, 1882) pointed out that the epilobes are

actually always present and when stated to be absent are really just re-

duced. The epilobes, along with the tooth on the mentum, were used more

extensively in previous treatments of Lebia than they are here. These

structures are usually difficult to see and are mentioned only when nec-

nessary.

The neck region behind the eyes is usually moderately constricted

in Lebia (fig. 2). However, in three species it is very strongly constricted

(fig. 3) and there is a strong sulcus across the neck in front of the occi-

pital suture.

In previous works on Lebia the shape of the pronotum was des-

cribed in detail, but this is not done here. The differences between

species are usually slight, the variation within a species is often exten-

sive, and better characters are available elsewhere for identification.

Illustrations of the pronotum are presented only for those species in

which the pronotum is not the typical transverse shape (fig. 6).

The wings of Lebia (fig. 13) show several useful characters which

in a few species allow reliable identifications to be made of either sex

where otherwise only males could be identified by an examination of the

endophallic armature. To examine the wings the beetle was relaxed in

near boiling water, the left elytron was then raised, and the left wing

broken off at the base with a fine pair of forceps. This wing was first

studied in water and then flattened out and glued on a card to be pinned

beneath the specimen. It could subsequently be studied on the card.

Madge

143

The veins are named (fig. 13) according to the system of Balfour-

Browne (1943). This system is preferred to that of Forbes (1922) be-

cause in respect to the cubital and anal veins (which are of taxonomic

importance in Lebia) it is in better agreement with the homologies based

upon the axillary sclerites as outlined by Snodgrass (1935). According

to Snodgrass the first anal vein (called postcubitus) in winged insects is

usually more closely associated with the base of the cubitus than with the

third axillary sclerite and the rest of the anals but in Neuroptera, Mec-

optera and Trichoptera it is grouped with the other anal veins. As the

Coleoptera are related to the Neuroptera probably the first anal vein in

beetles is also associated with the third axillary sclerite. On this as-

sumption the first anal vein in the Coleoptera is the vein which Forbes

called 2A3. Forbes' lA plus the branches 2A^ and 2A£ are considered

here as branches of Cu2.

The apical pinch of the elytron is a narrow flattened area along

the suture at the apex. This pinch is usually well developed (fig. 10) but

in two species, bivittata and bilineata , it is much reduced.

The basal ridge of the elytron is an extension of the lateral ridge

of the elytral disc across the base. If complete it extends across the

grooves on either side of the scutellum (here termed the parascutellar

grooves); if incomplete it ends at the brow of the groove. Some speci-

mens of species in which the basal ridge is typically complete lack it but

the opposite is never true as far as I know.

The lateral lobes of the abdominal sterna are shallow lateral ex-

tensions of the posterior margins of the sterna, best developed on the

fourth and fifth segments. The central part of the posterior margin,

flanked by the lateral lobes, is referred to as the central trough (figs.

11, 12).

Male Genitalia

Both the endophallic armature and the apex of the median lobe

afford taxonomic characters for the recognition of the species of Lebia.

In most species recognized in this study the armature of the endophallus

is distinctive. The shape of the apex of the median lobe is of diagnostic

value in a few species. Most species can be identified by external char-

acters alone but, in a few, reliable identification can be made only from

the male genitalia.

For the study of the endophallus the genitalia were removed from

a male beetle. The genitalia were then cleared in a hot 10% solution of

potassium hydroxide for about one minute. For small specimens 30 to

40 seconds was often enough while for large specimens a couple of min-

utes were required. After treatment with the KOH the genital structures

were washed in water. The endophallus could usually be everted by

squeezing the median lobe beginning at the base and progressing towards

the apex. It was usually necessary to complete the eversion by inserting

a hooked minuten needle into the endophallus to catch the tip and pull it

out. In specimens stored in alcohol before mounting the endophallus

could not be everted (it usually tore). This could be remedied by boiling

the cleared genitalia ina soapy solution for 10 to 15 minutes, after which

eversion could be accomplished in the usual way. The genital structures

144

Revision of Lebia

were stored in glycerine in a microvial, or glued on a small card, on the

pin beneath the beetle from which they were extracted.

Measurements

In a few instances measurements are useful for specific identi-

fication of species or for the analysis of intraspecific variation. Because

total length could not be measured satisfactorily and conveniently relative

size has been indicated by length of the elytra as measured from the

base of the humeral area to the apex. The range in length was obtained

from all specimens available while the mean elytral length for each

species was calculated from measurements made on a sample of 20 to

30 specimens (when available). This sample included the largest and

smallest specimens . To avoid bias as muchas possible specimens to be

measured were not picked individually but rather were picked in groups

(usually two or three rows of specimens in a unit tray). Width of the

pronotum was measured at the widest point and length was measured

along the midline. All measurements were made with a ruled eyepiece

in a stereoscopic microscope to the nearest half unit. At 25^, used for

measurements under 4. 8 mm, one unit is 0. 04 mm; at 12^, used for

measurements over 4. 8 mm, one unit is 0. 08 mm.

Illustrations and Maps

The drawings were made with the aid of an ocular grid in a ster-

eoscopic microscope. In the illustrations of the endophallus little im-

portance should be placed on indications of wrinkles, bulges and folds in

the endophallus except in a few cases which are noted in the descriptions .

For each species the everted endophallus has been drawn in the most ap-

propriate of four views to show the armature. These have been termed

apical, abapical, left and right views according to the position of the

apex of the median lobe when the median lobe is towards the top of the

drawing and the endophallus towards the bottom. In an apical view the

apex of the median lobe is in front of the endophallus (fig. 66); in an

abapical view, the opposite, the apex is hidden behind the endophallus

(fig. 67). In a left view the apex is to the left of the drawing and in a

right view the apex is to the right (figs. 64, 65). Using this nomenclature

the endophallus of a dissected specimen can be oriented with the drawings.

For the species of the subgenus Loxopeza where the endophallic armature

is very complex the endophallus has been drawn as if slit down the ab-

apical side and spread out. This allows for ready comparisons between

species. For the same purpose the groups of spines have been numbered

as in fig. 50. In all the other species the endophallus was drawn whole

in one or more of the four positions listed above. The simpler armature

of these does not require any nomenclatural system for the various groups

of spines.

Distribution maps aregivenfor all species except those with very

restricted ranges . On the maps dots are not placed for all records avail-

able (all records are listed separately) but rather only enough to show the

limits of distribution as I know them and to fill out the range. Dots re-

present counties or more restricted localities; stars are used when

only a state locality is available on the label.

Madge

145

In the lists of localities given for each species counties, if not

given, have been added where possible except for localities in Canada

where counties are not consistently used. In addition, no counties have

been given for non-restricted localities which are in two or more coun-

ties (mountain ranges, large lakes, national parks) unless this infor-

mation is given on the label.

Recognition of Moles and Females

As the male genitalia play an integral part in the identification of

at least some of the species of Lebia , it is important to be able to distin-

guish betweenmale and female specimens. Several characters facilitate

this. Males of all species have apreapical notch on the inner side of the

mesotibia (fig. 16), have a double row of papillate hair s on theunderside

of the first three protarsal segments (fig. 18), and lack the pair of in-

wardly placed setae on the apical abdominal sternum (fig. 11). Females

lack both thepreapical notch on the mesotibiae and the papillate hairs on

the underside of the protarsi, but have a pair of more inwardly set setae

on the apical abdominal sternum (fig. 12). In addition, males of the sub-

genus Loxopeza have the first three protarsal segments obliquely dilated

mesad (fig. 17) while in the females these segments are normal. Inmost

males the circumgenital ring protrudes from the end of the abdomen.

Of the three characters applying to the whole genus the presence

or absence of the preapical notch on the mesotibia is the easiest to use

as the apex of the mesotibia is usually visible in pinned specimens. The

undersurfaces of the protarsi are often folded close to the body and can-

not be seen without relaxing the specimen. The long setae on the last

abdominal sternum may be broken off leaving only small foveae. The

arrangement is then more difficult to discern. There is some variation

in the number of setae but this variation occurs only in the row common

to both male and female and the seta which indicates a female is always

present in this sex.

Synonymy

Several points in the specific synonymies need to be clarified.

Subgenera where used by an author are indicated in parentheses. If an

author made no distinctionbetween varieties and subspecies then varietal

names are listed as being subspecific. Otherwise they are not listed at

all. Type localities have been determined from the descriptions only and

may be more restricted on the label of the type specimen. Names of

journals are abbreviated according to the World List of Scientific Per-

iodicals, third edition.

Criteria for Species and Subspecies

Following the currently accepted definition, species are "groups

of actually or potentially interbreeding natural populations, which are

reproductively isolated from other such groups" (Simpson 1961). In

museum specimens the evidence on which reproductive isolation is judged

is necessarily based onmorphological and geographical characters. Two

forms were regarded as specifically distinct if they overlapped geograph-

ically and did not intergrade in the area of overlap in at least one mor-

146

Revision of Lebia

phological character. Sympatric forms differing only in color were con-

sidered conspecific. Sympatric forms which differed slightly in mor-

phology (and usually in color) but which tended to intergrade could be

either distinct species or polymorphic variants . In these cases the rea-

sons for the decisionmade are given in the discus sion under the species

concerned.

Allopatric forms were regarded as conspecific if there were in-

termediate forms in the intermediate geographical area or if the geo-

graphically nearest specimens approached each other in their distinguish-

ing characters. Allopatric forms not covered by the above statement

were regarded as specifically distinct if they differed in morphological

or color character s to the same extent as or more than other good species

did; or conspecific if they did not.

No subspecies have been recognized in this study. Intraspecific

variation is described and where possible dines are pointed out.

TAXONOMY

Genus Lebia Latreille

Description

Small to medium sized beetles. Color various and varied.

Head . Prognathous, slightly drooping; eyes usually prominent.

Labrum more or less truncate, with six setae across anterior margin.

Clypeus with a single seta on each side. Fronswith or without sculpture;

with two supraorbital setae above each eye. Mandibles moderately pro-

minent and with a distinct scrobe; labium with postmentum divided into

a mentum and a submentum; mentum with or without epilobes and a

tooth; ligula with paraglossae short and usually not extending beyond

glossae; palpi cylindrical, pointed or truncate apically, penultimate

segment of labial palpus usually bisetose. Antennae usually with seg-

ments one to three and basal third of segment four glabrous; extending

back to basal third or fourth of elytra. Neck usually moderately con-

stricted, sometimes strongly so, rarely rather stout.

Prothorax. Pronotum usually distinctly transverse in shape and

always with a basal lobe; lateral margins usually widened basally, oc-

casionally narrow throughout, with a seta just anterior to middle and at

basal corner; disc with variable sculpture.

Pterothorax . Wings fully developed; oblongum cell often reduced;

second branch of cubitus not forked. Metepimeron narrow.

Elytra. Apex obliquely truncate and usually slightly sinuate. Disc

usually somewhat flattened, with nine striae (usually distinct) and a scu-

tellar stria; intervals flat to strongly convex, third interval with two

dorsal punctures next to third stria; ninth interval with a series of um-

bilicate punctures, with one puncture at the outer apical corner set in-

ward and forming a jog in the series; base of disc with a strong groove

on each side of the scutellum; basal ridge complete or incomplete; api-

cal pinch usually large and well developed, rarely small.

Legs . Protibiae with or without an upper spur. Mesotibiae of

males with a preapical notch on inner side, rarely more than one. Fourth

Madge

147

tarsal segment of hind tarsus emarginate or bilobed. Tarsal claws al-

ways pectinate.

Abdomen. Venter with segments bearing shallow lateral lobes

along posterior margins. Pygidium with a mid- longitudinal keel.

Male genitalia . Parameres small, right smaller than left. Median

lobe with shape of apex various. Endophallus usually armed.

Discussion

Since its recognition the genus Lebia has undergone extensive

modification, first by the splitting off of the more distinctive groups in-

to separate genera and then by the absorption of some of these genera

back again as subgenera. Chaudoir (1870-71) recognized on a world

basis 22 genera (most of which had their species originally described as

Lebia ) as belonging to his group Lebiides and additional genera have since

been described. At the present time there is no generally accepted de-

finition of Lebia. While this taxon certainly does not include all the gen-

era which have been placed near it, it does seem to include many of

them. For the North American species the following seven characters,

when taken as a group, are regarded as being diagnostic and separate

Lebia clearly from the other lebiine genera in our fauna.

1. Pronotum lobed at base (fig. 5-9)

2. One umbilical puncture at outer apical corner of elytra set in,

thus forming a jog in the series (fig. 10)

3. Elytra "pinched" along the suture at the apex (fig. 10)

4. Pygidium (seventh abdominal ter gum) with a weak midlongitudinal

Carina

5. Abdominal sterna with shallow lateral lobes (figs. 11, 12)

6. Vein Cu^ not forked (fig. 13)

7. Males with a preapical notch on inside of mesotibiae (fig. 16)

In addition all species are probably parasitoids of chrysomelid leaf beet-

les.

Included with Lebia s.s. in this study, either as subgenera or sy-

nonyms, are the following groups: Aphelogenia Chaudoir, Dianchomena

Chaudoir, Lamprias Bonelli, Loxopeza Chaudoir, and Metabola Chaudoir.

Of the many exotic groups placed near Lebia , Lia Eschscholtz and

Lachnolebia Maindron have been seen and found to possess all seven of the

characters set down for Lebia. In addition G, E. Ball has kindly checked

examples in several European museums of most genera near Lebia for

the above characters except the fifth and sixth. The following groups

possess all five: Cymatographa Chaudoir, Ectomomesa Chaudoir, Grammica

Chaudoir, Helcosopha Chaudoir, Hemicycla Chaudoir, Lebidema Motschoulsky,

Metalebia Jeannel, Nematopeza Chaudoir, Orthobasis Chaudoir, Poecilostola

Chaudoir, Poecilothais Maindron and Promecochila ChdiUdoir . Four groups,

Lebistinida Peringuey, Rhopalostyla Chaudoir, Scythropa Chaudoir and Stephana

Chaudoir, possess the first four of the above characters but as the speci-

mens available were females the seventh character could not be checked.

In the species checked of Pachylebia Jeannel, Lebistina Motschoulsky and

Diacoptodera Alluaud the first four characters were present but males lacked

the preapical notch in the mesotibiae. I have s een one specimen of Lebistina

(a male) which had a series of very shallow preapical notches on the

148

Revision of Lebia

mesotibia. The loss (or reduction) of this character in these groups is

probably secondary. In the examples of Liopeza Chaudoir and Lionedya

Chaudoir the abdomen could not be seen to check the pygidial keel but

otherwise they were like Lebia as far as could be checked. All of the

above groups are probably Lebia as here defined. Arsinoe Castelnau,

Dromiotes Jeannel, Lebiomorpha Muller, Paralebia Peringuey and Scalidion

Schmidt- Goebel are probably not . Aristolebia Bates, Daer A. Semenov

and Znojko and Metabele Peringuey were not seen.

Key to the subgenera and species of Lebia in America north of Mexico

1 Upper protibial spur present 2

Upper protibial spur absent subgenus LEBIA 11

2(1) Frons and pronotum with many coarse setiferous punctures;

elytral disc metallic with the basal third pale

subgenus LAMPRl AS divisa LeC. , p. 165

Frons and pronotum without coarse setiferous punctures; ely-

tral disc either entirely metallic or entirely pale 3

3(2) Elytral disc entirely pale; proepisternum with longitudinal wrin-

kles suhgenns POLYCHELOMA lecontei new na.me, p. 164

Elytral disc entirely metallic; proepisternum smooth

subgenus LOXOPEZA 4

4(3) Frons dark (usually black) 5

F rons pale 6

5(4) Palpi and antennae pale; distribution - eastern half of United

States and adjacent Canada (fig. 126) tricolor Say, p. 156

Palpi and usually antennal segments 4 to 11 dark; distribution -

western half of United States and adjacent Canada except west

coast (fig. 141) atriceps LeC., p. 155

6(4) Palpi and usually antennal segments 4 to 11 dark; distribution -

eastern two thirds of United States and adjacent Canada (fig.

117) atriventris Say, p. 153

Palpi and antennae pale; distribution - eastern United States and

adjacent Canada, in the southwest to Arizona 7

7 (6) Elytral intervals strongly convex; elytral disc a dull green,

sometimes almost black; distribution - southeastern Arizona. .

pimalis (Csy. ), p. 159

Elytral intervals at most moderately convex; elytral coloration

variable 8

8(7) Anal margin of wing justdistadof vein 3A2 with sclerotized patch

strongly arched (fig. 14); armature of male endophallus as in

figs. 57, 58; distribution - eastern United States and in south to

Arizona 9

Anal margin of wing with sclerotized patch weakly arched (fig.

15); armature of male endophallus as in figs. 54, 55; distribution -

western Texas to Arizona 10

9(8) Distribution - eastern United States and adjacent Canada, in the

south possibly as far west as Davis Mountains, Texas; Texas

specimens with the third group on the endophallus large (fig. 58)

grandis Hentz, p. 161

Distribution - western Texas to Arizona; third group of spines

Madge

149

10(8)

11(1)

12(11)

13(11)

14(13)

15(14)

16(15)

17(16)

18(17)

19(18)

20(18)

21(13)

on the endophallus small (fig. 57) subgrandis, n. sp.,p. 160

Length of elytra 3. 80 to 4. 68 mm subdola n. sp.,p. 157

Length of elytra 6. 13 to 7. 33 mm deceptrix n. sp.,p. 158

Elytral disc metallic with pale fasciae 12

Elytral disc entirely metallic, entirely dark, or dark with pale

markings 13

Frons with coarse punctures and short erect hairs (best seen in

lateral view); third antennal segment distinctly hairy

pulchella Dej. (in part), p. 167

Frons without coarse punctures and short erect hairs; third

antennal segment with only a few scattered short hairs in addition

to the long distal hairs bitaeniata Chev. , p. 171

Elytral disc metallic (either blue or green) and pronotum pale .

14

Elytral disc dark, dark with pale markings or metallic but when

metallic, pronotum is dark 21

Pronotal margins narrow throughout (fig. 8) neck strongly con-

stricted abdominalis Chd. , p. 198

Pronotal margins widened baaally; neck not strongly constricted

15

Head metallic (blue or green); femora dark distally

viridipennis Dej., p. 170

Head darker pale but not metallic; femora entirely dark or pale

16

Basal ridge of elytra incomplete; distribution - Florida

lecta Horn, p. 213

Basal ridge of elytra usually complete; distribution - not in

Florida 17

Pterothoracic sclerites dark like abdomen; head dark (usually

black, reddish black in Montana, Alberta, and Saskatchewan

specimens) cyanipennis Dej. (in part), p. 176

Pterothoracic sclerites pale (except sometimes metepisternum),

contrasting with color of abdomen; head pale 18

Fourth segment of hind tarsus bilobed; distribution - southeas-

tern Texas or northeastern United States and adjacent Canada

(fig. 138) 19

Fourth segment of hind tarsus emarginate; distribution - south-

western Texas to southern California 20

Distribution - southeastern Texas; armature of male endophallus

as in figs. 66, 67 rufopleura Schfr. , p. 172

Distribution - northeastern United States and adjacent Canada;

armature of male endophallus as in fig. 11 ^pleuritica ~LeC , , p. 173

Metepisternum usually pale, occasionally dark; elytral intervals

usually moderately convex; microsculpture of frons usually dis-

tinct tuckeri (Csy. ), p. 174

Metepisternum infuscated; elytral intervals flat or weakly con-

vex; microsculpture of frons lacking or indistinct

arizonica Schfr. , p. 175

Head, pronotal disc and entire elytral disc either dark or metal-

lic 22

150

Revision of Lebia

22(21)

23(22)

24(23)

25(24)

26(25)

27(21)

28(27)

Elytral disc usually maculate, if entirely dark then pronotum

pale . 27

Pronotum bicolored, lateral margins pale and disc dark

marginicollis Dej. , p. 180

Pronotum. entirely dark, at most with tinges of red at sides. . . .

Frons with strong punctation and short erect hairs (the latter

best seen in lateral view); third antennal segment distinctly

hairy pulchella Dej. (in part), p. 167

Frontal punctation usually not strong and never with short erect

hairs; third antennal segment with only a few scattered short

hairs in addition to the long distal hairs 24

Lateral lobes of penultimate abdominal sternum each wider than

the central trough (fig. 12); third antennal segment usually pale;

basal ridge of elytra incomplete; elytral disc usually dark, or

if metallic, then legs pale pumila Dej., p. 215

Lateral lobes of penultimate abdominal sternum each equal to or

narrower than the central trough (fig. 11); if elytral disc dark

then basal ridge is usually complete and third segment is dark;

legs never pale 25

Frons and pronotum dark, elytral disc metallic; frons with fine

punctures but no fine striations; basal ridge of elytra usually

complete; distribution - southern British Columbia, Alberta, and

Saskatchewan to New Mexico, Arizona, and southern California

(fig. 129) cyanipennis Dej. (in part), p. 176

Not as above in color or if frons and pronotum black and elytra

metallic, then frons with fine striations (especially at sides) and

basal ridge of elytra incomplete 26

Frons and pronotum usually shiny black, sometimes with a met-

allic green tinge, elytra metallic; basal ridge of elytra incom-

plete; distribution - southern British Columbia to southern Cali-

fornia; endophallus with armature as in figs. 76, 77

perita Csy. , p. 182

Frons and pronotum concolorous with the elytral disc, either

metallic or black; basal ridge usually complete; distribution -

transcontinental; endophallus with armature as in figs. 72, 73.

viridis Say, p. 177

Lateral pronotal margins narrow throughout; head dark (usually

black) 28

Lateral pronotal margins widened basally; color of head var-

ious ... 29

Abdomen entirely pale; epipleuron dark and each elytron with

two pale vittae bivittata (Fab.), p. 195

Basal half of abdomen dark, apical half pale; epipleuron dark or

pale; each elytron usually with only one pale vitta, sometimes

two when epipleuron pale hilineata Mots. , p. 197

Neck strongly constricted; head pale (vertex sometimes slightly

infuscated); frons striated at least on lateral thirds 30

Neck not strongly constricted; head color and frontal sculpture

variable 31

29(27)

Madge

151

30(29)

31(29)

32(31)

33(32)

34(33)

35(31)

36(35)

37(35)

38(37)

39(38)

40(38)

Frons completely striated; elytra with a common sutural vitta

and two lateral spots; distribution - southern Arizona

miranda (Horn), p. 188

Frons striated on lateral thirds only; elytra with a common su-

tural vitta and two lateral vittae; distribution - eastern United

States and adjacent Canada (fig. 131) solea Hentz, p. 187

Femora dark at least distally 32

Femora entirely pale 35

Elytra with a dark sutural vitta and a lateral spot on posterior

half of elytra (sometimes joined to dark sutural vitta)

histrionica Bates, p. 192

Elytra with a dark sutural vitta and a dark lateral vitta, rarely

entire elytral disc (except apex and lateral margins) dark

33

Common sutural vitta furcate basally, rarely entire elytral disc

(except apex and lateral margins) dark and furcation obscured;

basal ridge of elytra usually complete vittata (Fab,), p. 189

Common sutural vitta not furcate basally, elytra not entirely

dark except apex and lateral margins; basal ridge of elytra in-

complete 34

Head black; distribution - Arizona . . . .nigricapitata n. sp. , p. 194

Head pale; distribution - eastern United States and adjacent

Canada pectita Horn, p, 19/^

Frons dark (usually black) and distinctly striated except a tri-

angular area above clypeus; abdomen pale 36

Not with above combination of characters 37

Pronotum distinctly striate (like frons) on anterior lateral re-

gions; apical pale marking on elytra interrupted by a fine dark

edging along suture analis Dej. , p. 184

Pronotum rugose on anterior lateral regions; apical pale marking

on elytra uninterrupted by a dark edging along suture

scalpta Bates, p. 186

Head with fine deep punctures on frons; typical elytral pattern

as in fig. 42; pronotal disc dark, margins pale

lohulata LeC. , p. 207

Head without fine deep punctures on frons; elytral patternnot as

above; coloration of pronotum variable 38

Elytral patternasin fig. 37 (note dark apex of elytra) or posterior

part of frons and vertex rugose-striate; mentum without a tooth

39

Elytral pattern not as above and frons and vertex not rugose-

striate; mentum usually with a tooth, lacking only in insulata . . .

40

Abdomen entirely dark; frons usually only rugose on lateral

thirds guttula LeC., p. ^99

Abdomen dark at sides, pale medially; frons usually entirely

rugose-striate ahdita n. sp. , p. 201

Basal ridge of elytra usually complete; pale apical spot of elytra

shaped as in figs. 38-41 or apex of elytra entirely dark ; dis-

tribution - Arizona 41

152

Revision of Lebia

Basal ridge of elytra incomplete; elytra with pale apical spot

shaped as in figs. 43-48 or absent; distribution - eastern half

of United States and adjacent Canada 45

41(40) Apex of elytra dark, pale marking usually restricted to humeral

area, sometimes extending three fourths of the elytra but never

much onto the mesal half; abdomen dark . . . scapula Horn, p. 183

Apex of elytra always pale, basal pale markings extending well

over onto the mesal half of the elytra; abdomen pale 42

42(41) Pale basal marking of elytra shaped as in fig. 38; distribution -

southeastern Texas; mentum without a tooth

insulata n. sp. , p. 2 02

Pale basal marking of elytra shaped as in figs. 39-41; distri-

bution - not in southeastern Texas; mentum with a tooth .... 43

43(42) Distribution - western Texas to Arizona 44

Distribution - not in above area (fig. 127) . . ,fuscata Dej. , p. 203

44(43) Elytral disc with at least a lateral dark spot, usually a complete

vitta (fig. 40); frons with a deep groove next to eyes

subrugosa Chd. , p. 205

Elytral disc without a lateral dark spot (fig. 41); frons without

a deep groove next to eyes perpallida n. sp. , p. 2 06

45(40) Elytra vaulted and patterned as in fig. 48; distribution - south-

eastern Texas; head pale bumeliae Schfr. , p. 213

Elytra flattened and not patterned as above; distribution - Texas

and eastern United States; head dark (reddish brown) in Texas.

46

46(45) Elytral disc entirely dark except for lateral margin; frons equally

dark; wing with a triangular remnant of oblongum cell; abdomen

pale, darkening apically collaris Dej., p. 214

Color of elytral disc not as above or if so then wing without a

triangular remnant of oblongum cell and abdomen pale through-

out 47

47(46) Elytral pattern as in fig. 47 (note shape of basal pale spot, that

basal dark marking is always present and that this marking joins

or approaches the middle dark fascia which extends forward along

the side of the elytral disc); endophallic armature of male as in

fig. Ill calliope Bates, p. 212

Elytral pattern as in figs. 43-46 or entirely dark except lateral

margin; endophallic armature of male as in figs. 108-110 ....

48

48(47) Distribution - eastern Texas; endophallic armature as in figs.

109-110; elytral disc (fig. 46) with dark circum-scutellar spot

usually not extending over to shoulder, when it does it usually

gradually becomes paler and is not divided by pale basal spot . .

esurialis Csy. , p. 211

Distribution - eastern United States (including eastern Texas)

and adjacent Canada; endophallic armature as in fig. 108; in

Texas elytra when largely pale (fig. 44) with a circumscutellar

and a humeral spot divided by an arm of the basal pale spot . . .

ornata Say, p. 208

Madge

153

Subgenus Loxopeza Chaudoir

Loxopeza Chaudoir 1870 : 138. Type species - Leb ia grandis Hentz (here de-

signated) .

Description

Characters in common among the species north of Mexico are

given in the following subgeneric description and are not repeated in the

species descriptions.

Head - Variable in color. Frons punctate-rugose, especially at

sides, microsculpture variable. Mentumwith a small tooth and epilobes;

ligula with paraglossae not extending beyond glossae. Palpi slender,

apex more or less pointed; penultimate segment of labial palpi bisetose.

Antennae variable in color, with segments one to three and basal third

of four more or less glabrous. Neck not strongly constricted.

Prothorax - Entirely pale (in species north of Mexico) , lateral mar-

gins of pronotum palest. Pronotum transverse in shape, lateral mar-

gins widened basally; disc rugose and with distinct microsculpture.

Pterothorax - Sterna, pleura, and scutellum pale (in species north

of Mexico). Wings with oblongum cell complete.

Elytra _ Disc metallic; color of epipleura dark or pale. Disc

with striae distinct, convexity of intervals variable; apical pinch well

developed; basal ridge complete.

Legs - Largely pale (in species north of Mexico). Protibiae with

an upper spur present. Mesotibiae of males with a single preapical

notch. Fourth segment of hind tarsus usually emarginate, sometimes

bilobed.

Abdomen _ Venter and pygidium dark.

Male genitalia - Median lobe stout, apex short and broad (fig. 49).

Endophallus strongly armed.

Discussion

Recognition - The diagnostic character s of the subgenus Loxopeza are:

upper protibial spur present, a complete oblongum cell in the wing, a

small tooth and epilobes on the mentum, elytra metallic, the male geni-

talia with a short broad apex to the median lobe, and the endophallus

heavily armed. The upper protibial spur and completely metallic elytra

readily distinguish the subgenus Loxopeza from the other subgenera of

Lebia north of Mexico.

Taxonomic status - Previous workers disagreed as to the status and

limits of Loxopeza, it being regarded as a distinct genus, a distinct sub-

genus within Lebia, or not distinct at all from Lebia. Sometimes similarly

colored species of Lebia s.s. were included within it. The species of

Loxopeza are clearly members of the genus Lebia as defined here. How-

ever, a& they occupy a primitive and isolated position within Lebia they

are regarded as forming a distinct subgenus.

1. Lebia (Loxopeza) atriventris Say

Lebia atriventris Say 1825 : 13. Type locality - not given. Dejean 1826:

454. LeConte 1848 : 193. LeConte 1863 : 5. Gemminger and

Harold 1868 : 136. Blatchley 1910 : 144. Leng 1920 : 65 {Loxopeza).

Csiki 1932 : 1316 (Loxopeza). Blackwelder 1944 : 53.

154

Revision of Lebia

Loxopeza atriventris \ Chaudoir 1870 : 142. Horn 1872 : 131. Bates 4RR3 T

220. Casey 1920 : 235.

Loxopeza enormis Casey 1920 : 257. Type locality - New York (near the

city). NHW SYNONYMY.

Lebi a enormis ; Csiki 1932 : 1316 {Loxopeza),

Description

Length of elytra - 3.40-5.40 mm; mean (22 specimens) 4,22 mm.

Head - Frons, vertex, clypeus and genae pale; microsculpture

on frons usually distinct. Mouth parts pale except dark palpi. Antennae

with basal three and a third segments pale, others usually dark or in-

fuscated, becoming paler apically.

Elytra - Disc metallic (usually ; blue) ; epipleura dark. Disc with,

intervals usually very weakly convex, almost flat.

Legs - Entirely pale except for infuscated tarsi. Fourth segment

of hind tarsus emarginate.

Male Genitalia - Armature of endophallus as in fig. 51 (note that some

of the spines of the fourth group are rounded and not pointed apically,

that the sixth group of spines is formed from short broad spines arranged

to form a vertical rectangle, and the seventh group is lacking). The

endophallic armature in five specimens was examined.

Discussion

Recognition - The only pale headed Loxopeza in the range of atriventris

is grandis which is larger, with the elytral intervals more strongly con-

vex, and the palpi pale. Superficially atriventris is very similar to pleuritica

but the latter lacks an upper protibial spur and has the epipleura pale

instead of dark.

Variation - Lebia atriventris shows only minor variation. The fourth

segment of the hind tarsus has been described in the past as being bi-

lobed or strongly emarginate but this does not seem to be the typical

condition. Specimens examined all had the fourth segment emarginate.

As in atriceps antennal segments 4-11 are sometimes pale, the elytral

disc is occasionally greenish, and the elytral intervals may be some-

what more strongly convex.

Synonymy - Casey's Loxopeza enormis , with its blackish antennae and

emarginate fourth tarsal segment, is clearly a synonym of atriventris .

Distribution - Lebia atriventris occurs in the eastern half of the United

States and adjacent Canada (fig. 117). Over 950 specimens were studied

from the following localities.

CANADA

MANITOBA - Makinack; Winnipeg. ONTARIO - Belleville; Grand Bend; Hillcrest; Leamington; Normandale; Ottawa;

Point Pelee National Park; Port Colborne; Prince Edward Co.; Ridgeway; Bondeau; Simcoe; Strathroy; Toronto;

Trenton; Vineland Station. QUEBEC - lanorale; Montreal; Saint Ailaine. SASKATCHEWAN - Saskatoon.

UNITED STATES

ALABAMA - Auburn (Lee Co.); Cheaha State Park (Clay Co.); Mobile (Mobile Co.). CALIFORNIA. CONNECTICUT -

Cornwall (Litchfield Co.); Lakeville (Litchfield Co.); Litchfield (Litchfield Co.); Lyme (New London Co.); New

Haven (New Haven Co.); Stamford (Fairfield Co.); Stores (Tolland Co.); Suffield (Hartford Co.). DELAWARE -

Newark (New Castle Co.). DISTRICT OF COLUMBIA. FLORIDA - Jacksonville (Duval Co. ).

ILLINOIS - Beverley Hills; Bowmanville; Chicago (Cook Co.); Downers Grove (Du Page Co. );

Edgebrook; Eldorado (Saline Co.); Evanston (Cook Co. ); Fox Ridge State Park (Coles Co.); Glenview (Cook Co. );

Grand Detour; Grand Tower (Jackson Co.); Illinois Beach State Park (Lake Co.); Joliet (Will Co.); Kickapoo State

Park (Vermilion Co. ) ; LaGrange (Cook Co.); Lake Zurich (Lake Co.); Lyons (Cook Co.); Macon Co. ; Palos Park

(Cook Co.); Quincy (Adams Co.); Riverside (Cook Co. ); Urbana (Champaign Co. ) ; Utica (LaSalle Co.). INDIANA -

Beverley Shores (Porter Co.); Dune Park; Fulton Co.; Gary (Lake Co.); Hammond (Lake Co.); Knox Co.; Lafayette

(Tippecanoe Co .) ; Lagrange Co. ; Long Lake; Marion Co.; Mineral Springs; Pine; Posey Co. ; Putnam Co. ; Tremont;

Madge

155

Vigo Co. IOWA - Ames (Story Co.); Council Bluffs (Pottawattamie Co.); Crawford Co. 1 Iowa City (Johnson Co. ) ;

Mount Pleasant (Henry Co. ) ; Polk Co.; Sioux City (Woodbury Co. ) Waukon (Allamakee Co. ). KANSAS - Chanute (Neosho

Co.); Kiowa Co.; Lawrence (Douglas Co.); Manhattan (Riley Co.); Mount Hope (Sedgwick Co. ) ; Onaga (Pottawatomie

Co.); Rage (Kingman Co. ) ; Saline Co.; Topeka (Shawnee Co.); Wellington (Sumner Co.). KENTUCKY. LOUISIANA -

Ruston (Lincoln Co.). MARYLAND - Baltimore (Independent City); College Park (Prince George# Co.); Forest Glen

(Montgomery Co. ); Frederick (Frederick Co. ) ; Hagerstown (Washington Co. ) ; Marshall Hall (Charles Co. ) ; Patuxent

Refuge (Prince Georges Co.); Plummers Island; Plum Point (Calvert Co.); Sparrows Point (Baltimore Co.); Suitland

(Prince Georges Co.); Travilah. MASSACHUSETTS - Arlington (Middlesex Co.); Boston (Suffolk Co.); Brookline

(Norfolk Co. ); Cambridge (Middlesex Co. ) ; Chicopee (Hampden Co. ) ; Framingham (Middlesex Co. ); Humarock (Ply-

mouth Co.); Lexington (Middlesex Co. ) ; Nahant (Essex Co. ) ; Needham (Norfolk Co. ) ; Revere (Suffolk Co. ) ; Saugus

(Essex Co.); Sherborn (Middlesex Co. ) ; Springfield (Hampden Co. ) . MICHIGAN - Alcona Co. ; Ann Arbor (Washtenaw

Co.); Beaver Island (Charlevoix Co. ) ; Bloomfield (Oakland Co. ) ; Cheboygan Co. ; Cooper Woods (Oakland Co. ) ; Detroit

(Wayne Co.); East Lansing (Ingham Co.); E. K. Warren Preserve, Sawyer (Barrien Co.); George Reserve (Livingston

Co.); Grand Ledge (EatonCo.); High Island (Charlevoix Co. ) ; Huron Mountain Club (Marquette Co.); Marquette (Mar-

quette Co.); Marysville (Saint Clair Co.)! Mason (Ingham Co.); Mecosta Co.; Milford (Oakland Co.); Mottawa; Nau-

binway (Mackinac Co.); Palmer Park (Wayne Co.); Paw Paw Lake (Van Buren Co.); Pigeon (Huron Co.); Port Huron

(Saint Clair Co.); Rochester (Oakland Co.); Poyal Oak (Oakland Co. ) ; Sanford (Midland Co.); Saugatuck (Allegan Co.);

Shiawasse Co. ; Silver Lake State Park (OceanaCo.); Southfield (Oakland Co.); South Fox Island (Leelanan Co.); South

Haven (Van Buren Co.); Sutton Farm (Lapeer Co.); Three Oaks (Barrien Co.); Whitefish Point (Chippewa Co.).

MINNESOTA - Big Stone Co.; Crookston (Polk Co.); Cyrus (Pope Co.); Frontenac (Goodhue Co.); Gray Cloud Island;

Hallock (Kittson Co. ) ; Houston Co. ; Mille Lacs Co.; Minneapolis (Hennepin Co. ); Mississippi Bluff (Houston Co. ) ;

Mora (Kanabec Co.); Olmsted Co.; Saint Paul (Ramsey Co.); Saint Peter (Nicollet Co.); Two Harbors (Lake Co.).

MISSISSIPPI - Camp Shelby (Forrest Co.). MISSOURI - Branson (Taney Co.); Kansas City (Jackson Co.); Saint Louis

(Independent City); Springfield (Greene Co. ) . MONTANA - Billings (Yellowstone Co. ) . NEBRASKA - Lincoln (Lancaster

Co.); Omaha (Douglas Co.); Saltillo (Lancaster Co. ) ; Waverly (Lancaster Co.). NEW HAMPSHIRE - Cornish; Exeter

(Rockingham Co. ) . NEW JERSEY - Arlington (HudsonCo.); Bergenfield (BergenCo.); Boonton (Morris Co. ) ; Collings-

wood (Camden Co.); Chester (Morris Co.); Clemton; Durham P. ; Emerson (BergenCo.); Fort Lee (BergenCo.);

Greenwood Lake; Hillsdale (Bergen Co.); Lavallette (Ocean Co.); Manasquan (Monmouth Co.); Morristown (Morris

Co.); Newark (Essex Co.); New Brunswick (Middlesex Co.); Phillipsburg (Warren Co.); Point Pleasant (Ocean Co.);

Riverton (Burlington Co. ); Snake Hille; South Orange (Essex Co. ); Westwood (Bergen Co. ) ; Woodbury (Gloucester Co.).

NEW YORK - Bear Mountain; Bronxville (Westchester Co.); Buffalo (Erie Co.); Callicoon (Sullivan Co.); Catskill

(Greene Co.); Chatham (Columbia Co.); Cranberry Lake (Saint Lawrence Co.); Florida (Orange Co.); Gpshen (Orange

Co.); Ithaca (Tompkins Co.); Kissing L. , L. I.; McLean Bogs (Tompkins Co.); New Rochelle (Westchester Co.);

New York City; N. Fairhaven; Ocean Beach, Fire Island (Suffolk Co.); Olcott (Niagara Co.); Oswego (Oswego Co.);

Peekskill (Westchester Co.); Pike Wyoming Co.); Ringwood Reserve, Dryden (Tompkins Co.); Sea Cliff (Nassau Co.);

Wayne Co. ; West Point (Orange Co.); W. Hebron; White Lake (Sullivan Co.); Wildwood State Park (Suffolk Co. ) .

NORTH CAROLINA - Black Mountains; Clayton (Johnston Co.); Faison (Duplin Co.); Laurel Springs, Upper Mountain

Research Station (Alleghany Co. ) ; Raleigh (Wake Co.). NORTH DAKOTA - Trail Co. OHIO - Ashtabula (Ashtabula

Co.); Athens (Athens Co.); Bedford (Cuyahoga Co.); Cincinnati (Hamilton Co.); Columbus (Franklin Co.); Flat Rock

Creek, Benton Twp. (Holmes Co.); Georgesville; Grove City; Holmesville (Holmes Co.); Jefferson (Ashtabula Co.);

Lockbourne (Franklin Co.); Lucas Co.; Mendon (Mercer Co.); Ottawa (Putnam Co.); Oxford (Butler Co.); Paulding

Co.; Put-in- Bay; Rock Creek (Ashtabula Co. ); S. Bass Island (Ottawa Co. ); Springfield (Clark Co. ) . OKLAHOMA -

Norman (Cleveland Co. ) . PENNSYLVANIA - Bethlehem (Northampton Co. ) ; Columbia Cross Roads (Bradford Co. );

Delaware Water Gap (Monroe Co.); Easton (Northampton Co.); Mt. Airy; Ohiopyle (Fayette Co.); Philadelphia (Phil-

adelphia Co.); Pittsburgh (Allegheny Co.); Tinicum (Bucks Co.); West Chester (Chester Co.). RHODE ISLAND -

Block Island (Newport Co.); Warwick (Kent Co.); Watchhill (Washington Co.). SOUTH CAROLINA - Beaufort Co. ;

Blackville (Barnwell Co.); Clemson (Oconee Co.). TENNESSEE - Elmwood (SmithCo.); Green Brier (Robertson Co.);

Knoxville (Knox Co.). TEXAS - Abilene (Taylor Co.); Carthage (Panola Co.); Columbus (Colorado Co. ) ; Dalhart,

Rita Blanca Lake (Dallam Co.); Tyler (Smith Co.). VIRGINIA - Blacksburg (Montgomery Co.); Falls Church (Fairfax

Co.); Fredericksburg (Spotsylvania Co.); Richmond (Henrico Co.); Warm Springs (Bath Co.). WEST VIRGINIA -

Fairmont (Marion Co.); Salem (Harrison Co. ) ; Sistersville (Tyler Co.); White Sulphur Springs (Greenbrier Co. ) .

WISCONSIN - Bayfield (Bayfield Co.); Brodhead (Green Co.); Madison (Dane Co.); Platteville (Grant Co.).

2. Lebia (Loxopeza) atriceps LeConte

Lebia atriceps LeConte 1863a : 5. Type locality - Nebraska. Gemminger

and Harold 1868 : 136. Leng 1920 : 65 (Loxopeza) , Csiki 1932 :

1316 (Loxopeza). Blackwelder 1944 : 53.

Loxopeza atriceps ; Chaudoir 1870 : 143. Horn 1872 : 132. Casey 1920 :

238.

Loxopeza nanulina Casey 1920 : 238. Type locality - Colorado (Boulder Co.).

NEW SYNONYMY.

Lebia nanulina ; Csiki 1932 : 1317 (Loxopeza).

Description'

Length of elytra - 3.67-5. 50 mm; mean (25 specimens) 4.49 rnm.

Head - Frons, vertex, and genae dark (frons usually black), cly-

peus pale; microsculpture of frons distinct. Mouth parts pale except

dark palpi. Antennae with basal three and a third segments pale, others

usually dark or infuscated but becoming paler apically.

Elytra - Disc metallic (usually blue); epipleura dark. Disc with

156

Revision of Lebia

intervals weakly to moderately convex.

Legs - Trochanters and femora pale; tibiae infuscated; tarsi

dark; fourth segment of hind tarsus emarginate.

Male Genitalia- Armature of endophallus as in fig. 52 (note that the

sixth group of spines is formed from short spines arranged to form a

transverse rectangle or square and that the seventh group is lacking).

The endophallic armature in nine specimens was examined.

Discussion

Recognition - North of Mexico there ar e only two black headed

species of the subgenus Loxopc2a : afrAeps and tricolor. The two are allo-

patric and differ in the color of their palpi and antennae (dark in atriceps ,

pale in tricolor .

Variation - Antennal segments four to eleven are sometimes as

pale as the basal segments, the elytral disc is occasionally greenish in-

stead of blue, and the elytral intervals are sometimes moderately con-

vex.

Synonymy _ Casey's Loxopeza nanulina is here considered a synonym

of atriceps as it differs only in size and other minor features. It occurs

within the range of atriceps .

Distribution - Lebia atriceps occurs from the Canadian prairies south

to Arizona, New Mexico and western Texas (fig. 141). Over 400 speci-

mens were studied from the following localities.

CANADA

ALBERTA - Cypress Hills; Edmonton; Lethbridge; Medicine Hat; Tilley. MANITOBA - Aweme. SASKATCHEWAN -

Saskatoon; Swift Current; Val Marie.

UNITED STATES

ARIZONA - Arivaca (Pima Co.); Bar Foot Park, Chiricahua Mountains (Cochise Co.); Brown's Canyon, Baboquivari

Mountains (Pima Co.); Calabasas Picnic Grounds, Ruby Road (Santa Cruz Co.); Canelo (Santa Cruz Co.); Cave Creek

Ranch, Chiricahua Mountains (Cochise Co.); Dragoon (Cochise Co.); Fairbank (Cochise Co.); Fort Grant (Graham

Co. ) ; Fort Huachuca (Cochise Co. ) ; Hereford, Carr Canyon, Huachuca Mountains (Cochise Co.); McNary (Apache Co.);

MormonLake (Coconino Co .) ; Nogales (Santa Cruz Co.); Palmerlee (Cochise Co.); Patagonia (Santa Cruz Co.); Pata-

gonia Mountains; Pena Blanca (Santa Cruz Co.); Portal (Cochise Co.); Prescott (Yavapai Co.); Ramsey Canyon,

Huachuca Mountains (Cochise Co.); Ruby (Santa Cruz Co.); Santa Rita Mountains; Southwest Research Station, Portal

(Cochise Co.); Texas Pass, Dragoon Mountains (Cochise Co.); Turkey Flat, Chiricahua Mountains (Cochise Co.);

White Mountains (Gile Co.); Winslow (Navajo Co.). COLORADO - Boulder (Boulder Co.); Cheyenne Mountains Museum

(El Paso Co.); Colorado Springs (El Paso Co.); Conejos Co.; Denver (Denver Co.); Fort Collins (Larimer Co.);

Pueblo (Pueblo Co.). IDAHO - Caldwell (Canyon Co.); Indian Cove (Owynee Co.); Mountain Home (Elmore Co.).

KANSAS. MONTANA - Bozeman (Gallatin Co. ) ; Crow Agency (Big Horn Co.). NEBRASKA- Glen (Sioux Co.); Mitchell

(Scotts Bluff Co.). NEVADA - Go Shu'.e Valley (White Pine Co.). NEW MEXICO - Bernalillo (Sandoval Co.); Coolidge

(McKinley Co. ) ; Hot Springs, Las Vegas (San Miguel Co .) ; lemez Mountains; Mescalero Indian Reservation (Otero

Co.). NORTH DAKOTA- Sentinel Butte (Golden Valley Co.). SOUTH DAKOTA - Hot Springs (Fall River Co. ) . TEXAS -

Davis Mountains; Fort Davis ( leff Davis Co. ) . UTAH - Farmington (Davis Co. ) .

3. Lebia (Loxopeza) tricolor Say

Lebia tricolor Say 1825 : 11. Type locality - "Pennsylvania . . . also on the

Missouri". Dejean 1826 : 453. LeConte 1848 : 192. LeConte

1863 : 5. Gemminger and Harold 1868 : 141. Blatchley 1910 :

144. Leng 1920 ; 65 {Loxopeza). Csiki 1932 : 1317 {Loxopeza).

Loxopeza tricolor I Chaudoir 1870 : 140. Horn 1872 : 131. Casey 1920 :

235.

Description

Length of elytra - 3. 72-5. 76 mm; mean (20 specimens) 4.71mm.

Head - Frons, vertex and genae dark (frons usually black), cly-

peus pale; microsculpture of frons usually distinct. Mouth parts pale.

Antennae entirely pale.

Madge

157

Elytra - Disc metallic (usually shiny green); epipleura dark.

Disc with intervals moderately convex.

Legs - Entirely pale. Fourth segment of hind tarsus bilobed.

Male genitalia - Armature of endophallus as in fig. 43 (note that the

first group of spines is very poorly developed, the sixth group is a loose

cluster of short broad spines, and that an eighth group is present). The

endophallic armature in five specimens was examined.

Discussion

Recognition - See under atriceps .

Variation - There appears to be no marked variation in tricolor .

Distribution - Lcbia tricolor occurs in the eastern United States and

adjacent Canada (fig. 126). Over 150 specimens were studied from the

following localities.

CANADA

ONTARIO - Ottawa; Prince Edward Co. ; Roseland; Toronto; Trenton. QUEBEC - Covey Hill; Montreal; Norway

Bay; Perkins Mills; Sherbrooke.

UNITED STATES

CONNECTICUT - Canaan (Litchfield Co.); Cornwall (Litchfield Co.); Litchfield (Litchfield Co.). DISTRICT OF COL-

UMBIA. FLORIDA - Levy Co. ; Marion Co. ; Tampa (Hillsborough Co. ). ILLINOIS - Chicago (Cook Co. ). INDIANA -

Bartholomew Co.; Gary (Lake Co.). KANSAS. LOUISIANA - Hart; New Iberia (Iberia Co.). MASSACHUSETTS -

A rlington (Middlesex Co. ) ; Boston (Suffolk Co. ) ; Brookline (Norfolk Co. ) ; Salisbury (EssexCo.); Springfield (Hampden

Co.). MICHIGAN - Cheboygan Co. ; Detroit ( Wayne Co .) ; Marquette (Marquette Co. ) ; Washtenaw Co. ; Whitefish Point

(Chippewa Co. ). MINNESOTA - Two Harbors ( Lake Co. ) . NEW HAMPSHIRE - Franconia (Grafton Co. ) ; Mount Wash-

ington (Coos Co.); Rumney (Grafton Co.). NEW JERSEY - Arlington (Hudson Co.); Hillsdale (BergenCo.); Lake Hopat-

cong; Manasquan (Monmouth Co.); Mountain Lakes (Morris Co.); Woodbury (Gloucester Co.). NEW YORK - Asps

Hill, L.l ; Bear Mountain (Rockland Co. ); Buffalo (Erie Co. ); Catskill Mt. , (Ulster Co. ); Dryden (Tompkins Co. );

Irving (Chautaugua Co. ) ; Ithaca (Tompkins Co.); Kingston (Ulster Co.); Lancaster (Erie Co. ) ; Lyons ( Wayne Co .) ;

Mount Whiteface (Essex Co. ) ; New Rochelle ( Westchester Co. ) ; New York City; Olcott (Niagara Co. ) ; Phoenicia (U Ister

Co. ); White Plains (Westchester Co. ). NORTH CAROLINA - Lake Junaluska (Haywood Co. ). PENNSYLVANIA - Arendts-

ville (Adams Co.); Milford (Pike Co.); Nanticoke ( Luze rne Co.); PhilacTelphia (Philadelphia Co. ) ; State College (Centre

Co.); The Rock. RHODE ISLAND - Warwick (Kent Co. ) . TEXAS. VIRGINIA - Mount Vernon (Fairfax Co. ) ; Rosslyn

(Arlington Co.). WEST VIRGINIA - White Sulphur Springs (GreenbrierCo. ). WISCONSIN - Milwaukee (Milwaukee Co. ) .

4. Lebia (Loxopeza) subdola new species

llolotype _ A male labelled as follows: Madera Cn, Sta. Rita Mts. , Sta.

Cruz Co. Ariz. VIII 3.60, 5000' - 5800' G. E. Ball family and

R. B. Madge collectors. To be deposited in the Canadian Nat-

ional Collection, Ottawa.

Paratypes are from the following localities.

ARIZONA - Carr Canyon, Huachuca Mountains (Cochise Co, ) (one male, California Academy of Sciences); Cave Creek,

Chi ricahua Mountains (Cochi se Co. ) (one female , California Academy of Sciences); Cave Creek Ranch, Chiricahua Moun-

tains (Cochise Co.) (three females, personal collection of G.E. Ball, University of Alberta); Chiricahua Mountains

(Cochise Co.) (one female, California Academy of Sciences; one female. United States National Museum); Huachuca

Mountains (Cochise Co. ) (one male California Academy of Sciences); Madera Canyon, Santa Rita Mountains (Santa Cruz

Co.) (two females, personal collection of G.E, Ball, U niversity of Alberta ; two males and one female, Cornell Univer-

sity); Mount Washington, Nogales (Santa Cruz Co. ) (two females, California Academy of Sciences); Palmerlee (Cochise

Co.) (one male. Museum of Comparative Zoology); Pinery Canyon, Chiricahua Mountains (Cochise Co.) (one male,

American Museum of Natural History; one male, Canadian National Collection); Southwest Research Station, Portal

(Cochise Co.) (three males and three females, American Museum of Natural History; one male, Canadian National

Collection); Turkey Flat. Chiricahua Mountains (Cochise Co.) (one male, California Academy of Sciences); White

Mountains (one male. Museum of Comparative Zoology) . TEXAS - Big Bend National Park (Brewster Co. ) (two females ,

personal collection of G. E. Ball, University of Alberta).

Description

Length of elytra - 3. 80 - 4. 68 mm; mean (22 specimens) 4. 21 mm.

Head - Frons, vertex, clypeus and genae pale; microsculpture of

frons usually lacking. Mouth parts pale. Antennae entirely pale. Neck

not suddenly constricted behind eyes.

Elytra - Disc metallic (usually bright blue); epipleura usually

dark. Disc with intervals weakly convex.

158

Revision of Lebia

Wings - The sclerotized patch just distad of vein SA^ weakly con-

vex (fig. 15).

Legs - Entirely pale. Fourth segment of hind tarsus emarginate.

Male genitalia - Armature of endophallus as in fig. 54 (note that the

first group of spines is small and poorly developed, the sixth group is

formed of short broad spines in a loose cluster, and the seventh is cres-

cent-shaped and lying between the sixth and first groups). The endo-

phallic armature in seven specimens was examined.

Discussion

Recognition^ This small Loxopeza is most likely to be confused with.

small specimens of suhgrandis , However, the two can usually be distin-

guished by the lack of microsculpture on the frons of suhdola. In addition

the small sclerotized patch in the anal region of the wing is shaped dif-

ferently in the two (weakly convexin suhdola ^ strongly convex in subgrandis,

figs. 14, 15).

Variation - In a few specimens the microsculpture of the frons is

more or less distinct. Otherwise there appears to be no major varia-

tion in suhdola .

Etymology- The name is derived from the Latin adjective subdolus -

subtle, deceiving - in reference to it being confused with Lebia subgrandis .

Distribution- This species is known only from southern Arizona and

western Texas; 29 specimens (type material) were studied.

5. Lebia (Loxopeza) deceptrix new species

Holotype - A male labelled as follows: Pena Blanca, Santa Cruz Co. Ariz.

4000' August 11, i960 at light G. E. Ball familyand R.B. Madge.

To be deposited in the Canadian National Collection, Ottawa.

Paratypes are from the following localities.

ARIZONA - Bear Valley, Tumacacori Mountain (Santa Cruz Co.) (one female, Museum of Comparative Zoology) ; Canelo

(Santa Cruz Co.) (two males and one female. University of Arizona); Cave Creek Ranch, Chiricahua Mountains (Cochise

Co.) (two females, personal collection of G. E. Ball, University of Alberta); Madera Canyon, Santa Rita Mountains

(Santa Cruz Co.) (three males, personal collection of G. E Ball); Pena Blanca (Santa Cruz Co.) (four males and six

females, personal collection of G. E. Ball): Southwest Research Station, Portal (Cochise Co.) (one female, American

Museum of Natural History). TEXAS - Davis Mountains (Jeff Davis Co.) (one male, California Academy of Sciences).

Description

Length of elytra^ 6. 13 - 7. 33 mm; mean (22 specimens) 6. 56 mm.

Head - Frons, vertex, clypeus and genae pale; microsculpture of

frons usually distinct. Antennae entirely pale.

Elytra - Disc metallic (usually bright blue) ; epipleura dark. Disc

with intervals weakly to moderately convex.

Wings - The sclerotized patch just distad of vein 3A£ weakly con-

vex (fig. 15).

Legs - Entirely pale. Fourth segment of hind tarsus emarginate.

Male genitalia - Armature of endophallus as in fig. 55 (note that the

first group of spines is small, and that the second, sixth and seventh

groups are not separate from each other). The endophallic armature in

five specimens was examined.

Madge

159

Discussion

Recognition - Lebia deceptrix may be confused with large specimens of

subgrandis and any specimens of grandis from western Texas. From both

it can usually be recognized by the smaller, less arched sclerotized

patch in the anal region of its wing. Positive identification is best ob-

tained from the endophallic armature of the male.

Variation - In the small series of deceptrix available variation in the

microsculpture of the frons and the color of the elytral disc was noticed.

Usually the microsculpture is present but occasionally it is reduced or

lacking as in specimens of subdola. The elytral disc is typically bright

blue, rarely with a greenish tinge.

Etymology- The name is derived from the Latin noun deceptrix -

she that deceives - in reference to the similarity of this species toother

Loxopeza , especially grandis .

Distribution - North of Mexico Lebia deceptrix is known from southern

Arizona and western Texas; 22 specimens (type material) were studied.

6. Lebia (Loxopeza) pimalis (Casey)

Loxopeza pimalis Casey 1920 : 237. Type locality - Arizona.

Lebis pimalis I Csiki 1932 : 1317 {Loxopeza).

Description

Length of elytra - 3.80- 5. 40 mm; mean (24 specimens) 4. 60 mm.

Head - Frons, vertex, clypeus, and genae pale, microsculpture

of frons distinct. Mouth parts pale. Antennae entirely pale.

Elytra « Disc metallic (usually a dull dark green); epipleura var-

ying from dark to pale. Disc with intervals strongly convex.

Legs - Entirely pale. Fourth segment of hind tarsus emarginate.

Male genitalia - Armature of endophallus as in fig. 56 (note that the

first group of spines is moderately large, the sixth group consists of

only one or two short spines and the seventh group lies in a fold at the

side of the first group). The endophallic armature in six specimens was

examined.

Discussion

Recognition - Lebia pimalis can be distinguished from our other species

of the subgenus Loxopeza by its very convex elytral intervals . Occasionally

there may be difficulty in separating some of the greenish specimens of

subgrandis in which case it is necessary to check the male genitalia.

Variation - The elytral disc varies in color from the usual dull

green to sometimes almost black while the epipleura vary from pale to

dark. Most specimens have the epipleura pale or partially so.

Distribution - Lebia pimalis is known north of Mexico only in southern

Arizona. Over 175 specimens were studied from the following local-

ities.

ARIZONA - Brown's Canyon, Baboquivari Mountains (Pima Co.); Canelo (Santa Cruz Co.); Cave Creek Ranch, Chiri-

cahua Mountains (Cochi se Co. ) ; Coyote Mountains; Douglas (Cochise Co.); Dragoon (Cochise Co.); El Mirador Ranch,

Sasabe, Baboquivari Mountains (PimaCo.); Fort Grant (GrahamCo.); Kits Peak Rincon, Baboquivari Mountains (Pima

Co.); Madera Canyon, Santa Rita Mountains (Santa Cruz Co.); Montezuma Pass, Huachuca Mountains (Cochise Co.);

Nogales (Santa Cruz Co.); Palmerlee (Cochise Co.); Patagonia Mountains (Santa Cruz Co.); Pena Blanca (Santa Cruz

Co.); Ruby (Santa Cruz Co.); Sabino Canyon, Santa Catalina Mountains (Pima Co.); Sierritas; Sonoita (Santa Cruz

Co.); Texas Pass, Dragoon Mountains (Cochise Co. ) ; Tombstone (Cochise Co. ) ; Tucson (Pima Co. ) .

160

Revision of Lebia

7. Lebia (Loxopeza) subgrandis new species

Holotype - A male labelled as follows: Pena Blanca, Santa Cruz Co.

Ariz. 4000' August 11, I960 at light G. E. Ball family and R. B.

Madge collectors. To be deposited in the Canadian National

Collection, Ottawa.

Paratypes are from the following localities.

ARIZONA - Brown's Canyon, Baboquivari Mountains (Pima Co.) (three inales and one female. Museum of Comparative

Zoology); Pena Blanca (Santa Cruz Co.) (one male and three females, personal collection of G. E. Ball, University of

Alberta): San Bernardino (Cochise Co.) (two males and one female. University of Arizona); Tucson (Pima Co.) (one

male and three females, California Academy of Sciences); Tucson Mountains, Desert Museum (Pima Co.) (one male

and one female. University of Arizona).

Description

Length of elytra - 4. 25 - 6. 25 mm; mean (22 specimens) 5. 22 mm.

Head - Frons, vertex, clypeus and genae pale; microsculpture

of frons distinct. Mouth parts pale. Antennae entirely pale.

Elytra - Disc metallic (blue or green); epipleura varying from

dark to pale. Disc with intervals usually moderately convex.

V^ings- The sclerotized patch justdistadof vein 3A2 strongly con-

vex (fig. 14).

Legs- Entirely pale. Fourth segment of hind tarsus emarginate.

Male genitalia ^ Armature of endophallus as in fig. 57 (note that the

first group of spines is moderately large, the third group is small, the

sixth group is formed of a dense group of long narrow spines and the

seventh group lies in a fold at the side of the first group). The endo-

phallic armature in 11 specimens was examined.

Discussion

Recognition - There are three other species of the subgenus Loxopeza

with pale heads which are sympatric with subgrandis north of Mexico:

pimalis, subdola, and dccepfnx . In addition, the range o£ grandis may over-

lap that of subgrandis in western Texas. In this area these two can be re-

liably separated only on the basis of differences in the endophallic ar-

mature. Separation of subgrandis from the other three species is best

done on the basis of the endophallic armature although there are some

external character s which can be used. The elytral intervals are not as

strongly convex in subgrandis as in pimalis and the two can usually be se-

parated on the basis of this character. The small sclerotized patch in

the anal region of the wing is usually more convex in subgrandis than in

deceptrix and subdo/a and this usually permits recognition. In addition, the

microsculpture of the frons is distinct in subgrandis and usually lacking in

subdola .

Variation - In addition to the considerable variation in size the

elytral disc varies from blue to green and the epipleura from dark to

pale. In a few specimens the elytral intervals are more strongly con-

vex and approach the condition found inpimalis.

Relationships - Lebia subgrandis is very closely related to grandis. The

two are largely allopatric but may overlap in western Texas. Because

the endophallic armatures are quite distinct where the two at least ap-

proach each other in Texas the two forms are regarded as distinct spec-

ies. The fact that the third group of spines in the endophallic armature

Madge

161

of grandis becomes smaller in northern specimens and thus approaches the

condition found in suhgrandis has no bearing on the question as the two are

then separated by hundreds of miles.

Etymology - The specific name is derived from the Latin prefix

sub _ a being, situated under and hence a being concealed behind some-

thing - and grandis in reference to it being confused with the closely re-

lated Lebia grandis .

Distribution - Lebia sub grandis occurs from western Texas to southern

Arizona. Over 250 specimens were studied from the following local-

ities.

ARIZONA - Arivaipa (Graham Co. ) ; Bear Valley, Tumacacori Mountains (Santa C ruz Co.); Brown's Canyon, Baboquivari

Mountains (PimaCo.); Canelo (Santa Cruz Co.); Carr Canyon, Huachuca Mountains (Cochise Co.); Cave Creek Ranch,

Chiricahua Mountains (Cochise Co.); Charleston (Cochise Co.); Cochise Stronghold, Dragoon Mountains (Cpchise Co.);

Cutter (Gila Co. ) ; F air bank (Cochise Co.); Fort Huachuca (Cochi se Co. ) ; Globe (Gila Co. ) ; Kits Peak Rincon, Babo-

quivari Mountains (Pima Co.); Madera Canyon, Santa Rita Mountains (Santa Cruz Co.); Nogales (Santa Cruz Co.);

Oracle (Pinal Co.); Palmerlee (Cochise Co.); Patagonia (Santa Cruz Co.); Patagonia Mountains (Santa Cruz Co.);

Pearce (Cochise Co.); Pena Blanca (Santa Cruz Co.); Portal (Cochise Co.); Prescott (Yavapai Co.); Rice; Ruby

(Santa Cruz Co.); Sabino Canyon, Santa Catalina Mountains (Pima Co.); San Bernardino (Cochise Co.); Southwest

Research Station, Portal (Cochise Co.); Sunnyside Canyon, Huachuca Mountains (Cochise Co.); Texas Pass, Dragoon

Mountains (Cochise Co.); Tucson (Pima Co.); Tucson Mountains (Pima Co.). CALIFORNIA. NEW MEXICO - Deming

(Luna Co.); Double Adobe Ranch, Animas Mountains (Hidalgo Co.). TEXAS - Alpine (Brewster Co.); Limpia Creek

Canyon, Davis Mountains (Jeff Davis Co.).

8. Lebia (Loxops ea) grandis Hentz

Lebia grandis Hentz 1830 : 258. Type locality - North Carolina. LeConte

1848 : 192. LeConte 1865 : 5. Gemminger and Harold 1868 :

139. Blatchley 1910:144. Leng 1920:65 (Loxopeza), Csiki 1932 :

1316 {Loxopeza) . Blackwelder 1944 : 54.

Loxopeza grandis ; Chaudoir 1870: 139. Horn 1872 : 131. Casey 1920: 235.

Loxopeza majuscula Chaudoir 1870 : 141. Type locality - Texas. NEW SY-

NONYMY. Horn 1872 : 131. Casey 1920 : 236.

Lebia majuscula I Leng 1920 : 65 {Loxopeza). Csiki 1932 : 1317 {Loxopeza).

Blackwelder 1944 : 54.

Loxopeza grandis rivularis Casey 1920 : 235. Type locality - Texas (Browns-

ville). NEW SYNONYMY.