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DEEL 121 1978

TIJDSCHRIFT

| VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

NEDERLANDSE ENTOMOLOGISCHE VERENIGING

BESTUUR (BOARD)

Voorzitter (Chairman) ............... R. H. Cobben

Vice-Voorzitter (Vice-President) ......... Th. van Leeuwen

Secretanisi(Secretany) EEE R. de Jong

A GOLOSO RN RN AS AE Rijksmuseum van Natuurlijke Historie,

Raamsteeg 2, Leiden

Penningmeester (Treasurer) . ........... H. Wiering

AGGRESSORI TE ONE Doorntjes 29, Bergen

Bibliothecaris (Librarian) ............. W. N. Ellis

Plantage Middenlaan 64, Amsterdam

Bedeni(MEembers) mens A. K. Minks, H. van Oorschot

TIJDSCHRIFT VOOR ENTOMOLOGIE

Redactie (Editorial Board) ............ P.J. van Helsdingen, R. de Jong, J. Krikken,

M. A. Lieftinck, C. van Achterberg

Ades AGIO Re Rijksmuseum van Natuurlijke Historie,

Raamsteeg 2, Leiden

The journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea.

Subscription rate: D.Fl. 170,— per year.

Afleveringen 1 —3 verschenen 19.vii.1978

Afleveringen 4—6 verschenen 31.xii.1978

ISSN 0040-7496

TIJDSCHRIFT VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

The article of De Jong (pt. 3) has its own index at the end of the article (p. 139), but nevertheless the pri-

mary references have been incorporated below. Of the article of Kielland (pt. 4) only the names of ge-

nera and new taxa have been included here.

* Een sterretje duidt een naam nieuw voor de wetenschap aan

* An asterisk denotes a name new to science

HYMENOPTERA

adempta 283, 288

Agaon 283, 290, 291

Alfonsiella 283, 288, 291,

[sqq

Allotriozoon 283, 287, 290

[sqq

arabicus 287

blandus 283, 290 sqq

Blastophaga 280 sqq

boschmai 290, 291

carinifrons 2

carlosi 290, 292

ssp. centralis

[(P. xanthognathus) 11

Ceratosclen 285, 287, 290

[sqq

ceylonicus 1, 2,9

clavigera 290

constabularis 283, 290

coronata 283

coronifrons |

cumanensis 283

Deilagaon 283, 287, 290 sqq

delhiensis 290, 292

Dolichoris 283, 291, 292

elisabethae 2

Elisabethiella 283, 290 sqq

errata 283

Eupristina 283, 290, 292

exiguus 2, 7

froggatti 291

*genalis 2, 4

gestroi 283

hamiferum 283

immaturus 291

imperialis 287, 291, 292

jacobsi 285, 287, 291 sqq

javana 283

*ssp. keiseri (P. maculatus)

PAY

leucognathus 10

longicaudus 291

longiscapa 290

Liporrhopalum 285, 287,

[290, 291

maculatus 2

ssp. madrasiensis

[(P. elisabethae) 2, 3

malabarensis 287

ssp. malayanus

[(P. carinifrons) 11

mandibularis 290 sqq

Maniella 283, 288, 290, 292

mariae 290, 292

masii 283, 291

matalensis 1, 2, 3

megarhopalus 283, 287 sqq

mindanaensis 287

ssp. modestum

[(A. hamiferum) 283

nervosae 287, 290

nietneri |, 2, 5

nitidus |, 2

orientalis 2

Paragaon 290, 291

Parapristina 283, 288, 290,

[292

Pegoscapus 283, 290, 292

pilipes 287

Platyscapa 283, 285, 287

plebejus 291

Pleistodontes 283, 287, 290

[sqq

prodigiosum 283, 287

Psen |

psenes 280, 287

Psenulus |

pulcherrimus 2,9

puncticeps 2, 6

quadraticeps 285, 287

regalis 291

rennellensis 291

rieki 291

stueckenbergi 288

sumatrana 283

sundaica 283, 287, 291

sycomori 287

Sycophaga 287

*ssp. taprobanensis

[(P. carinifrons) 1, 2, 10

Tetrapus 280, 287

Urostigma 283, 288

valentinae 287

vasculosae 283, 291, 292

verticillata 290

Waterstoniella 283, 287, 290

[sqq

xanthognathus 11

yoshimotoi 5

LEPIDOPTERA

Abantis 213

Abisara 192, 222

abscondita 49

Acada 215

aceris 246, 253

acetosae 18

Acleros 215, 216

Acraea 189—192, 222

Acticera 158, 208

Actis 202

*aegilopidella 269

agylla 28, 34, 36 sqq

Alaena 193

alceae 114, 134

Alenia 106, 108, 109, 114,

[117

296 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, 1978

Aloeides 157, 158, 199

Amauris 170

amenophis 53

Ampittia 214

*amygdali 264

Andronymus 216,217

anomalella 15, 16, 18

Antanartia 188

Anthene 147, 204—206

antonia 107

Apaturopsis 186

Aphelosetia 17

Aphnaeus 197, 221, 222

Aphniolaus 200

Aphysoneura 172

Appias 162, 163

argentipedella 13

Argiolaus 199

Ariadne 185

Artitropa 217

Aslauga 196, 197

asterodia 28, 34 sqq

Aterica 180

Athysanota 210

aurella 13, 18, 240

aurivillii 65

*australis 105, 128

Axiocerses 147, 198, 199

Azanus 208

*azaroli 261

Baliochila 195

bamptoni 38

basiguttella 243

Belenois 163, 164, 222

Bematistes 188, 189

bettoni 38

Bicyclus 130, 157, 170—172

bifida 98

Borbo 220, 221

Brusa 220

Byblia 185

Cacyreus 147, 207

Caenides 217, 218

Calleagris 212

Callimorpha 269

Caprona 213

Capys 204

caradjai 250, 269

Carcharodus 106, 113,114,

[115,117

carnea 72

Castalius 208

Catopsilia 169

Catuna 180

Celaenorrhinus 110, 211

Celotes 106, 109, 110, 114

centifoliella 13, 16, 18

cerricolella 243

Charaxes 147, 173—178,

[221

chenga 69

Chloroselas 198

Chondrolepis 217

Cnodontes 157, 195

Coeliades 210,211

Colias 158, 169

Colotis 157, 168

colotes 29, 54 sqq

confusa 29, 40, 55, 59 sqq

Crenidomimas 179

crenulatae 264

cribrellum 107, 110

Cupidopsis 206

cursoriella 13

Cymothoe 178

Cyrestis 184

Danaus 169, 170

delagoae 94 sqq

Deloneura 196

depauperata 31, 49, 96, 99,

[103 sqq

*deschkai 274

diomus 31, 42, 43, 45 sqq

Diopetes 202

Dixeia 164, 165

doris 29, 52, 53 sqq

dromus 32, 42, 43, 80 sqq

Eagris 211

Ectoedemia 15, 240 sqq

Eicochrysops 158, 208, 209

Elachista 17

elongata 80

*embonella 259

Epamera 200, 201

Epitola 196

Eretis 212

Eronia 169

Erynnis 114

Etesiolaus 201

Euchrysops 157, 158, 210

Euliphyra 196

Euphedra 179, 180

Euryphene 179

Euryphura 179

Eurema 169

Eurytela 185

Euxanthe 173

evanida 54, 66

*expansa 178

fasciata 60

Fedalmia 271

ferax 48, 49, 102, 128

Fomoria 274

Fresna 218, 219

freyella 246

Freyeria 210

galba 24, 32, 66 sqq

Gamia 217

gecko 52

Gegenes 221

geron 29, 52 sqq

Gomalia 106, 109, 212

Gorgyra 214, 215

Graphium 160, 161, 221

Gretna 217

groschkei 251

Hamanamida 180

hamirella 272

Harpendyreus 207

Heliopetes 106, 107, 110

hellas 98

Hemiolaus 201

Henotesia 172

hermona 52

Hewitsonia 196

higginsi 64, 128

Hypocopelates 202

Hypoleucis 216

Hypolimnas 186

Hypolycaena 201, 202

Hypomyrina 202

ignobilis 130

Iolaphilus 199, 222

lolaus 199

Isoteinon 214

Issoria 188

Johanssonia 18

Kallima 186

Kedestes 214

*kigoma 191, 193

kituina 28, 38 sqq

Lachnochnema 147, 157,

[197

Lachnoptera 188

lacreuzei 49

Lampides 206

Larinopoda 196

lavatherae 115

lentiscella 256

Lepidochrysops 157, 158,

[209, 210, 222

Leptosia 167

leucomelas 80

Libythea 192

Lipaphnaeus 198

Liptena 195, 196, 221

Loboela 114

Lycaena 158, 204

Lyonetia 15

machacoana 46

*macrolepidella 257

*mahale 192

malvae 110

mangana 31, 89 sqq

Mashuna 158, 173

melaleuca 80

Melanitis 170

meridionalis 80

Mesoxantha 185

mespilicola 242

Metisella 158, 214

Meza 216

Microsetia 15, 17

microtheriella 17

Mimacraea 157, 195

minimella 256

minuscula 68

minusculella 261

*montana 199

Monza 218

*muricatella 266

Mylothris 158, 165, 167

Myrina 197

nanus 31,91 sqq

Neita 172

Neocoenyra 147, 172

Nepheronia 169

Nepticula 13 sqq, 240 sqq

Neptidopsis 185

Neptis 147, 181—184, 222

Netrobalane 213

nivenburgensis 248

nora 55

oberthüri 65

Oboronia 147, 210

Obrussa 15

obscura 59, 128

occidentalis 31, 87

orbifer 71

Ornipholidotos 195

Osmodes 215

osthelderi 29, 52 sqq

Oxylides 197

paliurella 242

Palla 178, 221

Papilio 158—160

Pardaleodes 215

Pardopsis 192

Parnara 221

Parosmodes 215

paula 29, 62, 74 sqq

Pelopidas 220

Pentila 193

Phalaena 15, 17

Phalanta 188

Philiolaus 200

Philodeudoryx 202

phlomidis 30, 52 sqq

Phlyaria 207

Physcaenaura 172

Pinacopteryx 167

platani 253

Platylesches 219, 220

ploetzi 31, 84 sqq

Plutella 14

poggei 107

Precis 157, 186—188

promissa 250

Pseudacraea 180

Pseudargynnis 180

Pseudathyma 178, 221

Pseudonacaduba 206

pygmaeella 263

*pyrellicola 264

Pyrgus 25, 106 sqq

quadripunctaria 269

rebeli 85

*reducta 182

rehfousi 105

rhamnophila 245

rosella 14, 15

rosmarinella 271, 272

Salamis 186

Sallya 184, 185

samiatella 245

sanaos 130

Sarangesa 212

sataspes 31, 99 sqq

secessus 32, 76 sqq

Semalea 216

*semiconfluens 55, 56, 58

[sqq

septembrella 274, 276

sericopeza 13

sertorius 29, 71 sqq

sexguttella 17

shanta 69

*similis 163

Spalgis 197

Spialia 213

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, 1978 297

Spindasis 158, 198

spio 30, 41 sqq

Stigmella 13 sqq, 242 sqq

stipella 17

stoechadella 272

struvei 52

Stugeta 199

*styracicolella 267

suberis 251

superna 68

Syntarucus 147, 158, 207,

[208

Syrichtus 106, 108 sqq

szoecsi 246

Tagiades 211

*tanganyikae 161

Tarucus 208

tauromeniella 243

Telipna 193, 222

Teniorhinus 215

terebinthivora 251

Terimomima 195

therapne 71

Thermoniphas 210

thuringiaca 266

tiliae 269

Tinea 14, 15, 17

Toxichitona 195

transvaaliae 55, 56 sqq

Trifurcula 250 sqq

*trilobella 271

trimaculella 242

trimeni 77

Tsitana 214

*ufipa 174, 203

f. ulmicola (N. ulmivora)

[243

f. ulmifoliae (N. ulmivora)

[243

ulmivora 243

Uranothauma 147, 206, 207

Vanessa 188

Vanessula 188

vindex 41

Virachola 202—204, 222

viscerella 243

Weberina 253, 256

*wranghami 160

wrefordi 31, 73 sqq

Xanthodisca 215

Xenophanes 114

xylostella 14

Ypthima 147, 172, 173

Ypthimomorpha 158, 173

298 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, 1978

zaira 80

zebra 31, 38, 85, 96 sqq

Zenonia 220

Zizeeria 208

Zizula 208

zizyphi 15

Zophopetes 217

PLANTAE

Acer tataricum 246

Amelanchier parviflora 242

Amygdalis communis 266

Brachystegia 148, 151, 156,

[157, 186

Convolvulus althaeoides

[246

Convolvulus arvensis 246

Convolvulus lanatus 53

Cotoneaster racemiflora

[242

Crataegus azarolus 263

Crataegus monogyna 263

Crataegus oxyacanthae 263

Cryptosepalum 156

Cupressus 269

Euphorbia 156

Ficus carica 279

Ficus tuerckheimii 290

Hibiscus aethiopicus 43, 63,

[93, 101

Hypericum crispum 274,

[276

Hypericum hircinum 274,

[276

Laurus nobilis 269

Melhamia 98

Myrianthus 189

Myrthus italica 269

Paliurus spina-christi 242

Pavonia columella 43

Pinus halepensis 269

Pistacia atlantica 250

Pistacia lentiscus 256

Pistacia terebinthus 251

Platanus orientalis 253,

[269

Podocarpus 156

Populus italica 242

Prunus dulcis 266 :

Pyrus amygdaliformis 261

Pyrus spinosa 261

Quercus aegilops 259

Quercus coccifera 251

Quercus ehrenbergi 245,

[250

Quercus ilex 251

Quercus macrolepis 259,

[271

Quercus pubescens 250

Quercus suber 251

Rhamnus oleoides 245

Rhamnus pyrellus 264

Rhamnus rhodopaeus 246

Rhus coriaria 250

Rubus fruticosus 240

Salix alba 248

Salvia trilobella 272

Sanguisorba muricata 267

Styrax officinalis 267

Triumfetta tomentosa 81

Triumfetta rhomboidea 81

Ulmus glabra 243

Vitex agnus-castus 253

INHOUD VAN DEEL 121

Jong, R. de. — Monograph of the genus Spialia Swinhoe (Lepidoptera, Hesperiidae) ......

Kielland, J. — A provisional checklist of the Rhopalocera of the eastern side of Lake

hangar kali Ormes ey esp ee A ONO

Klimesch, J. — Beitrag zur Kenntnis der Nepticulidenfauna von Anatolien und der Insel

Rhodos (Lepidoptera, Nepticulidae) ..........................

Lith, J. P. van. — Psenini from Sri Lanka (Hymenoptera, Sphecidae, Pemphredoninae) . . ...

Ramirez B., W. — Evolution of mechanisms to carry pollen in Agaonidae (Hymenoptera

GChalcidord ea) RE se Dame en anne A it: Wy ec RE NE oat

Wilkinson, C. — On the Stigmella-Nepticula controversy (Lepidoptera) ..............

REIT carter ee ede ries eee ARE cal, eee fet nee arene TEN Dee RO TRI CIAO DI

295

serranda

Mare

aus

ea ts

AG 0 8 1973

DEEL 121 AFLEVERING 1 AUG a8 19/0 1978

SA STS

o cd a |

FRSITY

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

J. P. van LITH. — Psenini from Sri Lanka (Hymenoptera, Sphecidae, Pemphre-

doninae), p. 1—12, text-figs. 1—16.

| Tijdschrift voor Entomologie, deel 121, afl. I Gepubliceerd 19-VII-1978 |

PSENINI FROM SRI LANKA (HYMENOPTERA,

SPHECIDAE, PEMPHREDONINAE)

J.P. VAN LITH

Allard Piersonstraat 28C, Rotterdam

With 16 text-figures

ABSTRACT

This review of the Psenini from Sri Lanka is based mainly on material collected under the auspices of

the “Biosystematic Studies of the Insects of Ceylon” project of the Smithsonian Institution,

Washington, D.C. Three new forms are described: Psenulus genalis and P. maculatus keiseri from Sri

Lanka, P. carinifrons taprobanensis from Sri Lanka and South India. Supplementary descriptions of

previously recorded species are also given.

Thus far four species were recorded from Sri Lanka (formerly Ceylon), namely

Psen (Psen) matalensis, Psen (Psen) nitidus, Psenulus ceylonicus and Psenulus nietneri,

seven specimens in total. During the recent “Biosystematic Studies of the Insects

of Ceylon” project of the Smithsonian Institution, Washington, D.C., 51 fresh

specimens were collected. In addition I was able to study 19 Psenini from various

other museums, partly received through the kind offices of Dr. K. V. Krombein,

Washington, D.C., and Prof. Dr. J. van der Vecht, Putten, Netherlands. One new

Psenulus and two new subspecies of Psenulus are now described from Sri Lanka.

The new subspecies Psenulus carinifrons taprobanensis occurs both in South India

and in Sri Lanka, seven specimens being recorded from Sri Lanka, four from the

continent.

I am grateful to the Naturhistorisches Museum, Basle (NMB; via J. van der

Vecht), British Museum (Natural History), London, C. R. Vardy (BM), National

Colombo Museum, Colombo (NCM; via K. V. Krombein and J. van der Vecht),

Idaho University, Moscow, Idaho, A. R. Gittins (IU), Lund University, Lund (LU;

via J. van der Vecht), Oregon State University, Corvallis, G. R. Ferguson (OSU),

Entomology Research Institute, Ottawa, L. Masner and C. M. Yoshimoto (CNC),

H. and M. Townes, Ann Arbor (HT) and the National Museum of Natural

History, Smithsonian Institution, Washington, D.C. (USNM) for the loan of

material. My special thanks are due to Dr. Karl V. Krombein, Principal

Investigator of the project, for entrusting the study of the Psenini to me. Mrs. C.

van Driel-Murray kindly read and corrected the English text.

2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 1, 1978

The following species are at present known from Sri Lanka:

Psen (Psen) nitidus nitidus Van Lith

* Psen (Psen) elisabethae madrasiensis Van Lith

Psen (Psen) matalensis Turner

* Psenulus genalis sp. nov.

Psenulus nietneri Van Lith

* Psenulus puncticeps (Cameron)

* Psenulus exiguus Van Lith

* Psenulus maculatus keiseri subsp. nov.

Psenulus ceylonicus Van Lith

* Psenulus pulcherrimus (Bingham)

* Psenulus carinifrons taprobanensis subsp. nov.

(9°)

OQ; Oy

BB — SAN — BH UN — w

+O Oy +O 40 +0 40 40 40 40 40 40

All these species except P. nietneri are represented in the material recently

collected for the Smithsonian Institution. Those marked with an asterisk are new

to the fauna of Sri Lanka.

Psen matalensis, Psenulus ceylonicus and Psenulus maculatus keiseri are probably

endemic. Psenulus genalis and Psenulus nietneri are known from Sri Lanka only, but

the number of specimens is too small to enable any conclusions to be drawn. Most

of the material has been collected in the southern half of the island. Only twelve

males have been taken against 65 females; the males of three species and one

subspecies, all Psenulus, are still unknown.

Psen (Psen) nitidus nitidus Van Lith

Van Lith, 1959: 28— 30 (Java, Krakatau, Bangka, Sri Lanka): 1968: 105— 106 (South India, Sumatra).

Bohart & Menke, 1976: 166.

New records from Sri Lanka: Western Province, Colombo District, I 9,

Kalatuwawa Reservoir, 300 ft, 19 Sept. 1970, O. S. Flint Jr.; 1 9, Labugama

Reservoir Jungle, 13—14 Oct. 1973, K. V. Krombein, P. B. Karunaratne, P.

Fernando, J. Ferdinando: | ©, Labugama Reservoir, 16 Febr. 1975, K. V.

Krombein, P. B. Karunaratne, P. Fernando, S. Karunaratne (USNM). Central

Province, | ©, Katugastota, 1600 ft, 31 Aug. 1967, P. B. Karunaratne (CNC);

Kandy District, | ©, Kandy, Peak View Motel, 1800 ft, 15—24 Jan. 1970, Davis

and Rowe (USNM); I g, Kandy, Udawattekelle Forest, 23 Febr. 1974, P. J.

Chandler (BM); Matale District, 1 ©, Enselwatta, above 2500 ft elevation, 19—20

Oct. 1976, G. F. Hevel, R. E. Dietz, S. Karunaratne, D. W. Balasooriya (USNM).

Southern Province, Galle District, 2 © and 2 g, Kanneliya Jungle, 11—16 Jan.

1975, one of the & taken at white light, K. V. Krombein, P. B. Karunaratne, P.

Fernando, N. V.T. A. Weragoda (USNM).

Bingham (1897) recorded a female from Pundaloya which he identified as Psen

orientalis Cameron but which was later (Van Lith, 1959) recognized as P. nitidus.

The occurrence of the species in Sri Lanka has now been amply confirmed.

The nominate subspecies is also known from South India and has been found as

far eastward as Java. There seems to be no difference between the material from

VAN LITH: Psenini from Sri Lanka 3

Sri Lanka and that from Malang, Java, the type-locality. The labrum and the

greater part of the mandibles are reddish in the female only, in the male these

parts are dark.

Psen (Psen) elisabethae madrasiensis Van Lith

Van Lith, 1965: 31—32 (Psen (Psen) elisabethae subsp.); 1968: 107 (Psen (Psen) elisabethae madrasien-

sis: South India, Madras State). Bohart & Menke, 1976: 166.

First record from Sri Lanka: Sabaragemuwa Province, 1 9, Belihuloya,

Ougaldown Estate, 4000 ft, 6 Sept. 1928 (NCM).

The petiole of this female is 5.8 times as long as wide in the middle. In the

females from Madras — Nilgiri Hills and Anamalai Hills — the petiole 1s about

five times as long as broad.

The nominate subspecies is found in Java and Sumatra.

Psen (Psen) matalensis Turner

(figs. 1—3)

Turner, 1912: 362—363. Van Lith, 1965: 55-56. Bohart & Menke, 1976: 166.

Two females and one male have earlier been recorded from Sri Lanka, Matale,

2000 feet (Turner, 1912; Van Lith, 1965).

New records from Sri Lanka: Central Province, Kandy District, Kandy,

Udawattakele (Sanctuary), 2100 ft, 1 ©, 11 Febr. 1975, No. 21175A, K. V.

Krombein; 10 ©, 9—13 Febr. 1975, K. V. Krombein, P. B. Karunaratne, P.

Fernando, S. Karunaratne; 8 9, 5—15 July, 20—30 July and 2—13 Aug. 1976, S.

Karunaratne; Udawattakele, 1700 ft, 2 9, 29—30 May 1976, 1800 ft, 3 9, 3—5

June 1976, K. V. Krombein, P. B. Karunaratne, S. Karunaratne, D. W. Balasooriya

(USNM); Nuwara Eliya District, 1 9 and 2 g, Nuwara Eliya, 25 April 1923; 2 9,

Nuwara Eliya, Elk Plains, 5 May 1923 (NCM); | g, Kanda-ela Reservoir, 5—6 mi

southwest Nuwara Eliya, 6200 ft, 10—21 Febr. 1970, Davis and Rowe; | 9, Ohiya,

5500 ft, | June 1976, K. V. Krombein, S. Karunaratne, D. W. Balasooriya

(USNM): I ©, Hakgala, 9 May 1927 (NCM). Sabaragamuwa Province, Balangoda,

1 9,7 June 1935 (NCM). Southern Province, Galle District, 1 9, Kottawa Forest

Reserve, Hiniduma, 11 March 1972, K. V. Krombein and P. B. Karunaratne; 2 9,

Kanneliya Jungle, 13—16 Aug. 1972, K. V. Krombein and P. B. Karunaratne

(USNM). 1 g, without data (NCM).

Female. — Extent of red colour on gaster somewhat variable. Hind margin of

first tergite usually red on apical sixth of segment. Second tergite with large lateral

red marks and red sides, in one case almost entirely red. Second sternite entirely,

or basal %, red. Ventral plate of petiole entirely red or with red margin only. In

fresh specimens legs deep black except for fore and mid tarsi and foreside of fore

and mid tibiae. In some of the females second recurrent vein of fore wings

interstitial instead of ending in third submarginal cell. In one case this vein

interstitial in one wing only. Petiole slightly over 1% times length of first tergite, in

dorsal aspect.

4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 1, 1978

1 2

Figs. 1—3. Psen (Psen) matalensis Turner, &, genitalia, dorsal, posterior and lateral aspect.

Male. — At least fore and mid legs more reddish-brown than in female. In one

of the two males from Nuwara Eliya second tergite almost entirely red. Sternites

3—4 apically with golden-brown fasciculate hairs.

Genitalia (figs. 1—3) pale brown; stipes long, no distinct membraneous

extension on inner side of apical part, volsellae long.

This seems to be an endemic species.

Psenulus genalis sp. nov.

(figs. 4—9)

Female. — Length about 5 mm. Head and thorax black; mandibles before apex

reddish, palpi yellowish-brown. Underside of antennae including scape yellowish-

brown, dorsal side dark brown. Femora except apices black, fore and mid tibiae

and tarsi yellowish-brown, mid tibiae of paratype with black streak on outer side.

Hind tibiae black with yellowish-brown base, tarsal segments 2—4 brown. Petiole

including ventral plate, first tergite except for narrow reddish hind margin and

sternites 3—6 or 4—6 black, remaining parts of gaster dark red. Veins of wings

blackish.

Clypeus dull, broad anterior margin depressed and shining, medially narrowly

protruding, not distinctly emarginate or bidentate (fig. 4). Frontal carina

broadened between antennae into an elongated lozenge. This lozenge dorsally

reaches depression of anterior ocellus, ending below in a short transverse carina

(fig. 4). Frons densely rugoso-punctate, vertex punctate, interstices a few times

size of punctures. Occipital carina ending below in hypostomal carina, dorsally

strongly crenulate (fig. 5). Sides of head with sharp oblique carina, originating on

vertex and ending near mandibles (fig. 6). Genae with sharp transverse carinae on

VAN LITH: Psenini from Sri Lanka 5

either side of oblique carina. Upper part of sides of head densely punctate and

striato-punctate. Antennae short, clavate, third segment almost 1% times as long

as broad at apex, segments 4—8 about quadrate, segments 9—11 broader than

long, segment 12 more than 1% times as long as broad at base.

Pronotal corners rounded. Scutum shining, distinctly punctate, interstices from

once to twice as large as punctures. Prescutal sutures on posterior half not sharply

defined, almost reaching hind margin. Scutellum shining, sparsely punctate.

Metanotum finer, more densely punctate. Enclosed area of propodeum shining,

much concave with a few lateral oblique carinae (fig. 7), central part wide.

Propodeum dorsally with irregular, very fine oblique striae, lower half more

coarsely irregularly reticulato-carinate. Longitudinal groove on back of propo-

deum reduced to two large alveoles, which are distinctly separated. Propodeum

postero-laterally with broad alveolate band. Sides of propodeum posteriorly

coarsely reticulato-carinate, anteriorly finely striato-punctate. Mesopleura

shining, finely punctate, interstices slightly smaller to slightly larger than

punctures. Mesosternum laterally somewhat rugoso-punctate, medially with short

transverse rugae. Anterior plate of mesepisternum weakly rugoso-punctate.

Anterior oblique suture foveolate, widened dorsal part with a few transverse

carinae. Second submarginal cell dorsally open, upper side about ?/, or '/, of lower

abscissa. First recurrent vein ending in second submarginal cell near RS, second

recurrent vein ending well in third submarginal cell (fig. 8). Petiole almost

cylindrical, at least 1% times as long as first tergite in dorsal aspect, sides slightly

flattened, anteriorly a short latero-dorsal carina. First tergite much convex in

lateral view (fig. 9). Tergites shining, minutely punctate, no distinct pygidial area.

Sternites slightly stronger punctate.

Pubescence whitish, longer on vertex, on face and pronotum more silvery,

mostly appressed. Margins of sternites 2—3 with very short hairs, margins of

sternites 4—5 with some long, stiff, whitish hairs. Sixth sternite densely pale golden

pubescent. Epicnemial areas below with patch of dense, short, yellowish

pubescence. Petiole with latero-dorsal row of very fine and very short hairs,

laterally and ventrally a few long erect hairs.

Male unknown.

Sri Lanka: 19, holotype, Northern Province, Mannar District, 0.5 mi northeast

Kokmotte Bungalow, Wilpattu National Park, 21—25 May 1976, K. V. Krombein,

P. B. Karunaratne, S. Karunaratne, D. W. Balasooriya; | ©, paratype, Southern

Province, Hambantota District, Palatupana Tank, 3—4 Febr. 1975, Malaise trap,

K. V. Krombein, P. B. Karunaratne, P. Fernando, E. G. Dabrera (USNM).

P. genalis is closely related to P. yoshimotoi Van Lith (1969) from Borneo, which

has the same oblique carina on the sides of the head. However, its petiole has no

dorsal carinae and the second submarginal cell is not triangular.

Psenulus nietneri Van Lith

Van Lith, 1972: 167—168, 9 (Sri Lanka).

The single female known is labelled ‘“‘Ceylon, coll. Nietner” (ZMB), no further

data. Closely related to P. puncticeps. No fresh material collected.

6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 1, 1978

Figs. 4—9. Psenulus genalis sp. nov., 9, holotype, 4—6: head in frontal, dorsal and lateral aspect: 7,

propodeum, dorsal aspect: 8, second and third submarginal cells of fore wing: 9, petiole and first

gastral tergite.

Psenulus puncticeps (Cameron)

Cameron, 1907: 91 (Psen puncticeps: Bombay, India). Rohwer, 1923: 595—596 (Diodontus antennatus:

Singapore). Van Lith, 1962: 44—46 (Psenulus antennatus; Malaya, Java, Bali): 1973: 136—137 (Psenu-

lus puncticeps: Nepal): 1976: 99 (Laos, Flores). Bohart & Menke, 1976: 173.

First record from Sri Lanka: Northern Province, 1 ©, Vavuniya District,

Parayanalankulam Irrigation Canal, 25 mi northwest Medawachchiya, 100 ft,

20—25 March 1970, Davis and Rowe (USNM).

VAN LITH: Psenini from Sri Lanka U

This female is very similar to specimens from Java, but first gastral tergite red,

outer side of mid tibiae and base of hind tibiae more whitish than yellowish.

Antennae reddish, flagellum darkened dorsally. Vertex distinctly punctate,

between ocelli and oculi moreover distinctly striate.

P. puncticeps occurs in South India as well. I have seen a female from Nilgiri

Hills, Moyar Camp, 2900 ft, May 1954, P. S. Nathan (OSU, ex collection G. R.

Ferguson).

Psenulus exiguus Van Lith

Van Lith, 1976: 99— 101, © and g (Laos, Malaya).

First records from Sri Lanka: Northern Province, | ©, Mannar District, 0.5 mi

northeast Kokmotte Bungalow, Wilpattu National Park, 21—25 May 1976, K. V.

Krombein, P. B. Karunaratne, S. Karunaratne, D. W. Balasooriya. Eastern

Province (southern part), | ©, Amparai District, Lahugala Tank, 14—15 June

1976, Malaise trap, K. V. Krombein, P. B. Karunaratne, S. Karunaratne (USNM).

These females are somewhat more brightly coloured than the holotype

described from Laos. Pronotal tubercles yellowish-white instead of brownish-

yellow. Fore and mid tibiae not yellowish-brown but distinctly yellowish-white on

outer side, also basitarsi yellowish-white. Hind basitarsus yellowish-brown, tarsal

segments 2—4 dark brown. Petiole including ventral plate, greater part of first

tergite and of fourth sternite black. Petiole more slender. Length 5.5—6 mm.

P. exiguus is very similar to puncticeps but is much smaller (puncticeps length

7—7.5 mm) and its gaster is more slender.

Psenulus maculatus keiseri subsp. nov.

(figs. 10—11)

Female. — Length about 8 mm. Head black; mandibles except for tips, scape of

antennae except for small black spot dorsally, underside of pedicel and of

flagellum yellow, apical half of flagellum somewhat reddish below. Flagellum

dorsally dark brown. Thorax (fig. 10) black, following parts yellow: pronotum

dorsally, pronotal tubercles, tegulae partly, two small narrow marks along tegulae,

two triangular marks in front of scutellum, greater part of axillae, metanotum, four

large oblong marks on back of propodeum (fig. 11), a small mark in upper corner

of anterior plate of mesepisternum. Central part of scutellum reddish in holotype,

black or with small irregular yellow marks in paratypes; postero-lateral margins of

scutellum more or less yellow. Basal % of fore and mid femora black, at least on

posterior surface, on foreside at least a narrow yellow streak; trochanters, tibiae

and tarsi of fore and mid legs yellow. Hind trochanters mostly black, dorsal half of

femora black with reddish median line above, lower half of femora yellow. Hind

tibiae and tarsi reddish-brown, underside of tibiae slightly darkened in holotype,

more distinctly brown in paratypes which have also bases of hind tarsal segments.

brown. Petiole with yellowish-red base, apical half brown in holotype, almost

black in paratypes. Gastral segments reddish, base of first tergite dorsally brown.

8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 1, 1978

Figs. 10—11. Psenulus maculatus keiseri subsp. nov., @, holotype, thorax, dorsal aspect, propodeum,

posterior aspect. Figs. 12—15. Psenulus cevlonicus Van Lith, 9. 12—13, usual colour form, thorax,

dorsal aspect, mesopleura, lateral aspect: 14—15, same parts of paler form from Uda Walawe.

Clypeal margin distinctly bidentate. Interantennal carina sharp, a transverse

carina below antennae. Frons and vertex very finely punctate, scutum, scutellum

and mesopleura finely punctate, interstices mostly a few times size of punctures.

Base of hind tibiae with three long spines on outer side. First recurrent vein

VAN LITH: Psenini from Sri Lanka 9

ending in first, second recurrent vein ending in third submarginal cell.

Pubescence on face and temples silvery, mostly appressed, on vertex and on

thorax whitish.

Male unknown.

Sri Lanka: Southern Province, 1 9, holotype, Tanamalwila, 7 Jan. 1954, F.

Keiser (NMB), | ©, paratype, Katagemuwa, 26 March 1935 (NCM). Northern

Province, Mannar District, 1 ©, paratype, 0.5 mi northeast Kokmotte Bungalow,

Wilpattu National Park, 21—25 May 1976, 1 9, paratype, same locality, Malaise

trap, 22—25 May 1976, K. V. Krombein, P. B. Karunaratne, S. Karunaratne, D.

W. Balasooriya (USNM).

P. maculatus keiseri is easily recognized as a representative of the tuberculifrons

group because of the three long spines on the base of the hind tibiae. It shows the

same structure and sculpture as the nominate form from Malaya (Van Lith, 1962).

The colour-marking, however, is somewhat different. The underside of the

antennae is completely yellow or yellowish-red, at most the underside of the last

segment is darkened. The axillae are more yellow whilst the fore and mid femora

are darker than in the nominate subspecies. As in the latter form, the yellow

marking of the scutellum is subject to variation. The base of the petiole is much

paler.

Psenulus ceylonicus Van Lith

(figs. 12—15)

Van Lith, 1972: 195 (Sri Lanka; Kandy). Bohart & Menke, 1976: 172.

New records: Sabaragamuwa Province, 2 9, Belihuloya, 15—20 July 1968, P. B.

Karunaratne (CNC); | 9, Ratnapura District, Uda Walawe, 300 ft, in scrub-thorn

forest, 1 Aug. 1973, Malaise trap, Ginter Ekis (USNM). Western Province, | 9,

Colombo District, Padukka-Ingiriya Road, Kuruna Timber Reserve, 18 Jan. 1975,

K. V. Krombein, P. B. Karunaratne, P. Fernando, N.V.T.A. Weragoda (USNM).

In the female from Uda Walawe the yellow marking on the scutum is more

extended (fig. 14). Moreover, a large yellow mark behind the anterior oblique

suture is connected with a smaller yellow mark along the hind margin of the

mesopleuron (fig. 15). The usual form is somewhat darker (figs. 12—13). As a rule

the first recurrent vein of the fore wings is about interstitial; in the female from

Kuruna Timber Reserve this vein ends distinctly in the first submarginal cell. Hind

margin of sternites 4—5 densely yellowish pubescent.

Male unknown.

P. ceylonicus was described after two females from Kandy and is probably

endemic. It is the most western representative of the group of P. interstitialis,

which group extends eastwards to the Solomon Islands.

Psenulus pulcherrimus (Bingham)

(fig. 16)

Bingham, 1896: 443 (Psen pulcherrimus: Tenasserim). Van Lith, 1962: 101 (Psenulus pulcherrimus): 1969:

10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 1, 1978

200 (Vietnam); 1973: 140—141 (Nepal); 1976: 115—117 (Psenulus pulcherrimus pulcherrimus; Laos,

Thailand, Malaya, Singapore). Bohart & Menke, 1976: 173.

First record from Sri Lanka: Southern Province, Galle District, 1 ¢, Kanneliya,

200 ft, black light, 15—17 Oct. 1976, G. F. Hevel, R. E. Dietz, S. Karunaratne, D.

W. Balasooriya (USNM).

This male is somewhat darker than the males of the nominate form I have seen

from Nepal and southeastern Asia.

Marks along tegulae small. Posterior mark on scutum almost divided into two

square marks. Scutellum black except for narrow yellow hind margin. Marks on

back of propodeum reduced to two narrow oblong marks close to median

longitudinal groove (fig. 16).

Fig. 16. Psenulus pulcherrimus (Bingham), &, Sri Lanka, thorax, dorsal aspect.

Apical % of petiole and ventral plate of petiole blackish; narrow hind margin of

ventral plate reddish. Base of first tergite blackish-brown, bases of sternites dark

brown. Fore and mid femora dark brown on their posterior surface, hind femora

almost entirely black, apical % of hind tibiae and hind tarsi dark brown.

Punctation of scutum rather strong. First recurrent vein of fore wings ending in.

first submarginal cell, very close to RS, second recurrent vein ending well in third

submarginal cell.

Although the venation of the fore wings differs from the usual course, I believe

this specimen has been correctly identified. As the series from Nepal and Laos

show much variation in colour pattern, I hesitate to name the present material as a

distinct subspecies. It is easily distinguished from the closely related P. leucogna-

thus from South India (Van Lith, 1976).

Psenulus carinifrons taprobanensis subsp. nov.

Female. — Resembling nominate subspecies. Length about 7 mm. Basal % of

second gastral tergite either entirely red or with two separate red spots. Fore and

mid trochanters yellow, fore and mid femora yellow except for basal half which is

black. Fore and mid tibiae and tarsi yellow, last segment of mid tarsi somewhat

VAN LITH: Psenini from Sri Lanka Il

brownish. Hind trochanters and femora black, basal ¥ of hind tibiae in dorsal view

yellow; hind tarsi brown. Scape and underside of second antennal segment yellow,

following segments blackish-brown above, reddish-brown below.

Propodeum smooth behind enclosed area, back shining, very finely punctate,

dorso-laterally some fine or indistinct striation.

Male. — Gaster black, sometimes some indistinct reddish marking on base of

second tergite. Legs as in female but basal % of hind tibiae yellow in dorsal aspect.

Hind basitarsi yellowish, or pale yellowish-brown, tarsal segments 2—4 pale

brown, last segment brown.

Punctation of scutum coarser than in female. Declivous part of propodeum

rather coarsely reticulato-carinate.

Sri Lanka: Western Province, | ©, holotype, Yakkala, 18 mi NE Colombo,

locality 11, indoors, I—15 Febr. 1962; 2 3, allo- and paratype, Madinnagoda, 4 mi

east Colombo, at light, 15 Febr. 1962, A. Perera, all Lund University Ceylon

Expedition 1962, Brinck-Andersson-Cederholm (LU); 1 9, paratype, Colombo

District, Labugama, 400 ft, 9 May 1976, K. V. Krombein, P. B. Karunaratne, S.

Karunaratne, D. W. Balasooriya (USNM). Central Province, 1 9, paratype,

Ambacotta, 14 Dec. 1953, F. Keiser (NMB); | &, paratype, Kandy District,

Kandy, Udawattakele Sanctuary, 2100 ft, 16—31 Aug. 1976, S. Karunaratne

(USNM). Southern Province, | 9, paratype, Galle District, Udagama, Kanneliya

Jungle, 400 ft, 6—12 Oct. 1973, K. V. Krombein, P. B. Karunaratne, P. Fernando,

J. Ferdinando (USNM).

South India: Madras State, Coimbatore, | &, paratype, Sept. 1955, P. S. Nathan

(Ferguson collection, OSU) (earlier recorded as P. carinifrons subsp. nov.?, Van

Lith, 1966); 1 &, paratype, 1400 ft, April 1962 (CNC); | &, paratype, 1400 ft, Aug.

1972, T.R.S. Nathan (HT). Pondicherry State, Karikal, 1 G', paratype, Febr. 1962,

P.S. Nathan (IU).

This subspecies differs from P. carinifrons malayanus Van Lith, 1969, another

subspecies with black bases of fore and mid femora, in the reddish marked gaster

of the female. Thus far this character appears to be a constant one. The hind

basitarsi of the male of malayanus seem to be darker. The remaining known

subspecies of P. carinifrons, including the nominate subspecies, have entirely

yellow fore and mid femora.

P. carinifrons taprobanensis much resembles P. xanthognathus centralis from

Palawan and Mindanao. This latter form also has a rather flat and finely punctate

frons, but the dorsal side of fore and mid trochanters is brown whilst the yellow

marking of pronotum and scutellum is often reduced.

REFERENCES

Bingham, C. T., 1896. On some exotic fossorial Hymenoptera in the collection of the B. M. with des-

cription of new species and a new genus of Pompilidae. — JI. Proc. Linn. Soc. Zool. 25:

422—445.

—— 1897. The fauna of British India, including Ceylon and Burma, Hymenoptera, wasps and bees

1: 1—597.

Bohart, R. M., & A. S. Menke, 1976. Sphecid wasps of the world: 1—695. — University of California

Press, Berkeley, Los Angeles, London.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 1, 1978

Cameron, P., 1907. A contribution to the knowledge of the Hymenoptera of the Oriental zoological re-

Lith, J.

gion. — Ann. Mag. nat. Hist. (7) 20: 81—92. .

P. van, 1959. Contribution to the knowledge of the Indo-Australian Pseninae (Hymenoptera,

Sphecidae). — Zool. Verh. Leiden 39: 1—69.

, 1962. Contribution to the knowledge of the Indo-Australian Pseninae (Hymenoptera, Spheci-

dae), Part II. Psenulus Kohl, 1896. — Zool. Verh. Leiden 52: 1—118.

, 1965. Contribution to the knowledge of the Indo-Australian Psenini. Part III. New species of

the subgenera Psen Latreille and Mimumesa Malloch and a review of East-Asiatic and Indo-

Australian Psen Latreille, s.l. — Zool. Verh. Leiden 73: 1—80. |

, 1968. Contribution to the knowledge of Indo-Australian, South Pacific and East Asiatic Pseni-

ni. Genus Psen Latreille (Hymenoptera, Sphecidae). — Tijdschr. Ent. 111: 89—135..

, 1969. Descriptions of some Indo-Australian Psenulus and revision of the group of Psenulus pul-

cherrimus (Bingham) (Hymenoptera, Sphecidae, Psenini). — Tijdschr. Ent. 112:.197—212.

, 1972. Contribution to the knowledge of Oriental Psenulus (Hymenoptera, Sphecidae, Psenini).

— Tijdschr. Ent. 115: 153—203.

, 1973. Psenini from Nepal (Hymenoptera, Sphecidae). — Tijdschr. Ent. 116: 123—143.

, 1976. New species and records of Indo-Australian Psenini (Hymenoptera, Sphecidae, Pemph-

redoninae). — Tijdschr. Ent. 119: 79—122.

Rohwer, S. A., 1923. New Malayan Wasps of the subfamily Pseninae. — Phil. Jl. Sc. 22: 593—601.

BE 0-7 5082

DEEL 121 AFLEVERING 2 ;

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

C. WILKINSON. — On the Stigmella-Nepticula controversy (Lepidoptera), p. 13—22,

pls. 1—2.

Tijdschrift voor Entomologie, deel 121, afl. 2

Gepubliceerd 19-VII-1978

ON THE STIGMELLA-NEPTICULA CONTROVERSY

(LEPIDOPTERA)

CHRISTOPHER WILKINSON

Biological Laboratories, Free University, Amsterdam

With two plates

Ever since the genus Stigmella was first described by Schrank in 1802 and

Nepticula by Von Heyden in 1843, both generic names have been in common use

for the same group of leaf-mining micro-lepidoptera. This paper presents the

case for and against each name, but since it is clear from the International Code of

Zoological Nomenclature (1964) (subsequently referred to as the Code) which is

correct, it is also a plea for conformity in future amongst leaf-miner specialists.

Monographs have been completed on the moths of this genus occurring in

Canada and USA and are in preparation for Europe and South Africa. These

publications, together with the leaf-miner volume of the Moths of North America,

will adopt the recommendations made here.

My co-workers in this field concur with the principles of this paper and include

Dr. Don. R. Davis, Chairman Department of Entomology, National Museum of

Natural History, Smithsonian Institution, Washington D.C., USA, and my former

research assistants M. J. Scoble, Department of Entomology, Transvaal Museum,

Pretoria, South Africa and Philip J. Newton, Entomologist, Perifleur Ltd.,

Rustington, UK.

Although the name Stigmella (Schrank, 1802: 169) is older, Nepticula Heyden

(1843: 208) has been used for generic descriptions twice as often during the past

134 years. The great majority of described species were originally combined with

Nepticula.

In the early days Nepticula was probably used in ignorance of Stigmella, but

latterly attempts have been made to discredit Stigmella on the grounds that it is a

nomen dubium (e.g. Borkowski, 1972; Lempke, 1976) or a nomen nudum (e.g.

Busck, 1913; Braun, 1917).

Heyden’s description of Nepticula is very much more satisfactory and leaves one

in no doubt regarding the group of insects to which the name refers. He further

establishes the genus on a sound basis by assigning to it five species: aurella F.,

argentipedella Z., centifoliella Z., sericopeza A., and cursoriella Hdn. Unless

otherwise stated the first species cited was often taken as type and aurella was

subsequently designated type-species by Tutt (1899: 184).

This contrasts with the older Stigmella, the description of which is rather

inadequate and no species are cited. The German description (Plate 1) and English

translation are given because of their importance.

14 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 2, 1978

Schrank, 1802: 169

“Edelmotte. Fuhlhorner: borsten-

formig.

Stigmella. Schnauzen: fehlen.

Zunge: eingerollt.

Flugel: die obern

schmal, eingerollt.

Korper: Schmachtig.

Raupe: nackt.

sehr

Ich meyne, dass die mir nicht hinlang-

lich bekannte Motte, welche die

Rosenblatter gangweise minirt, hieher

gehore.

Uebrigens bedurfen alle diese Motten-

bestimmungen, die nur nach getrock-

neten Stucken gemacht sind, noch der

Berichtigungen nach frischen und wo

möglich, selbst erzogenen Stücken.

Translation

“Adult. Feelers: bristle-shaped.

Stigmella. Snouts: missing.

Tongue: rolled

Wings: rolled, the upper

ones very narrow.

Body: slender

Caterpillar: naked.

I suppose that the moth, not sufficiently

known to me, which causes gallery mines

in the leaves of Roses, belongs here.

Further, all these moth diagnoses,

which have been made only on dried

specimens, ought to be corrected

where possible on fresh material, and

where possible on bred specimens”.

The nearest Schrank goes to including a species here is his comment that the

moth causes gallery mines on rose leaves. At least he categorically states that it is a

moth and also refers to upper wings; therefore those who have said that the

description might equally refer to a dipteran leaf-miner are out of order. Thirty

pages earlier (page 139) Schrank, however, does name and describe a leaf-miner

which mines roses: Tinea rosella. He does not specifically link rosella to Stigmella

in print, but Walsingham’s paper (1907: 1008) is of paramount importance in this

connection. He says that the rose miner referred to in Schrank’s generic

description was the one he described earlier as T. rosella, and Schrank intended to

cite the species name under the genus but somehow it was overlooked. This

assertion looks highly probable. Plate 1 shows the relevant page of Schrank’s

generic description in the British Museum (Natural History) copy of “Fauna

Boica”. The hand written addition was made by Walsingham to show how Schrank

intended it to read. With the amendment it now conforms to Schrank’s usual

layout — compare with the previous description of Plutella xylostella. Whether

Walsingham knew somehow that Schrank had left out the type citation accidently

or simply assumed it, does not matter a great deal, since he has, in effect, assigned

a type-species by subsequent designation to an available genus and thus validated

the use of Stigmella once and for all. It must be appreciated that from the point of

view of nomenclature Srigmella Schrank is an available generic name originally

without associated nominal species; and the first such nominal species that were

subsequently and expressly referred to it are to be treated as the originally

included nominal species. (See Code, Articles 69 (a) and 69 (a)(ii) which provides

for the subsequent designation of type-species and Articles 11, 12 and 16 especially

11 (ci).

WILKINSON: Stigmella-Nepticula 15

Actually the first usage of Stigmella after Schrank, 1802, was by Oken (1815:

655, 677) and, in fact, he included the species T. rosella in the genus, but this work

has been rejected for the purposes of nomenclature by the International

Commission on Zoological Nomenclature, 1956, Opin. Decl. Int. Comm. Zool.

Nom. 14 Opinion 417: 3. The next usage of Stigmella seems to have been

Walsingham (1907) who not only included Phalaena anomalella Goeze, 1783 (=

rosella Schrank, 1802), but designated the former as the type-species. So whether

we like it or not, Stigmella is therefore valid and has priority over Nepticula. The

only legitimate way for the proponents of Nepticula to further their cause is to

make a case to the International Commission for the supression of Stigmella. Until

this body rules otherwise Stigmella should be used as the valid name for the taxon,

if Stigmella and Nepticula are considered to be congeneric.

Plate 2 is Schrank’s original account of T. rosella which, as can be seen, suggests

that he is providing a name for the species first described by DeGeer in 1752.

DeGeer’s account is very thorough and provides more information than is known

for the majority of leaf-miners. It includes figures of the moth, pupa, cocoon, larva

and leaf mines (see Plate 30 Figure 20 and Plate 31 Figures 13—21), but predates

Linnaeus’ tenth edition of Systema naturae (1758). Although DeGeer refers to his

species again in 1771 (p. 495), it was Goeze who first named it in 1783 (p. 168) as

Phalaena (Tinea) anomalella. Thus Schrank’s name T. rosella is a junior synonym of

anomalella. Therefore, anomalella is the legitimate type-species for Stigmella. Tutt

(1899: 209) was the first to synonymise these names. There can be no doubt that

Schrank and Goeze are referring to the same taxon because both cite DeGeer’s

rose leaf-miner as the insect they are naming.

It has already been suggested that Stigmella was not used in the early days

because of ignorance of its existence. In fact, Heyden did not know of the work of

DeGeer, Schrank or Goeze. If he had he would at least have included

anomalella/rosella in his publication, and indeed would probably never have

described Nepticula. In the early nineteenth century Tinea, Microsetia and Lyonetia

were used for Stigmella species. Even after Walsingham (1907) Stigmella has not

been used a great deal, although it is not true to say, as does Johansson (1971), that

Stigmella was lost and forgotten for 132 years. Between 1802 and 1934, the period

referred to, the name has been included in four significant contributions. In 1911,

Walsingham described Stigmella zizyphi (1911: 190), and Hampson (1918: 336) and

Fletcher (1929: 210) provided species lists under the name Stigmella. Johannson

attributed the reintroduction of Stigmella to Fletcher and wrongly cited his 1929

publication as 1934, also overlooking Walsingham’s papers (loc.cit.). Since that

time Stigmella has been used in at least six substantial works so any suggestion that

it is to be suppressed as a nomen oblitum is not tenable.

Until now Nepticula has always been used in North America, sometimes even for

Ectoedemia and Obrussa species (e.g. Freeman, 1962: 522). Busck (1913: 103) and

Braun (1917: 163) argued that Stigmella was a nomen nudum on the grounds that

there were no included species, and no-one appears to have questioned their

decision since. As previously pointed out the Code states that a uninominal genus-

group published before 1931 without associated nominal species is acceptable for

16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 2, 1978

nomenclatorial purposes. It also provides for the subsequent designation of a type-

species.

Janse (1948: 159) in South Africa adopted the name Nepticula, but his successor

Vari (1955: 331, 1963: 66) switched to Stigmella.

Recently in Europe Nepticula has been used in preference to Stigmella by

Klimesch (1951: 4), Johannson (1971: 241) and Borkowski (1972: 702). Klimesch

does not seem to like using Stigmella on the grounds that there are no included

species and objects to the citation of anomalella (Goeze) believing centifoliella

(Zeller) to be just as applicable to Schrank’s description.

Johannson’s and Borkowski’s preference for Nepticula is also based on

supposed inaccuracies in Schrank’s description. Johannson thinks that the moths

in the Nepticulidae differ from the description of Stigmella. He points out that the

palps are present and well developed whilst Schrank says they (Schnauzen) are

absent. I wonder if Schrank meant that they were absent in DeGeer’s figure or

were missing in his own material. Johannson is not right in saying that the tongue

is absent, nor Borkowski in saying that it is vestigial. Schrank described it (Zunge)

as rolled, and indeed it is present but minute and apparently functional (Downes,

1968).

Secondly Johannson says that “In the case [where] description and type

disagree the written definition is decisive.” This is not necessarily so. Article 61 of

the Code indicates that the type is the objective definition of a taxon and does not

change, whereas the limits of the taxon are subjective and liable to change. Each

case must be taken on merit.

Borkowski does not subscribe to the view that Stigmella is a nomen nudum, but

contests it on similar grounds to Johannson and says that Schrank’s second

paragraph (italics, page 14) is conjecture on Schrank’s part, and cannot be

taken as definitive. Indeed he believes the rose miner, or at least the Nepticulidae

leaf-miners, do not have characters agreeing with the generic diagnosis of

Stigmella. Thus he rejects Stigmella as a nomen dubium on the grounds that it is

not attributable to any known taxon. None of the early authors seem to have had

any doubt, and in any case, Stigmella has never been used to refer to any taxon

other than these leaf-miners. Schrank’s description of rosella, together with its

reference to DeGeer’s paper, and Walsingham’s designation of a type-species no

longer makes a nomen dubium admissable.

It is certainly true that Nepticula has been better defined and substantiated than

Stigmella, and these authors have rightly highlighted some of the problems.

However, they have all either been unaware of Walsingham’s (1907) paper or have

failed to realise its importance in validating Stigmella. To discard Stigmella stands

in contradiction to the rules of the Code and we are not free to select which name

we please.

It is my current view that Stigmella and Nepticula are congeneric. In this

circumstance, and in view of the foregoing evidence, I recommend that Stigmella

Schrank, 1802, be universally accepted for this group of leaf-miners and that

Nepticula Heyden, 1843, be reduced to a junior subjective synonym. The type-

species of Stigmella is anomalella Goeze, 1783.

WILKINSON: Stigmella-Nepticula 17

Turning now to the family name, prior to 1854 these leaf-miners were either

misplaced in other families, or simply referred to as the Nepticulae. Stainton

(1854a: 295, 1854b: 166) seems to have been the first person to give this group a

family name: Nepticulidae. Stigmellidae was not used until it was adopted by

Hampson in 1918: 387, as the family name based on the oldest genus, a procedure

which would have been correct in his day. However, applying the present Code

and the principle of priority to family-group names, we must adopt Nepticulidae.

For a variety of reasons, and sometimes the wrong ones, the name Nepticulidae

has met with greater universal acceptance than the generic name.

Following this, the use of Nepticuloidea as the superfamily name is the logical

consequence and has been demonstrated by Braun (1923: 17), Karsholt and

Schmidt Nielsen (1976: 17) and others.

Some of the principal users of Stigmellidae as the family name have been

Hampson (1918), Fletcher (1929), Beirne (1945), Vari (1963), also Klimesch (1948)

and Borkowski (1969, 1970) before they changed their minds about the generic

name.

. Before committing these views to print I have discussed the matter with three

entomological commissioners of the International Commission on Zoological

Nomenclature from both Europe and North America. They were unanimous in

their view that on the basis of the above evidence, Stigmella is the valid name for

the genus, and Nepticulidae is the correct family-group name.

ADDITIONAL NOTES

1. Zeller (1848: 301) cites Heyden as the author of Nepticula but calls it a new

genus. This may account for some writers giving Zeller as the author of

Nepticula (e.g. Stainton, 1854; Meyrick, 1895; Gerasimov, 1937).

2. Stainton published two relevant works in 1854. Although the preface in “List of

British Animals in the collection of the British Museum, XVI Lepidoptera”

gives 23 May, it is clear that Stainton did not expect it to be published earlier

than “Insecta Britannica, Lepidoptera Tineina VIII” which has the later preface

date of I July. This is assumed because Stainton gives in the “List”, page

numbers and references to taxa — even new species — described in “Insecta

Britannica”. The List was published on 13 August but no precise date has been

found for the other work. The List has, for some reason, often been overlooked

and therefore this problem has not always been appreciated. In this paper I give

it as 1854b to preserve the chronology which Stainton expected and until the

matter has been clarified.

3. Kirby (1897: 313) attributes Nepticula to Zeller (1848) and synonymises it with

Microsetia Stephens (1834), the older name. The problem is that Kirby gives

Nepticula microtheriella as the type of Microsetia, unaware that Westwood (1840)

had already designated a type, viz., Phalaena (Tinea) stipella Linnaeus sensu

Hübner (= Tinea sexguttella) (see Sattler, 1973: 224). Walsingham (1907: 1009)

linked this Linnaeus type with Aphelosetia (now = Elachista) but Microsetia

Stephens is now regarded as a Gelechiid genus. Tutt (1899) synonymised

Microsetia Stephens sensu Kirby with Nepticula.

18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 2, 1978

4. Beirne (1945: 197, 201) divided Stigmella into two genera, namely “Stigmella

Heyden” (wrong author given) with type-species anomalella and Nepticula

Heyden, type-species aurella Fabricius. This classification based on the male

genitalia has not been accepted by those who have made more extensive studies

(Borkowski, 1972; Emmet, 1976; Wilkinson and Scoble, in press).

5. Kloet and Hincks (1972: 2) retained Beirne’s division with Stigmella and

Nepticula as two separate genera. However, Borkowski (1972: 702) did not

agree with this and again amalgamated them under the name Nepticula. As

Stigmella is the older name it is recommended that this be reversed.

6. Although Zeller (1948) attributes the species centifoliella (see page 17) to

Heyden it is nevertheless an original Zeller description and species, and is

accepted as such in current check-lists.

7. The author of Nepticula acetosae (type-species of Johanssonia Borkowski) is

always given as Stainton 1854a: 303. However, this is not the first use of the

name. Shield, in the Zoologist (1853: 4153) writes. “Nepticula Acetosae Stainton

n.sp. larvae in the leaves of Rumex Acetosella July, October and November” in

“List of Micro-Lepidoptera taken in the vicinity of Dublin during the year

1853” and dated 12 November 1853. Stainton does not mention the species in

his paper a few pages earlier in the same journal (1853: 3952— 3960).

Considering this and note 2 it seems that the publication of Insecta Britannica

VIII must have been delayed.

ACKNOWLEDGEMENTS

I am most grateful to the following colleagues for reading the manuscript,

making suggestions and approving my conclusions on the basis of the evidence

presented:

Dr. I. W. B. Nye, Dr. K. Sattler, Department of Entomology, British Museum

(Natural History), London;

Dr. C. Sabrosky, Systematic Entomology Laboratory, U.S. Department of

Agriculture, Washington D.C.;

Dr. D. R. Davis, Chairman of Entomology, Smithsonian Institution, U.S.

National Museum of Natural History, Washington D.C.;

Mr. M. J. Scoble, Dr. L. Vari, Department of Entomology, Transvaal Museum,

Pretoria, South Africa;

Dr. E. Munroe, Biosystematics Research Institute, Ottawa, Canada.

I would also like to thank for helpful discussions and information:

Mr. S. Fletcher, Department of Entomology, British Museum (Natural History),

London;

Dr. J. Bradley, Commonwealth Institute of Entomology, London;

Col. A. M. Emmet, Saffron Walden, Essex:

Dr. D. Hardwick, Director, Biosystematics Research Institute, Ottawa, Canada;

Miss P. Gilbert, Entomology Librarian, British Museum (Natural History),

London.

WILKINSON: Stigmella-Nepticula 19

GENERIC SYNONYMY AND SELECTED REFERENCES

Stigmella Schrank, 1802: 169. Type-species by subsequent designation (Walsingham, 1907: 1008), Pha-

laena (Tinea) anomelella Goeze, 1783: 168. = Tinea rosella Schrank, 1802: 139.

Stigmella Schrank; Oken, 1815: 655, 677.

Stigmella Schrank; Walsingham, 1907: 1008.

Stigmella Schrank; Hampson, 1918: 387.

Stigmella Schrank; Fletcher, 1929: 210.

Stigmella Schrank; Gerasimov, 1937: 89.*

Stigmella ‘“Heyden”; Beirne, 1945: 197.} Incorrect author given by Beirne. Incorrectly synonymised

with Nepticula Heyden by Borkowski, 1972: 702.

Stigmella Schrank; Klimesch, 1948: 49.

Stigmella Schrank; Vari, 1955: 331.

Stigmella Schrank; Vari, 1963: 66.

Stigmella Schrank; Borkowski, 1969: 96.

Stigmella Schrank; Emmet in Heath, 1976: 213.

Nepticula Heyden, 1843: 208. Type-species by subsequent designation (Tutt, 1899: 184), Tinea aurella

Fabricius, 1755: 666 (syn. by Walsingham, 1907: 1008).

Nepticula Heyden; Zeller, 1848: 301.*

Nepticula Heyden; Stainton, 1853: 3952.

Nepticula Heyden; Stainton, 1854a: 295.*

Nepticula Heyden; Stainton, 1854b: 166.*

Nepticula Heyden; Herrich-Schaffer, 1855: 69.

Nepticula Heyden; Heinemann, 1862: 237.

Nepticula Heyden; Heinemann, 1871: 201.

Nepticula Heyden; Meyrick, 1895: 710.*

Nepticula Heyden; Tutt, 1899: 184.

Nepticula Heyden; Dyar, 1903: 545.

Nepticula Heyden; Spuler, 1910: 472.

Nepticula Heyden; Busck, 1913: 103.

Nepticula Heyden; Braun, 1917: 162.

Nepticula Heyden; Petersen, 1930: 1.

Nepticula Heyden; McDunnough, 1939: 107.

Nepticula Heyden; Beirne, 1945: 201.7

Nepticula Heyden; Janse, 1948: 159.

Nepticula Heyden; Klimesch, 1951: 4.

Nepticula Heyden; Johansson, 1971: 241.

Nepticula Heyden; Borkowski, 1972: 690.

Microsetia Stephens sensu Kirby, 1897: 313. Type-species subsequently incorrectly designated by Kirby

as Nepticula microtheriella Stainton, 1854a: 302. (see note 3). (syn. by Tutt, 1899: 184).

Johanssonia Borkowski, 1972: 702. Type-species by original designation and monotypy, Nepticula aceto-

sae Stainton, 1854a: 303. (syn. by Karsholt & Nielsen, 1976: 17 and 81, but retained as separate sub-

genus) (not syn. by Kloet & Hincks, 1972, see addenda).

* See additional note |

+ See additional notes 4 and 5

20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 2, 1978

WORKS CITED

BEIRNE, B. P., 1945. The male genitalia of the British Stigmellidae (Nepticulidae) (Lep.). — Proc. R. Ir. |

Acad. (B) 50(9): 191—218, 81 figs.

BORKOWSKI, A., 1969. Studien an Stigmelliden (Lepidoptera). Teil I. Zur Verbreitung, Biologie und |

Ökologie der Stigmelliden in den polnischen Sudeten. — Polskie Pismo ent. 39(1): 95—122, 20 |

figs.

———, 1970. Studien an Stigmelliden (Lepidoptera). Teil III. Beitrag zur Kenntnis der Stigmelliden |

fauna Polens. — Polskie Pismo ent. 40: 541—555, 29 figs.

—_——., 1972. Studien an Nepticuliden (Lepidoptera). Teil IV. Bemerkungen zur Nomenklatur und Sy- |

stematik der Familie Nepticulidae. — Ibid. 42(3): 689—709, 4 pls., 16 figs.

BRAUN, A. F., 1917. Nepticulidae of North America. — Trans. Am. ent. Soc. 43(2): 155—209, 4 pls.

ra of New York and Neighbouring States: 79—98, 10 figs. — New York.

Busck, A., 1913. Two Microlepidoptera injurious to Chestnut. — Proc. ent. Soc. Wash. 15(3): |

102—104, | fig.

Busck, A. & C., HEINRICH, 1921. On the male genitalia of the microlepidoptera and their systematic |

importance. — Proc. ent. Soc. Wash. 23(6): 145—152, 2 pls.

DEGEER, C., 1752. Memoires pour servir a l’histoire des insectes 1: 440—458, pls. 29—31. — Stockholm. |

‚ 1771. Ibid. 2(1).

Downers, J. A., 1968. A Nepticulid moth feeding at the leaf nectaries of Poplar. — Can. Ent. 100(10):

1078—1079.

Dyar, H. G., 1903. A list of North American Lepidoptera and key to the literature of this order of in-

sects. — Bull. U.S. Natn. Mus.: xix + 723 pp. — Washington.

Emmet, A. M. in J., HEATH, 1976. The moths and butterflies of Great Britain and Ireland: 343 pp. —

London.

FABRICIUS, J. C., 1775. Systema Entomologiae: 832 pp.

FLETCHER, T. B., 1929. A list of the generic names used for microlepidoptera. — Mem. Dept. Agric. In-

dia, Calcutta, Ent. Ser. 11: ix + 244 pp.

FREEMAN, T. N., 1962. A new species of Nepticula v. Heyd. on Birch (Lepidoptera: Nepticulidae). —

Can. Ent. 94(5): 522—523, 2 figs.

Gerasimov, A., 1937. Zur Systematik der Raupen von Stigmella Schrank (Nepticula Z.) und Tischeria Z.

(Lepid.). — Ent. Rundschau. 55: 89 —90.

Gorze, J. A. E., 1783. Entomologische Beyträge zu des Ritter Linné Zwölften Ausgabe des Natursy-

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Hampson, G. F., 1918. Some small families of the lepidoptera which are not included in the key to the

families in the catalogue of Lepidoptera Phalaenae. — A list of the families and subfamilies of

the Lepidoptera with their types and a key to the families. — Novit. zool. 25: 366—394.

HEINEMANN, H., 1862. Einige Bemerkungen über die Arten der Gattung Nepticula. — Wien. ent. Mo-

natschr. 6: 237— 268; 301—320.

———, 1871. Nachtrag zu den Bemerkungen über die Arten der Gattung Nepticula. — Berliner ent.

Ztschr. 15: 209— 223.

HERRICH-SCHAFFER, G. A. W., 1853-5. Schmetterlinge von Europa 5: 357.

HEYDEN, C. von, 1843. Amtlicher Bericht der Versammlung der Naturforscher zu Mainz: 208 pp.

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Nom. 31(2): 79—89.

JANSE, A. J. T., 1948. The moths of South Africa, Nepticulidae, 4 (3): 151—185, 13 pls. — Pretoria,

Transvaal Mus.

JOHANSSON, R., 1971. Notes on Nepticulidae (Lepidoptera) I. A. revision of the Nepticula ruficapitella

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WILKINSON: Stigmella-Nepticula 21

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Royal ent. Soc. Lond. 11 (2): 153 pp.

KL1MESCH, J., 1948. Zur Frage der verwandtschaftfichen Beziehungen einiger Stigmella — Arten auf

Grund des Baues des männlichen Kopulationsapparates (Lep. Stigmeliidae). — Zeitschr.

Wien. ent. Ges. 33: 49—82.

—, 1951. Zur Kenntnis der Genitalmorphologie einiger Nepticula Arten. — Zeitschr. Wien. ent.

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SCHRANK, F. von PAULA, 1802. Fauna Boica. 2(2): 412 pp. — Nürnberg.

SPULER, A., 1910. Die Schmetterlinge Europas. 2: 525 pp., 239 figs. — Stuttgart.

STAINTON, H. T. 1853. An introduction to the study of the Nepticulae. — Zoologist 11: 3952— 3960.

, 1854a. Insecta Britannica. Lepidoptera: Tineina: VIII + 313 pp., 10 pls. — London.

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STEPHENS, J. F., 1834. Illustrations of British Entomology: 436 pp. — London.

Tutt, J. W., 1899. A Natural History of the British Lepidoptera. 1: IV + 560 pp. — London.

VARI, L., 1955. South African Lepidoptera. I. Descriptions of new leafmining Tineina. — Ann. Trans-

vaal Mus. 22 (3): 331—351, 50 figs., 2 pls.

—, 1963. South African Lepidoptera. 3. Descriptions of new Stigmellidae. — Koedoe 6: 66—75, |

pl.

WALSINGHAM, T. (under Rt. Hon. Lord), 1907. Microlepidoptera of Tenerife. — Proc. zool. Soc. Lond.:

910—1028.

, 1911. Algerian Microlepidoptera. — Ent. Mon. Mag. 47 (2nd series 22): 187—193.

Westwoop, J. O., 1840. An Introduction to the modern classification of insects. Synopsis of the Gene-

ra of British Insects, 2: 1—58. — Londen.

ZELLER, P. C., 1848. Die Gattungen der mit Augendeckeln versehenen blattminierenden Schaben. —.

Linn. ent. 3: 248—343.

22 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 2, 1978

FRAME LA dark

Rofenblatt G. 1896, = = - T. rofella,

Degeer Ucbcrf, I. 4. Quart, 41. Tab. 31, Fig.

13-21. = Amma,

Wobnort: unter der Oberhaut der Rofenblitter,

welche die Raupe gangweife minirt,

Raupenzrit; Herbtt,

Amn. Die Raupe har 18 häutige Füfse, aber gar keie

ge hurnariigen B:uftiüifse.

Plate 1. Original account of Tinea rasella Schrank, 1802.

Mottenförmige Schmetterlinge, 169

Gabelmotte, Fübliörner : borftenförmig.

Plucella, Schnauzen: zwo, gabliz,

Zunge: eingerollt.

Körper : fchmächtig.

Flügel : eingerollt.

Raupe: nackt. - = © ©

Hieher gehört:

% Plutella xylottella.

Tinea xyloftella meiner Fauna n. 1854: p. 124

Edelmotte. Fablborner : borftenförmig.

Stigmella, Schnauzen: fehlen,

Zunge: eingerollt.

Flügel : die obern fehr fchmal, eingerollt,

Koper : fchmächtig,

Raupe: nackt. = = = -

Ich meyne, dafs die mir nicht hinlinglich bee

kannte Motte, welche die Rofenblätter gangweife

minirt, hieher gehöre. = T. nosdla TB.n.\8qo p 134

Uebrigens bedürfen alle diefe Mottenbeftim-

mungen, die nur nach getrockneten Stücken gee

macht find, noch der Berichtigungen nach fritchen,

und, wo möglich, felblt erzogenen Stücken,

Plate 2. Page 169 of the British Museum (Natural History) copy of Schranks “Fauna Boica”.

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LIBRARY

DEEL 121 AFLEVERING 3 AUG 281978 1978

HARVARD

UNIVERSITY

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

R. DE JONG. — Monograph of the genus Spialia Swinhoe (Lepidoptera, Hesperii-

dae), p. 23—146, text-figs. 1—-119, pls. 1—7.

Tijdschrift voor Entomologie, deel 121, afl. 3 Gepubliceerd 19-VII-1978

MONOGRAPH OF THE GENUS SPIALIA SWINHOE

(LEPIDOPTERA, HESPERIIDAE)

R. DEJONG

Rijksmuseum van Natuurlijke Historie, Leiden

With seven plates and 119 text-figures

ABSTRACT

The genus Spialia Swinhoe is distributed throughout the Afrotropical Region and in large parts of the

Palaearctic and Oriental Regions. Of the 26 species, 18 are confined to the Afrotropical Region. The

species are classified into seven species groups, which are mainly based on characters of the male and

female genitalia. Two subspecies are described as new, viz., Spialia colotes semiconfluens and Spialia

depauperata australis. A key to the species based on external characters is provided and drawings of

the male and female genitalia, photographs of the skippers and distribution maps of the species are given.

A number of characters is analysed as to their plesiomorphous or apomorphous conditions, resulting

in an outline of the evolutionary history and providing the phylogenetic arguments for the classification

adopted. Combined with known ecological changes in the past, this outline affords a means for the

reconstruction of the geographic history.

The genus is generally at home in dry open country. One species is restricted to the African forests;

it is shown that this habitat preference is apomorphous with respect to the preference for open country.

Due to the latter preference the development of the genus in Africa has been greatly influenced by the

alternation of dry and wet periods in the past causing changes in the extent and continuity of the

savanna area. Most, if not all, species existing at present originated during the Pleistocene. At various

times and to various degrees Spialia species succeeded in invading areas outside the Afrotropics, giving

rise to secondary radiation. Successful establishment in the Palaearctic was accompanied by a change

to other food plants. Only one, apparently recent, re-invasion into Africa can be recognized, viz., by

S. doris, a semi-desert species.

CONTENTS

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6. General outline of the evolution of Spialia .................................... 133

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INTERNES NS MINE Ae ANA nde rel ESE NA CE OCRA enn Oe A este e ns ees 138

1. INTRODUCTION

General

The genus Spialia has not yet been studied as a whole. This is probably due to

the fact that its members are distributed over three zoogeographic regions, viz. the

24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Palaearctic, Oriental and Afrotropical Regions. So we have reviews of the Palaearc-

tic species (e.g., Warren, 1926; Picard, 1947) and of the Afrotropical species

(Higgins, 1924; Evans, 1937). Especially the last ones are interesting as the bulk of

the species occurs in the Afrotropical Region. The last review, however, of the

Afrotropical species is 40 years old. Apart from the fact that new information has

become available, the female genitalia have never been described, so it does not

seem superfluous to give a revision of the whole genus.

Preliminary to this paper I published a paper on the Palaearctic Spialia species

(De Jong, 1974). As the variation and distribution of the Palaearctic species are far

better known and their evolutionary and geographic history can far better be

traced back than those of the Afrotropical species, a separate treatment seemed

appropriate. For the sake of completeness, however, the Palaearctic species will also

be dealt with in the present paper, although for details the reader is referred to my

earlier paper on Spialia. I may also refer to that paper for notes on the distinction

of subspecies and for methods and measurements.

The study of this genus is facilitated by the fact that many types are located in a

single collection: the types of 37 of the 59 names applying to species and subspecies

(including synonyms) are in the collection of the British Museum (Nat. Hist.), apart

from a number of types of seasonal and individual varieties.

It is attempted to give a classification that reflects the phylogenetic relationships.

In Chapter 4 the evolutionary history that forms the basis for the classification is

dealt with. It is unfortunate that many parts of Africa have still been badly explored

as to the Hesperiidae, thus making the study of the zoogeography of the genus

difficult. Moreover, literature records on the distribution cannot always be relied on,

as I found many misidentifications in the collections, even by Evans. This is un-

doubtedly partly due to the fact that good drawings of the genitalia do not exist, the

drawings by Evans only giving a vague idea that they represent genitalia. There-

fore, drawings of the male and female genitalia of all species are presented in the

present paper. Photographs have been added of the upper and underside of all

species, as I think a good photograph is more instructive than a page-long descrip-

tion. In this way I hope this paper will be a stimulus to fill the many gaps in our

knowledge.

Delimitation of the genus Spialia

The genus Spialia Swinhoe, 1913 (type species Hesperia galba Fabricius, 1793,

by original designation) comprises those black-and-white skippers that have the

median spots in spaces 4 and 5 of the fore wing upperside (if present) in line with

those in spaces 6 to 8, the median spot in space 2 of the fore wing upperside

central between the central cell spot and the median spot in space 3 or nearer the

latter, the termen of the hind wing evenly rounded without crenulation, no tibial

hair tuft on the hind legs in the male and usually no costal fold in the male.

For more information on the differences and relationship with other genera, I

may refer to a forthcoming paper on the generic classification of the Pyrgus group.

Wing markings

The wing markings of Spialia species are essentially the same as those of

DE JONG: The genus Spialia 25

Pyrgus species (see De Jong, 1972). In principle, each internervular space has

three spots, viz., a basal, median and submarginal spot. In addition, there is a basal

Figs. 1—3. Wing markings, male and female genitalia of Spialia. 1. Wing markings. bs, basal

spots; ccs, central cell spot; des, discoidal spot; ms, median spots; ss, submarginal spots. 2. Male geni-

talia, lateral aspect (left valva removed). ae, aedeagus; an, anellus; c, costa; cp, costal process; cu, cucul-

lus; f, fenestrula; fs, fultura superior; g, gnathos; s, saccus; t, tegumen; u, uncus. 3. Female genitalia;

a, ventral aspect; b, papilla analis. ap, apophysis posterior; as, area spinulosa; b, bursa; bl, basal lobe;

ds, dorsal sclerite of eighth abdominal segment; ob, ostium bursae; pa, papilla analis; vs, ventral sclerite

of eighth abdominal segment.

26 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

cell spot, a central cell spot and a discocellular spot covering the discocellular

veins. The markings may be very reduced, all basal spots may be absent and there

are no submarginal spots in spaces 8 to 12. In space 1b of the fore wing the spots

can often be seen divided into an upper and lower part, indicating the lost second

cubital vein (if we consider veins 2 and 3 branches of the first cubital vein). More-

over, in this space the median spot is double, being divided into an inner and outer

spot, which in their turn may be divided into an upper and lower part. So there

may be four median spots in space 1b of the fore wing, the relative development

of which is important in the distinction of some species. In Spialia one of these

spots is always absent.

The markings of the hind wing are simplified as compared with those of the fore

wing. The median spots on the underside of the hind wing are usually fused to-

gether and to the discocellular spot to form a more or less regular band (the median

band) parallel to the termen. In a few species these spots are separate and in a few

others the band is not parallel to the termen.

A diagram of the usual spots in Spialia is presented in fig. 1.

As the expression “median spot in space x” is unnecessarily long, it is abbreviated

in this paper to “median spot x”. The median spots in space lb of the fore wing

will be distinguished as the inner and outer median spots 1b and Ic, the latter de-

signing the upper part. Median spot la (between vein 1 and dorsum) is always

absent.

Genitalia

Male (fig. 2). Although relatively simply built, the male genitalia offer a good

means for the identification of most species and much of the relationships between

the species becomes clear when studying the male genitalia.

The uncus is rather short and entire, except in four species where it is bifid. The

apex of the uncus is pointed or inflated. The junction of the uncus with the tegumen

is often well visible as a narrow, more or less membranous suture and in some

species this suture is broadened mid-dorsally to a triangular or quadrangular

membranous area, called fenestrula (cf. Ogata et al., 1957). The gnathos is a simple

sclerotized fold or, usually, has various spined structures in the ventral part; dorsally

it is free or fused with the tegumen over some distance. The fultura superior is

smooth or finely spined and often passes imperceptibly into the central part of the

gnathos. The anellus is membranous or dorsally sclerotized and may bear strong

spines. The aedeagus is variously built, with a smooth surface or with spines, crests,

side-arms, etc., often characteristic of a species group.

The valvae have usually a little developed cucullus, that often forms a fold to

envelop partly the costal process. In some species the cucullus is spined. The most

strongly differentiated part of the valvae is the distal part of the costa that often

forms a process with spines or long hairs and is important for the distinction of

species and species groups.

Female (fig. 3). The eighth abdominal segment usually bears two dorsal sclerites,

but in a few species there is only a single sclerite. In most species the dorsal

sclerotization is free from the ventral sclerites. In principle, the ventral sclerotization

consists of two plates mid-ventrally separated by a membranous region which is

DE JONG: The genus Spialia 27

narrow near the ostium, but broadens caudally. This membranous area is often

slightly sclerotized and finely spinose in a more or less triangular patch. The spines

are especially visible along the caudal rim of the segment, where the membrane

folds inward. The slightly sclerotized spined area is called “area spinulosa” here. In

a number of species the area spinulosa is solidly sclerotized and fused with the

ventral sclerites to a single ventral sclerite. If strongly sclerotized, the area spinulosa

may lack the spines.

The ostium is situated close to the anterior edge of the eighth segment. The

narrow antevaginal region is usually membranous, but the ventral sclerites may be

fused here. The ductus bursae varies in length and is entirely membranous except

for some sclerotization near the ostium in some species. Usually it passes gradually

into the bursa copulatrix, which is without ornamentation, except in one species in

which the bursa bears a field of fine spines. The papillae anales are variously

shaped, in a number of species with a fold that partly or entirely covers the base

of the apophyses posteriores; this fold is called the “basal lobe” here.

Distribution

It is obvious that the distribution of the Spialia species is only imperfectly

known. Nevertheless, I am convinced that the distribution maps roughly indicate

the ranges of the species and additional material will mainly fill in the details

(especially in West Africa), rather than extending the known ranges.

Under the heading ”Localities” in the treatment of the species the countries in

the Afrotropical Region have been arranged from Sudan and Saudi-Arabia south-

ward through East Africa to South Africa and from there northward through

Central and Westcentral Africa to Senegal. The localities in each country have

been listed alphabetically.

Acknowledgements

I wish to express my sincere gratitude for the loan and gift of material and for

valuable information, to the following persons and institutions (with abbreviations

used in the text):

Dr. H.-E. Back (Zool. Forschungsinstitut und Museum A. Koenig, Bonn; ZMB),

L. A. Berger (Musée Royal de l’Afrique Centrale, Tervuren; MRAC), Dr. G.

Bernardi (Museum National d’Histoire Naturelle, Paris; MNHN), Dr. J. M. Burns

(National Museum of Natural History, Washington; USNM), G. Christensen

(Monemvasia, Greece), H. K. Clench (Carnegie Museum of Natural History, Pitts-

burgh; CMP), M. Clifton (National Museum, Nairobi; MN), Dr. W. Dierl (Zool.

Sammlung des Bayerischen Staates, München; ZSM), G. Ebert (Landessammlun-

gen fiir Naturkunde, Karlsruhe; LNK), B. Gustaffson (Naturhistoriska Riksmuseet,

Stockholm; NRS), Dr. H. J. Hannemann (Zoologisches Museum der Humboldt-

Universitat, Berlin; ZMHB), Dr. A. J. Hesse (South African Museum, Cape Town),

T. G. Howarth (British Museum, Natural History, London; BM), Dr. F. Kasy

(Naturhistorisches Museum, Wien), J. Kielland (Boröy, Norway), Dr. N. P. Kristen-

sen (Universitetets Zoologiske Museum, Copenhagen; ZMC), T. B. Larsen (London),

J. A. W. Lucas (Rotterdam; CL), Prof. Dr. Z. Lorkovic (Zagreb), J. H. Lourens

28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

(Amsterdam), Dr. I. Nakamura (Williamsville, USA), Dr. C. Naumann (München;

CN), Dr. M.-L. Penrith (State Museum, Windhoek; SMW), Dr. E. C. G. Pinhey

(National Museum, Bulawayo; NMB), Dr. F. H. Rindge (American Museum of

Natural History, New York; AMNH), E. Taylor (Hope Department of Zoology

(Entomology), Oxford; HDE), A. Tsvetajev (Moscow), Dr. L. Vari (Transvaal

Museum, Pretoria), Dr. Ir. N. L. Wolff (Universitetets Zoologiske Museum, Copen-

hagen; ZMC). The Rijksmuseum van Natuurlijke Historie (Leiden) is abbreviated:

(ML).

I am also greatly indebted to the Netherlands Foundation for the Advancement

of Tropical Research (WOTRO) for a grant which enabled me to study the East

African Spialia species in the field and to visit the important collection in the

National Museum, Nairobi.

2. KEY TO THE SPECIES AND CHECKLIST

The distinction of species groups proposed in this paper is based on the male and

female genitalia. These groups can be recognized only incompletely by external

characters. As it is quite well possible to identify the species by their external

characters, it seems best not to bother with species groups in the key, otherwise

the characters of the genitalia should be introduced in the key, diminishing its

practical value (viz., identification of species). To facilitate reference to the descrip-

tions in Chapter 3, the species group to which a species belongs, is mentioned in

the key. In cases of uncertain identification the genitalia should always be referred

to.

1. On fore wing upperside, central cell spot closer to discocellular spot than to

basal cell spot, basal spots in spaces 9 and 10 usually well developed (lying

over the central cell spot) (fig. 4); male with costal fold .............. 2

— On fore wing upperside, central cell spot not closer to discocellular spot than

to basal cell spot, or basal cell spot absent; no spots in spaces 9 and 10 4

2. Basal spot 7 on hind wing underside fused to median band along vein 7, so that

basal spots 7 and 8 together with the median band form a Y; on fore wing

upperside submarginal spots incomplete, often only 4 and 5 present (asterodia

group) NA DELLI MITI Oe ie BRE, EO EN kituina

— Basal spot 7 on hind wing underside not fused to median band; on fore wing

upperside submarginal spots well-developed ...................... 3

3. Median band on hind wing underside with irregular edges, parallel to the

series of basal spots; fringes not conspicuously spotted on underside (asterodia

group) Di iii VRE RE er Te e asterodia

— Median band on hind wing underside very irregular, along vein 2 pointing

basad and reaching series of basal spots; fringes conspicuously chequered

black and white on underside (asterodia group) ................ agylla

4. Submarginal spots 4 and 5 of hind wing not in line with other submarginal

spots, being placed more basad (fig. 10); male with costal fold .......... S

DE JONG: The genus Spialia 29

— Submarginal spots 4 and 5 of hind wing in line with other submarginal spots;

10.

maleswithoutcostalifoldi& ae RE eee ae 6

Median band on hind wing upperside composed of separate spots (colotes

STOUD) HE RE SAN re RS EDITED REIFE colotes

Median spots on hind wing upperside fused into a continuous band (colotes

PROUD) BORA Rs Dern SE GAS A confusa

Median spot 6 on hind wing underside absent or very small, not connecting

the large central spot to one of the spots in space 7 (figs. 11—13) .... 7

Median spot 6 on hind wing underside present, connecting the central spot (in

spaces 45) with one of the spots in space 7 (figs. 14—15) .......... 13

Median band on hind wing underside directed towards inner spot in space 7

(fig. 11); fore wing upperside without basal cell spot; Palaearctic species 8

Median band on hind wing underside directed towards outer spot in space 7,

or towards a point central between the inner and outer spots in space 7 (fig.

12); fore wing upperside usually with basal cell spot, but this spot sometimes

vaguely outlined; Palaearctic and Afrotropical species .............. 9

Ground colour of hind wing underside red in various shades, from ochreous to

a very vivid red; central spot of hind wing underside angular; Europe and

North Africa (serforius group) wensen cso coi eo cee ie sertorius

Ground colour of hind wing underside greenish or yellowish olive-grey (except

in the Asian ssp. carnea, where it is red); central spot of hind wing underside

rounded; E. Europe to E. Asia (sertorius group) ................ orbifer

On hind wing underside, inner spot in space 7 central between basal cell spot

and median spot 4—5 (= central spot); median band directed towards a point

between inner and outer spots in space 7; Palaearctic species (phlomidis

ETOUP)H Betere Ser TOG tere Mered «Tetons owes Sigel ey osthelderi and geron

On hind wing underside, inner spot in space 7 closer to basal cell spot than to

central spot; median band directed towards outer spot in space 7; Palaearctic

andeAtrotropicalispecCieS RR EE IO I 10

On hind wing underside basal spot lc absent, median spot 6 present, but

small; on fore wing upperside discocellular spot strongly curved and outer

median spot 1b usually present; a subdesert species (phlomidis group) doris

— On hind wing underside basal spot lc present, median spot 6 absent; on fore

11.

12.

wing upperside discocellular spot usually less strongly curved or straight, outer

median spot Ib usually absent |: ................................ 11

On hind wing underside median spots 1c—5, basal cell spot and basal spot Ic

shining white, median spots lc and 2 entirely fused, no spot between median

and submarginal spots in space lc (dromus group) ................ paula

All spots on hind wing underside plain white ...................... 12

No light spot at base of costa on fore wing upperside; hind wing underside

without a spot between median and submarginal spots in space lc, inner spot

in space 7 usually narrow and irregular, submarginal spot Ic double, the upper

part out of line, median spot 1c usually not joined to median spot 2 (sertorius

Acoli) Nob gama ondGanonoo se die 0 0 odo gece loon oo eos CO ODO odo mafa

— Usually a light spot at base of costa on fore wing upperside (fig. 7); hind wing

30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

underside with a spot between median and submarginal spots in space Ic,

inner spot in space 7 usually large and rounded, submarginal spot Ic usually

single, median spot lc joined to median spot 2 (spio group) .......... spio

13. Median spot 6 on hind wing underside joins central spot to inner spot in space

7; Palaearctic species (phlomidis group) .................. .. phlomidis

— Median spot 6 on hind wing underside joins central spot to outer spot in space

7; Afrotropical and Oriental species ....... a dels sole ORR oo ol 0 14

14. Fore wing upperside without basal cell spot, at most some white scales forming

a small dash against cubitus (fig. 5) ........ a ae EEN 15

— Basal cell spot on fore wing upperside present (figs. 6, 8,9) ........ diario

Talk

Figs. 4—g. Upperside of right fore wing of Spialia species. 4, S. asterodia. 5, S. zebra. 6, S.

wrefordi. 7, 8. spio. 8, S. diomus. 9, S. dromus.

Figs. 10-15. Underside of left hind wing of Spialia species. 10, S. colotes. 11, S. sertorius. 12, S.

osthelderi. 13, S. spio. 14, S. nanus. 15, S. secessus.

15:

20.

DIE

De JONG: The genus Spialia 31

Submarginal spots entirely absent on upper and underside of fore and hind

wings; hind wing underside without basal spots or a very vague one in space 7

(delagoae Sroup) jah Ree tao. des HONS SEAS mangana

On upper and underside submarginal spots at least partly visible ...... 16

Hind wing underside without basal cell spot; median spot 6 on hind wing

upperside absent or at most vaguely indicated .................... 17

Hind wing underside with basal cell spot; median spot 6 on hind wing upper-

side,presentorabsentstr. etsen Bee were teren ts. men ee: 18

Median band on hind wing underside winding (fig. 14) (delagoae group)

ect Ae Smelen TB nanus

Median band on hind wing underside straight (delagoae group)... delagoae

Submarginal spots on hind wing underside absent or very vaguely indicated

along inner edge of light region along termen; median band slightly winding

(delagoaeigtoup) Mr ENA IIND IE RN sataspes

Submarginal spots on hind wing underside present; median band straight 19

On hind wing upperside median spots reduced, usually only 2 and 4—5 visible;

length of fore wing usually less than 11 mm (delagoae group) ........ zebra

On hind wing upperside usually median spots 2—6 present in a more or less

straight band that is crossed by veins 3, 4 and 6; length of fore wing usually

more than 11 mm (delagoae group) .................... depauperata

On fore wing upperside, outer median spots 1b and 1c forming a bar from vein

1 to vein 2; inner median spot lc absent and 1b usually smaller than one of

the outer median spots (fig. 8); mid-tibiae spined (spio group) ...... diomus

On fore wing upperside, outer median spot 1b absent; inner median spot 1b as

large as or larger than outer median spot Ic, which may be absent; mid-tibiae

without spines, except for the apical pair (as in all other species of the

BONUS) PARA ap ect eR Gece OO NEU RE el APN AC pa Ps 21

Median band on hind wing underside ends on termen .............. EN 022

Median band on hind wing underside ends on dorsum ................ 23

Hind wing underside with a white streak from top of median band (median

spot 7) to end of vein 5 at termen, veins lighter than ground colour; on hind

wing upperside median band not divided by dark veins; on fore wing upperside

submarginal spots complete, forming a sinuous series (angled at vein 5)

(dromusseroup)i Sted Wir We Heer ploetzi ploetzi

— White streak from top of median band to end of vein 5 on hind wing under-

23:

side absent or scarcely visible and veins not lighter than ground colour; on

hind wing upperside median band divided by dark veins; submarginal spots

on fore wing upperside less conspicuous, not always complete, and series less

SINUOUSI(Z/CIMUS CLOUD) AEE TN NRN ole eee noe oe ploetzi occidentalis

Basal cell spot on fore wing upperside very large, about twice as long as

broad, much larger than central cell spot (fig. 6); on hind wing upperside

median spots very conspicuous, median spot 1c running from vein 1b to vein

2(dromusigroup)iita vergat. waor icon. alt RAVES EE wrefordi

Basal cell spot on fore wing upperside much smaller, rounded and not larger

than central cell spot; median spots on hindwing upperside smaller or less

32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

complete, median spot lc rarely running from vein lb to vein 2 ........ 24

24. Median spots on hind wing upperside incomplete, only 2 and 4—5 present

and of the submarginal spots only 2 and 3 well developed, sometimes also 4

and 5 present as vague dots, very rarely a few white scales at place of sub-

marginal spots 1b and Ic; median band on hind wing underside straight, outer

margin finely dentate (spots slightly extended towards termen along veins) and

along this margin ground colour darker (fig. 15) (dromus group)... secessus

— On hind wing upperside median spots and usually also submarginal spots better

developed; median band on hind wing underside more or less curved, outer

margin not finely dentate nor contrasted by darkening of ground colour .. 25

25. On fore wing upperside inner median spots 1b and lc well-developed, forming

a bar from vein 1 to vein 2; outer median spot 1b absent, 1c a small dot at

lower outer angle of median spot 2, or absent (fig. 9); median spots on upper-

side of hind wing fused into a continuous, characteristically curved white band;

Ethiopian species (dromus group) ...... IRE dromus

— On fore wing upperside inner median spot 1b and outer median spot Ic well-

developed, inner median spot lc and outer median spot lb absent (the latter

rarely indicated as a white point close to the outer median spot lc); median

spots on hind wing upperside separated by ground colour along the veins;

Oriental species (sertorius group)... ais See N galba

Checklist of Spialia species

asterodia group asterodia dromus group wrefordi

agylla paula

kituina secessus

spio group spio dromus

diomus ploetzi

phlomidis group phlomidis

osthelderi delagoae group mangana

geron nanus

doris delagoae

colotes group colotes zebra

confusa sataspes

sertorius group mafa depauperata

galba

sertorius

orbifer

3. THE SPIALIA SPECIES

The species groups recognized in this paper are pretended to be monophyletic

subdivisions of the genus. They are based largely on the male and female

genitalia. When studying the genitalia it soon became evident that a number of

evolutionary trends could be found in the genus Spialia, these trends being rep-

resented by apparently autapomorphous character conditions of certain parts of the

DE JONG: The genus Spialia 33

male and female genitalia. The intricacy of the genitalia makes them much more

convenient for the study of phylogenetic relationships than the wing markings.

Indeed, within the genus Spialia the latter vary mainly in the presence or absence

and relative development of the spots. The absence of a particular spot in various

species is a poor indication (or no indication at all) of relationship, as the loss of a

spot may occur more than once independently. The presence of an additional spot

would be a better indication, but in Spialia this seems to be realized in a single

case only.

Thus, the three species of the spio group of Higgins (1924), which was based

on the absence of median spot 6 on the underside of the hind wing, had to be

placed in three different species groups in this paper while a number of other species

groups recognized by Higgins could be united. The result of basing the genus groups

on genital characters is that the classification shows the (supposed) interrelationships

of the groups, while the exclusive use of characters of the wing markings results in

a number of groups of which the interrelationships are quite obscure.

This does not mean, of course, that all characters of the wing markings are use-

less for the classification and as will be shown below, the position of certain spots is,

in this respect, much more important than their presence or absence.

The phylogenetic implications will be dealt with in Chapter 4.

In the following, descriptions are given of all species groups as to their external

characters, male and female genitalia. In the descriptions of the species these general

features of the relevant species groups have not been repeated. Thus when a species

has been identified by means of the key, the characters of its genitalia can be

found in the combination of the characters of the relevant species group and of

the species concerned (and, of course, by reference to the figures).

a. The asterodia group

External characters. — Upperside of fore wing with basal cell spot; central cell

spot nearer to discocellular spot than to basal cell spot; basal spots in spaces 9 and

10 usually present as two white streaks lying over the central cell spot; submargin-

al spots present, but sometimes very incompletely developed. Underside of hind

wing with all median spots present, forming an irregular band more or less parallel

to the termen or fused with the basal spots in spaces 7 and 8 to form a Y; sub-

marginal spots 4 and 5 in line with the other submarginal spots. Male with a costal

fold (not very conspicuous).

Male genitalia. — Uncus deeply incised; junction with tegumen slightly marked,

no fenestrula. Gnathos dorsally joined to tegumen over short distance, ventrally

unspined, well sclerotized, the left and right parts ventrally connected by a (some-

times very narrow) membrane. Fultura superior smooth. Anellus membranous.

Juxta may be hairy. Aedeagus straight, surface smooth. Saccus relatively long.

Costal process of valva large, strongly spined and heavily sclerotized, more or less

detached from costa or even attached to cucullus. Cucullus without fold covering

costal process.

Female genitalia. — Segment 8 with a single, large, dorsal sclerite, separated by

a membranous gap from the ventral sclerotization that is continuous antevaginally,

34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

but shows a large postvaginal membranous gap. On distal edge of postvaginal

membrane no spines observable. Ductus long and imperceptibly passing into bursa.

Beginning of ductus may be sclerotized. Bursa without sclerotization. Papilla analis

without basal lobe; apophysis posterior longer than papilla, sharply bent at con-

junction with papilla.

Spialia asterodia (Trimen)

Pyrgus asterodia Trimen, 1864. — Trans. ent. Soc. London (3) 2 : 178 (Plettenberg Bay, Cape |

Province). Holotype (o ) in the British Museum (Nat. Hist.), London. |

Note. Trimen twice described this species as new, the first time in the above _

mentioned paper, the second time (1866) in his book ‘“‘Rhopalocera Africae

Australis”, Part II. In his first description Trimen quoted as type locality only

“Cape Colony”, in his second description he mentioned Plettenberg Bay. The type

specimen is labelled as coming from the latter locality.

External characters (pl. 1 figs. 1, 2). — The pronounced characters of the

asterodia group, especially the position of the central cell spot, make a confusion

with species of other groups improbable. The characters mentioned in the key will

suffice to distinguish asterodia from the other species of the group, but we may

add that agylla, the only species likely to be confused with asterodia, is on the

average smaller: asterodia © 9.8—12 mm, agylla & 9.2—10.5 mm.

Apart from variation in size there is a slight variation in the extension of the

spots and the ground colour of the underside of the hind wing may be darkened

along the white spots. According to Swanepoel (1953) the spring specimens

(August) have a darker underside of the hind wing than summer specimens

(January).

Male genitalia (figs. 16—18). — Two arms of uncus close together. Two parts

of gnathos ventrally very close together. Costa of valva without spines. Costal

process large, distally thickly beset with fine spines, more proximad the spines

becoming larger and in the proximal part the spines coarse and large and directed

downwards.

Female genitalia (figs. 25—-27). — In addition to the group characters, the

female of asterodia has the upper part of the ductus sclerotized. Antevaginal

sclerotization appears slightly stronger than in agylla.

Ecology. — According to Swanepoel (1953) asterodia is a species of grasslands

that ascends the Drakensberge to a height of more than 3000 m. It flies from

August till March. Murray (1959) mentions as food plants various species of

Hermannia (Sterculiaceae) and Pavonia macrophylla (Malvaceae).

Distribution (Map 1). — Confined to the southeastern part of Africa. In South

Africa widely distributed through the southeastern, eastern and northeastern parts.

Outside South Africa only known from Mozambique and Rhodesia.

Localities. — Rhodesia: Hillside (Bulawayo), Mashonaland, Salisbury (BM,

NMB). Mozambique: Rikatla (24 km N of Lourengo Marques) (BM). South Africa.

Cape Province: Addo, Assegaibosch, Brak River, Burghersdorp, Cape Town,

Cathcart, Dordrecht, Grahamstown, Herd’s Bay, Jefferey’s Bay, Keurbooms River,

DE JONG: The genus Spialia 35

Figs. 16—24. Male genitalia, lateral aspect, valva, and dorsal aspect of tegumen and uncus. 16—18,

S. asterodia (Natal). 19—21, S. agylla (Bloemfontein, S. Africa). 22—24, S. kituina (Voi, Kenya).

Kokstad, Plettenberg Bay, Port Elizabeth, Somerset East, Steynsburg, Swartkops,

Swellendam (BM, CMP, MN, MNHN, NMB; Swanepoel, 1953). Orange Free

State: Bethlehem, Bloemfontein, Kroonstad, Ladybrand, Lindley, Rouxville (MN,

NMB; Swanepoel, 1953). Natal: Balgowan, Cleopatra, Estcourt, Greytown, Lady-

smith, Loteni, Newcastle, Vrijheid (BM, MN, NMB; Swanepoel, 1953). Transvaal:

Barberton, Ermelo, Graskop, Haenertsburg, Johannesburg, Lydenburg, Messina,

Munnik, Pietersburg, Potchefstroom, Pretoria, Zoutpansberg (BM, CMP, NMB;

36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Swanepoel, 1953). Lesotho: Giant’s Castle, Mamanthes, Maseru, Mokhotlong

(NMB; Swanepoel, 1953).

Material examined. — 410 , 35 9 : 39 ” (1 © holotype), 34 9 , South

Africa (BM, CMP, MN); 10°, 19 , Mozambique (BM); 1 0° , Rhodesia (BM).

Map 1.

Spialia asterodia

Spialia agylla (Trimen)

Pyrgus agylla Trimen, 1889. — South African Butterflies 3 : 286 (Griqualand West). See further under

subspecies.

External characters (pl. 1 figs. 3, 4). — The characters mentioned in the key

and under asterodia will suffice to distinguish this species. It varies in the same

way as asterodia, but it is slightly smaller, 9.2—10.5 mm. One female in the

BM has the median spot on the upper and underside of the hind wing much

reduced. Swanepoel (1953) does not mention a seasonal variation.

Male genitalia (figs. 19—21). — Two arms of uncus less close together than in

asterodia, more or less diverging. Two parts of gnathos ventrally clearly diverging.

Costa of valva with spines in distal half along ventral edge. Costal process large,

DE JONG: The genus Spialia 37

HOP

Ss ra

fi a)

De: I

= N J 5

Map 2

Spialia agylla

more detached from costa than in asterodia, thickly beset with short spines of

uniform length, with a dorsal, heavily sclerotized excrescence ending in a single

very large spine, or one large and a few smaller spines, that all point towards the

base of the valva (junction with vinculum) and not downwards.

Female genitalia (figs. 28, 29). — The only difference with asterodia seems to

be a slighter sclerotization of the antevaginal region and the lack of sclerotization

in the upper part of the ductus.

Ecology. — This species is found on mountains, mountain slopes and grassy

plains, where it flies from August till April (Swanepoel, 1953). The food plants are

said to be the same as for asterodia (Murray, 1959).

Distribution (Map 2). — Confined to southern Africa, where it is mainly found

in the high uplands in the interior.

Geographic variation. — The recent discovery of this species in Namaqualand

has led to the erection of a new subspecies. As the material of the new subspecies

is limited and has been caught within three weeks, and as the differential characters

are liable to environmental influences in other Spialia species, I consider the

distinction of ssp. bamptoni tentative.

38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Spialia agylla agylla (Trimen)

Pyrgus agylla Trimen, 1889. — South African Butterflies 3 : 286 (Griqualand West, between Modder

River and Riet River). Holotype (0 ) in the British Museum (Nat. Hist.), London.

External characters. — Upper and underside of wings with size of spots more or

less as in asterodia. Slightly larger than ssp. bamptoni: ” 9.2—10.5 mm.

Distribution. — The main range of the species, the uplands in the interior of

South Africa. I also regard the specimens from Botswana as belonging to the

nominate subspecies.

Localities. — South Africa. Cape Province: Aberdeen, Boschberg Mts, Burghers-

dorp, Cradock, Griqualand West, Hopetown, Kimberley, Klaarstroom, Matjesfontein, |

Murraysburg, Nauwpoort, Port Elizabeth, Queenstown, Somerset East, Steynsburg

(BM, MN; Swanepoel, 1953). Orange Free State: Bethulie, Bioemfontein, Deel-

fontein, Ladybrand, Trompsburg (AMNH, BM, CMP, MN, MRAC; Swanepoel,

1953). Transvaal: Heidelberg, Johannesburg, Klipdrift, Standerton, Vaal River

(BM; Swanepoel, 1953). Lesotho: Maseru, Leribe (Swanepoel, 1953). Botswana:

Ghanzi (BM).

Material examined. — 38 © , 10 © : 11 & (1 © holotype), 3 © , Cape

Province (BM, MN); 25 °°, 69 , Orange Free State (AMNH, BM, CMP, MN,

MRAC, NRS); 1 ©, Transvaal (BM); 1 , 1 9 , Botswana (BM).

Spialia agylla bamptoni Vari

Spialia agylla bamptoni Vari, 1976. — Ann. Transvaal Mus. 30 : 123—124, figs. 3, 4, pl. 13 figs. 3,

4 (Hondeklipbaai, 1—18.xii.1974). Holotype (o ) in the Transvaal Museum, Pretoria.

External characters. — Spots on upper and underside of wings larger and more

whitish than in ssp. agylla, ground colour of underside of hind wing also more

whitish in the subbasal and submarginal areas. Slightly smaller than ssp. agylla,

cd’ 9—10 mm.

Distribution. — Only known from the type-locality, Hondeklipbaai (Namaqua-

land Distr.) at the west coast of South Africa (Vari, 1976).

Material examined. — None.

Spialia kituina (Karsch)

Hesperia kituina Karsch, 1896. — Ent. Nachr. 22 : 374—375 (Kitui, Kenya). Holotype (>) in

Zoologisches Museum der Humboldt Universität, Berlin.

Evans (1937) stated as type-locality “Zanzibar” and Dr. Hannemann kindly informed me that the

type is labelled in the same way, but from Karsch’ original description (‘‘aus dem Sansibar-gebiete, |

von Kitui”) it follows that the type-locality is not the present island of Zanzibar. At the time of Karsch

the Sultanate of Zanzibar covered large parts of East Africa.

Pyrgus bettoni Butler, 1898. — Proc. zool. Soc. Lond. 1898 : 415, pl. 32 fig. 1 (Maungu Inkubwa,

Kenya). Holotype (0 ) in the British Museum (Nat. Hist.), London.

External characters (pl. 1 figs. 5, 6). — An unmistakable species. It has the

position of the central cell spot, close to the discocellular spot, in common with

asterodia and agylla, but it will at once be distinguished by the Y-shaped band on

the underside of the hind wing. By this character it is also distinguished from all

other Spialia species.

DE JONG: The genus Spialia 39

Figs. 25—29. Lateral and ventral aspect of female genitalia, and papilla analis. 25—27, S. asterodia

(Natal). 28—29, S. agylla (Hopetown, S. Africa).

Male genitalia (figs. 22—24). — Higgins (1924) mentioned an undivided uncus,

but in fact the uncus is deeply indented and the apices of the two parts diverge.

Ventral parts of gnathos also diverging apically. Costa of valva with a few spines

pointing upwards in the distal half. Costal process detached from costa, attached to

undeveloped cucullus (at least, this process is supposed to be homologous with the

costal process in the other Spialia species), strongly sclerotized, oblong, with a

strong tooth at its proximal end and a very large tooth at its distal end protruding

beyond the dorsal rim of the costa.

40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Higgins (1924) mentioned “elaborate interlocking terminal teeth”. From his

figure (pl. 9 fig. 1) it is clear that one of these teeth is the distal tooth of the costal

process, while the other one is in fact the costa that is connected by a membrane

(omitted in Higgins’ figure) to the costal process.

Female genitalia. — Due to lack of material kituina is the only Spialia species

of which the female genitalia could not be studied.

Map 3 Ms

Spialia kituina

RES Gn

Ecology. — In the Shimba Hills (SE. Kenya) specimens have been found in

January, February, July, August and November (Sevastopulo, 1974). Food plants

unknown. I found this species together with Spialia zebra along a dry, hot road

near Ukunda (S. of Mombasa) in a rather open, more or less cultivated terrain in

the beginning of November.

Distribution (Map 3). — Confined to Kenya. Higgins (1924) stated that there is

a specimen in the British Museum (Nat. Hist.) from British Central Africa, but I

could not find that specimen. Neave (1910) recorded a number of specimens from

the Luangwa Valley (Zambia), which he hesitatingly placed as the wet-season form

of “Hesperia bettoni”. The only specimens in the Neave collection in the Hope De-

partment, Oxford, that come into consideration belong to Spialia confusa confusa.

DE JONG: The genus Spialia 41

Localities. — Kenya: Bura (Teita), Diani, Kaitir, Kangondi, Kibwezi, Kitui,

Landjoro (Pori), Lokichogio, Lukenia, Makueni, Maungu Inkubwa, Mombasa,

Mulango, Shimba Hills (Kakardara and Marere Forests), Taveta, Ukambani,

Ukazzi Hill (Garissa Rd), Ukunda, Voi (AMNH, BM, CMP, ML, MN, MNHN;

Higgins, 1924; Karsch, 1896; Sevastopulo, 1974).

Material examined. — 21 © (1 & holotype of bettoni), 5 9 , Kenya (AMNH,

BM, CMP, ML, MN, MNHN).

b. The spio group

External characters. — Upperside of fore wing with basal cell spot; central cell

spot midway between basal cell spot and discocellular spot; no spots in spaces 9 and

10; submarginal spots usually well-developed. Median spots on underside of hind

wing complete, forming a continuous band, or median spot 6 absent; submarginal

spots 4 and 5 in line with the other submarginal spots; an additional spot between

the submarginal and median spots in space Ic. Male without costal fold.

Male genitalia. — Uncus undivided, junction with tegumen well-marked and with

fenestrula or almost invisible. Gnathos free from tegumen by a very narrow mem-

branous gap; dorsally broad, ventrally narrower and with some spines, distally very

gradually passing into the more or less spinulose membrane that joins the left and

right parts of the gnathos. Anellus membranous, spinulose or smooth. Aedeagus

straight, without marked spines. Saccus slightly shorter than or about as long as

uncus. Costal process of valva a strongly spined structure in central part of costa

or a finger-like excrescence of dorso-distal part of costa. Cucullus spined dorso-

distally.

Female genitalia. — Segment 8 with two dorsal sclerites that extend far laterally;

ventrally a single sclerite or two sclerites separated by a narrow membranous gap

distally of the ostium. In the distal part the ventral surface bears a membranous or

sclerotized flap that partly overlaps the spinulose area. This area is well-developed

and not connected to the ventral sclerite(s). Antevaginally there is a strong scleroti-

zation which may be divided into a left and right part, recalling the sclerotized

“wings” in the phlomidis group. Upper part of ductus may be sclerotized. Bursa

without sclerotizations. Papilla analis with basal lobe; apophysis posterior long,

longer than papilla.

Spialia spio (Linnaeus)

Papilio spio Linnaeus, 1767. — Syst. Nat., Ed. XII, 1 (2) : 796 (Cape of Good Hope). Of the type

only figures exist. Linnaeus referred to figures by Clerck, which were not published before 1882,

when Aurivillius reproduced them. Only then it could be determined that the poor description by

Linnaeus referred to the species that was generally known as Papilio vindex Cramer, 1782. Al-

though this synonymy is undoubtedly correct, the figures of Clerck reproduced by Aurivillius are so

bad as to lead Oberthiir (1912) to consider spio and vindex separate species.

Papilio vindex Cramer, 1782. — Cramer, Uitlandsche Kapellen 4 : 122, pl. 353 figs. G, H (Cape of

Good Hope). Type presumably lost.

External characters (pl. 1 figs. 7, 8). — A variable species, that is, however,

usually not difficult to distinguish. To the characters mentioned in the key we may

add the following notes. In rare cases the median spot 6 on the underside of the

hind wing is faintly developed. The additional spot in space lc on the underside of

42 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Map 4

Spialia spio

= er tee “|

the hind wing is not always well-developed. A useful distinguishing character can

also be found on the upperside of the hind wing, where the submarginal spots 4

and 5 are fused to form a single large conspicuous spot in most specimens; this

spot usually distinguishes spio from all other Spialia species.

From the two other widely distributed and common Spialia species in Africa,

viz., dromus and diomus, spio is easily distinguished by the development of the

median spots in space lb on the upperside of the fore wing (figs. 7—9). In spio,

inner median spot ib and outer median spot lc are always present and subequal,

the other median spots in space lb are absent. In dromus the inner median spots.

lb and lc are always both present and usually fused, outer median spot lc is very

small or absent, outer median spot 1b is always absent. In diomus the inner median

spot lb is present, but usually small, inner median spot lc is always absent, the

outer median spots lb and lc are always both present and usually fused. In other

words, on the upperside of the fore wing, there is no white bar in space 1b extend-

ing from vein 1 to vein 2 in spio, such a bar is present and composed of the inner

median spots in dromus and also present but composed of the outer median spots

in diomus.

Further differences are the following. On the upperside of the hind wing median

DE JONG: The genus Spialia 43

spot 6 is always absent in spio (in very few specimens represented by a few white

scales), usually present but faintly developed in diomus and usually present as a

clear spot in dromus. The series of submarginal spots on the upperside of the fore

wing is much more sinuous in dromus (especially in spaces 2 and 3) than in spio

and diomus. The black bar in the fringes of the fore wing at the end of vein 5 is

always well-developed in dromus, variable, but often narrow in diomus and

narrow or (in most cases) absent in spio.

There is a large variation in size, the males varying from 8.7 to 13.0 mm; in the

development of the spots and in the ground colour of the underside of the hind

wing, which varies from pale yellow to dirty green. In addition, there exists some

variation in the wing shape. Usually the fore wings of the males are slightly more

pointed than those of the females, but males with rounded and females with pointed

wings also occur. Probably, temperature and humidity are largely responsible for

these varieties. The described variation is not geographical.

Male genitalia (figs. 30, 31). — A large fenestrula at junction of uncus and

tegumen. Gnathos free from tegumen, distally gradually passing into fultura superior,

which bears some small spines, ventrally forming a more or less triangular, slightly

spined “head”. Anellus membranous, smooth. Costal process of valva a strongly

spined structure near the centre of the costa. Cucullus dorso-distally pointed, strongly

sclerotized and with a strong tooth near the junction with the costa.

Female genitalia (figs. 35, 36). — Ventral sclerotization of segment 8 forming a

single, not sharply defined sclerite with a sclerotized projecting flap that partly

overlaps the area spinulosa which is pointed distally. Antevaginally with two

strongly sclerotized and folded, wing-like structures. Papilla analis with small basal

lobe, that scarcely covers the base of the apophysis posterior.

Ecology. — Although in South Africa spio is a species of open grounds (Swane-

poel, 1953), at least in East Africa it seems to prefer fringes of dense scrub, some-

times occurring along forest paths (own observations; experiences of my friend,

Mr. J. H. Lourens). Probably, it is a rather adaptable species, that can live where

the vegetation structure is not too dense (closed forest) or too open (subdesert,

desert). It may fly almost throughout the year, but this depends on local conditions.

Recorded food plants are several Hermannia species (Sterculiaceae) and Pavonia

macrophylla and columella (Murray, 1959), Hibiscus aethiopicus (Gifford, 1965),

and Sida (Van Someren, 1939) (all Malvaceae).

Distribution (Map 4). — One of the most widely distributed African Spialia

species, occurring throughout the Afrotropical Region in suitable habitats and

penetrating the Arabian peninsula in the extreme south. Although the species is not

known from large areas, this often appears to be the result of undercollecting rather

than of true absence.

Localities. — Yemen: Wadi Dhar (6 mls NW of San’a) (BM; Gabriel, 1954).

South Yemen: Hadramaut (BM). Somalia: Buran, 40 mls S of Buran, Holl-Holl

(MN, MNHN; Talbot, 1932). Sudan: Bamaka, Jebel Marra (Darfur), Niurnya

(BM). Ethiopia: throughout (BM, MNHN; Carpenter, 1935). Kenya: throughout

(AMNH, BM, CMP, ML, MN, MNHN, NMB, USNM, ZSM). Uganda: through-

out (AMNH, BM, CMP, ML, MN, MRAC). Tanzania: throughout (BM, ML, MN,

44 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Figs. 30—34. Male genitalia, lateral aspect, and inside of right valva. 30—31. S. spio (S. Africa). 32—33,

S. diomus ferax (Pretoria, S. Africa). 34, S. diomus diomus (Ukunda, Kenya).

MNHN, MRAC, ZSM; Aurivillius, 1910). Ruanda: Cyangugu, Rugege Forest

(Lake Kivu) (BM, ML). Burundi: Butoni, Kibimba, Kitega, Misumba, Usumbura

(MRAC). Malawi: throughout (BM, MNHN, NMB; Gifford, 1965). Zambia: Chin-

gola, Fort Jameson, Kabompo River, Mid Luangwa Valley, Upper Luangwa Valley

(BM, NMB). Rhodesia: throughout (MN, MNHN, NMB, USNM). Mozambique:

Augustino Farm (Revue River), Delagoa Bay, Inhanucarara, Serra Rotanda, Siluve

(Xiluvo) Hills (BM, MNHN, NMB, ZSM). Swaziland: Gollel (NMB). South Africa:

throughout (AMNH, BM, CMP, ML, MN, MNHN, MRAC, NMB, USNM,

ZSM; Swanepoel, 1953). Botswana: Kwaai River, Maun, 55 mls E of Maun, 25

mis E of Tsane (NMB). Namibia: Grootfontein, Okahandja, Omaramba-Oamatako

(SMW; Trimen, 1891). Angola: Banga, Barraca (Quanza River), Benguela, Biho

Distr., between Capelongo and Dongo, Cassualalla, Cubal River, Gauca, Huambo,

Kuvali River, N’Dalla Tando, Omrora, Pacolo (Luanda), Pungo Adongo (BM,

CMP, ML, MNHN; Trimen, 1891). Zaire: throughout (AMNH, BM, CMP, MN,

DE JONG: The genus Spialia 45

MNHN, MRAC, NMB, ZSM). Congo-Brazzaville: Fort Crampel (MNHN). Gabon:

Oyem (MNHN). Rio Muni: Makomo Campo (ZSM). Central African Rep.:

Boukoko M’Baiki, Oubangui (MNHN, MRAC). Chad: Bebedjia (Moundou) (ML).

Cameroun: throughout central and southern parts (BM, CMP, MNHN, USNM).

Nigeria: Afikpo, Akassa to Onitsha, Fumban, Kano, Lagos, Lokoja, Lower Niger,

Ogruga, Zaria, Zungeru (NMB, BM). Dahomey: Porto Novo (MNHN). Ghana:

Accra, Likpe, Gold Coast (BM, MRAC, USNM). Upper Volta: Pundu (ZSM).

Ivory Coast: Bayota Forest (Gagnoa), Dimbokro (BM, MN). Liberia: Monrovia

(Holland, 1896). Sierra Leone: Benikoro, Freetown, Sefadu, Yiraia (BM, ML,

ZSM). Guinea: Conakry, Dalaba, Macenta, N’zerekore (BM, MNHN). Gambia:

Gambia (BM). Senegal: Dakar, Niokola Koba (BM, MNHN; Condamin, 1969).

Material examined. — 692 o , 384 9 : 10°, Yemen (BM); 10 , S. Yemen

(BM); 3 7, Somalia (MN); 16 7, 9 © , Ethiopia (BM, MN); 3 © , 2 © , Sudan

(BM); 82 7, 63 2 , Kenya (AMNH, BM, CMP, ML, MN, USNM, ZSM); 37 ©,

22 © , Uganda (AMNH, BM, CMP, ML, MN, MRAC); 106 © , 54 © , Tanzania

(BM, ML, MN, MRAC, ZSM); 19 , Ruanda (ML); 21 © , 7 © , Burundi

(MRAC); 12 7,9 © , Malawi (BM); 37, 2 9 , Zambia (BM); 3 © , Rhodesia

(MN, USNM); 9, 3 © , Mozambique (BM, ZSM); 110 © , 79 © , South

Africa (AMNH, BM, CMP, ML, MN, MRAC, USNM, ZSM); 20,29,

Namibia (SMW); 100 , 3 9 , Angola (BM, CMP, ML); 204 © , 819 , Zaire

(AMNH, BM, CMP, MN, MRAC, ZSM); 1 9 , Rio Muni (ZSM); 1 © , Centr.

Afr. Rep. (MRAC); 10, 1 9 , Chad (ML); 41 % , 30 , Cameroun (BM,

CMP, USNM); 11 ,6 9 , Nigeria (BM); 2 ” ,2 © , Ghana (BM, MRAC,

USNM); 19 , 19 , Ivory Coast (BM, MN); 100 , 49 , Sierra Leone (BM,

ML, ZSM); 17, 29, Guinea (BM); 19 , Gambia (BM); 29 , 19 ,

Senegal (BM).

Spialia diomus (Hopffer)

Pyrgus diomus Hopffer, 1855. — Monatsber. K. Akad. Wiss. Berlin 1855 : 643 (Mozambique).

For synonymy, see subspecies.

External characters (pl. 2 figs. 9—12). — An unmistakable species, distinguished

by the following combination of characters:

1. On the upperside of the fore wing the outer median spots 1b and Ic are both

present, forming a bar that extends from vein 1 to vein 2; inner median spot 1b

present, lc absent; submarginal spots well-developed; basal cell spot present (fig. 8).

2. On the underside of the hind wing the median band is entire, not broken at

space 6. See also under Spialia spio.

Apart from the characters of the wing pattern diomus can be distinguished

from all other Spialia species by the presence of spines on the mid-tibiae (besides

the apical pair of spines).

There is a marked variation in the ground colour of the underside of the hind

wing, from bright creamy yellow to deep olive-brown (sometimes dirty green). (See

further under Geographic variation).

Male genitalia (figs. 32—34). — No fenestrula at junction of uncus and tegumen.

46 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Gnathos free from tegumen, distally passing gradually into a finely spinulose, folded

fultura superior, ventrally without differentiated “heads”. Anellus membranous,

spinulose. Costal process of valva a finger-like projection of the dorso-distal part

of the costa. Cucullus strongly spined dorso-distally (with serrated edge).

Female genitalia (figs. 37, 38). — Ventral sclerotization of segment 8 consisting

of two sclerites separated by a membranous region along the midline of the segment;

tongue-like flap membranous, partly overlapping the area spinulosa, which is not

pointed distally. Ventrally of the ostium there is a strongly sclerotized, irregular

antevaginal plate. Papilla analis with a large basal lobe, covering half of the base

of the papilla and the whole base of the apophysis posterior.

Ecology. — In many parts of its range a common species on open grounds.

Swanepoel (1953) even states: “You cannot wander over the South African veld

without meeting diomus almost everywhere .. ”. In more or less forested areas

diomus is scarce or absent and from own experience I think that it is much more

confined to open grounds than spio.

S. diomus may be found throughout the year where it is not too dry or too cold

seasonally. Recorded food plants are various species of Hermannia (Sterculiaceae)

and Pavonia macrophylla (Malvaceae) (Murray, 1959).

Distribution (Map 5). — Distributed throughout the Ethiopian region where

open grounds occur, but lacking in the extensively forested areas in the central and

western parts of the Ethiopian region. Possibly continuously distributed from South

Africa through Mozambique and Tanzania to Ethiopia, Sudan and Yemen. The

apparent gap in the known distribution in Malawi and Mozambique may be due to

undercollecting rather than to real absence. The same holds for the savanna-belt

from Sudan to West Africa, where diomus is unknown from large regions, pre-

sumably as a result of incomplete exploration. Although I have seen specimens

from various parts of West Africa, diomus was recorded neither by Berger (1962)

from Guinea and Ivory Coast, nor by Lindsay & Miller (1965) from Liberia.

Geographic variation. — There is a marked difference between the forms found

in the southern parts of Africa and in the rest of the Ethiopian region. The contact

zone is obscure by lack of data, but may be running through Zambia, Malawi and

Central Mozambique. The differences are found in the wing markings and male

genitalia. Some authors (e.g. Oberthür, 1912; Higgins, 1924) regarded the differ-

ences too large to unite the forms into a single species, but a close examination

reveals that the difference in the wing markings is not as large as would appear

at first sight. But apart from this, there is no need for a specific separation as there

is no evidence for geographic overlap of these forms.

Spialia diomus diomus (Hopffer)

Pyrgus diomus Hopffer, 1855. — Monatsber. k. Akad. Wiss. Berlin 1855 : 643 (Mozambique). The

type appears to be lost; it is not in the Zoologisches Museum der Humboldt-Universität, Berlin.

Pyrgus machacoana Butler, 1899. — Proc. zool. Soc. London 1899 : 426, pl. 25 (Machakos, Kenya).

Holotype ( S') in the British Museum (Nat. Hist.), London.

External characters (pl. 2 figs. 9, 10). — The main difference with ssp. ferax

is found on the underside of the hind wing, where the median band is directed

DE JONG: The genus Spialia 47

Figs. 35—38. Female genitalia, ventral aspect, and papilla analis. 35—36, S. spio (Natal). 37—38, S.

diomus ferax (Mozambique).

towards mid-dorsum by a strong curve of the band in space 2, caused by a shift of

median spot 2 towards the base of space 2; median spot Ic is well-developed, but

usually free from median spot 2, the additional spot in space 1c is well-developed,

usually larger than the median spot, sometimes connected to median spot 2, but

usually separated by dark scales along vein 2; the median band is broad in its

48 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

upper part, especially in space 8 where it may almost touch the basal spot. The

series of submarginal spots on the underside of the hind wing is more strongly

curved and more irregular than in ssp. ferax, leaving a narrow, light-coloured

region between the submarginal spots and the termen. The differences between the

subspecies found on the underside of the hind wing are only weakly reflected on

the upperside: only the shift of median spot 2 towards the wing base in ssp.

diomus is usually visible, but this shift is sometimes also found in ssp. ferax.

Male genitalia (fig. 34). — The cucullus is broader and dorsodistally more

pointed than in ssp. ferax.

Distribution. — The northern part of the range of the species, from West Africa

to Ethiopia and Yemen. The southern limit is not exactly known, it may run

through Zambia and Central Mozambique. In Mozambique ssp. ferax is known

from the extreme south only, while the holotype of diomus originated from Mozam-

bique. Lindsey & Miller (1965) concluded from the fact that Evans (1937) classified

all material from Mozambique in the British Museum (Nat. Hist.) as ssp. ferax,

while the type-locality of ssp. diomus is also Mozambique, that the correct name

for the northern populations should be machacoana, “unless there is evidence that

Hopffer’s type was mislabelled geographically”. As Hopffer clearly described and

figured (Hopffer, 1862) the same form as machacoana, the name diomus cannot

be replaced by machacoana, the latter being the junior synonym. Moreover, as all

material of ssp. ferax in the British Museum (Nat. Hist.) originates from the

extreme south, and the length of Mozambique (north-south) is about 2000 km,

there is no reason to suppose that the type of ssp. diomus was mislabelled.

It is unfortunate that the species is not yet known from Malawi (Gifford, 1965),

as it would be interesting to know whether ssp. diomus or an intermediate form

occurs there. From Tanzania only ssp. diomus is known. Neave (1910) recorded

“Hesperia diomus Hpff.” from Zambia, but the two specimens from his collection

(HDE) studied turned out to belong to ssp. ferax.

Localities. — Yemen: Hada (Gabriel, 1954). South Yemen: Aden, Dhala, Jebel

Jihaf, Lahej, Laudar, Wadi Dareija (BM; Gabriel, 1954). Somalia: Gala River,

Upper Sheikh (BM). Ethiopia: throughout (BM, MN, MNHN, MRAC; Carpenter,

1935). Sudan: Bahr-el-Ghazal, Darfur Prov., Dilling Forest Reserve, Gondokoro

(White Nile), Kulme, Kurmuk (BM, MN). Kenya: throughout (AMNH, BM,

CMP, ML, MN, MNHN, NMB, USNM, ZSM; Sevastopulo, 1974). Uganda: Arua,

between Jinja and Busia, Busoga, Kafu River, Kakindu, Kampala, Kotido, Madi

Opei, Metu Hills, Mtanda (AMNH, BM, NM, USNM). Tanzania: Dar es Salaam,

Iringa, Kasulu, Katoto, Kifumbu, Kilema, Kilimanjaro West, Kiloa, Lake Manyara,

Moshi, Mpapura, Mwanza, Ngorongoro, Pemba, Rulenge, Tange, Ukerewe,

Usambara, Zanzibar (BM, ML, MN, MNHN, NMB, ZSM). Burundi: Kitega

(MRAC). Zaire: Kadjuju, Luna River, Nioka, Nyamunyunye, Rutshuri to Kabali,

Usumbura to Schangugu (BM, MNHN, MRAC). Chad: Bebedjia (Moundou), Fort

Archambault (ML, MNHN). Nigeria: Bida, Farniso, Kano, Lagos, Muri Prov.,

O’okemeji, Wurkum, Yelwa (BM). Upper Volta: Pundu (ZSM). Liberia: Zorzor

(CMP). Guinea: Kouroussa, N’zerekore (BM, MNHN). Gambia: Gambia (BM).

DE JONG: The genus Spialia 49

Senegal: Einaye, Melon à Kaolack, Oussouye Sandiala, Tabi près Bignona (BM,

MNHN; Berger, 1968).

Material examined. — 134 7,67 2 :4d ,19 , South Yemen (BM); 19,

19 , Somalia (BM); 97 , 49 , Ethiopia (BM, MN, MRAC); 5 © , Sudan (BM,

MN); 56 , 289 , Kenya (AMNH, BM, CMP, ML, MN, USNM, ZSM);

10 7,42 , Uganda (AMNH, BM, MN, USNM); 22 0°, 89 , Tanzania (BM,

ML, MN, ZSM); 115,99 , Burundi (MRAC); 5 , 1 9 , Zaire (BM, MRAC);

43,29 , Chad (ML); 35 , 5 9 , Nigeria (BM); 1 © , Upper Volta (ZSM);

19 , Liberia (CMP); 10, 19 , Guinea (BM); 1 © , Gambia (BM); 20,

1 2 , Senegal (BM).

Spialia diomus ferax (Wallengren)

Syrichtus ferax Wallengren, 1863. — Wien. ent. Monatschr. 1863: 137 (Kuisip River). Holotype (7)

in the Naturhistoriska Riksmuseet, Stockholm. The type-locality is generally known as the Kuiseb

River, south of Swakopmund (Namibia).

The name was misspelled “ferox” by Holland (1896) and Oberthür (1912).

Pyrgus abscondita Plôtz, 1884. — Mitt. naturw. Ver. Neu-Vorpomm. 15 : 21 (Africa). Type lost.

Higgins (1924), who treated diomus and ferax as distinct species, considered abscondita a synonym

of diomus. This is, however, not in accordance with the rather bad reproduction of Plôtz’s figure by

Aurivillius (1925). Evans (1937) treated abscondita as a senior synonym of depauperata. This cannot

be correct, as Plötz mentioned a basal spot on the upperside of the hind wing, which is rarely or

never present in depauperata. Moreover, the reproduction of Plôtz’s figure by Aurivillius (1925)

shows the band of submarginal spots on the underside of the hind wing extending to the end of vein 8,

while in depauperata it expires on the end of vein 7, and the median band in its upper part placed

more basad than in depauperata, rather like diomus ferax.

Syrichthus lacreuzei Oberthiir, 1912. — Et. Lep. Comp. 6 : 117, fig. 1332 (Zoutpansberg, South

Africa). Holotype (o ) in the British Museum (Nat. Hist.), London.

External characters (pl. 2 figs. 11, 12). — Median band of hind wing underside

more regular, less broadening in its upper part and straighter than in ssp. diomus,

directed towards tornus; median spot 2 in line with median spot 3 and connected

with the additional spot in space Ic, which is obliquely extended towards the tornus;

median spot 1c usually present, but weakly developed and free from median spot 2.

Series of submarginal spots on the underside of the hind wing straighter and more

regular than in ssp. diomus; the light-coloured area between the submarginal spots

and the termen is narrow near the tornus, but broadens conspicuously towards the

apex and is widest in space 6. See further under ssp. diomus.

Male genitalia (figs. 32, 33). — The cucullus is narrower, more rounded and

more strongly spined than in ssp. diomus.

Distribution. — Southern Africa, north to Zambia and possibly Central Mozam-

bique.

Localities. — Zambia: mouth Chambezi to Mansya River and Lake Young, high

plateau S of Lake Tanganyika (HDE; Neave, 1910). Rhodesia: Buhera, Bulawayo,

Matetsi, Mtoko, Mutambara, Nyamandhlovu, Salisbury, Selukwe, Strathmore

Ranch (BM, CMP, MN, NMB). Mozambique: Delagoa Bay, Lourenço Marques,

Shinawane (Xinavane) (BM, ML, MNHN, SMW, ZSM; Hopffer, 1855). Swaziland:

Stegi (NMB). South Africa: throughout (AMNH, BM, CMP, ML, MN, MNHN,

MRAC, NMB, NRS, USNM, ZSM; Swanepoel, 1953). Botswana: Kachikau,

Kang Rd, 80 mls W of Kanye, Mafeking, Mahalapye, Maun, Nxai Pan, Okavango,

50 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Okwa, Sepopa, Tsane, Tsau (NMB, ZSM). Namibia: Kalidona, Kalkrand, Kano

vlei, Kuiseb River, Okahandja, Omaramba-Oamatako, Otawi, Ovamboland,

Tsumele (NMB, NRS, SMW, ZSM; Trimen, 1891; Wallengren, 1863). Angola:

Ehanda (Trimen, 1891).

Material examined. — 927, 56 9 : 29, Zambia (HDE); 3 © ,19 ,

Rhodesia (BM, CMP, MN); 50° , 9 9 , Mozambique (BM, ML, SMW, ZSM);

723 , 442 , South Africa (AMNH, BM, CMP, ML, MN, MRAC, NRS,

USNM, ZSM); 1 © , Botswana (ZSM); 9 , 2 © , Namibia (NRS, SMW, ZSM).

Map 5

Spialia diomus

l_—— nn

c. The phlomidis group

External characters (pl. 3). — Fore wing with basal cell spot on the upperside

usually present, but vaguely outlined; central cell spot midway between basal cell

spot and discocellular spot; submarginal spots usually all present and well-

developed. On underside of hind wing usually all median spots present, but in most

cases spot 6 not connected to inner or outer spot in space 7; submarginal spots 4

and 5 in line with the other submarginal spots. Male without costal fold.

DE Jong: The genus Spialia SI

Male genitalia (figs. 39—41). — Uncus undivided, junction with tegumen well

visible but without fenestrula. Gnathos free from tegumen, dorsally broad, ventral-

ly tapering to a small “head”; left and right part of gnathos joined by a more

or less folded, usually finely spinulose membrane (fultura superior). Anellus

membranous. Aedeagus with a strongly sclerotized and indented crest, latero-ven-

tral in the middle, or a long, unindented branch in the same place, and with an

extended apex. Saccus slightly shorter than uncus. Costa of valva broadening

distally, in most species spined dorsally; costal process usually present as a ven-

trodistal excrescence of costa with strong spines. Cucullus spined dorso-distally,

without fold covering part of the costal process.

Female genitalia (figs. 42, 43). — Segment 8 with two dorsal sclerites and two

large ventral sclerites, which occupy the larger part of the ventral surface or are

fused to a single large ventral sclerite. At the distal edge of the ventral side of

segment 8 the area spinulosa may be visible. To the left and right of the ostium

there is a heavily sclerotized and indented wing-like structure at the base of the

segment. Ductus rather short, sclerotized in its upper part, rapidly broadening to

Figs. 39—41. Male genitalia, lateral aspect, aedeagus, and inside of right valva of S. phlomidis (Turkey).

Figs. 42— 43. Female genitalia, lateral aspect, and papilla analis of S. phlomidis (Greece).

52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

the oblong bursa, which does not bear any ornamentation. Papilla analis with

basal lobe; apophysis posterior long, longer than papilla.

Identification. — S. doris is the only species of the phlomidis group that by its

distribution can be confused with African Spialia species (especially mafa and

spio, by the absence or poor development of median spot 6 on the underside of the

hind wing), as doris penetrates the Afrotropical region in Sudan, Ethiopia, Somalia

and Kenya. It is, however, recognized on the upperside of the fore wing by the

vague outline of the basal cell spot and the usual presence of both outer median

spots lb and Ic (in mafa and spio the outer median spot 1b is always absent)

and on the underside of the hind wing by the presence of a small median spot 6.

Note. — The species of the phlomidis group (phlomidis, osthelderi, geron and

doris), which are mainly distributed in the eremic parts of the southern Palaearctic

region, have been extensively dealt with in a previous paper (De Jong, 1974) and

there is no need to repeat that treatment here. This is, however, a good opportunity

to present photos of the species and to give the following corrections and addi-

tions.

Spialia phlomidis (Herrich-Schäffer)

Since my 1974 paper I could examine 90 additional specimens. They were

mainly taken within the boundaries of the known distribution area, but two spe-

cimens greatly enlarged our knowledge of the distribution of this species, as they

were taken in S. Iran and C. Asia, respectively. The additional localities are:

ssp. phlomidis (Herrich-Schäffer):

Bulgaria: Stanimaka (ZSM),

Greece: Kastoria, Mt. Chelmos, Zachlorou (near Kalavryta) (Christensen, LNK,

ML, ZSM),

Turkey: Aksehir, Bulu, Cilician Taurus, Konia (LNK, ZSM),

South Russia: Eriwan (— Yerevan) (ZSM),

Iran: Golhak (near Teheran), Qader Abad Pass (Fars) (LNK, ML);

Central Asia: Alai Mts (ZSM).

ssp. hermona Evans:

Lebanon: Djezin (ZSM).

Spialia osthelderi (Pfeiffer)

Although the number of additional specimens examined is scarcely less than the

number of specimens examined for my 1974 paper, only three localities of ssp.

gecko can be added, all in Turkmenia: Askhabad, Merw (both ZSM) and Kara

Kala (Kopet Dagh) (ML). These additions scarcely enlarge the known distribution

area.

Spialia geron (Watson)

The known range of ssp. struvei is much extended by the capture of some speci-

mens in E. Afghanistan, south of Safed Koh (4 ” , 2 9 , LNK). The male geni-

DE JONG: The genus Spialia 53

talia are typical of this subspecies, as are the fore wing lengths ( © 13.5 — 14.2

mm, ® 14.2 — 14.3 mm). Wing length can, however, not be taken as a reliable

character: smaller specimens of ssp. struvei do occur, as shown by two males

from Dscharkent (Ili region) (ZSM), which measure 12.0 and 12.8 mm, respecti-

vely. These specimens differ further in having a dull greenish instead of yellowish

underside of the hind wing. Their genitaiia are of the usual struvei type. It is

possible that on the whole the ground colour of the underside of the hind wing

is more greenish in Central Asian specimens and more yellowish in Afghan speci-

mens of ssp. struvei, but the material for comparison is too scanty to allow a

definite statement.

The occurrence of ssp. struvei in Afghanistan makes it possible that the taxa

known as ssp. geron and ssp. struvei are partly sympatric, as ssp. geron is

known to occur at Arbarp (10 miles W of Kabul) (De Jong, 1974). A female from

Tshehaltan (vicinity of Kabul; CN) measures 13.5 mm. For that reason it may

belong to ssp. geron, but it can also be a small specimen of ssp. struvei. Unfor-

tunately, I have not yet found characters to distinguish between ssp. geron and ssp.

struvei in the female genitalia.

As the distance between the known populations of ssp. struvei and ssp. geron

in Afghanistan is about 70 km, the sympatry of both forms cannot be said to be

established at the moment and therefore they will provisionally be considered con-

specific.

Also the range of ssp. geron is considerably more extensive than previously

thought. In Coll. Brandt (NRS), 14 © and 19 from Iran were arranged under

“osthelderi”. The series turned out to consist of 5 °° of S. osthelderi, 40 and

12 of S. doris, and 5 d of S. geron. Examination of the genitalia showed the

geron specimens to belong to ssp. geron. The specimens originate from the fol-

lowing localities:

Iran: Fars, Shiraz, ca. 1600 m; Fars, road Shiraz—Kazeroun, Fort Sine—

Sefid, ca. 2200 m; Balouchistan, Bender Tchahbahar (= Chah Bahar).

The fine spinules at the distal edge of the area spinulosa of the female geni-

talia in S. geron (cf. De Jong, 1974, fig. 11) is found in both subspecies. They

seem to be absent from the edge in the other species of the phlomidis group and

may help to distinguish S. geron from the very similar S. osthelderi.

Spialia doris (Walker)

Ssp. doris. On re-examination the only specimen from Kenya mentioned in my

1974 paper turned out to belong to Spialia mafa. Nevertheless, ssp. doris does

occur in Kenya, according to a male caught in the semidesert area of the North-

eastern Province along the Tana River between Garissa and Bura (AMNH). Fur-

ther new localities of this subspecies are: El Ryadh (Saudi Arabia) (ZMB, ZSM),

and the Mudugh region (Garoe, Galkayu) in Somalia (MN).

Dr. Nakamura (Williamsville, U.S.A.) kindly informed me that S. doris spreads

northward through Israel at least as far as Herzelia, north of Tel Aviv, along

the coastal sand dunes, while it also occurs in the central and northern parts of the

Negev Desert. Besides Convolvulus lanatus, the species feeds on at least two or

54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

three more species of Convolvulus in Israel. The Israeli population is remarkably

variable. I agree with Dr. Nakamura, that this may be due to the fact that ssp.

doris and ssp. amenophis are largely intergrading in this area.

Another zone of intergradation between the last named subspecies may be found

in North Sudan: a male and female from Ed Damer (Hudeiba; ZSM) are rather

pale above and below, with large white markings, though not as large as usual

in ssp. amenophis.

Ssp. evanida. The following two localities, both situated in the Prov. of Fars,

S. Iran, can be added: Shiraz, Tchouroum (road Kazeroun—Bouchir) (NRS).

d. The colotes group

External characters. — Fore wing upperside with basal cell spot; central cell

spot midway between basal cell spot and discocellular spot; no spots in spaces 9

and 10; submarginal spots well-developed. Median spots on underside of hind wing

all present, separated or forming a continuous band, which is directed towards

the dorsum; submarginal spots 4 and 5 out of line, about halfway median spots

4— 5 and termen. Male with inconspicuous costal fold.

Male genitalia. — Uncus undivided; junction with tegumen slightly marked, no

fenestrula. Gnathos dorsally joined to tegumen over short distance, ventrally

passing into the membranous fultura superior, which is finely spinulose and folded

lengthwise. Anellus with dorsal sclerotization. Aedeagus straight, surface smooth.

Saccus well-developed, but shorter than uncus. Costa of valva of rather even width;

costal process more or less ovoid, with spines in its ventral part, most spines point-

ing dorso-cephalad, but a smaller number of longer spines directed dorso-caudad.

Cucullus well-developed, with a fold that partly conceals the costal process. A

large, spinulose area in the membrane between costa and sacculus, joined to the

costa.

Female genitalia. — Segment 8 with two dorsal and two ventral sclerites. Later-

ally, the dorsal and ventral sclerites almost fused, the ventral sclerites showing some

small ridges at this place. Membranous postvaginal region with a slight, rather

triangular sclerotization, imperceptibly passing into the deeply indented area spinu-

losa, which may be devoid of spines. Wall of ductus corrugated shortly before

bursa, but without distinct sclerotization. No signum. Papilla analis slightly longer

than broad, without basal lobe; apophysis posterior short, about as long as papilla.

Spialia colotes (Druce)

Pyrgus colotes Druce, 1875. — Proc. zool. Soc. London 1875: 416 (Angola). For synonymy, see

subspecies.

External characters (pl. 2 figs. 13, 14). — This species is distinguished from all

other Spialia species, except confusa, by the submarginal spots 4 and 5 of

the hind wing not being in line with the other submarginal spots, but placed

halfway between the median spots 4—5 and the termen. From confusa it can

be separated by the median spots on the upperside of the hind wing being separated

by the ground colour along the veins and placed more irregularly. The same

DE JONG: The genus Spialia SS

difference is found on the underside of the hind wing, but in one subspecies of

colotes the median spots are fused into a band, thus suggesting confusa. This

band, however, has irregular sides, especially in spaces 2, 3 and 6, while it is

straighter and with almost straight sides in confusa.

Male genitalia (figs. 44, 45). — Distal end of costa of valva rounded, not

folding over costal process.

Female genitalia (figs. 47, 48). — Ventral sclerites of eighth abdominal seg-

ment touching antevaginally, but not fused. Distal rim of postvaginal area with

minute spines (area spinulosa).

Ecology. — This species is mainly met with in dry, more or less wooded savan-

na, but I also took it in Kenya in a grassy glade in the Karura Forest near

Nairobi. In Aden it has only been found above 2000 m, but in Africa it is usually

found at lower altitudes.

In S. Africa colotes is on the wing from January to May (Swanepoel, 1953),

but from localities further north I have seen specimens taken in all months, except

June and July.

No food plants have so far been recorded.

Distribution (Map 6). — From Transvaal north to Angola, Ethiopia and Aden.

Judging from the few localities known of this widespread species, it is a local

species that possibly has been overlooked in large areas. The only country from

where a reasonable number of localities are known, is Kenya.

Geographic variation. — The study of the geographic variation is hampered by

the lack of material from large areas and the unequal representation from diffe-

rent regions, but according to the material available the following division can be

made.

Spialia colotes colotes (Druce)

Pyrgus colotes Druce, 1875. — Proc. zool. Soc. Londen 1875: 416 (Angola). Holotype (7) in the

British Museum (Nat. Hist.), London.

Pyrgus nora Plôtz, 1884. — Mitt. naturw. Ver. Neu-Vorpomm. 15 : 7 (Loango, Angola). Type lost.

The unpublished coloured figure by Plôtz was copied by Swinhoe (1908). It clearly represents the

species described by Druce as Pyrgus colotes, as stated correctly by Swinhoe.

External characters. — Spots relatively small and widely separated; of the

median spots on the upperside of the hind wing only spots lc, 2 and 4—5 pre-

sent. According to Higgins (1924), the ground colour of the upperside is rather

paler brown than in ssp. transvaaliae, but I did not observe this difference. As

large as ssp. transvaaliae, 9.6—11.7 mm.

A good figure of this subspecies is given by Oberthiir (1912, fig. 1326).

Distribution. — Only known from Angola. Higgins (1924) mentioned this sub-

species from Mombasa (Kenya). This record supposedly refers to an intermediate

between ssp. transvaaliae and ssp. semiconfluens.

Localities. — Angola: Loango (Plôtz, 1884; as nora); Barraca, Cuanza River

(BM).

Material examined. — 6 0° (10 holotype), 1 © , Angola (BM).

56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Spialia colotes transvaaliae (Trimen)

Pyrgus transvaaliae Trimen, 1889. — South African Butterflies 3: 286 (Transvaal). Holotype (9)

in the British Museum (Nat. Hist.), London.

External characters. — Spots large and prominent; on the upperside of the

fore wing the inner median spots 1b and Ic fused into a single spot that is often

the largest spot of the fore wing; on the upperside of the hind wing the median

spots lc, 2, 3 (usually), 4—5 and 6 present. On the underside of the hind wing the

spots large, but well separated by the veins. © 9.6—11.9 mm.

pee

Map 6

Spialia colotes

Distribution. — From Transvaal to Kenya and Uganda. In Kenya ssp. trans-

vaaliae mainly inhabits the western and mountainous regions. In the drier eastern

and southern parts it is replaced by ssp. semiconfluens, but many intermediate spe-

cimens can be found. In Uganda ssp. transvaaliae occurs in the wetter areas; in

the dry northeastern part it encounters ssp. semiconfluens. Apparently intermediate

populations are listed after ssp. semiconfluens, but intermediate specimens can also

be found in other localities.

DE JONG: The genus Spialia 57

MILLY

Figs. 44—46. Male genitalia, lateral aspect, and inside of right valva. 44—45, S. colotes transvaaliae

(Sigor, Kenya). 46, S. confusa obscura (Arabuko forest, Kenya). Figs. 4749. Female genitalia, latero-

ventral aspect, and papilla analis. 47—48, S. colotes transvaaliae (Arusha, Tanzania). 49, S. confusa

confusa (Upper Luangwa Valley, Zambia).

58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Localities. — Kenya: Lake Baringo, Bura (Teita), Chawia Forest (Teita Hills),

Chepalungu, Chyulu Hills, Mt. Elgon, Escarpement, Fourteen Falls, Gilgil, Karen,

Karura Forest, Davirondo, Kima, Kisii, Kisumu, Kitui, Langata, Lumbwa, Me-

kueni, Maranga, Meru, Mombasa, Nairobi, Ngong, Shimba Hills, Sigor, Sultan

Hamud, Teita Hills (AMNH, BM, CMP, ML, MN, NRS, USNM). Uganda: Bu-

kedi, Bundibugyo, Jinja (AMNH, ML, NMB). Tanzania: Arusha Nat. Park, Chi-

mala River, Great Craters, 60 mls S of Kondoa Irangi, Mt. Meru, Mkoe, Mpanda,

Ruaha Nat. Park (BM, ML, MN, ZSM). Rhodesia: Chirundi, Dotts Drift, Fila-

busi, Gwanda, Matetsi, Matopos, Nyanadzi River, Odzi River, Rusape, Sabi Valley,

West Nicholson (MN, NMB). South Africa. Transvaal: Blaauwberg, Dendron, Jo-

hannesburg, De Kroon, Leydsdorp, Pietersburg, Potchefstroom, Potgietersrust,

Tubex, Vivo, Warmbaths (BM; Swanepoel, 1953). Botswana: Maun, 16 mls N of

Maun (NMB). Namibia: Grootfontein, Griinfelde, Kalidona, Okahandja, Otjitambi,

Tsumele, Waterberg, Windhoek (SMW, ZSM).

Material examined. — 1240 ,47Q :99% ,299 , Kenya (AMNH, BM,

CMP, ML, MN, NRS, USNM); 3 © , Uganda (AMNH, ML); 75 , 3 9, Tanzania

Material examined. — 124% , 47 © : 997, 29 9 , Kenya (AMNH, BM,

CMP); 10 5,12 2 , Namibia (BM, SMW, ZSM).

Spialia colotes semiconfluens subspec. nov.

External characters (pl. 2 figs. 13, 14). — On the upperside the spots are smal-

ler than in ssp. transvaaliae, particularly the inner median spots 1b and Ic of the

fore wing and the median spot lc of the hind wing, but contrary to ssp. colotes,

all spots of ssp. transvaaliae are present. On the underside of the hind wing the

median spots are also smaller than in ssp. transvaaliae, but they are fused into a

continuous band, usually with the exception of median spot 7 which is separated

by dark scaling on vein 7. By the continuous band this subspecies is suggestive of

Spialia confusa, but in that species the band is much more regularly shaped and

the median spots are also fused into a continuous band on the upperside of the

hind wing.

Ssp. semiconfluens is on the average slightly smaller than the other subspecies:

O 9.3—10.9 mm. In most specimens the ground colour of the upperside is rather

paler brown than in ssp. transvaaliae.

Distribution. — Aden, Ethiopia and Somalia and southwards into Kenya and

Uganda. In Kenya this subspecies flies especially in the dry east and southeast

and is replaced by ssp. transvaaliae in the less dry mountainous regions and

western part of the country. In Uganda ssp. semiconfluens is restricted to the

dry north-eastern regions.

Material examined. — Holotype: , Bihendula (Somalia), 4.xi.1949, K. M.

Guichard. Paratypes: 1 , Bihendula (Somalia), 1 v. 1949, K. M. Guichard;

1 © , Harar (Ethiopia), 23.xi.1939, R. F. Ellison; 10 , W. Errer Valley (Ethio-

pia), 28.xi.1939, R. E. Ellison; 10° , E of Errer Valley (Ethiopia), 19.xi.1939,

R. E. Ellison; 1 9 , Fich Shoa Prov. (Ethiopia), 8.v.1947, K. M. Guichard. All

types in the British Museum (Nat. Hist.), London.

DE JONG: The genus Spialia 59

Further material. — 49 © , 20 Q : 35 , 1 9 , Somalia (Bardera, Bihendula,

GanLibah, Haro-Bussar) (BM); 110° , 5 9 , Ethiopia (Dakotta Valley, Dire Dawa,

Errer Valley, Fich, Harar, Mulata Mts) (AMNH, BM); 17 , 2 2 , South Yemen

(Jebel Jihaf) (BM); 23 © , 10 © , Kenya (Kacheleba, Kenani, Kima, Kitui, Kulal,

Makueni, Mutomo, Namanga, Ukazzi Hill (AMNH, MN); 119 , 2 2 , Uganda

(Amudat, Kotido) (MN).

Material examined from populations intermediate between ssp. transvaaliae and

ssp. semiconfluens. — 229 ,4 © , Kenya (Besil, Kibwezi, Voi) (BM, ML).

Spialia confusa (Higgins)

Hesperia transvaaliae var. confusa Higgins, 1924. — Trans. ent. Soc. London 1924: 90 (SW shore

Lake Nyassa). For synonymy, see subspecies.

External characters (pl. 2 figs. 15, 16). — Readily distinguished from all

other Spialia species except colotes by the basad shift of the submarginal

spots 4 and 5 of the hind wing. From colotes it can be separated by the median

spots of the hind wing, which are fused into a continuous band, not only on the

underside, but also on the upperside. The band on the underside of the hind wing

is much more regularly shaped than in colotes semiconfluens. In rare doubtful

cases the genitalia have to be checked for identification.

Male genitalia (fig. 46). — Distal end of costa of valva slightly protruding and

folding over costal process.

Female genitalia (fig. 49). — Ventral sclerites of eighth abdominal segment

broadly fused antevaginally. Sclerotization in postvaginal area slightly stronger than

in colotes, without spines along distal edge.

Ecology. — Very little is known about this apparently rare species. Swanepoel

(1953) found it in Natal “playing along a road that leads through the bush”, and I

took it along very dry, sandy roads in the Arabuko Forest near Malindi, Kenya. It

is difficult to capture as it is a small and swift species, which frequently skips

from sun-lit places into shadows and then appears to be lost.

It may be on the wing throughout the year, but I have not seen specimens or

records from May and June.

According to Sevastopulo (1974) the larvae live on Melhamia (Sterculiaceae)

and Triumfetta (Tiliaceae).

Distribution (Map 7). — This species has been observed over a wide range, from

Natal to Kenya, in seven countries, but at relatively few localities.

Geographic variation. — There is some variation in the development of the

spots, but in view of the scarce material any division into subspecies can only be

provisional.

Spialia confusa obscura (Higgins)

Hesperia transvaaliae var. obscura Higgins, 1924. — Trans. ent. Soc. London 1924 : 92—93 (Nairobi).

Holotype (o ) in the British Museum (Nat. Hist.), London.

External characters (pl. 2 figs. 15, 16). — A dark form, with smaller spots. On

60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

the upperside of the fore wing the central cell spot vestigial or absent, on the upper-

side of the hind wing median spot Ic faint or absent. 7 8—10.2 mm.

Distribution. — The northern part of the range of the species, in Kenya and N.

Tanzania.

Localities. — Kenya: Arabuko Forest, Nairobi, Rabai, Sokoke Forest, Shimba

Hills (AMNH, BM, ML, MN; Sevastopulo, 1974). Van Someren (1939) mentioned

confusa in a report on his expedition to the Chyulu Hills, but his specimens (in

MN) turned out to belong to S. colotes transvaaliae with slight influence of ssp.

semiconfluens. Tanzania: Amani, Dar-es-Salaam, Paga Hill (BM, MN).

Material examined. — 170,39 : 140 (10 holotype), 2 © , Kenya

(AMNH, BM, ML, MN); 30”, 1 2 , Tanzania (BM, MN).

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Spialia confusa

Spialia confusa confusa (Higgins)

Hesperia transvaaliae var. confusa Higgins, 1924. — Trans. ent. Soc. London 1924 : 90 (SW shore -

Lake Nyasa, between Ft. Johnston and Monkey Bay). Holotype (0 ) in the British Museum (Nat.

Hist.), London.

Hesperia transvaaliae var. fasciata Higgins, 1924. — Trans. ent. Soc. London 1924 : 90—91

(between Ft. Mangoche and Chikala Boma, Malawi). Holotype (© ) in the British Museum (Nat.

Hist.), London. Evans (1937) incorrectly stated the type to be a male.

DE Jonc: The genus Spialia 61

External characters. — Spots larger than in the foregoing subspecies; on the

upperside of the hind wing median spot lc well-developed. 7 9.8—10.9 mm.

The type of fasciata is not clearly different from that of confusa, except for the

median band on the hind wing, which is rather narrower. There is no apparent

reason to rank fasciata otherwise than as a forma of ssp. confusa.

Distribution. — The southern part of the range of the species, from C. Tanzania

and N. Zambia to Natal.

Localities. — Tanzania: Mikumi Nat. Park, Mkami, Mkoe (ML, ZSM). Zambia:

Upper Luangwa Valley, Petauke (BM, HDE; Neave, 1910, as “Hesperia bettoni”).

Malawi: between Ft. Johnston and Monkey Bay, between Ft. Mangoche and

Chikala Boma (BM). Rhodesia: Sanyati Valley (= Umniati R.) (NMB). Mozam-

bique: Delagoa Bay, Inhaminga, Siluve (= Xiluvo) Hills (BM, NMB, USNM,

ZSM). South Africa. Natal: False Bay, Hluwehluwe, Messina, Zulu (NMB, NRS;

Swanepoel, 1953).

Material examined. — 18 7,5 9 :39 ,19 , Tanzania (ML, ZSM); 29,

19 , Malawi (10 holotype confusa, 1 © holotype fasciata) (BM); 55 ,39 ,

Zambia (BM, HDE); 6 , Mozambique (BM, USNM, ZSM); 1 3 , South Africa

(NRS).

e. The sertorius group

External characters. — Fore wing with or without basal cell spot on upperside;

if basal cell spot present, then central cell spot about midway between basal cell

spot and discocellular spot or slightly closer to basal cell spot; no spots in spaces 9

and 10; submarginal spots usually all present, but may be vague. Median spots on

underside of hind wing form a continuous band or spot 6 is absent; submarginal

spots 4 and 5 in line with the other submarginal spots. Male without costal fold.

Male genitalia. — Uncus undivided; at junction with tegumen in some species

with large fenestrula. Gnathos dorsally not fused to tegumen, well sclerotized,

ventrally forming enlarged and spined “heads”; left and right part of gnathos

ventrally united by a usually smooth membrane. Anellus membranous. Aedeagus

more or less straight, surface smooth. Saccus relatively short, shorter than uncus.

Costa of valva narrow; costal process well developed, more or less oval, ventrally

with spines which are directed dorso-distad. Cucullus dorsally excavated or with a

small fold that partly overlaps the costal process. A hairy fold in the membrane

between costa and cucullus.

Female genitalia. — Segment 8 with two dorsal sclerites, medially separated by a

membrane, and two ventral sclerites, united antevaginally by a narrow sclerotiza-

tion. Dorsal and ventral sclerites more or less fused. Postvaginally a membranous

region with a proximally narrow, distally broadening sclerite, which at the distal

edge of the segment passes into the spinoluse region, or without this sclerite so that

the area spinulosa can be seen as a dark patch in the membranous distal part of the

ventral surface. Ductus gradually broadening into bursa. Ductus and bursa without

ornamentation, or bursa with a finely spinulose area. Papilla analis with basal

lobe; apophysis posterior long, longer than papilla.

62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Superspecies Spialia mafa

External characters. — Fore wing with a basal cell spot on the upperside. Median

spots on the underside of the hind wing complete or spot 6 is absent.

Male genitalia. — A large fenestrula at the junction of uncus and tegumen.

Heads of gnathos larger than in superspecies Spialia sertorius. Costa of valva

continued beyond the costal process as a finger-shaped projection. Hairy fold at the

inside of the valva obvious, oblique, wavy or branched.

Female genitalia. — Dorsal sclerites of eighth abdominal segment fused to the

ventral sclerites along the proximal edge of the segment, leaving a wide membranous

area, or connecting sclerotization between dorsal and ventral sclerites more extensive,

enclosing a membranous area. Bursa copulatrix with or without ornamentation.

Ecology. — Food plants: Malvaceae.

Distribution. — Very widely distributed, from South Africa through East Africa,

Saudi-Arabia and India to Hainan. It replaces the superspecies Spialia sertorius in

the Old World Tropics.

Spialia mafa (Trimen)

Pyrgus mafa Trimen, 1870. — Trans. ent. Soc. London 1970 : 386, pl. 6 fig. 12 (Maseru, Basutoland).

For synonymy, see under subspecies.

External characters (pl. 4 figs. 25, 26). — From the other species of the sertorius

group, mafa is easily distinguished by the combination of the presence of a basal

cell spot on the upperside of the fore wing and the absence of median spot 6 on

the underside of the hind wing. The other Spialia species, which almost or entirely

lack median spot 6 on the underside of the hind wing and occur in the Ethiopian

region, are doris, paula and spio. The differences with these species have been

outlined in the key and it is unnecessary to repeat them here.

There is some variation in the extension of the spots. On the upperside of the

hind wing the median spots 1c—5 are usually present, 6 is always absent. The

ground colour of the underside of the hind wing is lighter or darker yellowish or

brownish.

See further under Geographic variation.

Male genitalia (figs. 50, 51). — At junction of uncus and tegumen a large

fenestrula. Spined heads of gnathos rather slender, less than half of entire gnathos.

Costal process of valva situated before distal end of costa. Distal part of costa looks

like a finger-shaped projection of the costa instead of being the costa itself. Cucullus

with short fold covering ventral part of costal process. On the inside of the valva,

between costa and cucullus, a large, branched fold, very thickly beset with short

hairs.

Female genitalia (figs. 54, 55). — Dorsal sclerites of segment 8 more or less

fused with ventral sclerites, enclosing a small, membranous area. Ventral sclerites

antevaginally conjoined by a narrow sclerotization, postvaginally separated by a

narrow membranous strip. This strip widens distally and passes into the non-

sclerotized area spinulosa, which has a more or less straight distal edge. Bursa with-

out ornamentation. Papilla analis rather narrow, apex rounded.

Ecology. — Like most other Spialia species, mafa prefers more or less open

DE JONG: The genus Spialia 63

country, but judging from its distribution it largely avoids low-lying areas, being

mainly found in hilly or mountainous regions. In Kenya I took this species in open

places in a more or less wooded area up to 2000 m (between Nanyuki and Naro

Moru) and Gabriel (1954) recorded specimens from Yemen captured up to more

than 2500 m. According to Swanepoel (1953) it is a scarce and local species, at

least in South Africa.

Figs. 50—53. Male genitalia, lateral aspect, and inside of right valva. 50—51, S. mafa higginsi

(Kenya). 52—53, S. galba galba (Ceylon).

In appropriate places mafa may be on the wing throughout the year, but in

many areas adults are absent or very scarce in June, July and August.

The only recorded food plant is Hibiscus aethiopicus L. (Malvaceae) (Murray,

1959).

Distribution (Map 8). — South and East Africa, not known from Angola, and

Zaire except for the extreme south. Distributed from Cape Town and SW Africa to

Ethiopia and Sudan, and extending into the Arab peninsula, where it occurs as far

east as Oman.

Geographic variation. — There is a marked geographic variation in the size of

the specimens and the development of the spots. Two subspecies are distinguished.

64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Spialia mafa higginsi Evans

Spalia mafa higginsi Evans, 1937. — Catal. Afric. Hesp. : 62 (Mumias, Kenya). Holotype (7) in

the British Museum (Nat. Hist.), London.

Map 8

Spialia mafa

External characters (pl. 4 figs. 25, 26). — The original description reads: ‘Below,

the markings are broader and on the hind wing the basal spot in space 7 and the

discal spot in space Ic are not detached as in mafa”. These differences are rather

slight, but more conspicuous differences can be found in the following characters:

1. ssp. higginsi is smaller than ssp. mafa, 9 (8.5—) 9.2—10.7 (—11.3)

mm, © 9.8—11.3 (—12.6) mm, as against ssp. mafa © 9.9—11.7 mm,

(9.1—) 10.2—12.5 mm. This difference is conspicuous if one can compare long

series of both subspecies;

2. in ssp. mafa the median spot 5 on the underside of the hind wing is almost

always extended basad, just above the continuation of vein 5 in the cell; this is

almost never, and certainly never conspicuously the case in ssp. higginsi.

Distribution. — The northern part of the range of the species. The southern

limit presumably runs through Malawi, Zambia and S. Zaire. As most specimens

DE JONG: The genus Spialia 65

examined from Zambia show characters of ssp. mafa, all specimens from this

country have provisionally been listed under that subspecies. Additional material will

possibly indicate a transition zone. The specimens examined from S. Zaire are

listed under the present subspecies, but they probably also belong to the transition

zone.

Localities. — Saudi-Arabia: Ashaira (BM). Oman: Khasab, Nizwa area, Rostaq

(Larsen, 1977). Yemen: Hada, San’a, Sug-es-Sabt (N of Ibb) (BM, ZSM; Gabriel,

1954). South Yemen: Dhala, Jebel Jihaf (BM; Gabriel, 1954). Sudan: Erkowit

(Prov. Kassala) (ZSM). Ethiopia: Addis Abeba, Burdji, Daroli, Dessié, Getri, Harar,

Jowaha, Negelli, Ft. Zuguala (BM, MN, ZSM). Kenya: Besil (= Bissil), Broderick

Falls, Bura (Teita), Campi-ya-Moto, Escarpment, Gilgil, Ilala, Kabarnet, Kedai,

Kima, Kitale, Mazoe Valley, Meru Distr., Migori Valley, Mirua Valley, Mt. Kenya,

Muruanysigar Mt. (Turkana), Nairobi, Naro Moru, Ngong Escarpment, Olkasale,

Rumuruti, Saboti Hill, Soi, Stony Athi, Subukia, Thomson Falls (AMNH, BM,

ML, MN, MNHN, USNM). Uganda: Entebbe, Koki Country, Siroko River, Tororo,

Queen Elisabeth Park (AMNH, BM, CMP, MN). Tanzania: Chukwe (Mpanda),

District of Great Craters, Njombe, Tendaguru (BM, ML). Zaire: Kapiri, Musonoie

(BM, MRAC). Malawi: Cholo (CMP).

Material examined. — 104 , 44 92 : 1 Saudi-Arabia (BM); 7% ,29,

Yemen (MB, ZSM); 4 ” , 4 © , South Yemen (BM); 2 à, Sudan (ZSM); 110,

30 , Ethiopia (BM, MN, ZSM); 64 © (1 5 holotype of higginsi), 29° ,

Kenya (AMNH, BM, ML, MN, USNM); 87 , 6 © , Uganda (AMNH, BM,

CMP, MN); 4 5 , Tanzania (BM, ML); 2 ” , Zaïre (BM, MRAC); 1 © , Malawi

(CMP).

Spialia mafa mafa (Trimen)

Pyrgus mafa Trimen, 1870. — Trans. ent. Soc. London 1870 : 386, pl. 6 fig. 12 (Maseru, Basuto-

land). Holotype (7) in the British Museum (Nat. Hist.), London.

Hesperia oberthüri Aurivillius, 1925. — In: Seitz, Macrolepidoptera of the World 13 : 565 (Kimberley).

Holotype (© ) in the British Museum (Nat. Hist.), London. This name was given to a figure of mafa

by Oberthiir, which was incorrectly named transvaaliae. The figured specimen is in the BM. Pre-

occupied by Hesperia oberthüri Leech, 1891 (now placed in the genus Pyrgus).

“ie aurivillii Shepard, 1935. — Lepid. Catalogus, Pars 69 : 496. Replacement for Hesperia oberthüri

urivillius.

External characters. — The differences with ssp. higginsi have been dealt with

under that subspecies.

Distribution. — The southern part of the range of the species, from S. Zaire,

Zambia and Malawi southwards.

Localities. — Zambia: Chilanga, Fort Jamieson, Livingstone, Luangwa Valley

(BM, MN; Neave, 1910). Malawi: Florence Bay, Ruo Valley (BM; Gifford, 1965).

Rhodesia: Bulawayo, Darwendale, Essexvale, Hope Fountain, Matetsi, Matope

Hills, Odzi, Salisbury, Selukwe, Umtali, X-mas Pass (BM, CMP, MN, NMB).

Mozambique: Rikatla (24 km N of Lourengo Marques) (BM). Botswana: Sepopa,

Tshabong (NMB). South Africa. Transvaal: Balfour, Barberton, Crocodile River,

Dendron, Germiston, Haenertsburg, Johannesburg, Limpopo River, Messina, Pieters-

burg, Potchefstroom, Pretoria, Rustenburg, Shilouvane (BM, CMP, MN, NMB;

Swanepoel, 1953; Trimen, 1870). Natal: Bergville, Estcourt, Frere, Greytown,

66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Howick, Impenza, Ladysmith, Loesskop, Middle Rest, Montello, Muden, Newcastle,

Pitzkop, Tugela River, Weenen (AMNH, BM, MN, MRAC; Swanepoel, 1953).

Orange Free State: Bethulie, Bloemfontein, Ladybrand, Vaal River (AMNH, BM,

MRAC, NMB; Swanepoel, 1953). Cape Province: Barkly District, Brakfontein,

Burghersdorp, Cape Town, Durbansville, Katzenberg, Kimberley, Lions Head,

Malmesbury, Mamre, Maseru, Melkbosch, Stellenbosch, Tijgerberg (BM, ML,

MN, MNHN, MRAC, NMB; Swanepoel, 1953). Lesotho: Koro Koro, Maseru,

Namanthes (BM, NMB; Swanepoel, 1953; Trimen, 1870). Namibia: Niangana,

Okahandja (SMW). Angola: Ehanda (Trimen, 1891).

Material examined. — 73 &, 569 :40,19 , Zambia (BM); 10 ,19 ,

Malawi (BM); 57, 12 , Rhodesia (BM, CMP, MN); 1 © , Mozambique (BM);

620 (1 © holotype mafa), 49 ” , South Africa (AMNH, BM, CMP, ML,

MRAC); 49 , Namibia (SMW).

Spialia galba (Fabricius)

Hesperia galba Fabricius, 1793. — Entomologia Systematica 3 : 352 (Tranquebar).

External characters (pl. 4 figs. 27, 28). — From the other species of the sertorius

group galba is easily distinguished by the combination of a well-marked basal cell

spot on the upperside of the fore wing and an unbroken, rather regular median

band on the underside of the hind wing. As galba is the only real Oriental species

of the genus, confusion with other Spialia species is unlikely. However, in the

extreme west of the Oriental Region distributional overlap with zebra and doris

does not seem impossible and it may be convenient to mention some differences.

From zebra, galba is distinguished by the presence of a well-marked basal cell

spot on the upperside of the fore wing, and from doris by the unbroken median

band on the underside of the hind wing.

There is some variation in the development of the spots and in the size ( © 9.4—

11.8 mm), but the variation does not appear geographic, except for the variation

mentioned under Geographic variation.

Swinhoe (1913) thought “Pyrgus evanidus Butler” to be “undoubtedly the

extreme cold-weather form of galba”. However, “Pyrgus evanidus” is considered

a subspecies of Spialia doris (cf. De Jong, 1974).

Male genitalia (figs. 52, 53). — Very large fenestrula at junction of uncus and

tegumen. Gnathos with large ventral “heads”, occupying more than half of the

entire gnathos. Costal process of valva placed before distal end of costa; continu-

ation of costa finger-shaped, longer than in mafa. Cucullus with narrow fold, just

covering ventral side of costal process. Between costa and cucullus a winding, but

unbranched hairy fold on the inside of the valva.

Female genitalia (figs. 56, 57). — Dorsal sclerites of segment 8 along proximal

edge fused with ventral sclerites. Ventral sclerites conjoined antevaginally by a

narrow sclerotization. Postvaginally there is a triangular membranous area, distally

enclosed by the ventral sclerites, which fuse while the sclerotization weakens. Distal

edge of ventral sclerites vague, sclerotization passing into membrane that forms a

pointed area spinulosa. Bursa with large area of small spines. Papilla analis rather

DE JoNG: The genus Spialia 67

Figs. 54—57. Female genitalia, ventral aspect, and papilla analis. 54—55, S. mafa higginsi (Kenya).

56—57, S. galba galba (Ceylon).

short and broad, less rounded than in mafa, slightly triangular; basal lobe large,

curving.

Ecology. — S. galba can commonly be found throughout the year in all sorts

of open country, but in some areas it is absent during the dry season. It ascends to

2400 m in the Himalayas (ZSM) and to 2600 m in South India (Wynter-Blyth,

1957), but in Ceylon it is rare above 1300 m (Woodhouse, 1950).

68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Recorded food plants are Sida rhombifolia, Hibiscus and Waltheria indica (all

Malvaceae) (Swinhoe, 1913; Woodhouse, 1950).

Distribution (Map 9). — Throughout India and Ceylon. In the west it extends to

Karachi and Chitral, in the east it penetrates Burma as far as the S. Shan States.

A single specimen is known from eastern Thailand. Still further east it occurs

isolated in the island of Hainan.

Geographic variation. — Over its large Indian area galba does not seem to vary

geographically, but in the east two subspecies can be distinguished, one confined

to Hainan, the other occurring in Burma. It is obscure how far the Burmese sub-

species comes into contact with the nominate form from India. The single known

specimen from Thailand cannot be distinguished from Indian specimens, so for the

time being it is listed under the nominate subspecies.

Spialia galba galba (Fabricius)

Hesperia galba Fabricius, 1793. — Entomologia Systematica 3 : 352 (Tranquebar).

According to Zimsen (1964) the type material includes three specimens. One of these specimens

(which are in the Universitetets Zoologiske Museum, Copenhagen), however, though bearing a label

with “H. Galba” in probably Fabricius’ handwriting, comes from West Africa (“Guinee”) and

belongs to S. diomus. The remaining two specimens do not bear locality labels, but as they belong to

the only Spialia species known to occur at the type locality of S. galba, one of these specimens

comes into consideration as type specimen. One specimen, labelled “Galba” in probably Fabricius’

handwriting, lacks the abdomen. The other specimen, a male labelled ‘““Mus. Seh. e T.L.” (i.e. “Mu-

seum Sehestedt e Tgnder Lund”) is best fitted for lectotype designation, as Fabricius mentioned the

specimen to be in the Sehestedt collection. I have labelled the specimen accordingly.

Pyrgus superna Moore, 1865. — Proc. zool. Soc. London 1865 : 792 (Calcutta). Holotype ( © ) in the

British Museum (Nat. Hist.), London.

External characters (pl. 4 figs. 27, 28). — On the upperside of the fore wing

median spot 4 conspicuously smaller than median spot 3, or absent; submarginal

spot 2 in line with the other submarginal spots. o 9.4—11.8 mm. Larger and

smaller specimens occur throughout the area. A very small specimen (+) from the

Himalayas (Simla) was named ab. minuscula by Reverdin (1933), a redundant

name; the specimen is in the ZSM.

Distribution. — The western subspecies, occupying the whole range of the

species west of Burma. As the single known specimen from Thailand cannot be

distinguished from this subspecies, it is provisionally listed here.

Localities. — Pakistan: Chitral, Karachi (BM, LNK). India: throughout, from

Kutch in the west to the Naga Hills in the east, and from Kashmir in the north to

Madras in the south (AMNH, BM, ML, ZSM). Ceylon: throughout (AMNH, BM,

CMP, ML, USNM). Nepal: Baklore (W. Terai), Bhainse Dobhan (Chisapani

Garhi), Katmandu, Katmandu Valley, Magarkot Manbu, Rangit Valley, Rapti

Valley, Sun Khosi Valley (AMNH, BM, ZSM). Sikkim: Sikkim (BM, UNSM).

Bhutan: Bhutan (BM, USNM). Thailand: Khun Kaen (USNM).

Material examined. — 229 © , 163 9 : 69,2, Pakistan (BM, LNK);

157 , 118 © , India (AMNH, BM, CMP, ML, ZSM); 34 © , 24 2 , Ceylon

(AMNH, BM, CMP, ML, USNM); 19 ” , 10 © , Nepal (AMNH, BM, NRS,

ZSM); 117,8 © , Sikkim (BM, USNM); 20 , 19 , Bhutan (BM, USNM);

1% , Thailand (USNM).

DE JONG: The genus Spialia 69

Spialia galba shanta Evans

Spialia galba shanta Evans, 1956. — Ann. Mag. Nat. Hist. (12) 9 : 750 (Hsipaw, S. Shan States).

Holotype (7) in the British Museum (Nat. Hist.), London.

External characters. — On the upperside of the fore wing median spot 4 well-

developed, as large as median spot 3; submarginal spot 2 in line with the other sub-

marginal spots. © 10.4—11.5 mm.

Distribution. — N. and S. Shan States, Burma.

Localities. — Burma: Hsipaw, Maymyo, Tilin Yaw, Poungadaw nr. Thayet-

myo, Myingyan, 40 km E. of Taunggyi (BM, NRS).

Material examined. — 7 © , 12 © , Burma (incl. “ holotype) (BM, NRS).

Spialia galba chenga Evans

Spialia galba chenga Evans, 1956. — Ann. Mag. Nat. Hist. (12) 9 : 750 (Kiung-Chow, Hainan).

Holotype (o ) in the British Museum (Nat. Hist.), London.

External characters. — On the upperside of the fore wing median spots 3 and 4

equal, as in ssp. shanta, submarginal spot 2 placed further from termen than in

7 ge}

Map 9

Be galba

70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

other subspecies, giving the submarginal series a much more sinuous aspect.

O 10.6—10.7 mm.

Distribution. — Hainan.

Localities. — Hainan: Cheng-Mai, Kiung-Chow, Porten, Youboi (BM).

Material examined. — 2 © ,3 © , Hainan (incl. + holotype) (BM).

Superspecies Spialia sertorius (Hoffmansegg)

This superspecies has extensively been dealt with in my previous paper on

Figs. 58—59. Male genitalia, lateral aspect, and inside of right valva of S. orbifer orbifer (Greece).

Figs. 60—61. Female genitalia, ventral aspect, and papilla analis of S. sertorius sertorius (Chiclana, S.

Spain).

DE JONG: The genus Spialia 71

Spialia (De Jong, 1974). It may suffice to mention here only the characters by

which the two constituting species are distinguished from other Spialia species

and to give some new information on the distribution of the species.

External characters (pl. 4 figs. 29—-32). — From the other species of the

sertorius group, viz., mafa and galba, sertorius and orbifer are easily distin-

guished by the absence of a basal cell spot on the upperside of the fore wing, while

this character in combination with the absence of median spot 6 on the underside

of the hind wing and the direction of the median band towards the inner spot in

space 7 separates sertorius and orbifer from all other Spialia species.

Male genitalia (figs. 58, 59). — No fenestrula at junction of uncus and tegumen.

Gnathos with relatively narrow heads. Costal process of valva at distal end of

costa. Cucullus dorsally hollowed to receive the costal process, but without definite

fold. Hairy fold between costa and cucullus on inside of valva curved or angled,

neither winded nor branched.

Female genitalia (figs. 60, 61). — Dorsal sclerites of segment 8 broadly fused to

ventral sclerites; the enclosed membranous area between the dorsal and ventral

sclerites found in mafa and galba reduced to a small pit at the top of a lateral

bump. Antevaginally ventral sclerites connected by sclerotization that is continued

over short distance into ductus. Postvaginally a wide membranous area, containing

a large oblong sclerotization, which starts from the ductus and distally expands

strongly before passing into the area spinulosa. Bursa without ornamentation.

Papilla analis slightly pointed.

Ecology. — Food plants: Rosaceae.

Distribution. — The combined distribution areas of sertorius and orbifer

comprise the greater part of the Palaearctic Region, from the Atlantic Ocean to

Korea and the Amur Region, southwards to NW. Africa, Israel and N. Baluchistan,

northwards to the Netherlands, C. Germany, C. Russia and into Siberia. The only

other Spialia species, which occur within this area, belong to the phlomidis group.

Spialia sertorius (Hoffmansegg)

In my 1974 paper on Spialia I referred to a paper by Kauffmann (1955), who

reported the occurrence of ssp. sertorius in Corsica, suggesting that ssp. sertorius

and ssp. therapne are sympatric in this island. Recently I could study the material

that prompted Kauffmann to the mentioned paper: 20 specimens in the ZSM, all

bearing the following data: Evisa, 15.VI.—2.VII.1954, Leinfest. They are insep-

arable from ssp. sertorius. The long series, all from the same date and locality,

suggests the presence of a prospering population. However, the prosperity of the

population makes it improbable that it was not found before (and after as well).

For this reason I have some doubts about the correctness of the labels.

Spialia orbifer (Hiibner)

1. Ssp. orbifer. In addition to the 10 ” and 6 © from Sicily mentioned in

1974, I have studied 4 and 1 © from this island (ZSM). They confirm the idea

72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

that there is no pure orbifer in Sicily, the female being similar to sertorius, the

males being closer to orbifer. Their sizes fall within the limits mentioned in 1974

for Sicilian specimens.

2. Ssp. carnea. The 55 additional specimens studied from Afghanistan dem-

onstrate that this subspecies is rather well distributed in this country, but it is

scarcely found below 2000 m. It appears that in most specimens the submarginal

spot in spaces 4 and 5 on the underside of the hind wing is as large as in other

subspecies of orbifer, contrary to what was stated in my 1974 paper. In a nice

series of 24 0° and 2 © from Badakhshan (CN) all males are dark due to almost

complete absence of submarginal spots on the upperside, while these spots are well-

developed in the two females. On the average the specimens are smaller than

carnea specimens from other localities, the males measuring 11.8—13.5 mm. The

ground colour of the underside of the hind wing is yellow or brown rather than

red, and more or less strongly overshaded by black scales. The spots on the under-

side of the hind wing are well-developed, contrary to the upperside, where in most

cases the median band is reduced to an exclamation mark: a white bar closing the

cell and a white point in space 2.

f. The dromus group

External characters. — Basal cell spot on fore wing upperside present; central

cell spot central between basal cell spot and discocellular spot; no spots in spaces

9 and 10; submarginal spots usually all present, but sometimes vague; outer median

spot lb absent, lc absent or present; inner median spot lb present, lc rarely

absent. Median spots on hind wing underside all present (except in paula, where

median spot 6 is absent), forming a continuous band of very diverging shape and

direction; submarginal spots 4 and 5 in line with the other submarginal spots. Male

without costal fold.

Male genitalia. — Uncus undivided; junction with tegumen slightly marked, no

fenestrula. Gnathos dorsally joined to tegumen over short distance, ventrally passing

into a semiglobular fold, which is finely spinulose, thus suggesting sclerotization;.

the fold is medially uplifted, so that there is a left and right part. Anellus dorsally

strongly sclerotized to form a roof or a winged structure over the aedeagus.

Aedeagus straight or slightly S-shaped, surface smooth. Saccus relatively long.

Costal process of valva well-developed, with long spines usually directed cephalad

(towards base of valva) or cephalo-dorsad (spines upturned). Cucullus with a fold,

that partly conceals the costal process.

Female genitalia. — Segment 8 with two dorsal and two ventral sclerites, sep-

arated by membranes. The ventral sclerites may touch each other, but are not

conjoined. Membranous postvaginal region with a slight triangular sclerotization

that usually imperceptibly passes into the area spinulosa. Ductus usually long and

gradually passing into bursa. Bursa without ornamentation. Papilla analis without

basal lobe; apophysis posterior usually longer than papilla.

DE JONG: The genus Spialia 73

Map 10

Spialia wrefordi

Spialia wrefordi Evans

Spialia wrefordi Evans, 1951. — Ann. Mag. Nat. Hist. (12) 4 : 1270—1271 (Kotido, Karamoja,

Uganda). Holotype (o ) in the British Museum (Nat. Hist.), London.

External characters (pl. 5 figs. 33, 34). — Easily recognized by the extensive

spotting on the upper and undersides of the wings. The very large basal cell spot on

the upperside of the fore wing (about twice as long as broad and much larger than

the central cell spot) is found in no other Spialia species. On the upperside of the

hind wing the most distinctive character, apart from the strong development of the

spots, is the median spot lc that runs from vein 1b to vein 2. On the underside of

the hind wing the median band is more or less divided by darker veins, especially

veins 4 and 7, so that the underside reminds of a strongly spotted S. colotes, but

the submarginal spots 4 and 5 are not out of line as in that species. ” 8.5—11.7

mm, © 10.5—13 mm.

Male genitalia (figs. 62, 63). — Dorsal sclerotization of anellus laterally upturned.

Aedeagus S-shaped. Saccus distinctly shorter than uncus. Costal process with

straight, horizontal spines, directed cephalad, not upturned. Near base of valva a

spinulose area in the membrane between costa and sacculus.

Female genitalia (figs. 70, 71). — Area spinulosa deeply indented, forming a gap

74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

that is closed by the distinct, triangular, postvaginal sclerotization. Papilla analis

short, broader than long; apophysis posterior short, slightly longer than length of

papilla, slightly shorter than width of papilla.

Ecology. — Unknown. The known specimens have been caught during the

months April, June and July. Apparently restricted to dry and hot areas.

Figs. 62—65. Male genitalia, lateral aspect, and inside of right valva. 62—63, S. wrefordi (Kotido,

Uganda). 64—65, S. paula (Essexvale, Rhodesia).

Distribution (Map 10). — Only known from four localities, viz., Kotido (Uganda,

Karamoja), Samburu Game Reserve (Kenya, Rift Valley Prov.), Mt. Kulal and Mt.

Marsabit (both Kenya, Eastern Prov.). Undoubtedly, this species is more widely

distributed in the badly known dry northern parts of Uganda and Kenya.

Material examined. — 19 , 12 © : 18 & (incl. holotype), 10 9 , Uganda

(AMNH, BM, ML, MN); 17 , 2 9 , Kenya (AMNH, BM, ML).

Spialia paula (Higgins)

Hesperia paula Higgins, 1924. — Trans. ent. Soc. London 1924 : 77, pl. 8 fig. 22 (Bulawayo,

Rhodesia). Holotype (o) in the British Museum (Nat. Hist.), London.

External characters (pl. 5 figs. 35, 36). — This species differs from the other

DE JONG: The genus Spialia 75

species of the dromus group in the absence of median spot 6 on the underside of

the hind wing. In this respect it may be confused with mafa and spio, though

these species, and especially spio, are usually much larger. However, the shining

white of most spots on the underside of the hind wing and the fusion of median

Figs. 66—69. Male genitalia, lateral aspect, and inside of right valva. 66—67, S. ploetzi ploetzi (Nam-

wamba Valley, Uganda). 68—69, S. ploetzi occidentalis (Warri, Nigeria).

spots Ic and 2 on the same wing make the distinction of paula not extremely

difficult.

In the few specimens studied the length of the fore wing varies in the male from

8.4 to 9.1 mm, in the female from 10 to 10.2 mm.

Male genitalia (figs. 64, 65). — Dorsal sclerotization of anellus laterally bent

downwards, forming a roof over the aedeagus. Aedeagus straight. Saccus slightly

shorter than uncus. Costa of valva in distal part indistinctly delimited. Costal process

narrow, curving upwards, so that the spines point cephalo-dorsad.

Female genitalia (figs. 72, 73). — Ventral sclerites of segment 8 very narrowly

conjoined antevaginally. Weak triangular sclerotization in membranous postvaginal

76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

area, separated from slightly indented area spinulosa by a narrow membranous

band. Papilla analis pointed, longer than wide; apophysis posterior much longer

than papilla.

Ecology. — In S. Africa paula is on the wing from August to April (Swane-

poel, 1953). It usually flies over grassy hill slopes. Food plants have not been

recorded. .

Distribution (Map 11). — A very restricted range, in Transvaal and Rhodesia.

Localities. — Rhodesia: Bulawayo, Essexvale, Filabusi, Gwanda, Penkridge,

Salisbury, Umtali (BM, MN, MNHN, NMB). South Africa. Transvaal: Munnik,

Pietersburg, Rustenburg, Swartruggens (NMB; Swanepoel, 1953).

Material examined. — 47,29 :4 (incl. holotype), 19 , Rhodesia (BM,

MN); 12 , South Africa (BM).

Spialia secessus (Trimen)

Pyrgus secessus Trimen, 1891. — Proc. zool. Soc. London 1891 : 102, pl. 9 fig. 22 (Omrora, Damara-

land). Holotype (7 ) in the South African Museum, Cape Town.

Map 11 St

Spialia paula

E me en

Hesperia secessa forma trimeni Aurivillius, 1925. — Hesperidae, in: A. Seitz, The Macrolepidoptera of

the World 13 : 564, pl. 75d (Rhodesia). The type of trimeni is probably lost; the only trimeni in the

Naturhistoriska Riksmuseet, Stockholm, is a male from Angola.

DE JONG: The genus Spialia 77

Figs. 70— 76. Female genitalia, ventral and lateroventral aspect, and papilla analis. 70—71, S. wrefordi

(Kotido, Uganda). 72—73, S. paula (Essexvale, Rhodesia). 74—75, S. ploetzi ploetzi (Namwamba

Valley, Uganda). 76, S. ploetzi occidentalis (Bingerville, Ivory Coast).

External characters (pl. 5 figs. 37, 38). — This species is unlike any other

Spialia species on the underside of the hind wing, where the median band is

straight and of even width throughout, with the outer margin finely denticulate (i.e.

slightly produced along the veins); this denticulation is accentuated by a darkening

of the ground colour along the outer margin of the band. The inner margin of the

band is less clearly defined. The effect is difficult to express in words, but the

photograph gives a good impression, and once having seen a real specimen, it is

improbable that one confuses this species with any other Spialia species. More-

over, there are two colour forms, one of which is particularly distinct, because the

band on the underside of the hind wing is brown instead of white; as the ground

colour is also brown, the band is mainly distinguishable by the darkening of the

ground colour along its outer margin.

The two colour forms have been named forma secessus Trimen (median band

on hind wing underside brown) and forma trimeni Aurivillius (median band on

78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

hind wing underside white or cream-coloured). Aurivillius (1925: 564) supposed

these forms to be the result of seasonal variation. This assumption is undoubtedly

Map 12

Spialia secessus

correct, as their ranges overlap completely and their flight periods are largely

separated. However, while Aurivillius considered f. trimeni the wet season form,

Evans (1937) mentioned f. secessus as such. I think Aurivillius was right. Almost all

specimens of f. secessus examined were collected in July and August, while the

bulk of f. trimeni examined date from October till February. In the distribution

area of the species the rains usually occur in the latter period, while July and

August are dry.

There are some exceptions, especially in f. secessus of which I have seen single

specimens from January, May, June, September and October, while f. trimeni may

appear as early as September and single specimens are known from April, May

and June. Supposing these specimens are correctly labelled, they may be the result

of microclimatic differences. The brown band of f. secessus occurs in various

shades and specimens intermediate between f. secessus and f. trimeni occur,

especially in April-May and September.

Male genitalia (figs. 77, 78). — Gnathos relatively narrow and double semi-

globular spinulose fold smaller than in other species of the group. Dorsal sclerotiza-

tion of anellus upturned laterally. Aedeagus rather thick in central part, laterally

DE JONG: The genus Spialia 19

slightly expanded in distal part, but apex narrow. Saccus well-developed, but shorter

Figs. 77—80. Male genitalia, lateral aspect, and inside of right valva. 77—78, S. secessus (Mpanda,

Tanzania). 79— 80, S. dromus (Tabora, Tanzania).

than uncus. Costa of valva distally expanded, mainly in ventral direction. Costal

process strongly upturned, spines pointing dorsad. Fold of cucullus not covering

base of costal process.

Female genitalia (figs. 81, 82). — Ventral sclerites of segment 8 widely sep-

arated near ostium. Sclerotization of postvaginal membranous region slight, but

distinct, proximally continued over a short distance in dorsal side of ductus, distally

passing into protruding, unindented area spinulosa. Papilla analis obtusely triangular;

apophysis posterior nearly 1.5 times as long as papilla.

Ecology. — A species of grassy slopes, flying in probably two, maybe more

generations, mainly from July to February, rarely from March to June. Food plants

not recorded.

Distribution (Map 12). — From Tanzania and S. Zaire to Transvaal and Natal.

Localities. — Tanzania: Butler South, Chala (Ufipa), Dunduma, Kampissa,

Kasoje, Kassaka River, Katari, Kotuma, Lubalisi, Mahale, Mbisi Forest, Mbosi,

80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Mpanda, Sibweza, Sitebi, Tungamu (Ngara), Wanzizi (BM, ML, MN, MRAC,

ZSM). Zambia: L. Bangweolo (Luwingu), High Plateau south of Lake Tanganyika,

Kafue Flats, Lofu River, Mporokoso, Ndola (BM, NRS, ZSM; Neave, 1910).

Rhodesia: Bulawayo, Essexvale, Gurbi River, Headlands (E. Mashonaland), Hope-

fountain (nr. Bulawayo), Inyanga Mts., Melsetter, Odzani River (Umtali), Salisbury

(BM, NMB). South Africa. Transvaal: Barberton, Graskop, Groenbult, Haenerts-

burg, Helpmekaar, Munnik, Nelspruit, Shilouvane, White River (BM; Murray,

1959; Swanepoel, 1953). Natal: Natal (BM). Angola: Bailundo, Bihe Distr.,

Capelongo-Dongo, Cauca, Chitau, Cubal River, Omrora (AMNH, BM, CMP,

MNHN, NRS; Trimen, 1891). Zaire: Bukama, Ditanto, Elisabethville, Kafakumba,

Kambove (150—200 mis W of Kambove), Kapanga River, Kapelekese, Karavia,

M’Pemba Zeo (Gandajika), Sando (BM, MRAC).

Material examined. — 102 , 42 9 : 519, 18 © , Tanzania (BM, ML,

MN, MRAC, ZSM); 5 ,39© , Zambia (BM, NRS, ZSM); 5 ©, Rhodesia (BM,

MN); 10 , Natal (BM); 3 ”, 7 9 , Transvaal (BM); 70° ,59 , Angola (AMNH,

BM, CMP, NRS); 30 7,99 , Zaïre (BM, MRAC).

Spialia dromus (Plötz)

Pyrgus dromus Plôtz, 1884. — Mitt. Naturwiss. Ver. Neu-Vorpomm. 45 : 6 (Congo). Type(s) lost.

Holland (1896) mistrusted the type-locality given by Plôtz, because he supposed dromus not to occur

“further north than Angola on the West Coast”, a very incorrect supposition.

Pyrgus zaira Plôtz, 1884. — Mitt. Naturwiss. Ver. Neu-Vorpomm. 45 : 6 (Congo). Type(s) lost.

Syrichthus melaleuca Oberthür, 1912. — Et. Lep. Comp. 6 : 113, fig. 1327 (Kitanga). Holotype (9)

in the British Museum (Nat. Hist.), London.

Syrichthus leucomelas Oberthiir, 1912. — Et. Lep. Comp. 6 : 118, fig. 1308 (Mpala, L. Tanganyika).

Holotype (3 ) in the British Museum (Nat. Hist.), London.

Hesperia dromus var. elongata Higgins, 1924. — Trans. ent. Soc. London 1924 : 95 (Makindu, Kenya).

Holotype (o ) in the British Museum (Nat. Hist.), London.

Hesperia dromus var. meridionalis Higgins, 1924. — Trans. ent. Soc. London 1924 : 95 (Zanzi-

bar). Holotype (7) in the British Museum (Nat. Hist.), London.

External characters (pl. 5 figs. 39, 40). — This species can be distinguished by

the following combination of characters:

a. Fore wing upperside. — Basal cell spot well-defined, rounded; central cell

spot midway between basal cell spot and discocellular spot; inner median spots 1b

and Ic present, forming a bar from vein 1 to vein 2, outer median spots 1b and Ic

absent or Ic visible as a small spot at the outer lower angle of median spot 2.

b. Hind wing underside. — Band of median spots continuous from vein 1b to

costa, sharply bent basad in space 2.

The characters of the upperside of the fore wing will usually suffice to distinguish

dromus from all other Spialia species.

There is much variation in the extension of the spots and some forms have

received names. In forma meridionalis Higgins, the spots are reduced on the upper-

side and the submarginal spots may be entirely wanting. A specimen without

median spots on the upperside of the fore wing except in space 1b, but with well-

developed submarginal spots, was described as a new species, melaleuca, by

Oberthiir (1912), but the genitalia of this specimen clearly showed it to belong to

dromus, of which it appears to be a unique aberration. Another aberrant male in

DE JONG: The genus Spialia 81

the BM, from Natal, has the spots on the upperside of the hind wing wanting

except for the basal cell and small submarginal spots, while most of the spots on

the upperside of the fore wing are abnormally large, reminding of forma taras of

Pyrgus malvae L.

A form with the band of median spots on the upperside of the hind wing distinct-

ly continued across space lc (as on the underside) was named “var. elongata” by

Higgins (1924). This character is also found in the forma leucomelas that was

described as a new species by Oberthiir (1912) and only differs from elongata in

the extremely broad band of median spots of the hind wing.

There is also much variation in the length of the fore wing, see under Geographic

variation. | ‘

Male genitalia (figs. 79, 80). — Dorsal sclerotization of anellus flat or medially

slightly depressed. Aedeagus straight, cylindrical. Saccus shorter than uncus.

Costal process of valva long, upturned, with long spines pointing ventrad and

cephalo-dorsad; fold of cucullus covers greater part of costal process, including its

base.

Female genitalia (figs. 83, 84). — Ventral sclerites of segment 8 well-defined,

separated from each other and from the dorsal sclerites by a membranous gap.

Postvaginal membranous region with triangular sclerotization that is especially

distinct in its narrow part near the ostium and that passes into the unindented area

spinulosa. Papilla analis rather triangular; apophysis posterior somewhat longer

than papilla.

Ecology. — S. dromus is more at home in wooded areas than in open savanna

and can particularly be encountered at forest fringes, along forest paths, etc.,

though not in dense forest. It has been found flying in all months of the year.

Recorded food plants are Triumfetta tomentosa Boj. and rhomboidea Jacq.

(Tiliaceae) (Murray, 1959; Gifford, 1965).

Distribution (Map 13). — One of the most widely distributed Spialia species in

Africa, occurring from Natal to Ethiopia and Senegal. Although it is apparently

less common than diomus in S. Africa, dromus appears to be the commoner

species in the rest of Africa. It reaches its greatest abundance in Central and East

Africa and it is outnumbered by spio in West Africa. The absence of records from

many regions (e.g. Liberia, Lindsey & Miller, 1965; Guinea and Ivory Coast,

Berger, 1962) is undoubtedly due to incomplete collecting.

Geographic variation. — There appears to be a slight clinal variation in the

length of the fore wing ( 7 9.1—13.8 mm) and the extension of the white spots.

In general, the larger and better marked specimens are found in the south, but there

is very much overlap in the length of the fore wing and the extension of the spots

in the various regions. Although specimens with large spots are most common in the

southern part of Africa (north to Tanzania), the form elongata is most common

in the western and northern parts of the distribution area. The large range of

overlap makes it impossible to delimit subspecies.

Localities. — Ethiopia: Cherosh Wonz, Fich (Shoa Prov.), Gimera to N. end of

L. Rudolf (BM; Carpenter, 1935). Sudan: Jebel Marra (W. Darfur), Talanya

(Vambio) (BM). Kenya: throughout (AMNH, BM, CMP, ML, MN, MNHN).

82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Figs. 81—84. Female genitalia, ventral aspect, and papilla analis. 81—82, S. secessus (Mpanda, Tan-

zania). 83—84, S. dromus (Natal).

Uganda: throughout (AMNH, BM, CMP, MN). Tanzania: throughout (AMNH,

BM, ML, MN, MNHN, MRAC, NMB, ZSM). Rwanda: Kitengi, Ruhengeri (ML,

MRAC). Burundi: Bukcba-Usumbura, Kitega, Muyaga (MRAC; Rebel, 1914).

Zambia: throughout (BM, NMB, USNM, ZSM; Neave, 1910). Rhodesia: through-

out (BM, ML, MRAC, NMB, USNM). Malawi: throughout (BM, CMP, NMB,

USNM;; Gifford, 1965). Mozambique: Amatonga Forest, Andrada, Delagoa Bay,

Dondo Forest, Gorongoza, Inhaminga, Lourengo Marques, Salone Forest, Serra

DE JONG: The genus Spialia 83

Rotanda (BM, ML, MNHN, NMB, ZSM). South Africa. Transvaal: Barberton,

Chuniespoort, Graskop, Konatipoort, Louis Trichardt, Lydenburg Distr., Malta

Forest, Mariepskop, Mica, Munnik, Nelspruit, Olifantsrivier, Pietersburg, Potgieters-

rust, Pretoria, Saltpan, Sibasa, Tubex, Warmbaths, Woodbush, Zoutpansberg

(NMB; Swanepoel, 1953). Natal: Amahlongwa, Durban, Eshowe, Greytown,

Hluhluwe, Howick, Isipingo, Ismont, Karkloof, Maritzburg, Umbloti, Umkomaas

(AMNH, BM, CMP, ML, MNHN, MRAC, NMB, ZSM; Swanepoel, 1953).

Angola: Ehanda (BM; Trimen, 1891). Zaire: throughout (AMNH, BM, CMP,

MNHN, MRAC; Holland, 1920). Congo-Brazzaville: Etoumbi (Berger, 1967).

Gabon: Gabon River (Trimen, 1889). Tchad: Fort Archambault (MNHN).

Cameroun: Bamenda (BM). Nigeria: Aba, Kaduna, Obuda Ranch (E. Nigeria)

(BM, MN, ZSM). Ghana: Likpe (BM, MRAC). Sierra Leone: Sierra Leone (BM).

Guinea: “‘Guinee française” (MNHN). Senegal: Dakar, Oussony, Sandiaba, Tabi

près Bignona (BM, CMP, MNHN; Berger, 1968).

Material examined. — 618 © , 242 9 : 19 , 1 © , Ethiopia (BM); 2 ©,

Sudan (BM); 102 © , 37 © , Kenya (AMNH, BM, CMP, ML, MN); 39 & ,

13 © , Uganda (AMNH, BM, CMP, MN); 155S , 529 , Tanzania (AMNH,

BM, ML, MN, MRAC, ZSM); 2 ”, 1 ® , Rwanda (ML, MRAC); 26 7,149,

ennn

Map 13

Spialia dromus

84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Burundi (MRAC); 2 7, Zambia (USNM, ZSM); 12 © , 4 © , Rhodesia (BM, MN,

USNM); 37 9, 169 , Malawi (BM, CMP, USNM); 24 © , 9 © , Mozambique

(BM, ML, ZSM); 55 ” , 449 , South Africa (AMNH, BM, CMP, ML, MRAC,

ZSM); 18 4, 79 , Angola (BM); 1120, 319 , Zaire (AMNH, BM, CMP,

MRAC); 19 , Congo-Brazzaville (MRAC); 14 , 59 , Nigeria (BM, MN,

ZSM); 37, 5 @ , Ghana (BM, MRAC); 95 , 39 , Sierra Leone (BM); 4 7 ,

Senegal (BM, CMP).

Map 14 ur

Spialia ploetzi ploetzi

en tt

Spialia ploetzi (Aurivillius)

Hesperia Ploetzi Aurivillius, 1891. — Ent. Tidskr. 12 : 227 (Cameroun). For synonymy, see subspecies.

The nomenclatural and taxonomic confusions about this species have been dealt

with in a previous paper (De Jong, 1977). The species shows a marked geographic

variation. The subspecies recognized have generally been considered separate spe-

cies and Higgins (1924) created a species group for them (“Hesperia plotzi

group”).

External characters. — On the underside of the hind wing the median spots in

spaces 3 to 7 and the submarginal spots in spaces lc and 2 conjoin to form an

DE JONG: The genus Spialia 85

oblique white bar from about the middle of vein 8 to the termen in space Ic or 2.

By this character S. ploetzi is easily distinguished from all other Spialia spe-

cies. On the upperside of the hind wing the marked median band which is sharply

broken along vein 3 by the direction of the median spot 4—5 and the basad shift

of median spot 2, is distinctive. Length of fore wing 11.4 — 13 mm.

Male genitalia. — Spinulose area of gnathos extending dorsad to slightly

beyond the middle of the gnathos. Dorsal sclerotization of anellus folded over

aedeagus like a roof, but laterally bent upwards. Aedeagus long, S-shaped, sur-

face smooth. Saccus about as long as uncus. Costa of valva extended distad. Cos-

tal process large, curving upwards, spines pointing cephalo-dorsad. Fold of cucullus

covering about basal third of costal process.

Female genitalia. — Ventral sclerites of segment 8 touching near ostium, but

not conjoined. Postvaginal membranous area wide, with slight triangular scleroti-

zation that imperceptibly passes into area spinulosa. Latter with straight or slight-

ly wavy distal edge. Papilla analis with obtuse apex; apophysis posterior less than

1.5 times as long as papilla.

Ecology. — S. ploetzi is the only Spialia species confined to the forest. Spe-

cimens are known from all months of the year. Food plants unknown.

Distribution. — Confined to the forest belt from Sierra Leone to Western

Kenya. The southern limit appears to run through N. Angola, S. Zaire and along

the north side of Lake Tanganyika. East of this lake, ploetzi does not appear

to occur south of Burundi. The northern limit presumably runs through the Cen-

tral African Republic.

Geographic variation. — There is a marked variation in the extension of the

spots, the male and female genitalia. Two subspecies can be recognized.

Spialia ploetzi ploetzi (Aurivillius)

Hesperia Ploetzi Aurivillius, 1891. — Ent. Tidskr. 12 : 227. Type locality: Cameroun, environs of

Douala, see De Jong (1977). Lectotype (7) in the Naturhistoriska Riksmuseet, Stockholm.

Hesperia zebra Rebel, 1914. — Ann. k.k. naturh. Hofmus. Wien 28 : 271—272, pl. 22 figs. 52, 53

(NW of Lake Tanganyika). Holotype (7) in the Naturhistorisches Museum, Vienna. Junior primary

homonym of Hesperia zebra Butler, 1888 (which is now also placed in Spialia).

Hesperia rebeli Higgins, 1924. — Trans. ent. Soc. London 1924 : 101. Replacement name for Hes-

peria zebra Rebel, 1914. Evans (1937) stated that the type of rebeli is in the British Museum (Nat.

Hist.), and he labelled, indeed, a specimen from Uganda as type (nr. H. 892). This was, however, an

incorrect action, for the type of rebeli Higgins should be the same as that of zebra Rebel, as Hig-

gins only proposed a replacement name.

External characters (pl. 6 figs. 41, 42). — On the whole, this eastern subspecies

is more extensively spotted than the western one. In detail, the differences are as

follows: i

1. on the upperside the spots are larger, especially the central cell spot on the

fore wing and the submarginal spots on both wings;

2. on the fore wing the series of submarginal spots is less sinuous;

3. on the upperside of the hind wing the submarginal spots 5 and 6 are strong-

ly developed, forming two white streaks (in the western form they are scarcely

visible);

86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

4. on the underside of the hind wing the submarginal spots 5 and 6 are fused

into a white streak from median spot 7 to the end of vein 5 (this streak is at

most faintly indicated in the western form);

5. in addition to this streak the veins are usually overlaid with light scales, so

that the underside of the hind wing has a striped appearance;

6. median spot 2 on the underside of the hind wing is usually indistinct and

sometimes absent (in the western form usually very distinct. and often more conspi-

cuous than the lower part of the median band).

According to Higgins (1924) the ground colour of the upperside is paler in the

eastern form. I cannot confirm this observation, but it may be true for fresh

material.

Eastern specimens of ssp. ploetzi generally have larger spots on the upperside,

especially on the hind wing, and often a lighter ground colour on the underside

of the hind wing, due to a denser light superscaling. There is, however, much over-

lap in these characters and it does not appear to be sensible to recognize further

subspecies.

Male genitalia (figs. 66, 67). — Costa of valva not extending beyond tip of

cucullus.

Female genitalia (figs. 74, 75). — Narrow, medio-proximal parts of ventral

sclerites (i.e. the parts nearest the ostium) at least two times as long as broad.

Distribution (Map 14). — The eastern part of the range of the species, from W.

Cameroun eastward. The westernmost localities known are Kumbo and Bamenda

in W. Cameroun, close to the Nigerian frontier and formerly Nigerian territory.

The easternmost localities are situated in the forest remains in the west of Kenya.

In the Nairobi Museum there is a specimen labelled “W. Atomo, Mombasa”; I

could not find W. Atomo on any map or in any gazetteer, the only name ap-

proaching it is Watamu, about 100 km north of Mombasa. It would be highly

interesting if ploetzi really occurred there.

Localities. — Kenya: Kaimosi, Kakamega Forest, Malaba (Kabras), ? Mom-

basa (W. Atamo) (MN, USNM). Uganda: Bwamba Forest, Budongo Forest, Bu-

funbo Forest, Entebbe, Hoima, Impenetrable Forest, Jinja, Kalinzu Forest, Kamen-

go, Kanaba Gap, Katera, Kayonza Forest (Kigezi), Kazi, Mafuga Forest (Kigezi),

Mbarara, Mt. Kokanjero, Muhende, Mulange, Namwamba Valley, Rutenga Fo-

rest (Kigezi), Toro (AMNH, BM, CMP, MN, MRAC, NMB). Rwanda: Cyangugu,

Kisaba Forest (BM, ML). Burundi: Bugarama, Bulumbura, Bururi, Kitega, Ruvuvu

River (BM, MRAC). Zaire: Bambesa, Bena-Tshiadi, Bokala a Busanga, Buta,

Dungu, Eala, Epulu Forrst, Gamangui, Goma-Rutshura, Itimbiri-Dingila, Itoko

a Sombe, Kabongo, W. of Kambove, Kaniama, Kapanga, R. Kapelekese, Katako-

Kahudi, Katako-Kombe, Lake Kivu, NW. of Lake Tanganyika, Lodja, Lowa-

Nduba, Lufupa River, Lukolela, Luluabourg, Lusambo-Batempa, Mbudi, Medje,

Moto, Nioka, Nyamunyunya (Mulungu), Omotumba, Paulis, Ruwenzori, Sandoa,

Stanleyville, Tshibinda, Tshikunia, Tshiole, Uele, Utshudi, Yakoma (AMNH, BM,

CMP, MN, MRAC, NMB, USNM; Heron, 1909; Higgins, 1924; Holland, 1920;

Neave, 1910; Rebel, 1914). Angola: N’Dalla Tando (BM). Congo-Brazzaville:

Etoumbi (MRAC; Berger, 1967). Gabon: Lastousville, Oyem (MNHN, MRAC).

DE JONG: The genus Spialia 87

Rio Muni: Nkolentangan (ZMHB). Central African Rep.: Boukolo M’Baiki

(Oubangui-Chari) (MRAC). Cameroun: Akom, Bamenda, Batouri Distr., Bitye (Ja

River), Buea, Bule Country, Chang, Ebogo s. le Nyong, Efulen, Ekutu, Elat, Kribi,

Lolodorf, Lomié, Metet, Penderu, Sakbayeme (Sanaga River), Zoatoupsi (20 km

W. of Yaoundé) (AMNH, BM, CL, CMP, MNHN MRAC, NMB, NRS, USNM,

ZWM; Condamin, 1960).

Material examined. — 262 7,69 9 :407,39 , Kenya (MN, USNM); 85

9,27, Uganda (AMNH, BM, CMP, MN, MRAC); 2 ©, Rwanda (BM, ML);

69,19 , Burundi (BM, MRAC); 67 7 , 21 2 , Zaïre (AMNH, BM, CMP, MN,

MRAC, USNM); 5 7, Angola (BM); 10, Congo-Brazzaville (MRAC); 20’,

Gabon (MNHN, MRAC); 15, Rio Muni (ZMHB); 10 , Centr. Afr. Rep.

(MRAC); 88 & , 37 © , Cameroun (AMNH, BM, CMP, MNHN, MRAC, NRS,

USNM, ZSM).

Spialia ploetzi occidentalis De Jong

Spialia ploetzi occidentalis De Jong, 1977. — Ent. Ber., Amst. 37 : 43 (Ghana). Holotype (o) in the

Rijksmuseum van Natuurlijke Historie, Leiden; paratypes in the British Museum (Nat. Hist.), London,

and the Naturhistoriska Riksmuseet, Stockholm. This subspecies is generally known in the literature

as the species Spialia ploetzi.

orem = |

Map 15 2:

Spialia ploetzi occidentalis

88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

External characters (pl. 6 figs. 43, 44). — Less extensively spotted than ssp.

ploetzi; for details, see under that subspecies.

Male genitalia (figs. 68, 69). — Costa of valva projecting well beyond tip of

cucullus.

Female genitalia (fig. 76). — Medio-proximal part of ventral sclerites about

as long as broad.

Distribution (Map 15). — The western part of the range of the species, from

Sierra Leone to E. Nigeria. Specimens of this subspecies recorded from localities

further east are probably incorrectly labelled, see De Jong (1977).

Localities. — Cameroun (?): “Cameroons” (MNHN). Nigeria: Afikpo, Anam-

bara Creek, Cape Coast Castle, Ebele, Ibadan, Iporia-Ilara Div., Kabba Prov.,

Lagos, Lokkoh, Oban, Ogruga, Old Calabar, Oshodi, Warri (BM, CMP, MN,

MNHN; Higgins, 1924). Dahomey: Porto Novo, Dahomey Interior (MNHN).

Togo: Missahohe (= Misa-hôhe) (ZSM). Ghana: Asuchari, Kumasi, Likpe,

Odumase Swamp, Sunyani (BM, ML, MRAC). Ivory Coast: Abengourou, Abidjan,

Adiopodoume, Bingerville, Issia, Seleu (BM, MN, MNHN, MRAC; Berger, 1962).

Liberia: Bigtown, Ganta, Harbel, Kpain, Maloubli, Monrovia, Penoke, Zorzor

(BM, CMP; Lindsey & Miller, 1965; Picard, 1950). Sierra Leone: Freetown, Ka

Yima, Moyamba, ‘“Sierra-Leone” (BM, MNHN, NRS, ZSM; Holland, 1896).

Guinea: Forecariah, Tondon (C. Dubreka) (BM, MNHN; Berger, 1962).

Material examined. — 108 © , 38 © : 1 cc, ? Cameroun (MNHN); 13 ©,

99 , Nigeria (BM, CMP, MN); 10 Togo (ZSM); 21 © (incl. holotype of occi-

dentalis) 5 9 , Ghana (BM, ML, MRAC); 290,99 , Ivory Coast (BM, MN,

MRAC); 109,39 , Liberia (BM, CMP); 315 , 129 , Sierra Leone (BM,

NRS, ZSM); 2 ” , Guinea (BM).

g. The delagoae group

External characters. — Fore wing without basal cell spot on upperside, at

most some white scales forming a small dash along the cubital vein; central cell

spot not remarkably close to end of cell; no spots in spaces 9 and 10; submar-

ginal spots present or absent. Median band on underside of hind wing straight or

winding, continuous from space Ic to space 8, more or less parallel to termen;

submarginal spots 4—5 (if present) in line with the other submarginal spots. Male

without costal fold.

Male genitalia. — Uncus undivided or bipartite apically; junction with tegumen

indicated by a narrow, less strongly sclerotized band, medially broadening to a

fenestrula. Gnathos dorsally joined to tegumen over a shorter or longer distance

or free from tegumen, the dorsal edge often folding outwardly; broadly sclero-

tized, ventrally usually with stronger sclerotization and some short spines; left

and right part of gnathos ventrally joined by a membrane that is usually

smooth, but may be slightly spinulose and folding upward medially. Anellus

with very strong dorsal sclerotization and spines. Juxta relatively large. Aedeagus

straight or slightly curved, with ventral, cylindrical, spined excrescence in distal

half; coecum short or absent. Saccus relatively short, i.e. shorter than uncus.

DE JONG: The genus Spialia 89

Map 16 vil. 1064,

Spialia mangana

Costa of valva narrow; costal process various, usually well-developed. Cucullus

excavated to receive costal process or forming a small fold, that does not cover

the costal process.

Female genitalia. — Segment 8 with two small dorsal sclerites, which are invi-

sible in ventral sight; ventral surface occupied by a single large sclerite covering

the whole surface or with small membranous areas distally from the ostium to

the left and to the right. In all but one species there is no noticeable sclerotization

antevaginally. Just behind the ostium the sclerite may protrude in two small

keels. Small spines of area spinulosa in some species visible at the distal edge of

the ventral sclerite. Ductus long, gradually broadening into bursa; ductus and

bursa without sclerotizations or spines. Papilla analis without basal lobe, short,

broader than long; apophysis posterior short, about as long as or shorter than

width of papilla.

Spialia mangana (Rebel)

Hesperia mangana Rebel, 1899. — Sber. Akad. Wiss. Wien 1899 : 360 (Ras Fartak, S. Yemen).

Holotype ( © ) in Naturhistorisches Museum Wien (Vienna, Austria).

90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Figs. 85—89. Male genitalia of S. mangana (Bihendula, Somalia). 85, lateral aspect. 86, inside of

right valva. 87, dorsal aspect of uncus and tegumen. 88, lateral aspect of aedeagus. 89, ventral aspect _

of aedeagus. Figs. 90—93. Male genitalia, lateral aspect, and inside of right valva. 90—91, S.

nanus (Sheldon, S. Africa). 92—93, S. delagoae (Muden, S. Africa).

External characters (pl. 6 figs. 45, 46). — An unmistakable species by the ©

complete absence of basal and submarginal spots on the upper and underside |

of both wings. On the upperside the ground colour is evenly dark brown, on the |

underside of the hind wing the ground colour of the basal part is usually of a

DE JONG: The genus Spialia 91

lighter shade than between the median band and the termen. A good figure was

given by Rebel (1907).

There is much variation in size, © 7.5 — 9.9 mm, 9 8.1 — 11 mm, but the

material is too limited to show whether this variation is individual, seasonal or

geographic.

Male genitalia (figs. 85—89). — At junction of uncus and tegumen a long,

triangular fenestrula. Gnathos connected with tegumen by a membrane; ventrally

a double, semiglobular, spinulose fold (cf. dromus group). Sclerotization in dor-

sal part of anellus forming a spined prong to the left and the right, ventrally

conjoined to the juxta. Aedeagus short and broad, with a strong, spined excres-

cence medio-ventrally beyond the middle. Just beyond the ventral excrescence the

aedeagus is constricted laterally and beyond this constriction it widens to a flat,

almost circular apex; coecum absent. Costal process small, with a few short

spines radiating distad. Cucullus excavated to form a narrow fold that scarcely

covers a part of the costal process.

Female genitalia (figs. 94, 95). — Ventral sclerites of segment 8 conjoined

antevaginally by a thick, bumpy sclerotization. Slight sclerotization in postvaginal

area, the membranous area separating the ventral sclerites. Area spinulosa slightly

indented at distal edge. Papilla analis dorso-distally rather pointed.

Ecology. — Nothing is known of the ecology of this possibly rare skipper.

The studied specimens date from all months, except February and March.

Distribution (Map 16). — Confined to the southern part of the Arab peninsula,

Ethiopia, Somalia, N. Kenya and N. Uganda.

Localities. — Yemen: Ta’izz (BM; Gabriel, 1954). South Yemen: Ahdaherma,

Dhala, Jebel Jihaf, wadi east of Jebel Ma’fari, Moola Mutar, Ras Fartak (BM;

Gabriel, 1954; Rebel, 1899, 1907). Ethiopia: Dire Daoua, Fich Shoa Prov., Meiso

(BM, MNHN; Carpenter, 1935). Somalia: Bihendula, Galkayu, Gan Linah, Har-

geisa, Mandera, Mudugh (BM, MN). Kenya: Kacheleba, Kenailmat (Suk)

(BM, MN). Uganda: Cholol (Karamoja) (MN).

Material examined. — 19 7,82 : 1 , Yemen (BM); 35 , 3 © , South

Yemen (BM); 4 7, 2 9 , Ethiopia (BM); 65 , 2 9 , Somalia (BM, MN); 39,

19, Kenya (MN); 2 © , Uganda (MN).

Spialia nanus (Trimen)

Pyrgus nanus Trimen, 1889. — South African Butterflies 3 : 290 (Malmesbury, Cape). Holotype

(+) in the British Museum (Nat. Hist.), London.

External characters (pl. 6 figs. 47, 48). — By the combination of the absence

of a basal cell spot on the upperside of the fore wing and a continuous band

of median spots on the underside of the hind wing, this species is easily recognized

as a member of the delagoae group. The basal cell spot on the underside of

the hind wing is absent or very faintly indicated, the median band is very winding

and runs from the end of space 8 through a double curve to about the middle of

the distal part of space Ic.

There is much variation in size (0° 8.5 — 11 mm) and spotting. The submar-

ginal spots on the upperside may be almost absent; the spots 2 and 4—5 and

92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

regularly also lc and 3 of the median spots are present on the upperside of the

hind wing. The ground colour of the underside of the hind wing varies from warm

yellow to brown. Although there are two generations per year, Swanepoel (1953)

does not mention seasonal variation.

Male genitalia (figs. 90, 91). — A triangular fenestrula at junction of uncus

and tegumen. Uncus with a longitudinal keel. Gnathos not fused to tegumen, con-

sisting of two sclerotized plates, which curve outside in the upper part, so that in

dorsal view the gnathos protrudes laterally. In the ventral part of the gnathos the

sclerotization is fainter and here is a finely spinulose area. Anellus with strong,

dorsally spined sclerotization, partly surrounding the aedeagus and membranously

connected with the juxta. Aedeagus straight, with short coecum and a relatively

long, ventral, spined projection; large cornuti present. Costa of valva narrow;

costal process much larger than costa, directed to base of valva, slightly bent

upwards, strongly spined. Cucullus excavated, forming a very narrow fold that does

not cover the costal process.

Figs. 94—99. Female genitalia, ventral aspect, and papilla analis. 94— 95, S. mangana (Dhala, S.

Yemen). 96—97, S. nanus (Paarde Berg, S. Africa). 98—99, S. delagoae (S. Africa).

DE Jong: The genus Spialia 93

Female genitalia (figs. 96, 97). — Ventral sclerites of segment 8 not con-

joined antevaginally, folded alongside the ostium to a few ridges. Postvaginally

there is a short and narrow membranous area to the left and to the right of the

ostium, enclosed by the ventral sclerites, which are fused and form small folds

overlapping the end of the membranous areas. The ventral sclerites include the

triangular postvaginal sclerotization. Area spinulosa deeply indented at distal

edge. Papilla analis with upper edge rather straight.

Ecology. — This species is at home in the Karroo bushveld of the Cape Province,

South Africa, where it can be found almost anywhere, on the flats and along

mountain slopes. There are two generations per year, the main flight periods

being September—October and March—April (Swanepoel, 1953). As food plants

the following species have been recorded (Murray, 1959): Hermannia species

(Sterculiaceae), Pavonia macrophylla and Hibiscus aethiopicus (Malvaceae).

Distribution (Map 17). — Confined to South Africa, where it has been found

in the Cape Province and the western Orange Free State.

Localities. — South Africa. Cape Province: De Aar, Addo, Beaufort West,

Bitterfontein, Calitzdorp, Cape Town, Carlton, Carnarvon, Clanwilliam, Coles-

berg, Cookhouse, Cradock, Deelfontein, Dunbrody, Graaf-Reinet, Kamieskroon,

Map 17 ee

Spialia nanus

Katzenberg Hill, Kimberley, Klaver, Malmesbury, Mamre, Matjesfontein, O’okiep,

Paarde Berg, Paarl, Prieska, Rosemead, Sheldon, Slang Hoek Mts., Somerset

East, Springbok, Steynsburg, Uitenhage, Vanrhijnsdorp, Worcester (BM, CMP,

ML, MN, MNHN, NMB; Swanepoel, 1953). Orange Free State: Bloemfontein

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Map 18

Spialia delagoae

le ca CRAN Je

(AMNH, BM, NMB; Swanepoel, 1953).

Pyrgus delagoae Trimen, 1898. — Trans. ent. Soc. London 1898 : 15 (Delagoa Bay). Holotype (4)

in the South African Museum, Cape Town.

External characters (pl. 7 figs. 49, 50). — This species closely resembles na-

nus by the absence of the basal cell spot on the underside of the hind wing, but

the straight median band is a reliable differentiating character. Of the other spe-

cies of the group, only mangana has no basal cell spot on the underside of the

hind wing, but that species is easily distinguished by the absence of submarginal

spots on upper and underside. The ground colour of the underside of the hind wing

iMasually dark olive-green. There is some variation in size (0° 9.9 — 11.6 mm)

and extension of spots. On the upperside of the hind wing the median spots 2 and

Material examined. — 27 ” , 10 © , South Africa (AMNH, BM, CMP, ML).

Spialia delagoae (Trimen)

DE JONG: The genus Spialia 95

4—5 are always present, often also 3, sometimes lc indicated by white scales,

6 always absent.

Male genitalia (figs. 92, 93). — Triangular fenestrula at junction of uncus and

tegumen. Uncus with a longitudinal keel, in dorsal view apex slightly expanded.

Gnathos dorsally strongly curved outward and projecting laterally; ventral

sclerotization stronger than in nanus, forming a strongly spined ventral projec-

tion. Fultura superior smooth. Dorsal sclerotization of anellus consisting of a

strongly spined, pointed structure to the left and to the right, the two parts con-

nected by a strongly incurving, slightly sclerotized fold that is connected dor-

sally to the gnathos and the fultura superior. Aedeagus straight; coecum longer

than thick; ventral projection large, 4—5 times as long as thick; apex flat, but not

expanded; a series of subequal cornuti present. Costa of valva very narrow, dis-

tally slightly expanded. Costal process very large, almost straight, at apex slightly

upturned, pointing towards base of valva, strongly spined. Cucullus hollowed to

receive the costal process, not forming a fold.

Female genitalia (figs. 98, 99). — Very much like nanus, but sclerotized

ridges near ostium smaller and no flaps of ventral sclerites overlying the end of the

membranous strips; area spinulosa possibly somewhat wider. Apophysis posterior

relatively long, longer than width of papilla.

Ecology. — In South Africa it is a species of the bushveld, that may be found

sipping from wet mud or fresh baboon dung (Swanepoel, 1953). There are no

records on its habitat in more northern areas. S. delagoae is on the wing

throughout the year, but in South Africa it is most abundant in February-March

and August-September. Food plants have not been recorded.

Distribution (Map 18). — From N. Kenya and N. Uganda to Transvaal and

Natal. At the moment the northern populations appear much isolated.

Localities. — Kenya: Kacheleba, Kedong (MN). Uganda: Amudat (MN).

Zambia: Solwezi (Higgins, 1924). Rhodesia: Bazely Bridge (SW. Umtali), Bula-

wayo, Chirinda Forest (Chipinga), Dotts Drift (Lower Sabi Valley), Essexvale,

Guzi Forest, Gwanda, Inyanga Mts, Matetsi, Mtoko, Nyamadzi R., Nyamandhlovu,

Odzi R., Salisbury, Sanyati Valley, Umtali, Vumba Mts, West Nicholson (BM,

MN, NMB, USNM). Malawi: Mlanje (Gifford, 1965; Higgins, 1924). Mozambique:

Delagoa Bay (BM, MNHN; Trimen, 1898). Botswana: Hothlotane, Macloutsi,

55 mls E. of Maun, Ngamiland, 55 mls W. of Tsane (BM, MNHN, NMB). Swa-

ziland: Gollel (NMB). South Africa. Transvaal: Bandolierkop, Barberton, Chu-

niespoort, Dendron, Haenertsburg, Munnik, Ohrigstad, Palabora, Pietersburg,

Rita, Shilouvane, Vivo, Warmbaths, Zoutpansberg (BM, CMP; Swanepoel, 1953).

Natal: Estcourt, Hluhluwe, Kearsney, Letsibele, Mhlosinga, Middeldrift, Muden,

Tugela R., Weenen (BM, CMP; Swanepoel, 1953). Cape Province: Lobabi (MN).

Namibia: Groot-fontein, Okawango, Otjitambi, Ovikokonva (SMW). Angola:

Dondo (Quanza River) (BM).

Material examined. — 49 7%, 19 © :397,19 , Kenya (MN); 1 , Uganda

(MN); 14 7, 49, Rhodesia (BM, MN, USNM); 2 © , Mozambique (BM,

MNHN); 1 , Botswana (MNHN); 24 & , 10 © , South Africa (BM, CMP,

MN); 3 7,4 9, Namibia (BM, SMW); 1 © , Angola (BM).

96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Spialia zebra (Butler)

Pyrgus zebra Butler, 1888. — Ann. Mag. Nat. Hist. (6) 1 : 207—208 (Koteer, Chittar Pahar). For

synonymy, see subspecies.

External characters (pl. 7 figs. 51, 52). — The presence of a basal cell spot and

submarginal spots on the underside of the hind wing distinguishes this species from

all other species of the group except depauperata. According to Higgins (1924)

zebra (under the name bifida) and depauperata are almost indistinguishable, but

in fact the identification is not very difficult if we pay attention to the following

differences:

1. on the average, zebra is smaller, 7 9.7—11.5 mm, as against depauperata

O 10.4—13.8 mm;

2. in zebra the median spots 2 and 4—5 on the upperside of the hind wing

are usually the only median spots present, although 3 may be more or less indicated;

in depauperata the median spots on the upperside of the hind wing form a more or

less straight band from vein 2 to vein 7, crossed by the dark veins 3, 4 and 6. I

never met with serious difficulties in distinguishing between zebra and depauperata,

but if one is not sure about the identification of a male, a clearing of the tip of the

abdomen will soon reveal the difference in the uncus.

There is some variation in the extension of the spots. The submarginal spots on

the upperside are sometimes very reduced, but always visible, at least in part. The

ground colour of the underside of the hind wing is of a greyish green tinge.

Male genitalia (figs. 100—103). — Uncus bifid at apex, more or less hammer-

like in dorsal (and ventral) view, left and right parts with slightly inflated apices. At

proximal end of distinct, longitudinal keel of uncus, a small and more or less oblong

fenestrula. Gnathos separated by very narrow membranous strip: ventrally slightly

more strongly sclerotized and with some small spines, passing into an inwards

sclerotized fold that forms a shallow sac and is flanked laterally by a large number

of very coarse spines; more ventrally this sclerotization forms two small pointed

processes, lying over the dorsal surface of the aedeagus, as in depauperata, but

much smaller. Aedeagus gently curved upwards; coecum slightly longer than wide;

ventral process small, inconspicuous, except for the short spines. Distal end of

costa of valva covering part of costal process. Latter not very conspicuous, with

radiating spines, in proximal part with some very coarse spines, but not such a

distinct, separate proximal part as in depauperata.

Female genitalia (figs. 104— 106). — Ventral sclerites of segment 8 fused into a

single sclerite, postvaginally with a low elevation that is pear-shaped in ventral

view. In the proximal part, to the right and to the left of the ostium, the sclerite is

excavated; laterally it has a few narrow, protruding ridges. No membranous post-

vaginal area. Area spinulosa slightly indented at distal edge. Ductus plus bursa

relatively short. Apophysis posterior not longer than width of papilla, relatively

straight.

Ecology. — In Kenya I found this species rather numerous along dry paths in

more or less open landscape, with scarce vegetation on coral rocks near Ukunda,

south of Mombasa. The specimens were flying especially during the hottest time of

the day and were not easy to collect, as they often flew very low, sometimes not

DE Jonc: The genus Spialia 97

Figs. 100—103. Male genitalia of S. zebra. 100—102, lateral aspect, inside of right valva, and dorso-

lateral aspect of uncus, of ssp. bifida (Ukunda, Kenya). 103, dorsolateral aspect of uncus of ssp.

zebra (Pakistan). Figs. 104—106. Female genitalia, ventral and lateral aspect, and papilla analis, of

S. zebra bifida (Ukambani, Kenya).

more than a few centimeters above the ground. In Kenya, specimens have been

found during all months of the year. In India, zebra appears to fly mainly in April,

May and June, but the material is scarce.

98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Sevastopulo (1974) records Melhamia (Sterculiaceae) as food plant.

Distribution (Map 19). — Highly discontinuously distributed, in East Africa/

South Yemen and Northwest Himalayas.

Geographic variation. — The African and Asian populations differ slightly in

the male genitalia. Externally both forms are similar.

Spialia zebra zebra (Butler)

Pyrgus zebra Butler, 1888. — Ann. Mag. Nat. Hist. (6) 1 : 207—208 (Koteer, Chittar Pahar).

Holotype (<7) in the British Museum (Nat. Hist.), London.

Hesperia hellas De Niceville, 1889. — Journ. Bombay Nat. Hist. Soc. 4 : 177, pl. B (Campbellpore).

Holotype (5) may be in the collection of the Zoological Survey of India, Calcutta, but I could not

get exact information.

Male genitalia (fig. 103). — Uncus apically slightly indented, the two parts of

the apex projecting laterally, so that in dorsal view the uncus is more or less T-

shaped.

Distribution. — Northwest Himalayas.

Localities. — Pakistan: Campbellpore; Koteer, Chittar Pahar; Punjab (Butler,

1888; De Niceville, 1889; Swinhoe, 1913; BM).

Material examined. — 6 + (incl. holotype), 1 © , Pakistan (BM).

Spialia zebra bifida (Higgins)

Hesperia bifida Higgins, 1924. — Trans. ent. Soc. London 1924 : 111—113 (Nairobi). Holotype (4)

in the British Museum (Nat. Hist.), London.

Male genitalia (fig. 102). — Uncus deeply indented apically, Y-shaped in dorsal

view.

Distribution. — East Africa and South Yemen.

Localities. — South Yemen: Wadi Tiban (NW of Jebel Jihaf) (Gabriel, 1954;

BM). Somalia: 20 miles S of Berbera, Bihendula (BM). Ethiopia: Bukuru Bridge

(BM). Sudan: Tembura (BM). Kenya: Athi Bridge, Besil (= Bissil), Chepalungu,

Diani, Elmenteita, Escarpment, Fourteen Falls, Gilgil, Hoey’s Bridge, Kacheleba,

Kakamega, Kangondi, Kibwezi, Kima, Kitale, Kitosh, Kitui, Landjoro, Lokichogio,

Lucania, Machakos, Makindu, Makueni, Malaba, Mombasa, Mount Elgon,

Mulango, Nairobi, Nairobi-Garissa Road, Nandi Plateau, Nanyuki, Ngong, Nyali,

Ologorsailie, Shimoni, Sotik, Soy, Suna, Taveta, Trans Nzoia, Twiga, Ukunda,

Upper Kuja Valley, Voi, Wangi, Watamu, Yala River (AMNH, BM, CMP, ML,

MN, MNB, MNHN, NRS, USNM); Shimba Hills (Sevastopulo, 1974). Uganda:

Kotido (MN). Tanzania: Dar-es-Salaam, Geita, Mount Meru, Oldeani, River Himo

(BM, ML, MN, MNHN).

Neave (1910) recorded zebra from Zambia (Fort Jameson, Luangwa Valley),

but this observation concerns depauperata, according to three specimens of Neave

in the collection of the Hope Department of Entomology, Oxford.

Material examined. — 126% , 56 © : 19 , South Yemen (BM); 2 ©,

Somalia (BM); 19 , 19 , Ethiopia (BM); 15 , Sudan (BM); 116 © (incl. holo-

type), 53 © , Kenya (AMNH, BM, CMP, ML, MN, MNHN, NRS, USNM);

19 , Uganda (MN); 57 , 19 , Tanzania (BM, ML, MN, MNHN).

DE JONG: The genus Spialia 99

Note. During a recent examination of five males from Mount Elgon (Kenya) two

males proved to have the uncus only slightly indented, thus being similar to ssp.

zebra. More material from the northern part of East Africa as well as from Paki-

stan is needed to establish the constancy of the difference between the subspecies

recognized.

Spialia sataspes (Trimen)

Pyrgus sataspes Trimen, 1864. — Trans. ent. Soc. London 1864 : 178—179 (Plettenberg, Cape Prov.).

Holotype (o ) in the British Museum (Nat. Hist.), London.

Trimen (1866) described the same species under the same name again as new.

External characters (pl. 7 figs. 53, 54). — The main differentiating character of

this species is the absence of submarginal spots on the underside of the hind wing.

This is also the case in mangana, but that species lacks the basal cell spot on the

underside of the hind wing. The lastmentioned character also distinguishes sataspes

from nanus and delagoae. Difficulties may arise with specimens of zebra and

depauperata, in which the submarginal spots on the underside of the hind wing are

faint, but in such cases the less straight median band on the underside of the hind

Map 19 Spialia zebra

100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Map 20 a

Spialia sataspes

wing in sataspes is helpful. In doubtful cases an examination of the genitalia is

necessary.

There is little variation in size (0° 9.1—11.5 mm) and in extension of the spots.

Sometimes the submarginal spots on the upperside are absent. On the upperside of

the hind wing the median spots 2 and 4—5 are always present and lc and 3 are

often indicated, while 6 is very rarely visible as a few white scales (contrary to

depauperata, which usually has a distinct median spot 6). The ground colour of

the underside of the hind wing is tawny.

Male genitalia (figs. 107—109). — Fenestrula at junction of uncus and tegumen

relatively small. Uncus with longitudinal keel and slightly inflated apex. Gnathos

fused to tegumen, not very much curving outward, ventrally somewhat extended to

form a rounded sclerotized flap with a few small knobs. Fultura superior smooth,

below gnathos passing into the extensive dorsal sclerotization of the anellus. This

sclerotization curves inward proximad to form a shallow sac, laterally strongly

spined, and then it folds roof-like over the aedeagus. Aedeagus slightly bent in the

middle; coecum scarcely longer than wide; ventral process short, about as long as

wide, with strong spines. Costa of valva expanding distally; costal process small,

mainly consisting of a series of spines radiating from the ventro-distal end of the

DE JONG: The genus Spialia 101

Figs. 107—111. Male genitalia, lateral aspect, aedeagus, and inside of right valva. 107—109, S.

sataspes (Cape Town, S. Africa). 110—111, S. depauperata australis (Muden, Natal).

costa. Cucullus broadly developed, dorsally excavated to receive the costal process.

Female genitalia (figs. 112—114). — Ventral sclerites of segment 8 fused into a

single sclerite, covering the ventral and lateral parts of the segment. No antevaginal

sclerotization. Just behind ostium two flap-like longitudinal outgrowths. Laterally a

few very small folds. Area spinulosa broad, distal edge slightly impressed. Papilla

analis much broader than long, apophysis posterior shorter than width of papilla.

Ecology. — Mainly found in grassy spots on mountain slopes, where it skips

about and feeds on flowers. It is on the wing from the middle of October to Feb-

ruary, and seems to be most abundant in November (Swanepoel, 1953). Food plants

are Hermannia species (Sterculiaceae), Pavonia macrophylla and Hibiscus

aethiopicus (Malvaceae) (Murray, 1959).

Distribution (Map 20). — Confined to South Africa, ranging from the Cape

peninsula eastwards to the Grahamstown district and with a few localities outside

this range. It was recorded by Butler (1901) from Mombasa, but this observation

probably relates to zebra.

102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Localities. — South Africa. Cape Province: Addo, Assegaibosch, Bathurst, Brak

R., Cape Town, Coega, Grahamstown, Knysna, Ladysmith, Lion’s Head, Llandud-

no, Muizenberg, Outeniqua Mts, Plettenberg, Port Alfred, Port Elizabeth, Sea Point

(Cape Town), Signal Hill, Simonstown, Springbok, St. James, Zwartkops (BM,

MN, NMB; Swanepoel, 1953).

Material examined. — 21 ” (incl. holotype), 119 , South Africa (BM, MN).

Spialia depauperata (Strand)

Hesperia ferax var. depauperata Strand, 1911. — Mitt. zool. Mus. Berlin 5 : 304 (Msamwia, between

Lake Rukwa and Lake Tanganyika, S. Tanzania; not Msamusia, as stated by Evans, 1937).

External characters (pl. 7 figs. 55, 56). — The largest species of the group; I saw

only one specimen (7) which was smaller than 10.8 mm, while the largest male

measured 13.8 mm. Its size, in combination with the presence of the basal cell spot

and the straight median band on the underside of the hind wing and the usual

presence of median spot 6 on the upperside of the hind wing, will usually suffice to

identify this species. See also zebra and sataspes. In doubtful cases a genital

examination is necessary, but this will seldom be the case.

Part of the literature suggests difficulties in distinguishing between depauperata

and diomus ferax and, actually, depauperata was originally described as a

“variety” of “Hesperia ferax”. There will be no problems if one pays attention to

the following points:

1. on the upperside of both fore and hind wing the basal cell spots are absent

(at most indicated by a few white scales) in depauperata, well-developed in diomus

ferax.

2. on the upperside of the fore wing only one of the outer median spots in space

lb (viz., spot Ic) is present in depauperata, sometimes scarcely visible; in diomus

ferax, both outer median spots are present and form a bar from vein 1 to vein 2.

3. on the underside of the hind wing the series of submarginal spots starts at the

end of vein 7 in depauperata, at the end of vein 8 in diomus ferax; the distal

edge of the median band reaches the termen at the end of vein 8 in depauperata,

but crosses vein 8 well before its end and runs to the costa in diomus ferax.

For variation, see subspecies.

Male genitalia (figs. 110, 111). — Uncus with high, longitudinal keel, apex usual-

ly more or less inflated. Relatively small fenestrula at junction of uncus and tegumen.

Gnathos dorsally fused to tegumen, dorso-distal edge protruding; ventral part of

gnathos somewhat more strongly sclerotized and with small, thick spines. Fultura

superior smooth, passing into dorsal sclerotization of anellus. This sclerotization

curves inward to form a shallow sac, and from there it extends as two strongly

sclerotized and spined, oblong processes with a pointed apex on either side of the

aedeagus. Aedeagus slightly bent in the middle, coecum shorter than wide, ventral

process relatively small, somewhat longer than wide. Costa of valva expands

distally and bears a twofold costal process. Distal part of costal process much as in

sataspes, with radiating spines, but better developed; proximal part very heavily

sclerotized, with serrated edge. Cucullus broadly developed, dorsally excavated to

receive the costal process.

DE Jong: The genus Spialia 103

1302

Figs. 112—116. Female genitalia, lateral and ventral aspect, and papilla analis. 112—114, S.

sataspes (S. Africa). 115—116, S. depauperata australis (Muden, Natal).

Female genitalia (figs. 115, 116). — Ventral sclerites of segment 8 fused to a

single sclerite. No antevaginal sclerotization. Postvaginally a narrow, V-shaped,

membranous strip, flanked by slightly protruding sclerotization. Laterally some

conspicuous, sclerotized folds. Distal edge of segment 8 straight. Papilla analis

104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Map 21

Spialia depauperata

rounded, nearly as long as broad; apophysis posterior slightly curving, about as

long as papilla.

Ecology. — Mainly found on grassy hill-sides, where the skippers feed on

flowers, but also found in the thorn bush. In S. Africa it is on the wing from July

to April in two generations (Swanepoel, 1953). This may also be the case in the

northern part of the range, but from there I have also seen specimens captured in

May and June. Food plants have not been recorded.

Distribution (Map 21). — From Uganda and Kenya through eastern Africa

south to Transvaal and Natal.

Geographic variation. — The width of the median band on the underside of the

hind wing varies geographically, the northern populations showing a broader band.

There is also some variation in the length of fore wing and the ground colour of the

underside of the hind wing, but much overlap exists in these characters.

Spialia depauperata depauperata (Strand)

Hesperia ferax var. depauperata Strand, 1911. — Mitt. zool. Mus. Berlin 5 : 304 (Msamwia, between

Lake Rukwa and Lake Tanganyika, S. Tanzania; not Msamusia, as stated by Evans, 1937). Holotype

(7) in the Zoologisches Museum der Humboldt-Universität, Berlin (labelled: S. Tanganyika, S. Ufipa,

25.1.09, Fromm.).

DE JONG: The genus Spialia 105

Syrichthus rehfousi Oberthür, 1912. — Et. Lép. Comp. 6 : 117, fig. 1331 (Mpala, W. shore of Lake

Tanganyika). Holotype (7 ) in the British Museum (Nat. Hist.), London.

For the use of the name abscondita (see Evans, 1937) instead of depauperata,

see synonymy of diomus ferax. |

External characters (pl. 7 figs. 55, 56). — Median band on underside of hind

wing broad, median spots 4—5 broader than width of ground colour in cell and

usually also broader than width of ground colour between median and submarginal

spots in spaces 4—5. This ground colour usually with a greenish tinge. & (10.4—)

11.3—13.8 mm; the largest specimens are found in Malawi and S. Tanzania.

Distribution. — The northern part of the range, Uganda and Kenya southward

to Zambia and Mozambique.

Localities. — Kenya: Kibwezi, Menfort, Mt. Elgon (BM, MN). Uganda: Ankole-

Toro Border, SE. Ankole, W. Ankole, Kampala, Kafu R. nr. Hoima-Kampala Rd,

Mbarara (S. Toro), Mulange (Mabira Forest), betw. Sezibwa R. and Kampala,

Simba (W. Lake Victoria) (AMNH, BM). Tanzania: Geita, Mpanda, Ngamo, Ufipa,

Wanzizi (BM, ML, MN). Zaire: Ditanto, Marungu Plateau, Mpala (= Makowiri),

Panda (BM, MRAC). Malawi: S. of Mangoche Mtn, Mlanje, Ncheu (BM, NMB;

Gifford, 1965). Zambia: Petauke, Solwezi (BM, HDE; Neave, 1910, as zebra).

Mozambique: Kola Valley, Tete to Ft. Jameson (Makanga Distr.) (BM, HDE;

Neave, 1910, as zebra).

Material examined. — 45 ,9 © : 4 , Kenya (BM, MN); 11° ,19,

Uganda (AMNH, BM); 14S , 4 9 , Tanzania (BM, ML, MN); 4 © (incl. holo-

type rehfousi), Zaire (BM, MRAC); 3 , 19 , Zambia (BM, HDE); 67,39,

Malawi (BM); 3 , Mozambique (BM, HDE).

Spialia depauperata australis subspec. nova

Holotype (© ): Muden (Natal), 15.III.1948. British Museum (Nat. Hist.), London.

External characters. — Median band on underside of hind wing narrow, median

spots 4—5 practically always narrower than width of ground colour in cell and

between median and submarginal spots in spaces 4—5. This ground colour usually

with a brownish tinge. 7 10.8—12.9 mm.

Distribution. — Transvaal and Natal, northwards probably to the Zambezi River.

Localities. — Rhodesia: Hunyani R. (Salisbury), Mtoko, Penkridge (Melsetter),

Redbank (W. Matabeleland), Umtali, Vumba (BM, MN, NMB). South Africa.

Natal: Estcourt, Ladysmith, Muden, Tugela, Weenen (BM, MN; Swanepoel, 1953).

Transvaal: Letaba, Lydenburg, Munnik, Pietersburg, Potgietersrust, Warmbaths,

Zoutpansberg (Swanepoel, 1953).

Material examined. — 44 7,21 9 :2 0,12 , Rhodesia (BM, MN). 42 ©,

20 © (incl. holotype), Natal (BM, MN, MRAC).

4. PHYLOGENY

Introduction

As said in the introduction to Chapter 3, the species groups recognized in this

paper are thought to be monophyletic groups. Such a statement would have little

106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

value, if it was not accompanied by arguments. The argument for considering a

group monophyletic (in the sense of Hennig, 1966) can only be the presence of an

autapomorphy. Thus, in this chapter we shall investigate phylogenetically the vari-

ous states of characters found in Spialia to find the arguments for the classification

proposed in the previous chapter.

In a future paper I shall show that only few of the many arguments used in the

literature to distinguish between plesiomorphous and apomorphous character states

and to find monophyletic groups are supported by the evolutionary theory. Our

present knowledge of Spialia allows the application of only three of these argu-

ments, viz., (1) the argument of outgroup occurrence, (2) the argument of differ-

entiation (complexity), and (3) the argument of vicariance. These arguments can

be defined as follows:

(1) If in a monophyletic group a character occurs in more than one state, the

state occurring also outside the group is considered plesiomorphous.

(2) If in a monophyletic group a phenocline of increasing differentiation or com-

plexity can be distinguished, the evolution is supposed to have taken place in the

direction of the strongest differentiation or the highest complexity, unless there are

indications that reduction took place as a result of a particular mode of life.

(3) If in a monophyletic group of three or more species two species are vicari-

ous, these species form a monophyletic subgroup (i.e. they are more closely

related to each other than to any other species).

In the following lines these arguments will be applied to a number of characters.

The outgroup is formed by the genera which are thought to be closely related to

Spialia, viz., Carcharodus, Gomalia, Celotes, Alenia, Syrichtus, Pyrgus and

Heliopetes. These genera, together with Spialia, are known as the Pyrgus group.

Where appropriate, the outgroup will be extended to cover the whole subfamily

Pyrginae or the whole family Hesperiidae.

In general the absence of a character in different species will not be taken as an

indication of kinship relationship. This is based on the following reasoning. If a

character arises, it can develop in a number of ways, each leading to a different

type and representing a different evolutionary line, that theoretically can be traced

back. A character can also be lost in different ways, each way representing a dif-

ferent evolutionary line, but as the result is the same in each way (loss of the char-

acter), the different evolutionary lines cannot be traced back.

Characters and character states

1. Wing markings

1.1. The central spot on the upperside of the fore wing is placed over or beyond

the origin of vein 3 in asterodia, agylla and kituina, and before the origin of vein

3 in all other species of Spialia. The first condition is also found in Syrichtus, the

latter in Pyrgus. So there is no clue to which condition is apomorphous and which

plesiomorphous.

1.2. On the upperside of the fore wing there are no spots in spaces 9 and 10,

except in asterodia, agylla and: kituina. These spots are absent in the outgroup,

DE JONG: The genus Spialia 107

except in Pyrgus and Heliotepes. In some Syrichtus species (antonia, cribrellum)

the spots may be vaguely outlined. Also in a number of other genera of the sub-

family Pyrginae there are spots in spaces 9 and 10. So there is no reason to con-

sider the presence of these spots in some Spialia species otherwise than a plesio-

morphous state. As said above, the absence of a character will not be considered

proof of kinship relationship. Consequently, this character is not very well suited for

our purpose.

1.3. The submarginal spots of the hind wing are in line, at equal distance from

the termen, in all Spialia species except colotes and confusa, where the sub-

marginal spots in spaces 4 and 5 are placed more basad. In the outgroup such a

displacement is only found in various degrees in some Syrichtus species. As it may

even vary within the species of Syrichtus (poggei), the presence of this character

state in Syrichtus and Spialia seems to be due to the phenomenon known as the

Law of Vavilov (cf. Vavilov, 1922), rather than to autapomorphy. Therefore, in

Spialia the shift of the submarginal spots in spaces 4 and 5 towards the base can

be considered apomorphous.

1.4. In the Pyrgus group the basal spots on the underside of the hind wing are

always detached from the median spots, except in two Spialia species, viz.,

phlomidis and kituina. In phlomidis it is the median band that bends to the basal

spot in space 7, in kituina it is the basal spot in space 7 that is elongated towards

the central spot of the median band. These unique arrangements of the spots can

only be considered apomorphous states that originated independently. Considering

one of these arrangements plesiomorphous would force us to recognize a large num-

ber of parallel changes in the arrangements of the spots, viz., towards a disconnec-

tion of basal and median spots. Only if the connection of basal and median spots

was also found outside the Pyrgus group, there could have been a reason for

considering the disconnection apomorphous.

1.5. In two Spialia species, viz., spio and diomus, a spot occurs between the

median spot and the submarginal spot in space 1c on the underside of the hind

wing. The spot does not occur in other species of the Pyrgus group and for that

reason its presence is considered apomorphous.

2. Legs

2.1. In all Spialia species the mesotibiae are unspined, except for the two

terminal spines, but in diomus the mesotibiae are provided with a double row of

spines. In the outgroup this character is only found in Syrichtus cribrellum. It is

possible that this is the result of common ancestry, in which case the spined meso-

tibiae constitute the plesiomorphous state in Spialia, unless we assume that cribrel-

lum is a descendant of diomus. There is no ground for the latter assumption. The

alternative explanation is a parallel development. Knowledge about the function of

the spines could support this explanation. Unfortunately, we do not have the slight-

est idea of their function. At the moment this character can be used in phylogenetic

considerations only in connection with other characters.

3. Secondary sexual characters

3.1. A costal fold in the fore wing, with specialized scales, is present in the

108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

males of five Spialia species, viz. asterodia, agylla, kituina, colotes, and confusa.

In the outgroup a costal fold is absent in Alenia, one species of Syrichtus and a

few species of Pyrgus. The character appears to be a characteristic of the sub-

family Pyrginae, though it is absent in many species. Its intricacy suggests that it is

an autapomorphy of the Pyrginae, that was lost independently in many cases. Thus

the presence of a costal fold is considered here a plesiomorphy of Spialia.

Table 1. Families of food plants of Spialia species.

Malvaceae

Sterculiaceae

Tiliaceae

Convolvulaceae

Rosaceae

unknown

asterodia

agylla

kituina

spio

diomus

phlomidis X

osthelderi

xx MK

xx KM

x xX

geron

doris X

colotes X

confusa X X

sertorius

orbifer

mafa

galba

wrefordi

paula

secessus

dromus X

ploetzi

mangana

nanus X X

delagoae

zebra

sataspes X

depauperata xX

x xX

x KK KKM

~*~ x

DE JONG: The genus Spialia 109

Table 2. Families of food plants of the genera of the Pyrgus group. The Sterculia-

ceae and Tiliaceae, food plants of a number of Spialia species and closely related

to the Malvaceae, have been omitted (see table 1).

Malvaceae

Rosaceae

Convolvulaceae

Labiatae

Cistaceae

Acanthaceae

Celotes

Carcharodus

Gomalia

Alenia X

Syrichtus X

Spialia

Pyrgus

Heliopetes

4. Ecology

4.1. All species of Spialia, as all species of the outgroup, are at home in open

formations, though they may penetrate forests along paths, etc. There is only one

exception, viz., ploetzi. This species is confined to the lowland evergreen forest. On

the basis of outgroup comparison this habitat preference is considered apomorphous.

4.2. The food plants of Hesperiidae are only very incompletely known, but the

little that is known appears to be interesting. Table 1 summarizes the families of

the food plants of the Spialia species as far as known and table 2 gives the families

of the food plants per genus of the Pyrgus group. From these tables it appears that

the Malvaceae are the original food plant family of the Pyrgus group, as species of

this family are used as food by species of almost all genera of the Pyrgus group.

This idea is supported by the fact that Malvaceae are food plants of small and

divergent genera like Gomalia, Celotes and Alenia. The divergence of these

genera suggests that they departed since long from their common ancestor. It is very

unlikely that they all independently adopted Malvaceae as food plants.

The Spialia species feeding upon Malvaceae usually also feed upon Sterculia-

ceae. These families are closely related and together with the Tiliaceae and some

other families they form the order Malvales. As the food plants are incompletely

known, the order Malvales instead of the family Malvaceae is considered here to

contain the original food plants of the Pyrgus group (and of Spialia). It follows

that the adaptation to Convolvulaceae and Rosaceae by some Spialia species is an

apomorphous character state.

110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

5. Male genitalia

5.1. A fenestrula is present in nine Spialia species, viz., spio, mafa, galba,

mangana, nanus, delagoae, sataspes, zebra and depauperata. In the outgroup it is

only found in Syrichtus cribrellum. This suggests that the presence of this character

is an apomorphy in the Pyrgus group and a plesiomorphy in Spialia. However,

a fenestrula occurs erratically outside the Pyrgus group, even in the subfamily

Hesperiinae. Thus it may be an apomorphous state in the Hesperiidae that was

secondarily lost on many occasions independently. The alternative explanation

implies a parallel development (e.g. due to the Law of Vavilov). However it may be,

the species of the Pyrgus group are all very similar, so that the idea that they are

closely related, seems a good working hypothesis. If in such a group of species a

character occurs in only few species, the presence of that character in these species

seems to be due to common ancestry rather than to parallelism. This may be inter-

esting for the relationship between Syrichtus and Spialia, but it implies that the

presence of a fenestrula in Spialia is a plesiomorphous state that is of little use in

the reconstruction of the phylogeny.

Though the presence of a fenestrula may have little value for the present study,

its further development may be useful. In Spialia galba and mafa, namely, the

fenestrula is very large, much larger than in any other species of the Pyrgus

group. Therefore this excessive development of the fenestrula is considered apo-

morphous.

5.2. The uncus of the Hesperiidae exhibits a great variety of shapes and dimen-

sions. In four Spialia species, viz., delagoae, zebra, sataspes and depauperata,

the apex is inflated. This character does not occur in any other species of the Pyrgus

group, nor in any other Hesperiid species examined by me. Therefore, I consider

the presence of this character the apomorphous state.

5.3. An indented or bipartite uncus occurs in many Hesperiidae, in the Pyrginae

as well as in the Hesperiinae. The same genus may have species with an entire and

with a bipartite uncus, e.g. Celaenorrhinus, Pyrgus. Both types of uncus may even

occur with a single species (Pyrgus malvae, see De Jong, 1972). The partition is

very variable, suggesting that it arose several times independently. In Spialia two

types of indentation occur:

1. in asterodia, agylla and kituina the uncus is bipartite, the two parts lying

more or less parallel. This is a usual type in the Pyrgus group, being found in

Pyrgus, Heliopetes and Celotes, though the parts are not always parallel. At the

moment there is no reason not to consider this type an apomorphous state within the

Pyrgus group, and thus a plesiomorphous state in Spialia;

2. in zebra the uncus is shallowly indented and the two parts are widely diverg-

ing, giving the uncus the shape of a Y or T. This type is unique and must be

considered an apomorphous state.

5.4. There is a keel running dorso-centrally over the uncus in five Spialia

species, viz., nanus, delagoae, zebra, sataspes and depauperata. This character

is absent in all other species of Spialia and the other genera of the Pyrgus group.

Therefore, its presence is considered to represent the apomorphous state.

DE JONG: The genus Spialia 111

5.5. The gnathos occurs in very different types in the Hesperiidae. Warren

(1957), studying the gnathos (“lateral apophyses”) of Pyrgus species, thought it to

be the original tenth abdominal segment with sclerites of the twelfth segment, but

his argumentation is little convincing. I very much doubt if the gnathos is homolog-

ous in all Lepidoptera where it occurs. If a gnathos is developed to some extent, it

appears to have a distinct function, being an extra grip during copulation. As it

apparently has adaptive value, it may be expected that adaptive radiation has

occurred and that more or less similar types originated more than once from the

same basic structure. |

Confining ourselves to the Pyrgus group, the following phenocline can be ob-

served:

a. sclerotization of fultura superior from right and left sides of tegumen down-

wards (towards aedeagus sheath);

b. differentiation of ventral parts of downward sclerotizations (sclerotization

stronger and better defined, development of spines, etc.);

c. sclerotization gets detached from tegumen (greater freedom of movement).

Stage c occurs only in conjunction with stage b. Applying the argument of

differentiation the phenocline can be read phylogenetically in the direction a > b

> c. Reading it in the reverse direction would imply that from a great variety of

forms (and how would they have originated?) reduction took place to a slight

sclerotization of the fultura superior and eventually to a fusion of this sclerotization

with the tegumen. Such a development would make little sense and is difficult to

understand.

In Spialia the following types are found:

1. stage a: colotes, confusa, paula, wrefordi, secessus, dromus, ploetzi;

2. stage b: 1. ventral parts strongly sclerotized but smooth: asterodia, agylla,

kituina; 2. ventral parts spined, no definite protrusions, dorsal parts broad:

sataspes, depauperata;

3. stage c: 1. ventral parts spined but no definite protrusions (“heads”): spio,

diomus, mangana, zebra, nanus; 2. ventral parts forming definite, spined protrusions

(“heads”): phlomidis, osthelderi, geron, doris, galba, mafa, sertorius, orbifer,

delagoae.

5.6. In some species of the Pyrgus group the fultura superior is densely set with

long, but weak spines (sometimes more looking like hairs), especially in the medio-

ventral part. This is found in a number of species of Syrichtus and Spialia, so that

it appears that it is a plesiomorphous state in Spialia. It is usually found in con-

junction with gnathos stage a, suggesting that it has the same function as the spined

heads of the gnathos found in stage b. The extension of the spines varies from an

undefined area to a well-defined area covering two semiglobular membranous folds

in the ventral part of the fultura superior, in connection with the rather undefined

gnathos. As the latter situation is a further differentiation, it is supposed here to be

apomorphous. It is found in paula, wrefordi, secessus, dromus and ploetzi, and to

some extent in mangana.

5.7. In some Spialia species sclerotization occurs in the dorsal parts of the

aedeagus sheath. This is unusual in the Pyrgus group. As the sclerotization is a

112 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

further differentiation and the two types of sclerotization found in Spialia do not

occur in the outgroup, they are supposed to have originated independently from the

unsclerotized stage. The two types are:

1. rather well defined sclerite dorsally over aedeagus. This sclerite is not flat, but

gutter-like (wrefordi, secessus, dromus, colotes, confusa) or roof-like (paula, ploetzi).

The relationship between these subtypes is not clear at the moment and they are

considered as belonging to the same type till more can be said about their relation-

ship;

2. dorso-laterally, to the right and to the left, a strongly sclerotized and spined

protuberance; the aedeagus sheath membrane between the right and left parts more

or less strongly sclerotized. This is found in mangana, nanus, delagoae, zebra,

sataspes, and depauperata. This type looks like a further differentiation of the first

type. Support for this suggestion must be looked for in the distribution of other

characters.

5.8. In the Pyrgus group the surface of the aedeagus is smooth or at most with

fine spines, except in some Spialia species. According to the argument of outgroup

comparison, the presence of outgrowths in Spialia is apomorphous. Three types

are found:

1. delagoae type — a short, ventral cylindrical, spined excrescence: mangana,

nanus, delagoae, zebra, sataspes, depauperata;

2. phlomidis type — a lateroventral, strongly sclerotized, dentate ridge: phlom-

idis, osthelderi, doris;

3. geron type — a lateroventral, long, cylindrical, smooth excrescence, giving

the aedeagus a forked appearance: geron.

The phlomidis and geron types look dissimilar, except that the position of the

excrescence (to the left) is identical. However, all other characters (see, e.g. the

female genitalia, below) point to a close relationship of the species exhibiting these

types of aedeagus and for that reason the long excrescence of the geron type of

aedeagus is considered here a further differentiation of the phlomidis type (see also

De Jong, 1974).

The delagoae type appears to be a development that is quite independent from

the other types. It is, indeed, difficult to imagine a relationship between these types,

not only the form but even the position of the outgrowth being dissimilar. In view of

the present knowledge we can only consider the delagoae type an independent line

of evolution.

5.9. In all species of the Pyrgus group the aedeagus is straight or very slightly

bent, except in the Spialia species wrefordi and ploetzi, where it is clearly bent

to a more or less S-shaped structure. Because of its distribution in the Pyrgus group

the bent condition is considered apomorphous.

5.10. In all species of the Pyrgus group the aedeagus has a well-defined coecum,

except in a number of Spialia species, where the coecum is shortened. These

species are: delagoae, nanus, zebra, sataspes, depauperata, and mangana. In the

last species the coecum is practically lost. As the differences in length of the coecum

between the species are often very slight, an arrangement of the species in a phen-

ocline would be rather arbitrary, except for the extreme condition found in mangana.

DE Jong: The genus Spialia 113

Because of the state of this character in the outgroup, the tendency to shorten the

coecum must be considered apomorphous.

5.11. The valvae have a clear function during copulation: they clasp the end of

the female abdomen. For a better grip the inside is often provided with prominent

outgrowths, spines, haires, etc. This is the general picture throughout the Lepidop-

tera. Often various types of ornamentation of the inside of the valvae can be found

within a group of genera or even within a genus. So three types can be found in

Spialia, all referring to the differentiation of the costa:

1. phlomidis type — distal end of costa more or less widened, ventrally usually

with a strongly spined outgrowth or dorsally with a finger-like excrescence: phlom-

idis, geron, osthelderi, doris, spio, diomus;

2. kituina type — ventro-distally a heavily sclerotized appendix with at least two

heavy spines: asterodia, agylla, kituina;

3. sertorius type — ventro-distally a more or less differentiated appendix that is

not more heavily sclerotized than the remainder of the costa and bears long, relative-

ly soft spines: remainder of Spialia.

The kituina and sertorius types are sufficiently differentiated to consider them

to represent two different, independent evolutionary lines: they are so much com-

plicated that a single origin of each of both types can be accepted and they differ

so much that a development of one type into the other is difficult to imagine. As

these types are unique in the Hesperiidae, they are both considered apomorphous,

but it is not easy to find the respective sister groups with the plesiomorphous state.

The phlomidis type is rather varied and relatively simple. It comes closest to what

is found in the other genera of the Pyrgus group, especially Syrichtus and Car-

charodus. For this reason it is supposed to have departed not far from the ancestral

type of Spialia. It is too varied to be simply considered the plesiomorphous state

of one or both of the other types. The species, however, with this type of costal

development can, as a group, quite well be considered the sister group of the group

of species exhibiting one of the other types.

5.12. Within the kituina and sertorius types mentioned in 5.11. a phenocline of

increasing complexity can be observed. As the primitive state of these types is

supposed to have been rather simple (see above) and there is no reason to suppose

that reduction took place (that would even be inconceivable in the kituina type,

see below), the phenoclines may represent phyloclines. The clines are as follows:

1. kituina type — a. costa smooth (except for costal process), costal process

relatively little differentiated (asterodia); b. costa with some short spines, costal

process more differentiated (agylla); c. costa with some short spines, costal pro-

cess gets loose from costa and joins cucullus (kituina). If asterodia had devel-

oped its costal process from the type found in kituina, i.e. through reduction of the

costal process of kituina, then the latter would have rejoined the costa before

further reduction, an inconceivable development. As the costal process in kituina

cannot be said to be more differentiated than that in agylla (it is only dissimilar and

has another position), it is quite well possible to suppose that kituina arose from

an asterodia-like ancestor, independent from agylla. In that case the spines on

the costa in kituina and agylla represent a parallel development.

114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

2. sertorius type — a. costal process relatively small, rounded, with spines

radiating to the ventral side (mangana, zebra, sataspes, depauperata); b. costal

process elongate towards base of valva, spines radiating to the ventral side, but

spines directed to the base of the valva predominant (wrefordi); c. costal process

elongated, straight or curved upwards, all spines directed to the base of the valva

(paula, secessus, dromus, ploetzi, nanus, delagoae). Starting from a), a second

phenocline of the sertorius type of costal process can be observed, in which the

connection between the costal process and the costa narrows, so that the costal

process looks more separate, while the spines are not radiating, but directed basad

and distad (colotes, confusa) or distad only (sertorius, orbifer, mafa, galba). As

both phenoclines show an increasing differentiation, they are considered to repre-

sent phyloclines.

5.13. There are some further developments of the costa, which for their unique-

ness in the Pyrgus group are considered apomorphous. They are:

1. In ploetzi the costa is extended distally over the cucullus.

2. In confusa the costa has grown fold-like over the costal process.

3. In osthelderi, geron and doris the costa is spined dorsally.

5.14. The cucullus is far less differentiated than the costa in Spialia. There are

only two remarkable differentiations:

1. In the species with a costal process, except in mangana, nanus, delagoae,

zebra, sataspes and depauperata, the cucullus is extended at the inside into a thin

fold that more or less covers the costal process. Outside Spialia a cucullus fold

occurs only in Carcharodus alceae, be it in another form. As C. alceae is in all

other aspects very different from Spialia, the common possession of a cucullus

fold is supposed to be a parallelism. Not only for the outgroup comparison, but also

for the fact, that the cucullus fold appears to be correlated with the costal process,

the presence of the fold is considered apomorphous. It would be interesting to know

the function of the fold. If the costal process has developed as an extra grip during

copulation, the cucullus fold must again reduce its effect.

2. In doris the cucullus is divided into two parts, apparently an apomorphous

condition as it is a further differentiation of the cucullus, that does not occur else-

where in the Pyrgus group. The division of the cucullus found in some members of

Syrichtus is of another type (see a forthcoming paper on the genera of the Pyrgus

group).

6. Female genitalia

6.1. The eighth abdominal segment is strongly modified in connection with the

genitalia, especially the ventral part. The dorsal part is much less modified and in

Spialia we can distinguish between two different types of sclerotization, viz., a) a

single sclerite (tergite) covering the greater part of the dorsal area (asterodia, agylla,

kituina), and b) two smaller sclerites, mid-dorsally separated by a membranous area

(all other Spialia species). The first type is found in the outgroup in Carcharodus,

Celotes and Alenia, and also outside the outgroup (e.g. Erynnis, Xenophanes,

Lobocla). For this reason, and because a single tergite most closely corresponds

with what must have been the original shape of the eighth tergite in females, the

first type is considered plesiomorphous.

DE JONG: The genus Spialia 115

6.2. The dorsal sclerites of the eighth abdominal segment are never fused to the

ventral sclerites in the Pyrgus group, except in the Spialia species sertorius,

orbifer, mafa, and galba, while in colotes and confusa the dorsal and ventral

sclerites practically touch. Thus, in Spialia there is a phenocline in the fusion of

the dorsal and ventral sclerites, with three stages, viz., a. not fused; b. touching;

c. fused. As the first stage is the usual one in the Pyrgus group (and outside), the

phenocline a — c is supposed to represent a phylocline in the same direction.

6.3. The development of the ventral sclerites of the eighth abdominal segment is

too varied to discover a certain line of evolution in it. If, e.g., the ventral sclerites

are fused postvaginally, it cannot be determined whether this is due to extension of

the sclerites or to an extending sclerotization of the postvaginal membrane. There-

fore, the characters of the ventral sclerites are left out of consideration here (see,

however, the discussion of the delagoae group).

6.4. In a number of Spialia species there is a distinct area spinulosa, viz., spio,

diomus, colotes, mafa, galba, sertorius, orbifer, wrefordi, paula, secessus, dromus,

and ploetzi, and in a few others there may be some indistinct spines postvaginally.

An area spinulosa is also found in the outgroup, in Syrichtus and Carcharodus.

Therefore I consider its presence a plesiomorphous condition. With increasing

sclerotization of the postvaginal area the spines seem to tend to disappear. As the

sclerotization probably evolved along different lines (see 6.3.), the absence of spines

in different species is not necessarily an autapomorphy.

6.5. A unique development in the Pyrgus group, and probably outside as well,

is the presence of strongly sclerotized, serrated “wings” at the proximal edge of the

ventral side of the eighth abdominal segment in the Spialia species phlomidis,

osthelderi, geron, doris, spio, and diomus. In the last two species there is only a

weak development of these structures. Thus, there is a phenocline as follows: a. no

“wings”; b. small, indistinct “wings”; and c. large “wings”. In view of the absence

of these structures outside Spialia their presence is supposed to be apomorphous and

the phenocline is considered to represent a phylocline in the direction a —> c.

6.6. In two Spialia species, viz. spio and diomus, the postvaginal area is

provided with a tongue-like outgrowth that partly covers the area spinulosa. Outside

Spialia a similar structure is only found in Carcharodus lavatherae. For the rest,

C. lavatherae and the Spialia species are so dissimilar that it seems most plausible,

that the outgrowth arose independently in Carcharodus and Spialia, just as

sclerotization in the same area must have arisen more than once. Consequently, the

presence of this outgrowth in Spialia is considered apomorphous.

6.7. The ductus bursae is smooth and more or less straight in all species of the

Pyrgus group, except in two Spialia species, viz., colotes and confusa, where it

is distinctly bent shortly before the bursa; the wall is corrugated at this place. In

view of the distribution of this character its presence is considered the apomorphous

condition.

6.8. In all species of the Pyrgus group the bursa copulatrix is without ornamen-

tation, except for Spialia galba, which has a large, finely spined area in the bursa.

Apparently an apomorphous character state.

6.9. Ten Spialia species have a basal lobe on the papilla analis. In the Pyrgus

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

116

a a a a

5 3 a 5 3

ro) O = O

3 u ep) ep)

I) a ic © © = ©

6.5 6.7 6.2

5.8 5.7 5.12

6.6 5.5 1.3 5.5

1.5 4.2 6.10

) 6.2 5.7 5.12

u 5.12 5.6 5.10

5.8

5.14 5.7

5.6

6.5 5.11

5.11 6.1

5.5 5.3

Fig. 117. Supposed evolution of Spialia. Black dots indicate apomorphous character conditions, the

plesiomorphous condition of the same characters is supposed to be present in the sister groups. The |

numbers refer to the paragraphs in the text where the relevant characters are discussed.

DE JONG: The genus Spialia 117

group a basal lobe is further found in Carcharodus (one species only), Alenia and

Syrichtus. If the presence of a basal lobe is not plesiomorphous in Spialia, then

the structure arose so often independently that it is of little use for phylogenetical

considerations.

6.10. Although the papillae anales are variously shaped in the Pyrgus group

and for that reason are difficult to interpret phylogenetically, an exception must be

made for the remarkably shaped papillae anales of the Spialia species mangana,

nanus, delagoae, zebra, sataspes, and depauperata, with their short and broad

body, and short, curved apophyses posteriores, giving the appearance of a quite

distinct type of structure. I cannot do otherwise than considering this type apo-

morphous in relation to the type found in the other Spialia species.

The above considerations are the arguments for the reconstruction of the phyl-

ogeny as shown in fig. 117. For the sake of surveyability the diagram has been

drawn up to the level of the species groups recognized in this paper, to show the

phylogenetic basis for the division into species groups. The further development will

be dealt with below for each species group separately and further diagrams will be

provided for the dromus and delagoae groups. For diagrams of the phlomidis

and sertorius groups the reader may be referred to my earlier paper (De Jong,

1974). The other species groups consist of so few species that the diagrams would

have no meaning at all.

The phylogeny of the species groups

The asterodia group

Starting from the supposed monophyly of Spialia (see Introduction and a future

paper on the genera of the Pyrgus group) the remainder of Spialia is considered

the sister group of the asterodia group. Autapomorphies of the latter are the

smooth, well-defined, heavily sclerotized gnathos and the peculiar shape of the

costal process of the valvae.

It is surprising that the characters by which the species can so easily be distin-

guished, viz., the position of the central cell spot on the upperside of the fore wing

and the presence of a costal fold at the fore wing in the male, cannot on the basis

of our present knowledge be considered autapomorphies of the group. By these

characters and by the single eighth tergite of the female abdomen and the bipartite

uncus in the male genitalia, the group stands out as having retained a relatively

large proportion of primitive character states, though it has developed some

pecularities of its own. In all, it gives the impression of a group of which the

ancestor has departed since long from the main stem of Spialia.

The development of the group seems to have been greatly influenced by the

division of the original distribution area into two parts. This resulted in two species

in South Africa (asterodia and agylla) that are very similar, and one divergent

species in East Africa (kituina). The close resemblance between asterodia and

agylla suggests that they originated after the separation of the East African pop-

ulation. Autapomorphies of kituina are the position of the costal process of the

valva and the peculiar Y-mark on the underside of the hind wing. For asterodia +

118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

agylla no autapomorphies can be listed at the moment. See also the discussion

under 5.12.

The sister group of the asterodia group

The monophyly of this group is based on two autapomorphies, viz., the un-

divided uncus of the male and the double tergites of the eighth abdominal segment

of the female. If this is correct, the group must have had a relatively simply built

valval costa, which only differentiated after the next cleavage of the phylogenetic

line. This cleavage gave rise to the ancestor of both the spio and phlomidis groups

and to the ancestor of the remainder of Spialia.

In the line leading to the spio and phlomidis groups the male genitalia remain-

ed relatively simple, the main differentiation affecting the gnathos, but a remarkable

development took place in the female genitalia, viz., the development of antevaginal,

free sclerotizations (“wings”). If such sclerotizations arose more than once in Spialia,

it can hardly be assumed that they would be of the same type. Therefore, the

monophyly of the spio + phlomidis group is considered rather well established,

though based on a single autapomorphy.

In the line leading to the remainder of the genus the main development took

place in the male genitalia, where the costa developed a special, spined structure,

viz., the costal process, and the gnathos became increasingly differentiated. The

costal process is basically so uniform in the different species, that it does not seem

too risky to accept it as an autapomorphy of the four remaining species groups.

One may wonder if it is not significant that the development of the female

genitalia in one line and of the male genitalia in the other both seem to serve a

better grip during copulation. If this is true, then it is understandable that in differ-

ent lines male and female genitalia evolve in an inversely proportional way. At the

moment this is pure speculation, but it may point to an important field of research,

viz., the exact use of the various parts of the genitalia during copulation.

The spio group

The species in this group, spio and diomus, are considered sister groups because

of two apomorphies which are both supposed to have arisen only once, viz., the

presence of a tongue-like postvaginal structure in the female and the presence of an

extra spot between the median and submarginal spots in space Ic on the underside

of the hind wing. Another shared character, which cannot however be called an

apomorphy, is their almost pan-Afrotropical distribution. For the rest they are rather

divergent, especially in the male genitalia, suggesting that their origin is not very

recent. As the male genitalia of spio most closely resemble those of the sister

group of the spio group, it appears that diomus has drifted farther away from the

ancestral condition. This being so, the spines on the mesotibiae of diomus are

rather a new development than a relic from ancient times.

This group is an example of the influence of phylogenetic considerations on

classification. Higgins (1924) thought the differences in the wing markings too

large to unite spio and diomus into a single group. Instead, he placed mafa and

paula with spio, as these species lack the median spot 6 on the underside of the

DE Jong: The genus Spialia 119

hind wing. As said before, absence of a character in different species should not be

taken as an indication of kinship relationship, as loss of a character may easily

occur more than once by different causes. Thus, the typologically based group spio

+ mafa + paula is replaced by the phylogenetically based group spio + diomus.

The phlomidis group

The monophyly of this group is well established by the shared possession of

four apomorphies, viz., food plants (Convolvulaceae), differentiation of gnathos,

ornamentation of aedeagus, and large, winged, antevaginal sclerotizations. The

further development of the group was treated in an earlier paper (De Jong,

1974) and there is no need to repeat the discussion here.

The sister group of the spio + phlomidis group

It seems that after the first cleavage of this group two sister groups originated

with highly different rates of evolutionary change. While the line giving rise to

the delagoae group apparently only split further after having obtained a num-

ber of new character conditions, so that the species share a large number of

apomorphies, the other line split already after a single new development, viz., an

excavation of the dorsal part of the cucullus resulting in a fold that partly covers

the costal process. As a result, the colotes + sertorius + dromus group looks

less homogeneous than the delagoae group. This is, of course, not an unexpected

statement, as the heterogeneity of the first group was the first reason for a division

into three species groups; it only shows the evolutionary background of this ar-

rangement. In fact the possession of a cucullus fold was not an entirely new de-

velopment of the colotes + sertorius + dromus group. It apparently started

with a hollowing of the dorsal part of the cucullus. This development must have

started before the cleavage with the delagoae line was finished, as the excava-

tion of the cucullus can also be seen in the delagoae group. But the character

attained full development only in the other line (see, however, the discussion on

the sertorius group).

After cleavage of the colotes + sertorius + dromus line the costal process

of the valva, still a rather undifferentiated excrescence with spines, begins to con-

tract to an oval shape in one of the sister groups, the colotes + sertorius group,

while also a tendency develops for the dorsal and ventral sclerites of the eighth ab-

dominal segment of the female to fuse. By these two new developments the mono-

phyly of the colotes + sertorius group is rather well established. There is,

however, one difficulty. If the reconstruction of the phylogeny given here is cor-

rect, then the dorsal sclerotization of the aedeagus sheath found in the colotes

and dromus groups must have arisen twice. Making, for the sake of this charac-

ter, the colotes + dromus group sister group of the sertorius group, would com-

pel us to accept an independent development of the differentiation of the costal

process and the tendency to fusion of the ventral and dorsal sclerites of the eighth

abdominal segment of the female in the colotes and sertorius groups. Just one

parallel development against two. I have thought it wisest to choose for parsimony

here and to attribute the occurrence of sclerotization of the aedeagus sheath in

two species groups to the Law of Vavilov.

120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Yet another character may seem to couple the colotes and dromus groups, viz.,

the spined fultura superior. In the colotes group, however, this character appears

to exist in a primitive stage, much like the condition it must have had before the

dromus group and even the delagoae group branched off, and different from the

large semiglobular folds characteristic of the dromus group.

It may be remarked that the above reasoning were superfluous if the colotes,

sertorius and dromus groups were united into one species group, as sister group

of the delagoae group. This would, however, only have moved back the problems

to the moment the development within the combined group had to be dealt with.

The colotes group

Leaving the dorsal sclerotization of the aedeagus sheath out of consideration,

the monophyly of the group is sufficiently established by the corrugation and

bend of the ductus bursae and the inward shift of the submarginal spots 4 and 5

of the hind wing. A diagnostic character of the group is further the presence of a

costal fold at the fore wing in the male. However, as said before, this character

has a very scattered distribution throughout the Pyrginae and the genetics ruling

its development are not yet understood, making it worthless at the moment for

phylogenetic studies.

The most remarkable new development within the group is the fold that from

the costa of the valva extends over the costal process in confusa. However, as the

group consists of only two species, colotes and confusa, the further development

of the group is of little interest from the phylogenetic point of view.

The sertorius group

Autapomorphies of this group are in the male the differentiation of the ventral

part of the gnathos into spined heads and the differentiation of the costal process

with spines directed only distad, and in the female the fusion of the dorsal and

ventral sclerites of the eighth abdominal segment. These apomorphies appear

enough to establish the monophyly of the group firmly. It is therefore surprising

to find that in two of the four species of the group, viz., sertorius and orbifer, the

cucullus fold is not much developed, suggesting a rather primitive state of this

character, dating from before the cleavage with the delagoae group. If on the

ground of this character sertorius and orbifer were left out of the group and

placed as a group that originated before or shortly after the cleavage that gave

rise to the delagoae group, then all the autapomorphies of the “old” sertorius

group, of the sertorius + colotes group and of the colotes + sertorius +

dromus group were based on parallel developments and no true autapomorphies.

It seems more plausible that the cucullus fold was reduced in sertorius and

orbifer, as a secondary development.

The group falls apart into two apparently monophyletic groups, viz., 1. ser-

torius + orbifer — autapomorphies: larval food plants belong to Rosaceae,

almost complete fusion of dorsal and ventral sclerites of eighth abdominal seg-

ment in the female;

2. mafa + galba — autapomorphy: very large fenestrula. The dorsal and

DE JONG: The genus Spialia 121

ventral sclerites of the eighth abdominal segment of the female are only partly

fused, not over their whole length.

For further details on the development of the group (also subspecific) the reader

is referred to De Jong (1974).

A remarkable feature of this group is the allopatry of all species (sertorius

and orbifer may overlap to a small extent). This is partly due to colonization

from Africa across the Saharo-Arabian eremic zone, which made a subsequent

sympatry very difficult. This will be dealt with in the next chapter.

> The dromus group

The autapomorphies of this group are found in the male genitalia: the large,

semiglobular, spined folds of the fultura superior and the elongated costal pro-

cess. The female genitalia do not yield characters that at the moment can be inter-

preted phylogenetically, with exception of the very deeply incised area spinulosa

in wrefordi, a unique and for that reason supposedly apomorphous character

state.

For the reconstruction of the phylogeny of the group the following characters

can be used:

a. costal process — plesiomorphous: wrefordi; apomorphous: paula, secessus,

dromus, ploetzi (see above, 5.12);

b. costa of valva — the distal end of the costa is usually rounded in Spialia,

but in secessus and dromus it is more or less angulated, and in ploetzi it is

elongated (see 5.13). This is a phenocline leading to an increasing differentiation

and, for that reason, considered to represent a phylocline. In paula the dorsal

ridge of the costa slopes gently down distally and the costa is only vaguely out-

lined ventrally;

c. area spinulosa — this part of the female genitalia shows some variation

in the Pyrgus group, especially with regard to the distal rim being arched, straight

or slightly indented. In view of this variation the situation in wrefordi (an almost

bipartite area spinulosa) can only be considered apomorphous;

d. habitat preference — the only Spialia species (virtually the only species of

the Pyrgus group) that is confined to the forest, is ploetzi. This preference is

considered apomorphous (see 4.1);

e. seasonal dimorphism — although in species of the Pyrgus group with more

than one generation per year the adults may vary seasonally, the dimorphism

found in secessus, in which the white of the spots on the underside of the hind

wing is entirely replaced by brown in the dry season, is unique. For this reason the

dimorphism of secessus is considered apomorphous.

On the basis of these characters the reconstruction of the phylogeny of the

group in fig. 118 has been made. The following comments may be added:

1. The aedeagus is distinctly bent in wrefordi and ploetzi. This was considered

apomorphous (see 5.9.). On the basis of the other characters, however, we must con-

clude that it is a parallel development in these species.

2. The monophyly of secessus and dromus is not very well established, as

it is not based on autapomorphies. Although these species have abundantly differ-

122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

secessus

dromus

ploetzi

Cle

paula

Fig. 118. Supposed evolution of the Spialia dromus group. Only apomorphous conditions have been

indicated (see also fig. 117). Numbers and letters refer to the relevant paragraphs in the text.

ent wing markings, their male and female genitalia are much alike and on the

basis of the present knowledge it cannot be stated whether this is due to symplesio-

morphy or synapomorphy. At the moment the reconstruction of the phylogeny as

given in fig. 118 seems the most probable one. It may be remarked that, if it turns

out that the similarity of secessus and dromus is due to symplesiomorphy, this does

nota priori imply that secessus and dromus are no sister groups.

The delagoae group

The species of this group share relatively many apomorphies, indicating that

their ancestor had already departed far from the ancestor of the colotes + ser-

torius + dromus + delagoae group before it split further to the present

situation. For the reconstruction of the phylogeny of the group the following

apomorphous character states can be used:

a. apex of uncus more or less inflated: nanus, delagoae, zebra, sataspes, depau-

perata (in nanus more rounded than inflated) (see 5.2).

DE Jong: The genus Spialia 123

b. uncus bifid: zebra (see 5.3.).

c. uncus with a longitudinal, dorsal ridge: nanus, delagoae, zebra, sataspes,

depauperata (see 5.4).

d. ventral differentiation of gnathos. The kind of gnathos found in the dela-

goae group is unique. There appears to be a phenocline of increasing differentiation

from a dromus-like gnathos (mangana) through the ventrally still undifferen-

tiated gnathos of nanus to a stronger differentiation in the ventral parts. Two

types can be distinguished: 1. gnathos ventrally strongly sclerotized with some

tubercles or short spines (zebra, sataspes, depauperata), and 2. gnathos ventrally

with a strongly spined excrescence (delagoae). Both types may have arisen from

the same undifferentiated nanus-like type.

e. costal process. The costal process occurs in two forms in the delagoae group,

viz., as a rather undifferentiated outgrowth of the costa with radiating spines

(mangana, zebra, sataspes, depauperata), and as an elongated excrescence of the

costa, directed towards the base of the valva (nanus, delagoae). As explained in

5.12 the first type represents the plesiomorphous type. The simplest development

of the costal process is found in mangana. In view of the occurrence of other

plesiomorphous character states in mangana (gnathos, see above; female genitalia,

see below), we may suppose that the simplicity of the costal process of mangana

is not due to reduction, but represents a very primitive state. The elongated form

of the costal process of nanus and delagoae reminds of the costal process of the

dromus group. This can, however, only be due to convergence, unless all apomor-

phous characters shared by nanus and delagoae with the other species of the

delagoae group, are due to convergence. Moreover, the costal process of nanus

and delagoae extends from a very narrow, reduced costa, as in the other species

of the delagoae group, and not from a well developed or even expanded costa as

in the dromus group. Thus, the elongated costal process of nanus and delagoae

is considered an apomorphous state.

A remarkable development is found in depauperata: apart from the usual costal

process there is an extra, strongly sclerotized outgrowth with a serrated edge at

the ventral side of the costa, close to the costal process. Such a unique development

can only be considered apomorphous, no similar structures being known.

f. Under 6.3. it was stated that the sclerotization of the ventral side of the

eighth abdominal segment of the female is too varied to be of use for phylogene-

tic considerations. An exception, however, must be made for the delagoae group.

In all Spialia species (as in other species of the Pyrgus group) some part of the

postvaginal area is membranous. In the delagoae group there is an increasing scle-

rotization of the postvaginal area, starting from a central sclerotization, as in

mangana that fuses with the ventral sclerite, as in nanus, delagoae, zebra and

sataspes, and leads to an almost entirely sclerotized ventral side of the eighth

abdominal segment, as in depauperata, where the area spinulosa is entirely incor-

porated in the sclerotization and has lost its fine spines. In view of the develop-

ment of the female genitalia in the other Spialia species and outside Spialia, the

progressing sclerotization of the postvaginal area in the delagoae group must be

considered to represent a progressing apomorphy.

124 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

per)

“= E

= È È g

O 2 O) © m 3

| es © = ©

= = D N u D

@ 6.10

@ 5.12

@ 510

5.8

5.7

Fig. 119. Supposed evolution of the Spialia delagoae group. Only apomorphous conditions have been

indicated (see also fig. 117). Numbers and letters refer to the relevant paragraphs in the text; t refers to

the general discussion of the evolution of this group.

The reconstruction of the phylogeny of the group in fig. 119 is based on the

above considerations. It may seem surprising that no apomorphies have been indi-

cated for mangana, suggesting that this species has remained unchanged after

the ancestor of the other species split off. This is improbable; there are several

characters that in mangana undoubtedly occur in an apomorphous state, but

this state becomes apparent only after mangana has been placed as sister group

of the remainder of the delagoae group and, for that reason, cannot be used as

DE JONG: The genus Spialia 125

argument for the drawing-up of the phylogeny. Such apomorphies are: 1. gnathos

free from tegumen; 2. apex of aedeagus flattened and wide, spoon-like; 3. coe-

cum virtually absent (see 5.10); 4. strong antevaginal sclerotization; 5. complete

lack of submarginal spots on all wings.

Also the supposed monophyly of zebra and sataspes calls for explanation. It

is difficult to give a single autapomorphous character state of these species. They

are, however, so similar in a number of characters, that there is a fair chance

that at least one of the characters occurs in an (aut-)apomorphous state in zebra

and sataspes. Such characters are: 1. extension and shape of sclerotiza-

tion in dorsal part of aedeagus sheath and in membrane between aedeagus sheath

and gnathos; 2. short, slightly bent aedeagus; 3. short ventral process of aedea-

gus; 4. length of coecum; 5. apparent absence of cornuti; 6. general shape and

development of valva. Further, if the monophyly of zebra, sataspes and depauperata

is correct, the geographic vicariance of zebra and sataspes may be an indication

of their close relationship, but the same applies to sataspes and depauperata.

5. GEOGRAPHIC HISTORY

As there are many factors that may influence the distribution history of a spe-

cies, it is usually little rewarding to study the history of a single species if it is not

possible to compare it with the history of the fauna the species belongs to. Thus

in Europe, where the history of the fauna is relatively well-known, a study of the

distribution history of a single species may yield some results. In tropical Africa,

however, where the knowledge of the faunal history is extremely fragmentary, we

can only hope to get some results by studying geographic patterns of distribu-

tion and evolution. A pattern, however, implies repetition and thus, it emerges only

when many species can be compared. For that reason the genus Spialia, inclu-

ding only 26 species of which 20 occur in tropical Africa, does not seem an

appropriate subject for a study of the geographic history. It appears, however, that

even from the relatively few species a pattern of distribution and speciation arises

that is worth discussing, as it may have some bearing on the general history of the

fauna of open formations (the usual habitat of Spialia species).

Distribution patterns

In Chapter 4 it was supposed that the preference for forest in Spialia ploetzi

is a recent development, coinciding with other recent developments in the same spe-

cies. The other Spialia species have, as a group, occupied all areas with open

formations throughout Africa, as far as there is some vegetation, as well as the

warmer parts of the Palaearctic and part of the Oriental Region. The ten spe-

cies occurring outside the Afrotropics (including SW. Arabia) belong to three spe-

cies groups. Their distributions are as follows:

phlomidis group. Three species from SE. Europe to C. Asia, one species, viz.,

doris, from Kenya through the Middle East to Pakistan, and isolated in Morocco.

sertorius group. Two species in the Palaearctic, viz., sertorius and orbifer, form-

ing a superspecies; one species in the Oriental Region, viz., galba, forming a super-

126 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

species with the fourth member of the species group, mafa, which occurs from S.

Africa through E. Africa to E. Arabia (Oman).

delagoae group. Six species from S. Africa to Kenya; one of these, zebra, in E.

Africa and Pakistan, and another, mangana, extending from E. Africa into

Arabia as far east as Ras Fartak, halfway along the outher coast.

The pattern that emerges from these distribution data shows a contact be-

tween the Afrotropics and the areas outside through Arabia and not via West

Africa. The Arabian connection is still more obvious, if we consider that three more

species (spio, diomus, and colotes) extend from Africa into SW. Arabia, though

without crossing the Arabian desert belt. This pattern agrees well with what is

found in other butterflies (De Jong, 1976), but it does not explain why there is

no connection between the Afrotropics and the Palaearctic in West Africa. In prin-

ciple there are two possible explanations for the absence of a West African con-

nection in Spialia, viz., (1) Spialia species once crossed the Western Sahara, but

became extinct in Northwest Africa because of adverse climatic conditions, and (2)

Spialia species never crossed the Western Sahara. The following arguments are

in favour of the latter explanation:

1. In the West African open formations only three Spialia species occur, viz.,

spio, diomus and dromus. These species also occur in East Africa, where nine

more species occur. So if only the number of species is considered, the chance of a

West African connection is smaller than that of an Arabian connection.

2. The Spialia species that occur in West Africa have a very wide distribution

in Africa, indicating that they are not very dainty in the choice of their habitats.

Nevertheless, there are no traces of a crossing of the eastern part of the Saharo-

Arabian desert belt by these species. If they were not able to traverse the desert

belt in the east, where many other species succeeded, why should they have succeeded

in the west?

3. There is no differentiation between the populations of spio, diomus, and

dromus in West and East Africa, though there have existed important geographic

barriers between West and East African open formations (see below). It means

that the species came to West Africa only recently and too late to make use of the

opportunities provided by the humid Postglacial Climatic Optimum to cross the

Western Sahara, or that they are so eclectic that the barriers mentioned are no

real barriers to them. In the latter case, however, we should expect that they

made use of the opportunities to cross the Saharo-Arabian desert belt in the east

(see also under Speciation patterns).

It may be thought that the occurrence of two other Spialia species, viz., serto-

rius and doris, in Northwest Africa made it impossible to spio, diomus and dro-

mus to establish themselves in that area, but as the food plants of sertorius and

doris are quite different from those of the Afrotropical Spialia species, this is

very improbable.

As said above nine of the twelve Spialia species in East Africa, and more pre-

cisely in Uganda and Kenya, do not occur northwest of that area. This can only

be understood by supposing the existence of an effective barrier between the open

formations in East Africa and north of the belt of lowland evergreen forest. It

DE JONG: The genus Spialia 127

must have been so effective that only three species succeeded in crossing it and so

eventually reached West Africa, apparently in a recent time (see above). If we con-

sider that the lowland forest extends as far east as West Uganda, and may have

extended to West Kenya in rather recent times, it seems most plausible to look

for the barrier in the area between the lowland forest and the Ethiopian high-

lands, the area that connects the East African plains with the open formations

north of the forest belt. Here we find at the present time a very extensive swamp

area, the Sudd region (Sudd is an Arabic word meaning “‘blockage’’, and relates

to the difficult passage for boats), extending west and east of the White Nile in

Southern Sudan and covering about 40.000 km? of permanent swamp, while about

twice that area is flooded by seasonal inundation (Beadle, 1974; Rzoska, 1976). It

is through broad water courses and lakes (Lake Albert, Lake Kioga) connected

to Lake Victoria and the lowland forest belt. Such an area must be an effective

barrier to the drought-loving Spialia species. Moreover, at least during the last

40.000 years the level of Lake Turkana (= Lake Rudolf) in North Kenya has

changed repeatedly and has been up to 80 m above the present level, causing an

overflow to the northwest, to the present Lotigipi Swamp (Livingstone, 1976). This

swamp may have drained into the White Nile by way of the present Kenamuke

Swamp and Pibor River, thus much extending the barrier to the Spialia species

at times.

The age of the swampy region is, however, uncertain. According to Livingstone

(1976: 27), “the deposits of the existing “Sudd”, the Upper Nile Swamps, though

presumably dating only from times since the end of the last interpluvial, deserve

careful stratigraphic investigation.” Moreover, at present a zone of deciduous sa-

vanna woodland extends to the southwest of the flood region (Rzoska & Wick-

ens, 1976). Although a more humid period could easily have led to a connection

between the lowland forest and the swamps, it is difficult to understand how con-

ditions drier than at present, or even the present condition can have brought about

the development of a barrier to expansion to the northwest. So for the time being

we must be content with the presumption that the swamps of Southern Sudan, in

combination with the Ethiopian highlands, the lowland forest belt, and unknown

factors have acted as a barrier to Spialia species for such a long time, that the

evolution of the genus took mainly place in East and South Africa. It is interest-

ing to note that in the area concerned (i.e. Southern Sudan) overlap occurs of

the faunas of the Sudanese and Somali zones (Carcasson, 1964). It implies that

the area acted as a barrier to other animals as well, but it does not solve the

problem of what exactly formed the barrier.

Speciation patterns

Starting from a speciation sequence as described in Chapter 4, a distinct geo-

graphic pattern of speciation arises from the north—south vicariance of sister groups

at various taxonomic levels. The following vicariances can be distinguished:

1. at the species group level: phlomidis group north of the Saharo—Arabian

desert belt (with slight penetration into the Afrotropics by doris), spio group

south of the desert belt;

128 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

2. at the species level: asterodia + agylla in southern Africa, kituina in East

Africa; mafa + galba south and east of the Saharo-Arabian desert belt, ser-

torius + orbifer north of it; mafa in Africa, galba in the Orientalis; wrefordi

in East Africa, its sister group further south (with secondary expansion to the

north); paula in southern Africa, its sister group further north (with secondary

expansion to the south); mangana in East Africa and SW. Arabia, its sister group

in southern Africa (with secondary expansion to the north); nanus in South

Africa, delagoae further north; sataspes in South Africa, zebra in East Africa

and Pakistan;

3. at the subspecies level: diomus ferax in southern Africa, diomus diomus

further north; colotes transvaaliae from South to East Africa, colotes semicon-

fluens in Northeast Africa; confusa confusa from southern Africa to Tan-

zania, confusa obscura in N. Tanzania and Kenya; mafa mafa from South to Cen-

tral Africa, mafa higginsi in East Africa; depauperata australis in southern Afri-

ca, depauperata depauperata in East Africa.

This pattern suggests a repeated fragmentation of the open formations, and

agrees with the current idea of speciation by geographic isolation (allopatric

speciation). It is only remarkable that the pattern is still so well discernible, though

the isolation of the fragments may have been initiated long ago. However, as long

as our knowledge of the exact ecological requirements of the species is so frag-

mentary, there is little use in speculating on the reasons for particular distributions,

and we can only speak in general terms. It is, moreover, plausible that during

Speciation ecological requirements change in the same way as morphological

characters, so that the barrier to range extension can be different before and

after the speciation has taken place. Only two kinds of barriers will have been

effective for all Spialia species (except for the forest species ploetzi), viz., desert

and dense forest.

The Afrotropical Region

As no past or present east—west extension of desert conditions across the whole

African continent south of the Sahara is known, a major factor in the disruption

of the distribution of Spialia species in the Afrotropics must have been the exten-

sion of forests across the continent during wetter and/or colder periods in the

Pleistocene (Carcasson, 1964; Moreau, 1966; van Zinderen Bakker, 1967). If only

the forest formed the barrier, the subsequent retreat of the forest must have al-

lowed range extensions of the Spialia species, obscuring the area of the pre-

vious isolation. We can imagine that in this way the following sister group origi-

nated: spio—diomus, colotes—confusa, and secessus—dromus, and possibly also

colotes group—sertorius group, all broadly sympatric because the barriers to their

distribution are only or mainly dense forest and desert. The restricted range of

some other species, however, indicate that not only dense forest, but also some

kinds of open formations act as a barrier. At least in East Africa the species con-

cerned are restricted to dry areas (kituina, wrefordi, mangana, and zebra),

while their sister groups are, or presumably have been, confined to grasslands in

southern Africa. If we suppose that the ancestors of these groups of sister species

DE Jong: The genus Spialia 129

had similar (though probably not identical) ecological requirements, the existence

in the past of a corridor of dry savanna from East to South Africa must be pre-

sumed. Such a condition can have existed during dry periods. Hamilton (1974) sup-

posed that a connection was possible for the last time 25.000 — 12.000 BP. In

view of the further differentiation of the southern sister groups, especially those

of mangana and wrefordi, dry conditions enabling the north—south connection

must have also existed much earlier.

It must be noted here that the disjunction concerned shows some resemblance to

the well-known Somalia-SW. Africa disjunctions of arid flora and fauna. Indeed,

at least two species, viz. mangana and wrefordi, are confined to the dry

thornbush habitat. However, the sister groups of the northern species are not res-

tricted to or do not at all occur in the arid SW. African (Namibia) habitat. Thus,

the ancestors of the pairs of sister groups showing north—south vicariance may

have become adapted to a richer vegetation than found in the arid Somali zone,

with greater possibilities for north—south connections. In that case the present ha-

bitat preference of wrefordi and mangana would be a secondary development.

In favour of this idea is also the supposed further development of their respective

sister groups. In both cases the sister groups not only differentiated, but also

extended northwards, eventually reaching the less differentiated wrefordi and

mangana. Possibly, the northern species could not keep up the competition with the

new arrivals and were chased away into less hospitable land.

There are only five Spialia species that show subspecific variation in Subsaharan

Africa. In three of them, viz., diomus, mafa, and depauperata, the zone of con-

tact between the northern and southern subspecies is situated somewhere in Zambia-

Malawi. This pattern may indicate a common history of the species concerned,

which was influenced by the extension of lowland forest to the east and/or the

extension of montane biota in East Africa during the late Pleistocene (cf. Moreau,

1966). For mafa, however, it is not likely that montane biota acted as a barrier,

as this species is well capable of living in the present montane belt in Kenya.

The fact that there is a clear north—south vicariance of subspecies in diomus

that does not occur in the two other wide-ranging Spialia species, spio and dromus,

is a reflection of differences in habitat preference: diomus prefers more open and

drier conditions than the other species and is for that reason more liable to range

fragmentation during humid periods. It is therefore remarkable, that not only spio

and dromus, but also diomus do not show subspecific differences between the

populations of West and East Africa, though the swampy Sudd region in South-

ern Sudan must have acted as a barrier during humid periods. Moreover, further

west another, probably still more effective, barrier existed during humid periods,

viz., a much extended Lake Chad (Megachad), at its greatest extension stretching

from the lowland forest to the Tibesti Mountains in the Central Sahara. The last

expansion of the lake occurred no longer than 5.400 years ago (Beadle, 1974).

The absence of traces of the isolating effect of these barriers in the populations

of diomus in West and East Africa can only be understood by supposing that

diomus entered West Africa only after the blockade caused by these barriers was

removed.

130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

A special problem is posed by the origin of the forest species Spialia ploetzi, as

it is difficult to understand the advantage of an adaptation to a completely differ-

ent habitat, except for reasons of competition. Unfortunately, the problem of

change of habitat preference from savanna to forest (and vice versa) has not yet

attained attention from the evolutionary point of view, thougn it must have acted

a part in many more lepidopterous genera: of the 63 larger genera of African but-

terflies listed by Carcasson (1964), 33 have representatives in the forest as well as

in open formations. For the time being we must be content with signalizing the

problem.

Another point is the subspecific differentiation of ploetzi. The boundary between

the two subspecies is at or close to the Cameroun—Nigerian border, apparently west

of the Cameroun highlands. A similar situation is found in the Satyridae Bicyc-

lus ignobilis (Butler) and B. sanaos (Hewitson) (Condamin, 1973) and will

undoubtedly occur in many other Lepidoptera as well. The origin of this type of

differentiation is uncertain and may be complex. The Cameroun highlands,

though not acting as a barrier now, may have done so during a colder period, but

that does not explain why the highlands are now populated by the eastern form.

On the other hand, a dry period would have reduced and fragmented the West

African forest area, so that there was no forest left between the River Niger and

the Cameroun highlands (see, e.g., Booth, 1958). According to van Zinderen Bak-

ker (1967) a dry, as well as cold period in West Africa was at least partly coeval

with the Last Wiirm Maximum. It is therefore plausible to suppose that the sub-

specific differentiation of ploetzi dates from that time. We shall leave the spe-

culation on the history of this species at that, as it falls outside the scope of the

general history of Spialia.

The Saharo-Arabian desert zone

The Saharo—Arabian belt of desert has exercised much influence on the history

and evolution of Spialia. According to Moreau (1952, 1966), the climate of the

Suez and Red Sea areas and Arabia seems to have been similar to that of the

present day since post-Miocene times, with a somewhat more abundant vegetation

during humid periods in mountainous areas. I have not found observations on how

far the Nile Valley can have acted as a corridor for north—south exchange since

the Miocene, but as the Red Sea area was probably never better than semi-

arid, the Nile Valley may never have been more than a relatively narrow corridor

of riverine forest with a belt of wooded savanna passing into semidesert with in-

creasing distance from the river. Consequently, during the period we may suppose

Spialia to have evolved to the present situation, the contact between the African

and Eurasian faunas of open formations more humid than semidesert was repeat-

edly broken and re-established, and there are no indications of a connection over

a broad front. A broad front connection existed in the Western Sahara up to

about 5.000 years ago, but apparently Spialia did not play a part in it (see

above).

Leaving the Western Sahara out of consideration, three corridors leading out of

and into the Afrotropics were open during humid periods:

DE JONG: The genus Spialia 131

1. Nile Valley and mountains west of the Red Sea. For an interesting discus-

sion of climatic and faunistic changes, see Rzoska e.a. (1976).

2. Mountains east of the Red Sea. Higher and more contiguous than the

mountains west of the Red Sea. For the relative importance of this corridor, see

De Jong (1976).

3. Elevated southern rim of the Arabian Peninsula. According to Moreau (1966)

this area may have had a savanna-like vegetation during humid periods. The im-

portance of this corridor is illustrated by the fact that almost half of the species

of Rhopalocera occurring in Eastern Oman have an Afrotropical or Indo-Afro-

tropical distribution (Larsen, 1977). The general lowering of the sea level during

the glacial periods must have drained the shallow Persian Gulf, allowing a faunal

exchange over land between the Afrotropical and Oriental Regions.

The contact across the eastern part of the Saharo—Arabian desert zone is illus-

trated by the following pairs of vicarious taxa: spio group — phlomidis group,

superspecies mafa — superspecies sertorius, mafa — galba; and by the occur-

rence of two species, viz., zebra and doris, north as well as south of the desert

belt. The divergence of the degree of differentiation after severing of the contact,

ranging from hardly any differentiation to species group level, suggests that the

desert belt was crossed several times, in agreement with what is found in other

butterflies (De Jong, 1976).

In view of the present distribution of the relevant taxa, the corridors were equal-

ly important. The first two corridors were used by the ancestor of the serto-

rius group, probably by the ancestor of the spio + phlomidis group, and by

doris and mafa, while the third corridor was not only used by zebra and the

ancestor of superspecies mafa, but to some extent also by mafa and mangana.

It is interesting to note that the present distribution of doris and mafa clearly

shows that these species used the corridors east as well as west of the Red Sea.

As mafa availed itself also of the opportunity given by the third corridor, it can

be called the most expansive Spialia species. The ancestors of superspecies mafa

and of the entire sertorius group were apparently also expansive, so that one won-

ders whether such a trait could be hereditary.

As mafa and mangana have not completed the crossing of the desert belt,

the direction of their expansion is easily detectable: apparently they came from

Africa. For the other groups concerned the direction of spread cannot be inferred

directly from the present distribution and some reasoning is needed to find plaus-

ible solutions, as follows. The sister group of zebra, and all successive sister

groups when going down along the phylogenetic tree of the delagoae group (fig.

119) are African. If we do not conclude from this that the occurrence of zebra

in Pakistan is due to an invasion from Africa, we are forced to assume that almost

all species of the delagoae group originated from invasions from the east, suc-

cessively mangana, nanus + delagoae, depauperata, and sataspes, while a fifth

invasion brought zebra to Africa. This is less plausible than the assumption of an

African origin of zebra. Reasoning along the same lines leads us to the conclu-

sion that doris (phlomidis group) is a Palaearctic intruder in Africa, while galba

and superspecies sertorius (both sertorius group) originate from a northward

132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

traverse of the desert belt (or rather from two African intrusions, the first bring-

ing the ancestor of the sertorius group to the Palaearctic, where it differentiated

into the ancestor of superspecies sertorius). For the further geographic and evolu-

tionary history of the phlomidis and sertorius groups the reader is referred to

De Jong (1974).

The last pair of vicarious taxa (spio group — phlomidis group) can also

be supposed to have an African origin, for if the origin was Palaearctic, at least

one more crossing of the desert belt would be needed to explain the cleavage of

the spio + phlomidis group and its sister group (fig. 117). Moreover, starting

from the assumption that the original food plants of Spialia belong to the Mal-

vales (see Chapter 4), the change to Convolvulaceae in the phlomidis group, being

an adaptation to Palaearctic conditions, would have been reversed in the spio

group, if the latter originated from a Palaearctic intrusion. This is not a plausible

explanation of the present situation. Also when applying the rule of parallelism

of chorological and morphological progression, the phlomidis group, having more

apomorphous characters than the spio group, must be considered the chorologi-

cally most progressed group. I quite agree with Hennig (1960), that the appli-

cability of this rule is limited, but at least it does not contradict the other argu-

ments.

Humid periods that opened the corridors occurred more than once in the past,

for the last time during the Postglacial Climatic Optimum, which ended about

5.000 years ago. From this last opportunity may date the occurrence of mafa in

Oman. Though mainly living in the highland grasslands in East Africa, the

occurrence of mafa as far north as Erkowit in Sudan and Ashaira in Saudi-

Arabia, and as far east as Oman, where it lives in an Acacia desert landscape,

indicates that a relatively small climatic change may enable the species to avail

itself of the corridors. For doris, which can live in wadi’s as far as its food plant

(Convolvulus lanatus) occurs, a still slighter increase of the humidity was needed

to open the corridors. One may even wonder how far the present conditions prevent

doris from crossing the desert belt.

In view of the differentiation of galba, superspecies sertorius and the phlomidis

group, their ancestors must have left Africa during one or more humid periods be-

fore the Last Glacial Period. Supposing that speciation in superspecies sertorius

and the phlomidis group took place under the influence of the glaciations in the

Western Palaearctic by way of isolations in refugial areas, both taxa must have

experienced at least two or three glaciations (see De Jong, 1974). It cannot be

stated at the moment whether the three glacial phases of the Last Glacial (Wiirm)

were enough for this development, but together with the two glacial phases of the

preceding Riss glacial period (see, e.g., De Lattin, 1967; Moreau, 1955; West,

1968), they certainly were, so that the occurrence of Spialia in the Palaearctic

does not need to date further back than some 200.000 years.

Contrary to the ancestor of superspecies sertorius and probably the ancestor

of the phlomidis group, the ancestor of galba apparently used the eastern route,

across the Persian Gulf, as did zebra. The best opportunities were provided

during glacial periods, when the Persian Gulf was drained. Therefore, the occur-

DE JONG: The genus Spialia 133

rence of these species in India and Pakistan, respectively, is probably not of post-

glacial age. The wide distribution and differentiation of galba suggests that its

ancestor arrived in Asia much earlier than zebra, as said above.

6. GENERAL OUTLINE OF THE EVOLUTION OF SPIALIA

In Chapters 4 and 5 the phylogeny and geographic history of Spialia have

been discussed. In the present chapter we shall combine the conclusions into a gen-

eral outline of the evolution of the genus.

The origin of the genus could not be discussed in the foregoing chapters, as that

can only be done by searching for sister groups (according to the method of Hennig,

1966) and, for that reason, needs a comparison with other genera. In advance of

a future revision of the genera of the Pyrgus group, it may suffice here to

state that the ancestor of Spialia probably came from the steppes of Central

and Western Asia. When and how it arrived in Africa is unknown. Its subsequent

development and differentiation took place mainly in Africa. It indicates that there

was a rather effective barrier to northward expansions from the very beginning of

the genus Spialia. Apparently this barrier was the main factor underlying the

geographic isolation that led to the origin of the genus. The only possible barrier

that comes into consideration is the Saharo-Arabian arid zone. While, however,

the main differentiation of Spialia took place in the savannas of Eastern and

Southern Africa, some other barrier or a combination of barriers must have pre-

vented Spialia from occupying the savannas north of the region of the low-

land evergreen forest up to a rather recent time. A possible barrier was (and is) the

swampy condition of the Upper Nile region (Sudd), but other, unknown, factors

must have played a part as well.

The differentiation led in the first place tot the development of three types of

male genitalia, especially with regard to the development of the costa of the valva,

coinciding with three types of female genitalia, which are, however, less well defi-

ned. The three types are represented in the present taxa by the asterodia

group, the spio + phlomidis group, and the remainder of the genus, respecti-

vely. Each type is distributed at least in Southern and Eastern Africa, obscuring

the geographic isolation underlying the origin of the differentiation. The early de-

velopment of the genus took place in a savanna habitat under not too dry condi-

tions. The present occurrence of some Spialia species in dry savanna or semi-

desert appears a rather recent development, as undoubtedly is the adaptation of one

species (ploetzi) to the lowland evergreen forest. The preference for open forma-

tions made the ancestors of the present Spialia species liable to fragmentation

of their ranges by expansion of the forest biota across the African continent. As

the occurrence of such expansions can be considered well established and similar

expansions of the arid biota are not known, the differentiation of Spialia must,

in the first place, be attributed to pluvial and/or cold periods that enabled a large-

scale extension of the lowland evergreen forest or the montane forest.

The expansions of the forest resulted mainly in the open formations being divi-

ded into northern and southern parts. This pattern is still visible in the vicariance

134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

of a number of apparent sister groups in Spialia, which have been listed in

Chapter 5. The fact that the vicariances occur at different taxonomic levels, from

subspecies to species group, and the fact that the present distributions of the vica-

rious groups point to barriers at various places, indicate that the fragmentation of

the open formations has occurred repeatedly.

Although periods with large-scale extension of forests are known to have occur-

red during the Pleistocene, data on the Tertiary climate are very uncertain. The

highest number of successive cleavages needed to understand the present differen-

tiation, is eight (viz., in the line leading from the ancestor of Spialia to the sis-

ter groups zebra and sataspes). It implies that the development of Spialia

may mainly, if not entirely, have taken place in the Pleistocene and there is cer-

tainly no reason to look for the origin of Spialia in a period prior to the Plio-

cene.

The barriers that kept Spialia in Eastern and Southern Africa remained

rather effective. The correctness of the assumption, that the arid condition of the

Saharo-Arabian desert belt was a main factor inhibiting northward range exten-

sion, is indicated by the fact that only species capable of living under dry condi-

tions succeeded in escaping the boundaries of the Afrotropics. Opportunities for

such escapes were provided several times during the Pleistocene by increased humi-

dity, for the last time only 7.000—5.000 BP (Postglacial Climatic Optimum). With

regard to the opportunities there is no reason to suppose that the occurrence of

Spialia species outside the Afrotropics dates back from a time prior to the

Pleistocene.

The northward expansions took place after the initial three types had differen-

tiated into the ancestors of the present species groups (the differentiation took mainly

place in one of the types), a further reason to suppose that only in rather recent

times, i.e. in Late Pleistocene, the Palaearctic and Oriental Regions were invaded.

If there were earlier invasions, they have left no traces. For an explanation of the

differentiation outside Africa it is not necessary to assume that the relevant inva-

sions occurred earlier than the last or at most the one-but-last interglacial period,

as the climatic changes since then were sufficient to be responsible for the deve-

lopment of the present differentiation. Consequently there are three kinds of evi-

dence for a geologically spoken, recent occurrence of Spialia outside Africa,

viz., (1) the advanced state of the differentiation in Africa, (2) the opportunities

given, and (3) the differentiation outside Africa.

It is remarkable that the establishment of Spialia in the Palaearctic was ac-

companied by a change of food plants, from Malvaceae or related families to Rosa-

ceae and Convolvulaceae. This may be due to competition. The Malvaceae are

supposed to be the original food plants of the Pyrgus group. As the ancestor

of Spialia most probably came from the Palaearctic, a return to the Palaearctic

can be supposed to have caused competition, unless the food plants were changed.

In this connection the common and wide-spread Palaearctic Carcharodus alceae

(Esper), a multivoltine species of the Pyrgus group and living on various Mal-

vaceae, comes in mind, but we shall not speculate here further on this subject, lea-

ving it for a future discussion on the history of the Pyrgus group. Only one more

DE JONG: The genus Spialia 135

remark about food plants. The faunal exchange between Africa and the areas out-

side was largely unidirectional as far as Spialia is concerned, viz., out of Africa.

Only one successful re-invasion of Africa took place, viz., of doris. The success

of the establishment of doris in Africa south of the desert belt may be due, apart

from its preference for dry conditions, to the food plant (Convolvulus) being en-

tirely different from the food plants of African Spialia species.

The connection between Africa and the areas outside was established only by

way of the Nile Valley, the Red Sea area and the southern rim of the Arabian

Peninsula. This is the more remarkable as the eastern part of the Saharo-Arabian

desert belt is supposed to have been always drier than the western part, which was

covered by a mediterranean type of vegetation only 7.000—5.000 years BP. The

absence of a western connection is probably due to the late arrival of Spialia in

the West African savannas, as indicated by the absence of differentiation between

the populations in East and West Africa of the three species (spio, diomus, and

dromus) that occur in West Africa, while one of these species (diomus) does

show subspecific variation in Subsaharan Africa, and by the number of cleavages

of the evolutionary line since the beginning of the genus, needed for dromus (se-

ven), showing that this species itself is young and thus its occurrence in West Afri-

ca cannot be old. The main factor that kept Spialia so long out of West Africa,

probably was the swampy condition of the Upper Nile region, connecting the

lowland evergreen forest and the Ethiopian highlands, but other, unknown factors

must also have played a part.

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Woodhouse, L. G. O., 1950. The butterfly fauna of Ceylon. 2nd. ed. — Ceylon Government Press,

Colombo.

Wynter-Blyth, M. A., 1957. Butterflies of the Indian Region. — The Bombay Natural History Society,

Bombay.

Zimsen, E., 1964. The type material of J. C. Fabricius. — Munksgaard, Copenhagen.

Zinderen Bakker, E. M. van, 1967. Upper Pleistocene and Holocene stratigraphy and ecology on the

basis of vegetation changes in Subsaharan Africa. In: W. W. Bishop & J. Desmond Clark (ed.),

Background to evolution in Africa: 125—146. — Univ. of Chicago Press, Chicago and London.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978 139

INDEX

Generic names have been added to specific names not belonging in Spialia. Page numbers in bold type

refer to descriptions.

abscondita 49

agylla 28, 34, 36, 38, 106, 108, 110, 111,

113, 114, 117, 128

alceae (Carcharodus) 114, 134

Alenia 106, 108, 109, 114, 117

amenophis 53

antonia (Syrichtus) 107

asterodia 28, 34, 36, 38, 106, 108, 110, 111,

113, 114, 117, 128

aurivillii 65

australis 105, 128

bamptoni 38

bettoni 38

Bicyclus 130

bifida 98

Carcharodus 106, 113, 114, 115, 117

carnea 72

Celaenorrhinus 110

Celotes 106, 109, 110, 114

chenga 69

colotes 29, 54, 58, 59, 107, 108, 111, 112,

114, 115, 120, 126

confusa 29, 40, 55, 59, 107, 108, 111, 112,

114, 115, 120, 128

cribrellum (Syrichtus) 107, 110

delagoae 94,99, 110, 111, 112, 114, 117,

122, 128, 131

depauperata 31,49, 96, 99, 102, 110, 111,

112, 114, 117, 122, 128, 131

diomus 31, 42, 43, 45, 102, 107, 113, 115, 118,

126, 128, 135

doris 29, 52, 53, 62, 66, 111, 112, 114, 115,

125, 126, 127, 131, 132

dromus 32, 42, 43, 80, 111, 112, 114, 115,

120, 126, 135

elongata 80

Erynnis 114

evanida 54,66

fasciata 60

ferax 48, 49, 102, 128

galba 24, 32, 66, 110, 111, 114, 115, 120,

125, 128, 131, 132

gecko 52

geron 29,52,111,112,113,114,115

Gomalia 106, 109

Heliopetes 106, 107, 110

hellas 98

hermona 52

higginsi 64, 128

ignobilis (Bicyclus) 130

kituina 28, 38, 106, 107, 108, 110, 111, 113,

114, 117, 128

lacreuzei 49

lavatherae (Carcharodus) 115

leucomelas 80

Lobocla 114

machacoana 46

mafa 29,52, 62, 110, 111, 114, 115, 118,

120, 126, 128, 131, 132

malvae (Pyrgus) 110

mangana 31, 89, 94, 99, 110, 111, 112, 114,

117, 123, 126, 128, 131

melaleuca 80

meridionalis 80

minuscula 68

nanus 31,91,94,99, 110, 111, 112, 114,

117, 122, 128, 131

nora 55

oberthüri 65

obscura 59, 128

occidentalis 31,87

orbifer 71, 111, 114, 115, 120, 125, 128

osthelderi 29,52, 111, 112, 113, 114, 115

paula 29, 62, 74, 111, 112, 114, 115, 118,

120, 128

phlomidis 30,52, 107, 111, 112, 113, 115

ploetzi 31,84, 109, 111, 112, 114, 115,

120, 125

poggei (Syrichtus) 107

Pyrgus 25, 106, 107, 108, 110, 111

rebeli 85

rehfousi 105

sanaos (Bicyclus) 130

sataspes 31,99, 110, 111, 112, 114, 117,

122, 128, 131

secessus 32, 76,111, 112, 114, 115, 120

semiconfluens 55, 56, 58, 59, 128

sertorius 29, 71,111,114, 115, 120, 125,128

shanta 69

spio 30, 41, 52, 62, 107, 110, 111, 113, 115,

118, 126, 135

struvei 52

superna 68

Syrichtus 106, 108, 110, 111, 113, 114,

115,117

therapne 71

transvaaliae 55, 56, 58, 128

trimeni 77

vindex 41

wrefordi 31, 73, 111, 112, 114, 115, 120, 128

Xenophanes 114

zaira 80

zebra 31,38, 85, 96, 110, 111, 112, 114,

117, 122, 126, 128, 131, 132

140 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Plate 1.

Figs. 1—8. Upper (left) and underside (right) of Spialia species. 1—2, S. asterodia (Natal). 3—4, S.

agylla (Bloemfontein, S. Africa). 5—6, S. kituina (Ukunda, Kenya). 7—8, S. spio (Naro Moru, Kenya).

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978 141

Plate 2.

Figs. 9—16. Upper (left) and underside (right) of Spialia species. 9—10, S. diomus diomus (Ukunda,

Kenya). 11—12, S. diomus ferax (Sheldon, S. Africa). 13—14, S. colotes semiconfluens (Bihendula,

Somalia). 15—16, S. confusa obscura (Arabuko Forest, Kenya).

142 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Plate 3.

Figs. 17—24. Upper (left) and underside (right) of Spialia species. 17—18, S. phlomidis hermona

(Mt. Hermon). 19—20, S. osthelderi gecko (Kopet Dagh, Turkmenia). 21—22, S. geron struvei

(Kotgai, E. Afghanistan). 23—24, S. doris evanida (Karachi, Pakistan).

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978 143

Plate 4.

Figs. 25—-32. Upper (left) and underside (right) of Spialia species. 25—26, S. mafa higginsi (Naro

Moru, Kenya). 27—28, S. galba galba (Haldumulla, Ceylon). 29—30, S. sertorius sertorius (Switzer-

land). 31—32, S. orbifer orbifer (Sarajevo, Jugoslavia).

144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Plate 5.

Figs. 33—40. Upper (left) and underside (right) of Spialia species. 33—34, S. wrefordi (Samburu

N. P., Kenya). 35—36, S. paula (Essexvale, Rhodesia). 37—38, S. secessus f. trimeni (Mpanda,

Tanzania). 39— 40, S. dromus (Mpanda, Tanzania).

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978 145

Plate 6.

Figs. 41—48. Upper (left) and underside (right) of Spialia species. 41—42, S. ploetzi ploetzi (Bitje,

Cameroun). 43—44, S. ploetzi occidentalis (Warri, Nigeria). 45—46, S. mangana (Dire Daoua,

Ethiopia). 47—48, S. nanus (Sheldon, S. Africa).

146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978

Plate 7.

Figs. 49—56. Upper (left) and underside (right) of Spialia species. 49—50, S. delagoae (Muden,

Natal). 51—52, S. zebra bifida (Ukunda, Kenya). 53—54, S. sataspes (King Williamstown, S. Afri-

ca). 55—56, S. depauperata depauperata (Mlanje, Malawi).

PAR

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MUS. COMP. ZOOL. 1978

| DEEL 121 AFLEVERING 4 LIBRARY

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DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

J. KIELLAND. — A provisional checklist of the Rhopalocera of the eastern side of

Lake Tanganyika, p. 147—237, text-figs. 1—36, pls. 1—14.

INHOUD

i | Tijdschrift voor Entomologie, deel 121, afl. 4 Gepubliceerd 31-XII-1978 |

A PROVISIONAL CHECKLIST OF THE RHOPALOCERA

THE EASTERN SIDE OF LAKE TANGANYIKA

JAN KIELLAND

4916 Boroy, Norway

With 36 text-figures and 14 plates

INTRODUCTION

The following list is in the first place based on collections made by the author, and

mainly concerns the districts of Kigoma, Mpanda and Ufipa. However, in order to

make the data of butterflies known from this area more complete, it includes as many

records as possible, made by other collectors. These are as follows: The Kyoto Univer-

sity African Primate Expedition, that collected for several years at Mukuyu and

Mihumu about 60 to 80 km to the south of Kigoma; Miss Goodall, who collected at

Gombe Stream north of Kigoma; T.H.E. Jackson’s African collector, Watulege, who

collected in some places around Mpanda and Kigoma, including the Mahale mountain

range; and finally Major Graham’s African collector, who trapped Charaxes in the

Mukuyu area.

In the present paper 733 species of butterflies have been listed, but more thorough

collecting will probably produce many more. I would estimate the total number of

species existing in this area at between 800 and 850. About 170 are new records for

Tanzania and probably over 30 of them are new for East Africa.

In order to study the genitalia, specimens have been dissected of the majority of the

more difficult groups, such as Syntarucus, Axiocerses, the Teriomima group, Lach-

nochnema, the Iolaus group, Oboronia, Anthene, Uranothauma, Cacyreus and others

of the Lycaenidae, Neptis (Nymphalidae) and Ypthima and Neocoenyra (Satyridae) and

several groups of the Hesperiidae and Pieridae. Much help with the identification has

earlier been received from Dr. R. H. Carcasson; some Lycaenidae have been checked

by Dr. H. Stempffer and Dr. G. E. Tite. Dr. M. Condamin has checked most of the

material of the genus Bicyclus (Satyridae). Many of the Charaxes have been examined

by the late Dr. van Someren.

147

148 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

For the nomenclature, Wallace Peters’ checklist (1952) has been followed to a great

extent, but whenever appropriate, newer works have been consulted.

DESCRIPTION OF THE RESPECTIVE AREAS (text-figs. 1 and 2, pls. 1-3)

The region dealt with in the present paper is illustrated in text-figs. 1 and 2. Two

main conditions have been taken into account in drawing the boundaries for the

recorded areas namely topography and geology. These are two important factors for

the lepidopterological fauna; in this respect reference can also be made to my two

papers in “Atalanta” (later “Atalanta Norvegica”) of 1968 and 1969.

The area between Malagarazi river (south of Kigoma) and Burundi, is not very well

known to me, and except for Gombe Stream, there are few records of butterflies here.

For that reason I have not taken the trouble to divide the area into numbered zones

and the collecting places from this area will be named in full. The same applies to the

Ufipa district and the very rarely investigated parts of eastern Mpanda.

From the area north of Kigoma, I have only records of part of Goodall’s and of my

own collection at Gombe Stream near Lake Tanganyika and scattered collecting

from the Uwinza-Kibondo road. The Gombe Stream area, with mountains rising

rather steeply up from the lake-shore, contains scattered riverine forests in the valleys

and Brachystegia woodland on the ridges and mountain slopes. The highest ridge,

Mil T

f Kast ====:== Secondary roads

Topographic _

area boundaries

District

boundaries

+4444

"SU

Lake

Tanganyika

0 \Rukwa

Sumbawanga

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Fig. 1. Topographical areas; for explanation, see text.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 149

extending parallel to the lake-shore, is mainly covered by open grassland. There are

some interesting finds of butterflies. The hills further inland are mostly devoid of

woodland and forest, due to the trees being cut down for firewood or on behalf of cul-

tivation. This part, except the Park itself, is much more densely populated than the

country to the south of Kigoma.

Area 1. The Mahale ridge, including Mt. Kungwe, is the highest mountain range

on this side of Lake Tanganyika. Area 1 which comprises this mountainous penin-

sula, is a basement area of uniform gneiss (Mahale gneiss), bordering the complex

basement formations to the east. The Mahale gneiss is very poor in mica and garnet,

while in the other basement areas mica bearing schists with garnets are predominant.

This is a very interesting area with great variation in flora; there is forest in

patches, from the lakeshore, as at Kasoge, to an altitude of more than 2300 m at

Sisaga Mt. and Kungwe Mt.; there is also open montane grassland and mountain

shrubs and woodland in lower altitudes. Numerous rivulets descend from the steep

mountain sides. À wild and magnificent landscape. I have made 14 short safaris to

this mountain, but I am sure that my collections are far from complete. The mountain

area can only be reached on foot, or by boat from the lake.

Area 2a and b. This is low, undulating country situated between Lake Tanganyika

and Mahale on one side and the Wanzizi-Kakungu scarp on the other. Area 2a is the

moister part with more permanent water courses, heavily flanked by riverine forest

and large areas of bamboo. The vegetation of 2b is largely woodland of savanna-like

character. There are some concentrations of bamboo in the moister area close to the

mountain range of 3a and 3b.

Area 3a and b. This is a mountainous area stretching in the Kigoma and Mpanda

district, and including the Kakungu-Mweze-Wanzizi escarpment, and the Lubalizi

and Utinta valleys. Utinta cuts through the highlands at an altitude of 1200-1300 m

and is therefore a natural border between 3a and 3b. Area 3a is the highest part, with

mountain tops of 1800-1900 m, and a large area, viz., Sitwe, Ngondo and the

K arobwa-Kakungu Mts., is covered with montane grassland and solid bamboo with

more or less scattered, stunted, deciduous trees and shrubs. There are numerous small

permanent water courses with rich riverine vegetation. Part of the Mweze Mt. itself

and also Karobwa Mt. are clothed with evergreen montane forest up to the top.

The Mweze-Sitwe grassland enjoys a very rich Lycaenid fauna and here the col-

lecting is very good after the grass is burnt in August to September and perhaps in

October if the rain is late.

Area 3b is a hilly country, most of which is heavily covered with Brachystegia

woodland. Some rare Lycaenids emerge here after the rain has started in October to

November. The only place with open grass, is the very top of the ridge of the Wanzizi

hills at 1550-1600 m. Here are also large patches of forests climbing the hillsides and

bordering the grassland on top.

Area 4a and b. The Luegele valley, situated in 4a, is the more interesting area, with

rather extensive forest remnants. Area 4b, the Katuma valley, is largely dominated

150

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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

9, 25 a È

2022,20 29 VE ©

EN i Be

-Kigëmax7 CE 31

2 0.--

“Mpandaz 33

Lake

Tanganyika

Sumbawanga

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ezine

Fig. 2. Localities.

Gombe Stream National Park 800— 1600 m.

. Muhanga 900 m.

. Mihumu 900—1000 m.

. Mukuyu 800—900 m.

. Kararumpeta Mt. ridge 1500—1600 m.

. Helembe 800 m, Kefu forest 900 m, Kasha

forest 1000 m.

. Kapalamasenga, Mugondozi rivers

900—1000 m.

. Mugombazi river ca. 1000 m.

. Bulimba 800— 1000 m.

. Luegele river ca. 1100 m, Luntampa, Kapapa

1100—1200 m.

. Ntakatta forest 1200— 1500 m.

. Logoza-Mgambo 800 m.

. Kasoje (K asoge) forest 800— 1000 m.

. Lubaliziriver 1100— 1300 m.

. Karobwa-Kakungu Mts. 1700— 1900 m.

. Kasja 1100— 1200 m.

. Ujamba 2000 m, Mahale ridge 2000— 2500 m.

. Lukandamira 1600— 1800 m.

. Masaba ca. 1200 m.

. Kibweza-Karia 800 m.

. Ugaraba, Yampehu ca. 1100 m.

. Kabakaranga 900 m.

. Ngondo 1700— 1800 m.

. Mweze Mt. 1900 m, Sitwe 1600— 1800 m.

25:

26.

Als

28.

29:

30.

31.

32.

33:

34.

35:

36.

Sil

38.

39.

40.

41.

42.

43.

44.

45.

46.

47.

48.

49.

50.

Mkulya river 1400—1500 m.

Luega river 1200 m.

Lower Luega 800— 1000 m.

Wanzizi hills 1400— 1600 m, Mabwe 1200 m.

Lugalla hills 1200— 1400 m.

Sibwezaca. 1100m.

Nkungwe 1100— 1200 m.

Karema, Sangu 800 and 900 m.

Kansanga 1100— 1200 m.

Mt. Ipumba 1900 m.

Mt. Sitebi 1900— 2000 m.

Kampisa river 1500— 1600 m. Ntobo river,

Upper Katuma river 1600 m.

Katuma village 1100 m.

Iloba 1000 m.

Katuma-Mpanda road ca. 1300 m.

Busondo ca. 1400 m.

Mishamu ca. 1200 m.

Nyamanziriver ca. 1200 m.

Uruwira plateau forest islands ca. 1300 m.

Useqya ca. 1000 m.

Kanindi, Mamba ca. 1000 m.

Rungwa ca. 900 m.

Mbuzi Mt. 2000 m.

Chulwe Mt. ca. 1900 m.

Mbizi forest 2000— 2200 m.

Ikola, 800 m.

- KIELLAND: Rhopalocera eastern side Lake Tanganyika 151

by heavy and open woodland, with riverine forests descending the gullies from the

sandstone ridge. Most of the original forest along the Katuma river was destroyed by

the local population in previous times. At present, most of the area is uninhabited.

Area 5a and b. The greater part of this area is occupied by a highland country situat-

ed inside the sandstone area, at the junction of this and the basic formations to the

south, at 4a and b (only the Kapalamagulu Mt. near the lake is not sandstone but

noritic gabro). The conspicuous feature of this section is the high ridge commencing

close to Lake Tanganyika at Mt. Kapalamagulu, stretching eastwards towards

Mt. Sifuta and Mt. Sitebi (2000 m) at 5b, which is the higest part of the ridge. The

lower stretch, closer to the lake, varies between 1200-1500 m and is mostly covered

with woodland and some forest. Mt. Kapalamagulu, however, is covered with grass-

lands and for the greater part devoid of trees. The gullies and valleys of the eastern

section of 5a are more heavily clothed with riverine forests and the largest of these is

the Ntakatta impenetrable forest and thickets which cuts diagonally through the

mountain range. But the lower part of this forest has been reduced by the local in-

habitants since I first saw it over 20 years ago. The highest part of the sandstone ridge

with bare, grass covered mountain slopes and ridges, is poor in butterflies during the

rains, but ideal collecting ground for Lycaenids after the grass is burnt, in August to

September. So far, the most important collecting ground the writer has found on this

high sandstone area, is the Kampisa valley and Mt. Sitebi. The forests are easily ac-

cessable on foot, as this is far from any human occupation and big game have tram-

pled plenty of paths inside.

Area 6. The northern and western part of area 6 is characterized by Brachystegia

clothed hills passing into savanna and plains further south-east. Here we approach a

dryer region with little or no evergreen trees.

Area 7. This is also a typical woodland and savanna country with low hills merging

with plains which mostly become flooded during the rainy season. It is a hot and arid

place during the dry spell.

Area 8a and b, and 9a and b. This part consists of sandstone formations with

numerous mesas and ridges reaching a height of up to 1600 m. The general direction

of these ridges is NE-SW, approximately parallel to the lake. From the lake-shore at

780 m, the country rises abruptly to a low ridge of about 1000 m; behind the ridge it

drops again to about 900 m, and rises gradually toward the east, broken by many

gullies and ridges. The coastal sections 8a and b are the best known parts of the area.

The most important collecting places are invariably the small remnants of evergreen

forest, which for the greater part are situated along water courses, of which the Lugufu

valley is a good example and probably the most interesting. Close to the coast near

Mukuyu and Helembe, there are also several evergreen forests away from the streams,

covering hills and valleys to the extent of one or two square miles, in patches. These

coastal forest remnants consist of shorter, more stunted trees and more heavily

covered with lianas than is usual further inland. These remnants and the forest relics

on top of many of the mountains are a good indication of a former well developed

primeval forest, covering most of the country to the east of Lake Tanganyika. Their

decline is mainly due to the present drier climate. As the trees are not fire resistant, as

152 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

is the woodland vegetation, they easily perish in the frequent grass fires during the dry

season.

The inhabitants of this part of Tanzania apply shifting cultivation, that is, they

seldom stay in one spot for more than two to five years, then shift to another place

where they again cut down and burn another stretch of forest for cultivation. These

evergreen forests hardly ever grow up again after they have been cut down and aban-

doned. This is first of all due to the yearly grass fires that destroy all regrowth of these

types of trees. The only spots where I have seen partial or complete regrowth of ever-

green forest in the areas of Mpanda, Kigoma and Ufipa, are clearings inside the

K asoge forest at Lake Tanganyika, west of Mt. Kungwe. The humidity here is higher

than elsewhere and grass fires are less frequent.

The coastal evergreen forests of 8a and b become very arid in the dry season and

towards the end of it, in September and October, very few butterflies are seen. But

during the rains and early dry season, June-July to August, they abound.

Lugufu is the largest river south of Malagarasi and its tributaries from Mihumu

upstream are richly bordered with riverine trees. Mihumu is situated at 900 m, sur-

rounded by hills up to 1300 m. These hills are all covered with woodland of the same

type as the rest of area 8a, 8b. A fairly high mountain area, the Kararumpeta range

(1600 m) stretches south-west of Mihumu. This range rises very steeply from the

Wazizi river, which runs to the north on the east side. The western sides slope less

steeply towards the low hills parallel to the lake. There is no, or very little forest

growth along the Wazizi; it was probably destroyed by earlier cultivation, but high

up in the Kararumpeta range, evergreen trees clothe the river gullies. The vegetation

on top is partly grass, partly bamboo and woodland. Where the mountain slope is not

too steep for trees, woodland is dominant.

South of Kararumpeta, in area 8b, stand Mkuyu Mt. (Mkuyu Mt. as indicated on

maps, is not correct; the local name is “Sifumbi”, but I have not found this name on

any map). Its vegetation is poor looking stunted woodland on sandstones and shales.

Further on lies a flat country covered by equally small-growing, scattered, deciduous

trees. This flatland between Mkuyu Mt. and the long sandstone ridge at 5a, 5b lies at

an average altitude of 1100 m and is crossed by small rivers running west towards the

lake. Some of these rivers are bordered by a rather well-developed forest vegetation

with a butterfly fauna that is similar to that of the forests to the north, though some-

what poorer in species.

Area 9a, 9b, east of 8a, 8b west of the Uwinza road, consists of rather high country

of mesas and numerous gullies covered with more or less heavy woodland of a more

robust character than further west. The rainfall is probably higher and the soil not so

poor. The rivers are furnished with some forest, where it has not been destroyed by

man. At present, people are only cultivating some places along the road and the vast

area between this and the lake shore is devoid of roads and tracks and almost com-

pletely uninhabited, mainly due to poor soil, and partly to inaccessibility.

East of the Mpanda-Uwinza road. The country here has been very little explored.

The western part consists mainly of sandstone formations and is hilly with steep

gullies. The only section I have collected in, is along the Mpanda-Tabora road. The

area is completely uninhabited and the only people who travel here outside the road, are

occasional hunters and honey gatherers. Tsetse flies abound and water is scarce,

KIELLAND: Rhopalocera eastern side Lake Tanganyika 153

making this part of the country difficult for collectors.

Southern Mpanda and Rukwa. South of area 7 and east of the Rukwa escarpment

are large sedimentary plains with savanna woodland and open grass, often flooded in

the rainy season. There is no riverine forest as far as I have seen. It is a poor habitat

for butterflies.

The Ufipa Plateau. This plateau culminates in the Liambalamfipa mountain chain

crowning the steep escarpment, dropping down into the Rukwa depression. To the

west, the plateau slowly descends towards Lake Tanganyika. The southern-most part

of Ufipa is lower country dominated by smaller hills clothed with open woodland. In

this southern area I have not collected so far. The high plateau is not so richly pro-

vided with permanent water sources as is the Mahale-Mweze-Sitebi area and there

are only few forest remains. This is most certainly due to the early destruction of

riverine vegetation and forest as a whole by the local inhabitants. Indeed, this is a

situation which threatens the entire area, if the present rate of forest destruction is

allowed to continue. The main forest remains are those at Mbizi Mt., near Sumba-

wanga and Mbuzi Mt. near Chala. The Chala Mt. itself, is sparingly clothed with

riverine forest on top.

The areas in which comparatively thorough collecting has been done, are as follows

(the best known areas first): 6, 3b, 3a, 5a, 5b, 1, 2a, 2b, 4a, 4b. Area 7 is little known,

but has more or less the same ecology as the neighbouring areas 6 and 2b, with hardly

any difference in the fauna. Areas 8 and 9 are both poorly collected, but 8 has recently

been better explored and many new finds for Kigoma have turned up. The area north of

8 and 9 has been little examined by the writer except for the Gombe Stream National

Park north of Kigoma. Ufipa has been investigated sporadically in the Chala and

Table 1. Composition of the butterfly fauna of Western Tanzania and its distribution over the different

areas. Areas 7, 9a and 9b left out for reasons given in the text. U = Ufipa, G = Gombe; D = doubtful

locality. For other explanation, see text.

Area ] 2a 2b 3a 3b 4a 4b 5a 5b 6 8a 8b U G D

Papilionidae 17 15 12 17 13 14 11 16 17 13 16 16 IO. 9

Pieridae 40 31 27 43 38 39 26 39 34 44 33 38 35 27

Danaidae 7 5 3 7 4 7 3 8 6 3 8 7 3 3

Satyridae 31 24 19 29 24 29 19 28 26 22 25 27 14 15

Nymphalidae 119 83 SZ 75 116 52. 01135. Oz OA 122, 123 49 76 l

Acraeidae 36 17 14 28 29 Il 7 34 22 25 36 32 19 14

Libytheidae 1 | 1 1 1 I 1 1 1 I 1

Riodinidae ]

Lycaenidae 98 64 sy II 82 76 De) NOS VIZO MS 90 109 67 54 3

Hesperiidae 81 57 32 84 68 67 22 VE) 91 88 54 78 42 52 2

Total EN 297 202 aS SA SO NOS) AAS ANS) SO ED dI) 239 246 6

154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Sumbawanga area and also to some extent below the escarpment near Lake Rukwa.

The number of species occurring in areas 2b and 4b cannot very well be compared

with those of the other localities, as only sporadic collecting has been done here while

walking through them to get to more interesting places. They are, however, certainly

not so rich as most of the other localities. The distribution of the recorded butterfly

fauna over the different areas is summarized in table 1.

Every species collected in Ufipa, at Gombe and in area 6 have been recorded, but I

may have forgotten to make a note of some species observed in other areas. It is very

difficult to remember each time all the species which has been taken in each locality,

as very numerous safaris have been made.

GENERAL ECOLOGY

A great problem for the continued existence of the fauna of evergreen forests of

western Tanzania is the old habit of the native population of shifting cultivation and

their reluctance to use other habitats. It would be less serious if the forest vegetation

grew up again after the cultivated land has been abandoned, but unfortunately, this

df IN Kasulu

Kigoma N I

KE Ve

ore ee

= ——

LK: +

i = LS Kätavi >

Nr G SE

<Ufipa i

Lake

Tanganyika

To Sa

== Deciduous woodland Evergreen forest

Montane grassland Solid-stemmed bamboo

777

CA marshland and savanna, Straight, hollow bamboo

flooded during the rains

—— Gg 3 I \ Rukwa

YSumbawanda ::: à |

Fig. 3. Map of vegetation types. White areas along the lake shores are open or cultivated areas.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 155

happens very seldom and only in certain circumstances. The only place where I

have seen regrowth of evergreen forest is in the Mahale area, particularly in places

where cleared farm land is surrounded by forest, and fire and elephant grass is pre-

vented from getting in. Also the high humidity of the Mahale area stimulates forest

growth.

It is generally presumed that the grass fires prevent the regrowth of forest vegeta-

tion. However, this is certainly not the only reason. During 20 years of observation

in this area, I have had the opportunity to notice that a cleared forest area hardly ever

returns to its original state. In a very short time it is covered by huge elephant grass.

The only places where this may not happen, are the cleared areas surrounded by forest

where the elephant grass more or less is prevented from intruding. Regrowth of forest

also occurs in the higher altitudes of the Mahale ridge, from 2000-2500 m, which are

far less suited for elephant grass than the lower altitudes and where the virility of the

montane forest vegetation is stronger than that of the grasses.

When I first visited the Kabezi (1500-1600 m) at Lukandamira 20 years ago, that

valley was covered by forest with only small cleared patches. Since then, a large part

of the forest has been cleared and then left to bush. This area is now covered by enor-

mous elephant grass and is moist all the year round. The greater part of this grassland

is never burnt, but there is no sign of forest growth. This is not surprising. The root

system of this particular grass covers the ground like a thick carpet and, in my

opinion, must prevent any attempt of forest growth.

The comparatively great density of species of Lepidoptera in the western part of

Tanzania, is mainly due to its earlier connection with rainforests of Zaire before the

formation of Lake Tanganyika, and the influence of elements from north, south and

east. Another favourable condition is the variation of environment (see Kielland,

1968). There are four main types of habitat, viz., forest, woodland, montane grass-

land and marshy ground (text-fig. 3).

Forest. The forest of Ufipa, Mpanda and Kigoma and for that matter the whole of

Tanzania, is in no way of great extension. It consists of scattered remnants of ever-

green, or rain forests. The largest remnant in western Tanzania is at Kasoge, at the

foot of Mt. Kungwe. There are several types of evergreen forests:

1. Lowland forest: The low level forests of this region starts at just below 800 m

which is the level of Lake Tanganyika. The upper boundary of low altitude forest of

tropical Africa reaches approximately 1600 m. This boundary descends, if one moves

away from the equator. In South Africa it reaches see level just north of the Cape

Province. The sylvan fauna of Western Tanzania is most abundant at 800-1000 m,

and here most of the lowland forest species occur. From 1000 m the species density

decreases slowly. At about 1300-1400 m a gradual change in the vegetation occurs

and consequently, the lepidopterous fauna changes.

As I pointed out previously (Kielland, 1969) the lowland forests of the sandstone

area north of the Luegele-Katuma rivers, differ markedly from those of the complex

basement formations to the south. Through investigation, it was noted that a con-

siderable number of butterflies common to the forests of the sandstone area, did not

occur on the other geological formations or were very rare, and vice versa. Again, the

lowland forests on the west side of the Mahale Mt. contained species of its own.

The similarity of a large part of the lepidopterous fauna to that of the lowland

156 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

forests of Zaire, suggests a similarity in the flora as well. This has lately been shown,

to a certain extent, by the discovery of several plants of typical West African status in

Western Tanzania.

2. Highland forest. The typical montane forest of the Mahale range, Sitebi Mt.,

Kakungu-Mweze highlands and the highland of the Ufipa plateau, commences at

about 1400-1600 m after a gradual change of lowland forest. This intermediate stage

from 1400-1600 m contains a mixture of both low and high level vegetation. Above

1600 m the lepidopterous fauna is markedly different and reduced in numbers of

species and this reduction continues upwards.

The character of the montane forests is remarkably homogenous in the three dis-

tricts, as is the case in the whole of Central Africa. They differ, however, in certain

aspects. Very noticeable in the Ufipa montane forests (Mbizi forest, Chala forest) are

the dominating, very large Euphorbia’s, while this particular species does not occur in

the Mpanda-Kigoma area. At the Mahale ridge, above 1800 m, a large bamboo is asso-

ciated with all the forests. The Mbizi and Chala forests do not contain bamboo, but in

the forests of Mbuzi Mt. a hollow bamboo occurs, that is different from that on Mt.

Mahale. There is some introduced bamboo at Sumbawanga.

A conspicuous tree of the Sitebi-Sifuta range above 1800 m is a Podocarpus

species. (According to members of the Japanese expedition there are also a few

Podocarpus on Mt. Mahale). Here also a long, extremely slender bamboo, quite dis-

tinct from that of Mahale abounds, from 1500 m upwards. Curiously enough, this

species only occurs within a few square miles around the Sitebi Mt.

3. Dry evergreen forest. Apart from the above two types of evergreen forest, a

third kind must be mentioned, which is encountered in a few places in Mpanda and

Kigoma. It consists of a dryer kind of evergreen trees and large shrubs, often quite

densely entwined with lianas. The canopy trees are much smaller than those of the

common rain and riverine forests, and consists to a large extent of distinct species.

I have so far only encountered these evergreen thickets or forests in the sandstone

formations. The best example is on the sandstone plateau just east of Uruwira, (see

text-figs. 2 and 3). Here, these thickets are scattered over a flat area in pockets of up to

a few hundred yards in diameter, surrounded by Brachystegia woodland. These dry

evergreen forests have an affinity to the Zambezian ““Mateshi” which is related to

Cryptosepalum forest of that country and Angola (Cottrell, 1966). According to

Cottrell, the Zambia dry evergreen forests probably harbour one species of butterfly

endemic to this vegetation. So far, I have seen no indication of endemism in the dry

evergreen forest of the Uruwira plateau, but the area is small and the existence of

endemic species is not likely.

There are indications of a similar type of vegetation on the plateau near Mishamu

on the Uwinza-Mpanda road. At Mihumu, between some of the numerous riverine

forests, the whole area is clothed with semi-evergreen thickets and small trees, related

to the Cryptosepalum forest. Also at Luntampa, in area 4a, isolated patches of similar

dry evergreen forest occur. This is slightly inside the basic formations. The semi-

evergreen vegetation contains comparatively few creepers and lianas and differs

somewhat from the Uruwira forests.

The dry evergreen forests discussed above, are first of all the home of butterflies

confined to a mixed habitat of forest and woodland. True forest species do not seem

KIELLAND: Rhopalocera eastern side Lake Tanganyika 157

to breed in this kind of vegetation.

Woodland. This type of habitat covers at least 90% of the three districts, Mpanda,

Kigoma and Ufipa. It contains a large number (200-300) of tree species of deciduous

character. The woodland is normally open, poor in lianas, but with a rich growth of

grasses varying in height from 2 to 4 m or more in the savannas and valleys, to 30 cm

high or so on the hills. At higher altitudes as at Sitwe, about 1600-1700 m, and Ufipa

at 1700-1900 m, there is often a kind of deciduous vegetation of stunted trees and

shrubs, of mostly the same species or species related to those occurring at lower al-

titudes, but much smaller.

On the Ufipa plateau there are some kinds of vegetation which do not occur in the

Mpanda-Kigoma highland and vice versa. The butterfly fauna also differs to some

extent.

The species density of butterflies in woodland is lower than in the evergreen

forests. On the other hand, particular species are often more numerous and it is in the

open country or woodland that we get most of the migratory species, which may occur

in millions.

There is a pronounced seasonal variation in species, due to several factors, or

several factors that influence each other. Every year most of the woodland grasses

are burnt early in the dry season, in June-July. This provokes a sudden germination

of flowering herbs of great variety and also the emergence of several species of

Lycaenids which may breed on these plants or visit their flowers. These insects can be

found for only one or two months, e.g. several species of Lepidochrysops, Euchrysops

and Aloeides. After the rain starts in October-November and the grasses begin to

grow, other short-lived species of Lycaenids emerge.

The great contrast of the dry and rainy seasons in woodland country, stimulates

two distinctly different broods in a species, a wet and a dry season form (w.s.f. and

d.s.f., respectively), sometimes so unlike each other as to be taken for two different

species, such as: Precis octavia sesamus w.s.f. natalensis and P. octavia sesamus d.s.f.

sesamus, many species of the Satyridae, particularly the genus Bicyclus and also the

Pierid genus Colotis. This seasonal variation in species is far more pronounced in

woodland than in true forest, because in the latter, the environmental condition is

much more homogenous due to the heavy evergreen canopy which helps to keep the

humidity fairly high, even in the dry season.

The woodlands of hills and savannas differ slightly. Certain species of trees, shrubs

and grasses are found on the hills, while other types prefer the savannas. Many but-

terflies frequent the hills and are seldom or never met with down in the valleys and on

the savannas. The following are some examples typical for Brachystegia clothed hills:

Lepidochrysops solwezi, L. anerius, L. parsimon, L. gigantea, L. dollmani, L.

intermedia, Mimacraea marshalli, Cnodontes vansomereni and Precis touhilimasa.

Although many butterflies occur in savanna country, very few, if any are really

confined to this habitat. Nearly all are met with in other habitats as well.

Open montane grassland. There are numerous indications that this kind of grass-

land has developed in place of a former, much more extensive montane forest. The

Ufipa plateau is by far the most extensive area of open montane grassland. This pla-

158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

teau is comparatively densely inhabited by people engaged in both agriculture and |

cattle farming. By burning and clearing, the forest has greatly diminished in favour

of open grass. Species typical for montane grassland are: Euchrysops subpallida,

E. messapus, Lepidochrysops mpanda, L. cupreus, Eicochrysops mahallakoaena,

E. nandianas, Lycaena abbotii, Acticera stellata and Colias electo.

On the high ridges of the Mahale Mt. open grassland has developed comparatively

recently. This can easily be seen, as most of the grassy ridges still carry numerous

scattered forest trees, in clumps or isolated, even in the most exposed places far away

from the main forest.

The Lycaenid fauna is very poorly developed in this Mahale grassland. The genera

Spindasis, Lepidochrysops and Aloeides, so well represented in the Sitwe-Mweze high-

land, are conspicuously absent here. The Sitwe-Mweze grass country is older and

much more mixed with herbs and scattered, dwarfed, deciduous trees and shrubs

which seem to be advancing into the grassland. The ridge of the Wanzizi hills is

clothed with open grassland of the same character as that of Sitwe, but the Lycaenidae

are not as richly represented, probably due to the area being much smaller and

isolated from Sitwe.

Swamps. Only few butterflies, whose foodplant grow in water or mud, are confined

to this type of habitat. The most common species seen in swamps and other wet

places, near riverbanks etc., are Ypthimomorpha itonia and Metisella midas, both in

high and low level swamps, Mylothris bernice rubricosta in low to medium altitude .

swamps, the less common Syntarucus pulchra along swampy lake shores and rivers

and in the highlands of Ufipa; Mashuna upemba was taken on swampy ground.

For the area dealt with in the present work, there is no noticeable “short rainy |

season”, as in the northern part of Tanzania. The rain normally starts in the middle |

of October to early November and carries on until late April to early May, with some- |

what less rain in January-February, but seldom with more than two weeks pause in |

January. The Mahale Mt. enjoys higher rainfall than the surrounding areas and the |

rain often starts earlier and lasts longer, with scattered showers even in the dry |

season.

SYSTEMATIC PART

Family Papilionidae

Subfamily Papilioninae

In continental Africa this subfamily is represented by two genera, viz., Papilio —

Linnaeus and Graphium Scopoli. In the area dealt with, 13 species of Papilio and 9 spe- |

cies of Graphium are known.

Papilio antimachus Drury, 1782. This species has not been taken so far, but I have |

good reason to believe that this giant occurs in the forests of the Sitebi mountain area

of Mpanda. Observed and described by one of my African collectors.

Papilio dardanus dardanus Brown, 1776, with several female forms. A forest |

species, flying in most of the riverine forests of Mpanda and Kigoma. Record: All |

KIELLAND: Rhopalocera eastern side Lake Tanganyika 159

the investigated areas, except very high mountains (800-1700 m).

Papilio jacksoni kungwe Cottrell, 1963. High level forests of Mpanda and Kigoma.

Record: 1, 3a, 3b, 5a, 5b (1400-2200 m). General distribution: Ssp. jacksoni Sharpe,

highland forest of Kenya and Uganda; ssp. ruandana Le Cerf, Kivu province of Zaire

to Ruanda and western part of Uganda; ssp. nyika Cottrell, Nyika Plateau of Malawi;

ssp. hecki Berger, Nioka in Zaire; ssp. kungwe is endemic to mountains of Mpanda

and Kigoma.

Specimens differing from ssp. kungwe have been collected by me at Mbuzi Mt.,

Chulwe Mt. and Mbizi forest in Ufipa and by Carcasson in the Mbizi forest (1900-

2200 m).

Papilio zoroastres joiceyi Gab., 1945. Forests of Kigoma and Mpanda. A fairly

wide range in altitude. P. zoroastres joiceyi generally replaces P. jacksoni in lower

altitude forests; however, this is only partly the case in our area. In the Wanzizi hills

both species fly together at all levels where they occur, from 1400 m to the ridge at

1600 m. At Lukandamira, Mahale, ssp. joiceyi reaches the 1700 m level. Record: 1,

2a, 3a, 3b, 4a, 5a, 8a, 8b, Gombe (800-1700 m).

Papilio cynorta cynorta Fabricius, 1793. So far, the only Tanzanian records are

from forests of the sandstone formations between Kigoma and south to Ntakkatta on

the sandstone ridge, except for a single specimen taken at Kasoge. Record: 1, Sa, 8a,

8b (800-1400 m). General distribution: West Africa to Zaire and Uganda and an isolat-

eed race in Ethiopia.

Papilio nobilis Rogenhofer, 1891. One male taken at Mkulya riverine forest, a few

taken at Kampisa and at Ntakatta. Record: 3a, 5a, 5b(1300-1700 m).

Papilio hesperus Westwood, 1843. Not uncommon in forests and at riversides of

Mpanda and Kigoma. Rarely going up to 1500 m, as at Kampisa, and 1700 m near

Mt. Sitebi. Record: 2a, 3a, 4a, 5a, 5b, 6, 8a, 8b (900-1700 m).

Papilio constantinus mweruanus Joicey & Talbot, 1927. One 6 taken on the shore

of Lake Tanganyika at Kibweza Point. At Ikola by T. H. E. Jackson’s collector

Watulege. Record: 1, 2b, observation at Gombe (780 m). General distribution: ssp.

constantinus Ward, Kenya coast, Morogoro in Tanzania and Zambia; ssp. monticolus

Le Cerf, Kenya Rift Valley at Kedong; ssp. mweruanus J. & Ta., Victoria Falls in

Zambia.

Papilio mackinnoni Sharpe, 1891. Ssp. intermediate between ssp. theodori Riley and

ssp. benguella R. & T. Local in montane forests of Mpanda and Kigoma. Taken in

the Mweze forest, Wanzizi and at Kampisa. Record: 3a, 3b, 5b (1400-2000 m).

General distribution: ssp. theodori, Nyika Plateau in Malawi; ssp. mackinnoni, Uganda

and Kenya; ssp. benguellae, Rhodesia and K atanga.

Papilio phorcas congoanus Rothschild, 1896. ® forms congoanus and thersan-

droides Aur. Common in forests and heavy woodland of Mpanda and Kigoma. Re-

cord: widely distributed (800-1800 m).

Papilio nireus nireus Linnaeus, 1758. Common in forests and woodland of Mpanda,

Kigoma and Ufipa, often visiting flowers in open grassland. Record: widespread

(800-2000 m).

Note. Ssp. /yaeus Doubleday occurs in eastern and northern part of Tanzania.

Papilio bromius Doubleday, 1845. Ssp. interjacens Storace, and transition to chrap-

kowskoides Storace: common in forests of Mpanda and Kigoma; record: wide-

160 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

spread (800-2000 m). Ssp. ufipa Carcasson, 1960: montane forests of Ufipa only; re-

cord: Chala Mt., Mbizi Mt., Mbuzi Mt. (2000-2200 m). The author examined two

males kindly presented by Prof. Lindroth from Lund Univeristy, Sweden, taken 9. v. —

1951, at Milepa, Rukwa. This is at a much lower altitude.

Papilio demodocus demodocus Esper, 1798. Common in most habitats, but prefer-

ing woodland and open grassland. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b,

9b and Chala to Sumbawanga in Ufipa, Gombe (800-2000 m).

Papilio ophidicephalus ophidicephalus Oberthür, 1878. Riverine forests of Mpanda,

Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, 9b and Chala

in Ufipa (800-2000 m at Mahale).

Graphium ridleyanus White, 1843. Low level forests of Kigoma. Local. Record:

1 at Kasoge, 2a at Masaba, 4a, 8a, 8b and just south of Kigoma (800-1200 m).

Graphium angolanus angolanus Goeze, 1779. Savanna and woodland throughout

Mpanda, Kigoma and Ufipa. Record: widespread (800-1700 m).

Graphium taboranus Oberthir, 1886. Savanna and woodland of Mpanda. Scarce

in the western area, more common further east. Record: 5b, 6, 9a, 9b and west of Uru-

wira (1000-1600 m).

Graphium tynderaeus Fabricius, 1793. Riverine forest of Mkulya in Kigoma, close

to boundary of Mpanda. Only one record, 1954 (1450 m). General distribution:

Nigeria to the eastern part of Zaire, Ituri forest. The recorded specimen is the only one

known from East Africa.

Graphium leonidas leonidas Fabricius, 1793. Common in woodland of Mpanda,

Kigoma and Ufipa. Record: widespread (800-1700 m).

Graphium almansor kigoma Carcasson, 1964. Common in forests and forest

margins of Kigoma as far north as Malagarazi river and the north-eastern part of Mpan-

da. One specimen was taken at Sibweza. This specimen is similar to the nominate

race, but is probably the d.s.f. of ssp. kigoma. Record: 1, 3a, 4a, 5a, 5b, 6, 8a, 8b

(800-1500 m).

Since Carcasson described ssp. kigoma, a large number of specimens has been

collected throughout its range and they clearly show that ssp. kigoma is represented

by two distinct forms, a d.s.f. and a w.s.f. As the description of ssp. kigoma was

based on the w.s.f. (pl. 4 figs. 13, 14), it would seem appropriate to describe the d.s.f.

here:

Graphium almansor kigoma Carcasson, dry season form (pl. 4 figs. 15, 16). Usually

smaller, fore wing narrower and more incised at the distal margin; all white markings

of fore wing considerably reduced, appearance rather similar to the nominotypical race.

The white discal spot widely separated from the spot in 3 and not noticeably tapering

towards costa, almost rectangular in shape; also the apical spot less tapering towards

costa than in the w.s.f.; usually there is a small white spot near apex of cell. Hind wing

pale median area much larger than in the w.s.f., reaching end of discoidal cell; distal

margin very irregular and smudged; postdiscal and submarginal streaks absent, except

for faint streaks in space 5. Fore wing spots in la and lb and hind wing pale

area creamy, not white as in the w.s.f..

Graphium almansor wranghami ssp. nov. Corresponds with ssp. kigoma Carc. and

differs from ssp. almansor H. and ssp. uganda Lathy in the great difference between

the two seasonal forms.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 161

Wet season form (pl. 5 figs. 17, 18). All pale markings ochreous as in ssp. uganda,

but inner cell spot of the fore wing closer to cubitus and the spot in 3, but not contig-

uous with it as in the w.s.f. of ssp. kigoma; the cell spot is divided into 3 or 4

spots, tapering towards costa, but much smaller than in the w.s.f. of ssp. kigoma; ap-

ical spots well developed, but slightly smaller than in ssp. kigoma; submarginal, in-

ternervular spaces paler than the ground colour. In other races there are no

distinct paler areas of the submargin, apart from the white dots; spots in la and 1b

as in ssp. kigoma. Hind wing pale median area narrow as in ssp. kigoma; distal

margin irregular, but well defined and not smudged; a row of clearly defined, double

discal streaks and a row of indistinct, submarginal streaks. In ssp. uganda the sub-

marginal streaks are clearest, the discal ones are seldom developed at all. Size as in

ssp. kigoma.

Dry season form (pl. 5 figs. 19, 20). Usually smaller; shape of fore wing narrower

and more deeply incised at the distal margin as in the d.s.f. of ssp. kigoma; pale

markings of all wings white; no paler submarginal areas apart from the very small

white dots in 2 and 3; sometimes there is also a small discal spot in 2. The cell spot

is contiguous and rectangularly shaped, almost reaching costa, but separated from

cubitus and the spot in space 3. There is a clearly defined white dot near apex of

the cell, that is missing in the w.s.f. Hind wing pale median area white, but otherwise

as in the d.s.f. of ssp. kigoma, much wider than in the w.s.f. of ssp. wranghami;

discal and submarginal streaks absent; sometimes there are traces of streaks in area S.

Underside pale area more whitish than in the d.s.f. of ssp. kigoma and the

distal part of the wing grayer and not rufous as in kigoma.

Female unknown.

Habitat. Riverine forest and along the lake shore. Record: Gombe Stream north of

Kigoma (800 m).

Holotype (w.s.f.): Tanzania, Kigoma, Gombe Stream Nat. Park, December 1972, J.

Kielland. Paratypes (w.s.f.): Same data and collector, 5 & . Paratypes (d.s.f.): Same

data46.

I have the pleasure to dedicate this new race to Mr. Richard Wrangham, Ass. Di-

rector of Gombe Stream Research Station, who gave me accommodation and per-

mission to collect in the Park area. Holotype and 2 paratypes in the National Mu-

seum, Nairobi, 2 paratypes in the British Museum (Nat. Hist)., London, and 5 para-

types in J. Kielland collection.

Graphium antheus Cramer, 1775. Forests, woodland and savanna throughout

Mpanda and Kigoma. Record: widespread (800-1700 m).

Graphium policenes Cramer, 1775. Forests and woodland of Mpanda and Kigoma.

Very common in forests. Record: widespread (800 tot 1700 m).

Graphium porthaon tanganyikae ssp. nov. (pl. 6 figs. 21-24). Male: Differs from

the east coast races in the darker ground colour, and in the reduced submarginal

spots of the hind wing; in one 6 there are no spots or only a very faint dot in space

2; in another, 3 submarginal spots in 2, 3, and 4. The distance from the discal long spot

in 2 to the submarginal spot in 2 is greater than in the coastal races. The fore wing

subbasal streak is constantly narrow and the submarginal streaks are straight. Female:

Same differences as in the male. Underside darker; red and black markings better devel-

oped than in the coastal races. A very local race, flying sparingly in the forests of

162 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Kasoge, Kefu, Mihumu and Mukuyu in Kigoma. Found in glades and flying along

footpaths of low level forest. Record: 1, 8a, 8b (800-1000 m). General distribution:

Ssp. porthaon Hew., Mozambique; form vernayi van Son, 1936, coastal Kenya and

Tanzania, inland to Morogoro and Mikumi Nat. Park. There is very little difference

between porthaon and vernayi.

Holotype Ó . Tanzania, Kigoma, K asoge, 27.x.1969, J. Kielland; Allotype ® : idem,

but Kefu forest, 12.x1.1972. Paratypes: Tanzania, Kigoma, Kasoge, 2.xi.1969, J. Kiel-

land, 1 6 , idem but November, 1969, 1 & , idem, but Kefu, 12.iv. 1971, 10 é . Holo-

type and allotype in the British Museum (Nat. Hist.), London, paratypes in J. Kiel-

land collection.

Family Pieridae

Subfamily Pierinae

Appias sylvia luvuensis Joicey & Talbot, 1927. Common in forests of Mpanda and

Kigoma. Record: 1, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, 9a, 9b, Gombe (800-1500 m). Form

nyasana Butler, 1896: forests of Mpanda and Kigoma, flying together with /uvuensis.

Appias phaola phaola Doubleday, 1847. Forests of Kigoma at Mukuyu, Mihumu

and Kefu near Helembe and upper part of the Ntakatta forest, in the sandstone

area only. Record: Sa, 8a, 8b, Gombe (800-1000 m, 1500 m at Ntakatta).

Appias sabina sabina Felder, 1865. Forests of Helembe and at the sandstone ridge.

Record: 1, 4a, Sa, 8a, 8b, Gombe (800-1600 m).

O ssp. raffrayi

* ssp. similis

@ ssp. extendens

Fig. 4. Distribution of subspecies of Belenois raffrayi Sh.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 163

Appias sabina udei Suffert, 1904. Forests of Mpanda and Kigoma, south of the

sandstone ridge. Ssp. udei mixes with ssp. sabina at Luntampa and at Mahale; probably

the two races meet there (800-1500 m).

Appias epaphia epaphia Cramer, 1779. Forests and woodland of Kigoma at

Mukuyu and Kefu forest at Helembe. One specimen taken at Sibweza, April 1971,

and one at Ntakatta. Rare. Record: 5a, 6, 8a, 8b (900-1500 m).

Belenois (Anaphaeois) gidica westwoodi Wallengren, 1853. Open country and wood-

land of Mpanda and Kigoma. Very common. Record: widespread (800-1700 m).

Belenois (Anaphaeois) creona severina Stoll, 1781. Open habitats and woodland of

Mpanda, Kigoma and Ufipa. Very common. Record: widespread (800-1600 m).

Belenois (Anaphaeois) aurota aurota Fabricius, 1793. Common in woodland and

savanna of Mpanda, Kigoma and Ufipa. Record: widespread (800-1700 m).

Belenois raffrayi similis ssp. nov. (pl. 7 figs. 25, 26). Male. Closest to ssp. raffrayi

Obth. from Ethiopia and Sudan, from which it differs in the fore wing central white

area, which is slightly more extended by reduction of the basal blue dusting and the

apical black patch. The apex of the fore wing is slightly more pointed and outer mar-

gin a little concave at vein 3 and 4; in most specimens there is a submarginal

blue spot in 7. Underside fore wing, white area extended to the black marginal line in

internervular space in 1b, the outer part lightly sprinkled with grayish black scales; in

most specimens the apical black area does not reach the discoidal cell, which is out-

lined in black. The specimens from the dry season have the distal black area in both

wings more reduced.

Female. Hardly distinguishable from the nominotypical subspecies.

Habitat. Montane forests and forest margins of Mpanda and Kigoma.

Record: Mt. Sitebi-Ipumba-Ntakatta; Mweze Highland and Mt. Mahale (1500-2000

m) (text-fig. 4).

Holotype 6 : Tanzania, Mpanda, Mt. Sitebi, 2000 m, 1.v. 1970, J. Kielland;

Allotype 2 : Same place and collector, 2.v.1970; Paratypes: Same, but from 1 to 3 May,

1970, 64, same, but Mt. Ipumba, 5 and 7.v.1970, 29 ; Kigoma, Mweze, August

1970, 12; Kigoma, Lukandamira, August 1970, 1 2; Sitebi, May 1970, 19 ; Site-

bi, September 1970, 16; Mahale, August 1970, 16 ; Mweze, Augut 1970, 26.

Holotype and allotype in National Museum, Nairobi, paratypes in the National

Museum, Nairobi, the British Museum (Nat. Hist.), London and in J. Kielland

collection.

Belenois zochalia agrippinides Holland, 1896, form tanganyikae Lanz, 1896. Open

country and woodland of Mpanda, Kigoma and Ufipa. Common. Record: 3a, 3b, 5b,

6, 8a, 8b, Chala, Mbuzi Mt. (1000-1900 m).

Belenois crawshayi Butler, 1893. Open woodland and forest margins of Kigoma

and Mpanda. Very common in the sandstone area, less common elsewhere. Record:

3a, 4a, 5a, 6, 8a, 8b (800-1400 m).

Belenois victoria Dixey, 1915. Rather common, but local in forests of Mpanda

and Kigoma. The males gathering in numbers on wet sand. Female extremely rare.

Record: 1, 3a, 4a, 5a, 5b (1100-1700 m).

Belenois subeida instabilis Butler, 1888. Woodland and savanna of Mpanda, Kigoma

and Ufipa. Record: 3a, 3b, 4a, 5a, 5b, 6, Gombe, Chala (1000-1800 m).

Belenois calypso welwitschi Rogenhofer, 1889. Common in woodland of Mpanda,

164 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Kigoma and Ufipa. Record: widespread (800-1700 m).

Belenois calypso butyrosa Talbot,1943. This race seems to be confined to forest and

forest margins, while welwitschi occurs in open habitats and woodland. Kigoma in the

sandstone area. Record: 8a, 8b (800-1000 m). General distribution: Forests of Uganda.

Belenois thysa thysa Hopffer, 1855, forms macularia Aur., 1910, and alarmi Suf-

fert, 1904. Rather common in woodland and savanna of Kigoma and Mpanda. Record:

1, 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-1700 m.)

Belenois rubrosignata kongwana Talbot, 1943. Woodland and savanna of Mpanda,

Kigoma and Ufipa. Common. Record: 2a, 3a, 3b, 4a, 5a, 6, 8b, Chala, Gombe (800-

1900 m).

Belenois theora concolor Aurivillius, 1898, and © f. sulfurea Talb., 1943. Wood-

land and forest margins of Mpanda and Kigoma. Not common. Record: 3b, 4a, 5a;

6, 8a, 8b, 9b (900-1500 m).

Belenois solilucis Butler, 1874. Forests of Mpanda and Kigoma and northwards

to Biharamulo and Geita. Record: 1, 2a, 3a, 3b, 4a, Sa, 5b, 8a, 8b, Biharamulo,

Gombe (800-1600 m). The Mpanda-Kigoma specimens approach the large W. African

race. Specimens from Biharamulo and Geita are smaller and similar to the Uganda

forms.

Dixeia dixeyi Neave, 1904. Two males from the Luntampa forest in Kigoma and

one male from Mantena near Sibweza. Very rare. Record: 4a, 6 (ca. 1200 m).

Dixeia pigea pigea Boisduval, 1836. Chala in Ufipa. Record: Chala (ca. 1800 m).

| N f Kasulu O rhodope

poms FR DEA Ù @ asphodelus

Uwinza Z-

È de

~ 1

Kigoma\

Mpanda

\Ufipa

à | à 7

Lake \

Tanganyika È \ Rukwa|

\ )Sumbawanga AC

L i Toll Zambia |

Fig. 5. Distribution of Mylothris species.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 165

Dixeia pigea astarte Butler, 1899. Common in woodland and savanna of Mpanda

and Kigoma. Record: 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6 (800-1700 m).

Mylothris dollmanni Riley, 1921. One male taken at Chala in Ufipa, August 1973. A

rare species. Record: Chala (1800 m).

Mylothris chloris agathina Cramer, 1779. Woodland, savanna and forests of Mpan-

da, Kigoma and Ufipa. Very common. Record: Widespread (800-2000 m).

Mylothris citrina Aurivillius, 1898. Recorded by T. H. E. Jackson. Taken by his

African collector, Watulege, at Mahale Mt. Not seen by the present author. Record:

1. Altitude not specified.

Mylothris poppea rhodesiana Riley, 1921. Woodland, savanna and forest margins

of Mpanda, Kigoma and Ufipa. Rather common. Record: 1, 3a, 3b, 4a, 5a, 5b, 6,

Chala, Mbuzi Mt. (1000-2000 m).

Mylothris sulphurea basalis Aurivillius, 1907. One male taken in the Kasha forest

near Kefu, January 1973. Record: 8b (1000 m). There is one female from Katanga in

the National Museum, Nairobi. The species also occurs in eastern Tanzania.

Mylothris rhodope rhodope Fabricius, 1775 (text-figs. 5, 9). Kefu forest at Helembe,

Kigoma Distr. Very local, but common at Helembe. Record: 8b (900 m). General

distribution: W. Africa to Zaire, ssp. rhodope; Uganda and Kenya, ssp. uniformis Tal-

bot, 1944.

Mylothris asphodelus Butler, 1888 (text-figs. 5, 10). Local. In the Kasoge forest, Lun-

tampa and forests around Helembe. Record: 1, 4a, 8b (800-1200 m).

Mylothris spec. (text-figs. 8, 11). One female very similar to females of hilara

Karsch was taken in a riverine forest at Mihumu, one male taken in the Kefu forest.

There is another female in the National Museum, Nairobi, from Mukuyu taken by the

Japanese expedition. Record: 8a, 8b (900 m). Probably a new species.

Mylothris ertli Suffert, 1904, stat. nov. (Mylothris yulei Butler, 1897, ssp. ertli

Suffert 1904) (text-figs. 7, 11). Common at Kampisa, Mt. Sitebi and upper reaches of

the Katuma river. Also taken at Mt. Mahale, Luntampa forest and Ntakatta. Record:

1, 4a, 5a, 5b, Gombe (1200-2000 m).

Differs from M. yulei in a much better developed apical patch on fore wing upper-

side; a slightly more extended and stronger basal yellow area in both wings; underside

apical area of fore wing and entire hind wing with more or less strong yellow

suffusion; the marginal spots better developed in both wings. The female has less

strong, but more extended yellow basal patch on the wings and more rounded wings

than the male.

Genitalia 6 : Tegumen-uncus longer and narrower than in M. yulei; aedeagus slightly

more slender; valva differs in having a double central process (ampulla), a smaller lobe

is attached ventrally to the large lobe, in M. yulei the process is single.

The double lobed ampulla is characteristic for the M. rhodope group and would

suggest that ertli is more closely related to this group than to M. yulei.

Mylothris yulei yulei Butler, 1897 (text-figs. 6, 11). Common in forest margins

of Kigoma and Mpanda. Record: widespread (800-1800 m). . -

The Mylothris rhodope—yulei group is very complex and highly in need of revision.

Mylothris sagala narcissus Butler, 1888. Montane forests of Mpanda, Kigoma and

Ufipa. Record: 1, 3a, 3b, 4b, 5b, Mbizi forest, Chala Mt., Mbuzi Mt. (1500-over

2200 m). Specimens from Mahale, Sitebi and Ufipa differ from each other and

166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Central process (

ampulla)

Figs. 6—10. Male genitalia of Mylothris species. a, tegumen-uncus; b, valva; c, aedeagus. 6. M. yulei

yulei Butl., Kampisa, Mpanda, no. 1048. 7. M. ertli Suffert, Ujamba, Mahale, Kigoma, no. 1167. 8. M.

spec., Kefu forest, Helembe, Kigoma, no. 1031. 9. M. rhodope F., Kefu forest, Kigoma, no. 1019. 10.

M. asphodelus Butl., Kefu forest, Helembe, Kigoma, no. 1027.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 167

may constitute separate races. There is some variation in the female signa.

Mylothris bernice rubricosta Mabille, 1890. Along rivers and on swampy ground of

Mpanda, Kigoma and Ufipa. Common. Record: 2a, 2b, 5b, 6, Chala (800-1700 m).

Leptosia medusa marginea Mabille, 1890. Forests of Mpanda and Kigoma.

Common. Record: 1, 2a, 2b, 3a, 3b, 4a, 6, 5a, 8a, 8b, 9b, Gombe (800-1500 m).

Leptosia wigginsi Dixey, 1916. Forests of Kigoma from Malagarazi to Luegele

rivers. Rather common. Record: 2a, 3a, 4a, 5a, 8a, 8b (800-1200 m). General distri-

bution: Uganda.

Leptosia alcesta alcesta Stoll, 1782. Forests and heavy shrubs and woodland of

Mpanda and Kigoma. Common. Record: 1, 2a, 3a, 3b, Sa, 6, 9b (800-1200 m).

Leptosia nupta Butler, 1873. Very common in the Ntakatta forest. Also taken spo-

radically at Kefu and at Kasoge. Record: 1, 5a, 8b (800-1600 m).

Leptosia hybrida Bernardi, 1951. One female taken in the Kefu forest in Kigoma. Re-

cord: 8b (900 m). General distribution: From W. Africa to Zaire and Uganda. The

author has also collected this species in the Ngara District of north-western Tanzania.

Pinacopteryx eriphia eriphia Wallengren, 1857. Open habitats and savannas of

Ufipa, Mpanda and Kigoma. Uncommon. Record: 2a, 4a, 6, 8b, Chala, Sumbawanga

(900-1900 m).

(m al

| SN # Kasulu O yulei

Kigomad \ __, MEO oR

LR Uwinza 7- Won?

pra — SS sa

O aN °) à

\ om

\ Ve N N

©) À 5

J e

/ oe

O © O x ey

\ 5 Ù A

o d Ao

\ D si

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Ni 7

"Mpanda

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Tanganyika

\ Rukwa,

Sumbawanga

Toll Zambia

Fig. 11. Distribution of Mylothris species.

168 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Subfamily Teracolinae

Colotis calais (?) crowleyi Sharpe, 1898. Open habitats and savanna of Mpanda and

Ufipa. From northern Mpanda at Sibweza, one specimen only. In the Rukwa de-

pression and Ufipa very common. Record: 6, Mpimbwe, Rungwa near Lake Rukwa

(800-1200 m).

Colotis aurigineus aurigineus Butler, 1883. Open habitats of Ufipa and Mpanda

in the south. Record: Chala, Sumbawanga, Rukwa basin (800-2000 m).

Colotis vesta mutans Butler, 1877. Open habitats of Ufipa and Mpanda in the south.

Record: Chala, Mpimbwe, Sumbawanga, Rungwa (800-1700 m).

Colotis celimene anima Hewitson, 1862. Woodland and open habitats of Ufipa and

Mpanda. Rare. Record: 2b, 6, Chala (800-1700 m).

Colotis ione Godart, 1819. Savanna of southern Mpanda. One male only,

October 1970. Record: Mpimbwe (ca. 900 m).

Colotis regina Trimen, 1863. Woodland of Mpanda. Kigoma and Ufipa. Common.

Record: 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Chala (800-1600 m, 1800 m at

Chala).

Colotis hetaera Gerstaecker, 1871. One record from Nkungwe, 1955, and one from

Sibweza, 1971. Record: 6 (1100-1200).

Colotis elgonensis nobilis Carcasson, 1960. Forests and forest margins of Mpanda

and Kigoma. One record from Mbuzi Mt., Ufipa. Not uncommon in the two first

areas. Record: 1, 3a, 3b, 5a, 5b, Mbuzi Mt. (1300-2200 m). Rather high altitude spe-

cies.

Colotis hildebrandti Staudinger, 1885. Open habitats and woodland of Mpanda

and Ufipa. Not common. Record: 1, 2b, 4b, 5b, 6, Chala (900-1700 m).

Colotis danae annae Wallengren, 1857. Woodland of Mpanda, Kigoma and

Ufipa. Not very common. Record: 2a, 2b, 3a, 3b, 4a, 4b, 6, 7, Chala, Rukwa basin

(800-1700 m).

Colotis eucharis incretius Butler, 1881. Woodland and savanna of Mpanda and K igo-

ma. Common. Record: 2a, 2b, 3a, 3b, 4a, 4b, 6, 7 (800-1400 m).

Colotis antevippe zera Lucas, 1852. Open habitats and woodland of Mpanda,

Kigoma and Ufipa. Common. Record: widespread in Mpanda and Kigoma, Chala

in Ufipa (800-1900 m).

Colotis evenina xantholauca Sharpe, 1904, d.s.f. xantholauca, w.s.f. sipylus Swin.,

1884. Common in woodland of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b,

4a, 4b, Sa, 5b, 6, 8a, 8b, Chala, Rukwa basin, Gombe (800-1800 m).

Colotis evippe omphale Godart, 1819. Open habitats and savanna of Mpanda,

Kigoma and Ufipa. Record: widespread (800-1900 m).

Colotis pallene Hopffer, 1855. Recorded from 35 miles north of Mbeya.

Colotis evagore antigone Boisduval, 1836. Open habitats and woodland of Mpanda,

Kigoma and Ufipa. Common. Record: 3a, 3b, 4b, 6, Chala, Rukwa basin (800-

1800 m).

Colotis eris eris Klug, 1829. Open habitats and woodland of Mpanda, Kigoma and

Ufipa. Common. Record: widespread (800-1800 m).

Colotis subfasciatus ducissa Dognin, 1891. Open habitats and woodland of Mpanda

and Kigoma. Common. Record: 3a, 3b, 4a, 4b, 5b, 6, 8a, 8b (800-1300 m).

KIELLAND: Rhopalocera eastern side Lake Tanganyika 169

Eronia cleodora cleodora Hübner, 1822. Savanna-like woodland in Mpanda at

Nkungwe and Mpanda township. Rare. Record: 6 (1000-1200 m).

Eronia leda Boisduval, 1847. Woodland and forest of Mpanda and Kigoma.

Common. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6 (800-1600 m).

Nepheronia pharis Boisduval, 1836. Forests of Kigoma in the sandstone area only,

north of Mugombazi. Record: 8a, 8b at Mukuyu, Mihumu, Helembe forests (800-

1000 m).

_ Nepheronia thalassina Boisduval, 1836. Forest and woodland of Mpanda and

Kigoma. Rather common. Record: widespread, 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b,

(800-1700 m).

Nepheronia argia argia Fabricius, 1775. Forests and dense woodland of Mpanda

and Kigoma. Common. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe

(800-1800 m).

Subfamily Coliadinae

Colias electo pseudohecate Berger, 1940. Open grassland on mountains of Ufipa,

Mpanda and Kigoma. Common. Record: 1, 3a, 5a, 5b, Chala to Sumbawanga

(1500-2300 m).

Catopsilia florella Fabricius, 1775. Almost any habitat of Mpanda, Kigoma and

Ufipa. Record: everywhere (800-2200 m).

Eurema hecabe senegalensis Boisduval, 1836.

Open habitats and woodland of Mpanda, Kigoma and Ufipa. Common. Record: wide-

spread (800-2000 m).

Eurema brenda Hewitson, 1847. Heavy woodland and forest of Mpanda and

Kigoma. Common. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-

2000 m).

Eurema brigitta brigitta Cramer, 1780. Open habitats and woodland of Mpanda,

Kigoma and Ufipa. Common. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b,

Chala, Gombe (800-2000 m).

Eurema hapale Mabille, 1885. Forest and swampy ground of Mpanda, ISSUE and

Ufipa. Very common. Record: widespread (800-2000 m).

Eurema desjardinsii regularis Butler, 1876. Very common in woodland, savanna

and forest margins of Mpanda, Kigoma and Ufipa. Record: widespread (800-2000 m at

Mt. Sitebi). In my opinion E. mandarinula Holland, 1892, is a form of E. desjardinsii,

restricted to lower altitudes.

Eurema punctinotata Butler, 1895. Forests and open habitats of mountains in Ufipa

and Kigoma. Record: 1 at Mahale Mt.; Mbizi forest in Ufipa; local (1700-2200 m).

E. punctinotata is found only in the more humid, high altitudes above 1600 m. There is

no great difference between dry and wet season forms.

Family Danaidae

Danaus chrysippus Linnaeus, 1758. Forms chrysippus, alcippus Cramer, 1777, do-

rippus Klug, 1845, and albinus Lang, 1896. Common and widespread everywhere

in open and wooded habitats. Record: widespread (800-2200 m).

170 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Danaus limniace petiverana Doubleday & Hew., 1847. Forest and woodland

throughout Mpanda, Kigoma and Ufipa. Record: everywhere except high mountains

(800-1600 m).

Danaus formosa formosa Godman, 1880. Forests of Kigoma and Mpanda. Also

met with in open habitats. Record: 1, 5b, 8a (800-2000 m). In the Mahale area

it is found at both low and high altitudes. Up to 1000 m at Mukuyu and Mi-

humu. In the Sitebi Mt., only taken at 1900-2000 m.

Amauris niavius niavius Linnaeus, 1758. Common in forests of Mpanda and Kigo-

ma. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 8a, 8b, 9b, Gombe (800-1600 m).

Amauris tartarea tartarea Mabille, 1876. Forests of Mpanda and Kigoma. Com-

mon. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe (800-1700 m).

Amauris egialea mukuyuensis Carcasson, 1964. An uncommon species, in forests of

Kigoma. Record: 1, 3a, 4a, 5a, 8a, 8b (800-1200 m, 1600 m at Lukandamira).

Amauris echeria mpala Talbot, 1940. Mahale area in Kigoma, Mweze and Kam-

pisa in Mpanda. In forests. Common. Record: 1, 2a, 3a, 4a, 5a, 5b (1200-2000 m).

Amauris albimaculata magnimakula Rebel, 1914. In the Mihumu forest of Ki-

goma. Very rare. Record: 8a (ca. 900 m).

Amauris crawshayi oscarus Thurau, 1903. A rare species in forests of Mpanda

and Kigoma. Record: Sa, 8a, 8b (900-1500 m).

Amauris ansorgei junia Le Cerf, 1920. Rather common in the Mbisi Forest,

Ufipa (2000-2300 m), February 1978.

Family Satyridae

Melanitis leda africana Frihstorfer, 1908. Very common in woodland of Mpanda

and Kigoma. Record: widespread (800-1500 m, 2000 m at Ujamba, Mahale).

Melanitis libya Distant, 1882. Forests and woodland of Mpanda and Kigoma. Com-

mon. Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, 9a, 9b (800-2200 m).

Common at Mahale and surrounding areas. Further east more scarce in the lowland,

but plentiful at high altitudes. At high altitude of Mahale flying exclusively in the

bamboo mixed forest.

Melanitis parmeno parmeno Doubleday & Hewitson, 1851. Common in forests of

Mpanda and Kigoma. In Mpanda mainly at higher levels, in Kigoma common at

all altitudes. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, at Kigoma (Miss

Goodall) (800-2000 m). General distribution ssp. parmeno: Uganda, Ghana, Angola.

Melanitis chelys Fabricius, 1793, forms chelys and minchini Heron, 1909. Com-

mon in dense forests of Kigoma and Mpanda. Record: 1, 4a, 5a, 8a, 8b (800-

1600 m).

Bicyclus sebetus sebetus Hewitson, 1877. Dense, low level forest of Kigoma and

Mpanda in the north-west. Record: 1, 3a at Lubalizi, 4a at Luntampa, 5a at Ntakatta,

8a, 8b (800-1400 m). Very common at Kasoge and in the sandstone area. Elsewhere

scarce.

Bicyclus sambulos Hewitson, 1877. Ssp. intermediate between ssp. sambulos and

ssp. cyaneus Condamin, 1961. Forests of Kigoma. Not rare in the Kasoge and

Kefu forests. Elsewhere scarce. Record: 1, 3a, 4a, 5a, 8a, 8b (800-1400 m). General

distribution: Uganda and Zaire.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 171

Bicylcus mandanes Hewitson, 1876. Common in forests of Kigoma and Mpanda to

Biharamulo Distr., Kahama Distr. Record: 1, 2a, 3a, 4a, 5a, 8a, 8b, Biharamulo

and Kahama Districts (800-1700 m).

Bicyclus auricrudus fulgidus Fox, 1963. Common in forests of Kigoma. Flying

together with B. mandanes. Record: 1, 2a, 3a, 4a, 5a, 8a, 8b, Gombe (800-1500 m).

General distribution: Zaire to Uganda and Kenya; ssp. auricrudus in Nigeria.

Bicyclus mesogena uganda Riley, 1926. In the K asoge forest only. Record: 1 (800-

1000 m). General distribution: Uganda and Kenya.

Bicyclus sandace Hewitson, 1877. Forests of Kigoma and Mpanda. Sometimes also

in heavy woodland. Common. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Kibon-

do, Gombe (800-1700 m). General distribution: Nigeria, Zaire to Uganda.

Bicyclus jefferyi Fox, 1963 (= addendus Cond. & Fox, 1964, = miriam Fab., 1793).

Forests, riversides and heavy woodland of Mpanda and Kigoma. Common. Record:

1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b (800-1800 m, 2000 mat Mt. Sitebi).

Bicylcus safitza Westwood, 1850. Almost any wooded habitat and forest of

Mpanda, Kigoma and Ufipa. Record: widespread (800 to 1800 m).

Bicyclus campus campus Karsch, 1893. Forest and woodland of Mpanda and

Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8b (800-2000 m).

Bicyclus dentata Sharpe, 1898. Montane forests of Mpanda and Kigoma. The

Mahale Mt. and the Sitebi-Ipumba range. Locally very common. Record: 1, 5b (1500-

2200 m).

Bicyclus cooksoni Druce, 1905. Forest and woodland of Mpanda, Kigoma and Ufi-

pa. Rather common. Record: 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Chala (1000-1800 m).

General distribution: Zambia, Katanga.

Bicyclus cottrelli cottrelli van Son, 1952. Common in woodland and forest in

Mpanda and Kigoma. Record: 3a, 3b, 4a, 5b, 6, Gombe (1000-1500 m). General distri-

bution: Zambia to Kahama in Tanzania.

Bicyclus ena Hewitson, 1877. Woodland of Mpanda and Kigoma. Not common.

Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8b, 9a, 9b (800-1700 m).

Bicyclus angulosus selousi Trimen, 1895. Common in woodland of Mpanda, Kigo-

ma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8b, Chala, Gombe (800-

2000 m). Some specimens are transitional to ssp. angulosus.

Bicyclus campinus campinus Aurivillius, 1901. Woodland and forest of Mpanda and

Kigoma. Mainly lower altitudes. Record: 1, 2a, 2b, 3a, 3b, 4a, 6 (800-1500 m).

Bicyclus vansoni Condamin, 1965. Very common in woodland and forest margins

of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, Sa, Sb, 6, 7,

8a, 8b, 9a, 9b, Chala, Gombe (800-2000 m).

Bicyclus anynana anynana Butler, 1879 (= vicaria Thurau, 1903). Common in

woodland of Mpanda and Kigoma. Record: all investigated areas in Mpanda and

Kigoma (800-1600 m).

Bicyclus vulgaris Butler, 1868. Common in forests and woodland of Mpanda and

Kigoma. Record: 1, 2a, 3a, 4a, 5a, 5b, 8a, 8b, 9a, Gombe (Miss Goodall) (800-

1500 m). General distribution: Uganda.

Bicyclus funebris funebris Guerin, 1844, d.s.f. nebulosa Felder, 1867, w.s.f. agraphis

Karsch, 1893. Common in forests and heavy woodland of Kigoma and north-western

part of Mpanda. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 8a, 8b, Gombe (800-1500 m,

172 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

1700 m at Lukandamira). The southern-most record is the Wanzizi Hills.

Bicyclus saussurei saussurei Dewitz, 1879. Taken in a riverine forest near Kibondo.

One male only. General distribution: Uganda, Katanga, Zambia, and Ngara Distr.

of Tanzania.

Bicyclus similis Condamin, 1963. Dry and wet season forms taken in the Mahale |

mountain area. First taken by T.H.E. Jackson’s African collector, Watulege. Record:

1(1600-2300 m). This species is endemic to the Mahale area and no trace of it has

been found outside this mountain. It is common at high levels, becoming rarer down

to 1600 m.

Henotesia perspicua Trimen, 1873, w.s.f. perspicua, d.s.f. maevius Staudinger,

1887. Very common in woodland and savanna of Mpanda and Kigoma. Also Ufipa.

Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, 9b, Chala, Gombe

(800-2000 m).

Henotesia simonsii Butler, 1877, w.s.f. victorina Westwood, 1881, d.s.f. simonsii.

Common in woodland and savanna of Mpanda, Kigoma and Ufipa. Record:

widespread (800-1800 m).

Henotesia spec. Common, but confined to the Mahale area at high elevations.

Frequents montane forests and forest margins. Wet and dry season forms occur.

Record: 1 (1500-2200 m). The description of this new species may be undertaken later

on, as a revision of the whole genus is needed. It is closely related to H. ubenica Thurau,

1903, but the genitalia differ. A species with similar genitalia occurs at Kitcheche and

Ankole — Toro, Uganda. However, this seems to constitute a separate race, distinct

from the Mahale population. Specimens similar to the Uganda form have been taken

at Ngara in northeast Tanzania by the present author.

Aphysoneura pigmentaria latilimba Le Cerf, 1919. Forests and forest margins of _

Mpanda, Kigoma and io High altitude species. Common, but local. Mainly in

bamboo country. Record: 1, 5a, 5b, Mbizi forest, Chala, Mbuzi Mt. (1400-above

2000 m).

Physcaenaura pione Godman, 1880. Common in woodland and savanna of Mpan-

da and Kigoma. Wet season only, from late December to April. Record: 2b, 3a, 3b, 4a,

4b, 6 (800-1300 m).

Neita victoriae Aurivillius, 1898 (= Neocoenyra victoriae Aur.). Occurring in wood-

land at Sibweza, Mpanda Distr. Also taken near Uwinza and at Biharamulo north

of Kibondo. Uncommon. Record: 6, 9a, Biharamulo (1050-1400 m).

Neocoenyra Butler, 1885.

This genus is represented by two or three gregorii-like species in this area. One is

N. kivuensis Seydel, 1929, described from Kivu in Zaire. The genitalia in this genus

are of the ”grayhound” type and the species are very difficult to separate from each

other. The whole genus is in need of revision.

Ypthima Hübner, 1818.

At present the writer makes a special study of this genus and a revision of the

continental African group will be submitted later. For the time being the following

species from western Tanzania are briefly listed.

Ypthima granulosa Butler, 1883 (= cataractae van Son, 1955). Occurring at Kasulu

and at Mihumu. Record: 8a, Gombe, Kasulu (900 at Mihumu, 800-1200 at Gombe,

not certain at Kasulu).

KIELLAND: Rhopalocera eastern side Lake Tanganyika 173

Ypthima impura Elwes & Edwards, 1893. Very common in woodland and savanna

of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 3a, 3b, 4b, Sa, 5b, 6, 8b, Sum-

bawanga, Chala, Gombe (800-1800 m).

Ypthima pupillaris Butler, 1888 (= gazana van Son, 1955). Common in Brachyste-

gia woodland and savanna of Mpanda, Kigoma and Ufipa. Preferring higher altitudes.

Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, Chala, Sumbawanga (1100-2000 m).

Ypthima albida albida Butler, 1888. One male taken at a small river just north of

Kibondo. Common at Bukoba and Ngara. Record: Kibondo.

Ypthima rhodesiana Carcasson, 1964. Common in woodland and savanna of Mpan-

da and Kigoma. Record: 2a, 3a, 3b, 4b, 5b, 6, 8a, 8b (1000-1500 m).

Ypthima antennata van Son, 1955. In rocky places and grassland of the Sitebi Mt.

and near Uwinza. In sandstone area only. Record: 5a, 5b, Uwinza (1200-2000 m).

Ypthima spec. Savanna and woodland of Mpanda, Kigoma and Ufipa, in the

Rukwa basin and at Chala. Record: 2a, 3a, 3b, 4a, 4b, 6, 8a, 8b, Kansindi near

Usewia, Rukwa basin, Chala, Mbuzi Mt. (800-1200 m, 2000 m in Ufipa). This new

species will be described in another paper.

Ypthimomorpha itonia Hewitson, 1865. Common in savannas, particularly swampy

places and riversides of Mpanda, Kigoma and Ufipa. Record: widespread (800-

1800 m).

Mashuna upemba Overlaet, 1955. Two males and one female were taken flying low

over marshy ground near Chulwe Mt. in Ufipa. Record: Chulwe Mt. (ca. 1900 m).

General distribution: Upemba Park in Katanga and Angola Highland.

Family Nymphalidae

Subfamily Charaxidinae

Euxanthe crossleyi ansorgei Rothschild & Jordan, 1900. Widespread, but not

very common. In forests and along rivers of Mpanda and Kigoma. Record: 1, 2a, 3a,

3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe (800-1600 m). General distribution: Nigeria

to Uganda and western Kenya.

Charaxes varanes vologeses Mabille, 1876. Common in open woodland and sa-

vanna of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7,

8a, 8b, Chala, Gombe (800-2000 m).

Charaxes fulvescens monitor Rothschild, 1900. Very common in forests and

riverine thickets of Mpanda and Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a,

8b, Gombe (800-2000 m).

Charaxes acuminatus Thurau, 1903. Observed at Mbuzi Mt. in Ufipa (2000 m).

Charaxes candiope candiope Godart, 1824. Common in forests and riverine

forests of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b,

Mbuzi Mt., Gombe (800-2000 m).

Charaxes protoclea Feisthamel, 1850; ssp. azota Hewitson, 1877; ssp. catenaria

Rousseau-Decelle, 1934. In the Mpanda area this species occurs commonly on

Brachystegia clothed hills. In the better forested areas further north, as in Kigoma

area, it is mostly a forest dweller and is seldom met with in woodland. Record:

1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe, Ufipa at foot of escarpment (800-

174 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

1700 m, 2000 m at Mt. Sitebi). Specimens agree with both races and must be regarded

as transitional.

Charaxes boueti macclouni Butler, 1895. Exceedingly common in areas with bam-

boo, the larval foodplant, but also taken at Sibweza more than 10 miles from nearest

bamboo growth. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Namanyere

and Chala in Ufipa (800-2000 m). The high level Mahale specimens are larger and |

brighter than those in the lowland.

Charaxes cynthia mukuyu van Someren, 1969. This race is endemic to the area |

between Burundi and the sandstone ridge at Ntakatta. Fairly common at Mihumu, |

but extremely scarce elsewhere. Record: 5a, 8a, 8b (800-1500 m). General distri- |

bution: Other races from eastern Kenya, Uganda across central Africa to Ivory Coast.

Charaxes lucretius maxima van Someren, 1971. Very common in the Kasoge forest,

very rare elsewhere. Record: 1, 3a, 4a, 5a, 8a, 8b, Gombe (800-1400 m).

Charaxes jasius saturnus Butler, 1865. Common in open woodland and savanna of

Mpanda, Kigoma and Ufipa. Seldom met with in forest. Record: widespread

(800-2000 m).

Charaxes castor castor Cramer, 1775. Common in open woodland and forests of

Mpanda and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 5b, 8a, 8b, Gombe

(800-2000 m).

Charaxes castor flavifasciatus Butler, 1876. In the Rukwa valley in Ufipa and South

Mpanda. Record: Muze (ca. 1000 m). Ssp. flavifasciatus probably merges with ssp.

castor further north.

Charaxes brutus natalensis Staudinger, 1886. Widespread, but not very common in

Mpanda and Kigoma. Preferring forest but one specimen also taken in open wood-

land at Nkungwe. Record: 1, 2b, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-

1700 m).

Charaxes ansorgei kungwensis van Someren, 1967. Very rare. Records from the

montane forests of Mt. Mahale only. Record: 1 (2000-2200 m). During 14 trips

to the Mahale Mt., only a few males and one female were obtained in banana traps.

Most were taken at the remote Sisaga Mt. General distribution: Other races from

Kenya to Uganda, at the northern end of Lake Tanganyika and Northern Malawi.

From Tanzania, races have been recorded from Kilimanjaro, Usambara, Uluguru and

Njombe to Mbeya. Ssp. kungwensis was first taken by T. H. E. Jackson’s collector

Watulege.

Charaxes ansorgei ufipa ssp. nov. (pl. 8 figs. 27-30). Male. Upperside discal band

of fore wing rather narrow and with outer border straighter and less incised at the

veins than in other races, much less than in ssp. kungwensis van Som. and ssp. levicki

Poult. (the latter comes from Southern Highlands in Tanzania). The discal band is not

as dark as that of ssp. levicki, but slightly darker than in ssp. kungwensis;

basal area as dark as that of ssp. levicki, slightly darker than in ssp. kungwensis.

Hind wing submarginal spots well developed in 4 to 7, but no spots in 2 and 3. In the

nominotypical race, which it otherwise rather closely resembles, the hind wing sub-

marginal spots are faint, but present also in 2 and 3, except for one specimen in

the National Museum, Nairobi. Length of fore wing: 41 mm.

Female. Differs from ssp. kungwensis in a darker brown, basal area of the fore wing;

the creamy, discal band is broadening towards the hind margin, not parallel in area

KIELLAND: Rhopalocera eastern side Lake Tanganyika 175

1 and 2 as in ssp. kungwensis; the shape of the fore wing is more falcate and

the outer margin less concave. Hind wing with pronounced submarginal spots in 5 to 7;

the green marginal stripe at tornus is hardly discernible. Underside wings as ssp.

kungwensis.

Habitat: Montane forests of Mbuzi Mt. in Ufipa.

Holotype é : Tanzania, Ufipa, Mbuzi Mt., 2000 m, October 1971, J. Kielland.

Allotype £ : Same, but May 1972; Paratypes: 1 6 and 1 May 1972; 16 October

1971.

Holotype and allotype are deposited in the National Museum, Nairobi, paratypes

in J. Kielland collection.

Charaxes pollux pollux Cramer, 1775. In riverine forests of Mpanda and Kigoma.

Rather common. Record: 1, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-2000 m).

Charaxes druceanus proximans Joicey & Talbot, 1922. Not common and local in

open country and forest margins of Mpanda, Kigoma and one record from Ufipa.

Record: 1, 2a, 3a, 4a, 5a, Gombe, Mbuzi Mt. (1200-2000 m).

Charaxes eudoxus mechowi Rothschild,1899 (according to M. Plantrou). Taken

in banana traps at Mihumu, Kasoge, Luntampa, Ntakatta and Kampisa forests.

Rather rare and local. Record: 1, 3a, 4a, 5a, 5b, 8a, 8b, Biharamulo (800-1500 m).

General distribution: Zaire to Uganda and Kenya. Also taken in Zambia.

Charaxes numenes aequatorialis van Someren, 1972. Very local in low level forests

of Kasoge, Luntampa, Lubalizi, Mukuyu, Kefu, Ntakatta and Mihumu. Not taken

south of Lubalizi. Record: 1, 3a, 4a, 5a, 8a, 8b, Gombe (800-1500 m). General distri-

bution: W. Africa to Zaire, Ethiopia, Uganda and Kenya.

Charaxes bohemani Felder, 1859. Common in open woodland and savanna

throughout Mpanda and Kigoma. Not in forest. Record: widespread (800-1600 m,

2000 m at Mt. Sitebi).

Charaxes smaragdalis kigoma van Someren, 1964. In forests of Kigoma at

Mukuyu, Kefu, Ntakatta and north of Kigoma to Burundi. South of Kigoma

very rare. Record: 5a, 8a, 8b, Gombe (Miss Goodall), Burundi (800-1500 m). General

distribution: represented by a number of races, scattered from western Kenya and

Uganda, over Zaire and W. Africa, to Ivory Coast.

Charaxes tiridates tiridatinus Rober, 1936. In forests of Mpanda and Kigoma. Rare

south of Luegele river, where it occurs as far south as Sibweza, in area 6. It is

common north of Luegele. Record: 1, 3a, 4a, 5a, 6, 8a, 8b, North of Kigoma (800-

1500 m).

Charaxes bipunctatus ugandensis van Someren, 1972. Fairly common in the

Ntakatta forest, but not found outside this forest. Record: 5a (1200-1500 m). General

distribution: W. Africa to Zaire, Uganda and western Kenya.

Charaxes xiphares brevicaudatus Schultze, 1914. Taken in two forest remains of the

Sitebi Mt., upper reaches of Katuma and at Ihassa Mt. near Ntakatta. All in banana

traps. Rare. Record: 5a, 5b (1600-2000 m). General distribution: races are known

from South Africa to northern Kenya and from Cameroun. Ssp. maudei J. & Tal.,

1918, occurs at the east coast of Tanzania; ssp. kilimensis van Som. at Kilimanjaro;

ssp. kiellandi Plantrou, 1976, at Mt. Oldeani, and there are earlier records of ssp.

brevicaudatus from Iringa and Mbeya.

Charaxes ameliae amelina Joicey & Talbot, 1925. A forest species, but often flying

176 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

in open woodland. Rather common in Kigoma and the north-western part of Mpanda.

Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe (800-1600 m, 2000 m at

Mt. Sitebi).

Charaxes imperialis ugandicus van Someren, 1972. Very rare and local in forests

of Mpanda and Kigoma, at Kasoge, Ujamba river, Kampisa, Lubalizi, Kefu, Mi-

humu and Ntakatta. Record: 1, 3a, 5a, 5b, 8a, 8b (800-1500 m). General distribution:

W. Africa, Zaire to Uganda.

Charaxes pythodorus pythodorus Hewitson, 1873. Uncommon in forests of Kigoma

and Mpanda. Wanzizi, Kampisa, Sitebi, Ntakatta, Luntampa, Ujamba riv. at 1300 m,

Lukandamira, Kefu, Mihumu. Record: 1, 3a, 3b, 4a, 5a, 5b, 8a, 8b (800-1600 m, 2000

m at Mt. Sitebi). General distribution: from Kenya, Uganda to the eastern part

of Zaire, western Zambia and Angola; ssp. occidens van Someren, 1963, W. Africa;

ssp. pallida Carpenter, 1934, central Tanzania, and nesaea G. Smith, 1889, east coast

of Kenya and Tanzania.

Charaxes eupale latimargo Joicey & Talbot, 1921. Riverine forests of Mpanda and

Kigoma. Not uncommon, but difficult to trap with bait. Record: 1, 5a, 5b, 8a, 8b,

Gombe (800-1500 m).

Charaxes dilutus dilutus Rothschild, 1898. Forests and riverine forests of Mpanda

and Kigoma. Not uncommon. One record from Sibweza. Same habitats as C. eupale.

Record: 1, 3a, 4a, 5a, 5b, 6, 8a, 8b (800-1700 m).

Charaxes jahlusa kigomaensis van Someren, 1975 (= kigoma v. Som., 1974). In

forests of Kasoge, Mihumu and Mukuyu in Kigoma, Ntakatta in Mpanda. Un-

common. Record: 1, 5a, 8a, 8b, Gombe (800-900 m, 1500 m at Ntakatta). General

distribution: Ethiopia and East Africa to Mozambique, Rhodesia and South Africa.

Charaxes pleione bebra Rothschild, 1900. Taken in the Mukuyu forest, Kefu

and Ntakatta. Very local. Record: 5a, 8a, 8b (800-900 m, 1500 m at Ntakatta). Gen-

eral distribution: W. Africa to Zaire, east to Uganda and West Kenya.

Charaxes zingha Stoll, 1780. Low altitude forests. Not taken south of Kefu forest.

Very local. Record: 8a, 8b (800-900 m). General distribution: W. Africa to Zaire, east

to Uganda.

Charaxes etesipe etesipe Godart, 1823. Kasoge and Kefu forest in Kigoma. Un-

common. Record: 1, 8b(900 m).

Charaxes etesipe tawetensis Rothschild, 1894. This subspecies was taken in the Lun-

tampa forest, and Lubalizi and probably meets ssp. etesipe in this area. Record:

3a, 4a(1200 m).

Charaxes penricei tanganyikae van Someren, 1966. Rare in woodland and savanna

of Mpanda. In the Kigoma area taken in forests only. Rarely attracted to banana

bait. Record: 1, 3a, 4a, 4b, 5a, 6, 8a, 8b, Ugala river, Gombe (800-1300 m). At

Mihumu ® form caerulescens van Someren, 1969, first taken by Zuzuki. General

distribution: ssp. tanganyikae from Mbeya along the east side of Lake Tanganyika to

south Burundi, thence west to Mwanza south of Lake Victoria.

Charaxes achaemenes achaemenes Felder, 1867. Very common throughout Mpanda,

Kigoma and Ufipa, in open woodland and savanna. Record: widespread (800-1500 m,

2000 m at Mt. Sitebi).

Charaxes anticlea adusta Rothschild, 1900. Forests of Kigoma at Kasoge, Lu-

balizi, Mihumu, Kefu, Luntampa, Ntakatta and Mukuyu. Not common. Record:

KIELLAND: Rhopalocera eastern side Lake Tanganyika 177

1, 3a, 4a, 5a, 8a, 8b, Gombe (800-1500 m).

Charaxes baumanni whytei Butler, 1893 (cline to ssp. baumanni). Forests and riv-

erine forests of Mpanda, Kigoma and Ufipa. Rather high level species. Local, but

common at Wanzizi and Kampisa. Record: 1, 3a, 3b, Sa, 5b, Chala, Mbuzi Mt.

(1200-2000 m, rare below 1500 m). General distribution: several races from Uganda

in the north to Rhodesia in the south.

Charaxes guderiana guderiana Dewitz, 1879. Very common in woodland and sa-

vanna throughout Mpanda, Kigoma and Ufipa. Record: widespread (800-1800 m).

Charaxes etheocles carpenteri Poulton, 1919. Forests of Kigoma and at Wanzizi in

Mpanda. Very rare in Mpanda, but more common in Kigoma. Record: 1, 2a, 3b, 4a,

5a, 5b, 8a, 8b, Mbuzi Mt. in Ufipa (800-2000 m).

Charaxes grahamei van Someren, 1969. Not uncommon in the forests of Kasoge,

Kefu, Ntakatta and Mihumu. Not yet taken south of Kasoge, and north of Kigoma.

Rarely obtained on banana bait, but males frequently on animal droppings.

Females difficult to obtain. Record: 1, 5a, 8a, 8b (800-1000 m, 1500 m at Ntakatta).

This species, first obtained by the Japanese Primate Exp. from Kyoto, is most certainly

endemic to the recorded area. At Mihumu form /acteata van Someren, 1969.

Charaxes howarthi Minig, 1976. The commonest of the black Charaxes in this area.

In woodland and forest throughout Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a,

3b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b, 9a, 9b, Chala, Gombe (800-1500 m, 2000 m at

Sitebi).

Charaxes pseudophaeus van Someren, 1957 (= Ch. manica® f. pseudophaeus van

Som. & Jackson, 1957). Uncommon in woodland of Mpanda and Kigoma. Record:

2a, 2b, Sa, 5b, 6, 7 (900-1500 m). I am not certain of the validity of this species.

Charaxes (?) aubyni Poulton, 1925. A male was taken in a banana trap at Gombe

Stream Nat. Park north of Kigoma, December 1972. Record: Gombe (900 m). Under-

side much darker than in aubyni; possibly a new species.

Charaxes fionae Henning, 1977. Open woodland of Mpanda, Kigoma and Ufipa.

More common in the south. Record: 2b, 5b, 6, Ufipa, Kigoma, Gombe (800-1600 m,

2000 m at Mbuzi Mt.).

Charaxes cedreatis Hewitson, 1874. forms vetula R., 1900, cedreatis H., inexpec-

tata v. Som., 1969. Rather common in open woodland and forests of Kigoma and

Mpanda. Record: 1, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b (800-2000 m).

Charaxes chepalungu van Someren, 1969. A few records from Kigoma and

Mpanda. One male also taken south of Biharamulo. Record: 4a, 6, 8a, 8b, Bihara-

mulo (900-1200 m). General distribution: east of Lake Victoria.

Charaxes ethalion nyanzae van Someren, 1967. forms ethalion B. and rosae B.,

1895. Common in open woodland, savanna and forest of Mpanda and Kigoma.

Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe (800-1700 m).

Charaxes ethalion nyasicus van Someren, 1975 (= nyasana van Som., 1967). Occur-

ring in the Ufipa area and probably the southern part of Mpanda. Record: Rukwa basin

(ca. 1000 m).

Charaxes nichetes pantherina R.-Decelle, 1934. Uncommon in forests of Mpanda

and Kigoma. Record: 1, 2a, 3a, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-1500 m).

Charaxes zelica toyoshimai Carcasson, 1964. Known from forests of Mukuyu, Mi-

humu, Kefu and Ntakatta in the sandstone area. Record: 5a, 8a, 8b (800-900 m,

178 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

1500 mat Ntakatta). First obtained by the Japanese Expedition from Kyoto.

Charaxes laodice Drury, 1782. Very local. Taken in low level forests at Kasoge,

Mukuyu, Mihumu, Kefu, Luntampa, Ntakatta and Lubalizi. Also a tattered male

taken at Gombe. Record: 1, 3a, 4a, 5a, 8a, 8b, Gombe (800-1200 m, 1500 m at

Ntakatta).

Charaxes zoolina zoolina Westwood, 1850. A male (w.s.f.) was taken in the Rukwa

basin in savanna at Muze (ca. 1000 m).

Palla publius kigoma van Someren, 1975. Flying in forests of Kigoma, Mukuyu and

Kefu forests. Uncommon and very local. Record: 8a, 8b (800-900 m).

Palla ussheri interposita Joicey & Talbot, 1925. Flying in low level forests of the

Kigoma area; Mukuyu, Mihumu and Kefu forests. Sandstone area only. Uncommon.

Record: 8a, 8b (800-950 m). The two Palla species were first collected by the

Japanese Expedition.

Subfamily Nymphalinae

Cymothoe theobene theobene Doubleday, 1848. The commonest Cymothoe species,

in riverine forests and heavy woodland and shrub throughout Mpanda and Kigoma.

Also one female taken near Chala in Ufipa at 1800 m. Some melanistic females are

nearly uniformly brown above. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Chala,

Gombe (800 to over 1800 m).

Cymothoe egesta confusa Aurivillius, 1887. Forests of Mpanda and Kigoma;

rather uncommon. Record: 1, 3a, 3b, 4a, 5a, 8a, 8b (800-1500 m).

Cymothoe lurida azumai Carcasson, 1964. Riverine forests of Mpanda and Kigoma.

Rather common. Record: 1, 2a, 3a, 4a, 5a, 8a, 8b (800-1500 m).

Cymothoe herminia johnstoni Butler, 1902. Rather common in forests of Mpanda

and Kigoma. Record: 1, 3a, 3b, 4a, Sa, 8a, 8b (800-1500 m).

Cymothoe coranus Grosse-Smith, 1889. A few specimens taken in forests of Mpanda

and Kigoma. Rare. One record from Wanzizi, from Kasoge, Ntakatta, Mihumu, Kefu

and Gombe. Record: 1, 3b, Sa, 8a, 8b, Gombe (800-1450 m).

Cymothoe caenis Drury, 1773. Low to rather high altitude forests of Mpanda and

Kigoma. Not uncommon. Record: 1, 3a, 5a, 6, 8a, 8b (800-1600 m). At Kasoge

2 f. rubida Holl., 1920, and conformis Aur., 1898; at Lukandamira ® f. caenis;

at Sibweza 9 f. rubida.

Cymothoe sangaris Godart, 1823; ssp. near luluana Overlaet, 1945. Forests of

Mpanda and Kigoma. More common in the sandstone area than south of the

Luegele river. At Wanzizi one record only. Record: 3a, 3b, 4a, 5a, 5b, 8a, 8b

(800-1500 m).

Pseudathyma plutonica expansa ssp. nov. (text-fig. 29, pl. 9 figs. 31-33). Several

males and females were taken, November 1971, along a forest river in the Ntakatta

forest. A few more specimens were taken in February and in November 1972. Record:

5a (1400-1500 m).

Male. Upperside fore wing discal white spots wider than in ssp. plutonica, partic-

ularly the spots from 2-6; all pale markings creamy, with only slight greenish tint;

hind wing band generally wider and its outer border incised at the ribs, making a wavy

outline; in ssp. plutonica the border is smooth, the inside less rounded, more uneven.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 179

Underside white markings as above, otherwise like the nominate race. Length of fore

wing 23 to 24 mm. Genitalia as in ssp. plutonica.

Female. As the male, but larger and the pale markings white and more extended.

Length of fore wing 26-27 mm. Genitalia as in ssp. plutonica.

Holotype 6 : Tanzania, Mpanda, Ntakatta forest, 1500 m, November 1971, J. Kiel-

land. Allotype : Same data and collector. Paratypes: 6 © and 6 6 ‚same data.

Holotype, allotype and paratypes are deposited in the National Museum, Nairobi.

Other paratypes in the British Museum (Nat. Hist.), London, and J. Kielland col-

lection.

This race has so far only been taken in the Ntakatta forest (Mpanda), close to the

Kigoma border, in sandstone formations. Early rainy season only.

Euryphura spec. A few specimens taken in the following places: Kasoge, Ntakatta,

Kefu, Kasha and at Mihumu. Rare. Record: 1, 5a, 8a, 8b (800-1400 m).

Occurrence February, April, September, October, November. The species was not re-

presented in the National Museum, Nairobi, and is probably undescribed.

Crenidomimas concordia Hopffer, 1855. Common in open woodland of Mpanda

and Kigoma. Record: 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b, 9a, 9b (800-

1600 m).

Euryphene itanii Carcasson, 1964. Flying close to the ground in dense forests of

Kigoma. Probably also in Mpanda in the north. Common. At low levels. Record: 1, 2a,

3a, 4a, 5a, 8a, 8b (800-1500 m).

Euphaedra spatiosa Mabille, 1877. Forests of Kigoma and Mpanda in the north-

west. Common. Record: 1, 2a, 3a, 4a, 5a, 5b, 8a, 8b, Gombe (800-1400 m).

Common in low forests, rapidly becoming scarcer at higher levels.

Euphaedra medon fraudata Thurau, 1903. Common in forests and forest margins

of Mpanda and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b, Gombe

(800-1700 m). |

Euphaedra sarcoptera nipponicorum Carcasson, 1965. Forests of Kigoma. Rare.

Records from Mihumu, Kapalamasenga river, Mugondozi river, Kefu forest, upper

part of Lukoma river near Mt. Karobwa. Record: 2a, 8a, 8b (900-1100 m). Collected

in March, June, September, October, November.

Euphaedra phosphor Joicey & Talbot, 1921 (= ceres Fabr. ssp. phosphor).

Low altitude riverine forests of Mpanda and Kigoma. Common. Record: 1, 2a, 3a, 4a,

5a, 8a, 8b, Gombe (800-1200 m, 1700 at Mahale).

Euphaedra nigrobasalis Joicey & Talbot, 1921. Probably an undescribed race.

Not uncommon in forests of Mpanda and Kigoma. Record: 1, 2, 3a, 3b, 4a, 4b, 5a, 5b,

6, 8a, 8b, Gombe (800-1500 m). Recorded as cooksoni katangensis Talbot, 1927,

by Carcasson (1966). According to M. Clifton (National Museum, Nairobi) this is E.

nigrobasalis.

Euphaedra cyanea Holland, 1920 (= E. xypete Hew. f. cyanea). So far only a few

males taken in the Kefu forest at Helembe. All in one year, 1970. Not seen since.

January and March. Record: 8b (950 m). Differs from typical cyanea and is probably a

new race.

Euphaedra coprates Druce, 1875 (= E. eleus Drury f. coprates). Forests of

Kigoma. Uncommon. Mukuyu, Mihumu, Kefu, Zanza river, Lubalizi, Luntampa,

Ntakatta. Record: 3a, 4a, 5a, 8a, 8b (800-1200 m, 1500 m at Ntakatta). Mention-

180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

ed by Carcasson (1966) as Najas eleus. This species differs considerably from eleus

and probably is a new race of coprates. Confirmed by M. Clifton (Nairobi).

Euphaedra zaddachi Dewitz, 1879. Forests of Mpanda and Kigoma. Not un-

common. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-1700 m).

Euphaedra mardania Fabricius, 1793 (= Euryphene mardania Fabr.), a new sub-

species. Common in low level forests and forest margins of Mpanda and Kigoma.

Record: 1, 2a, 2b, 3a, 4a, 5a, 5b, 8a, 8b, Gombe (800-1200 m, 1500 m at

Kampisa). This race is recorded from Ukerewe Island in Lake Victoria and at

Abercorn in Zambia.

Euphaedra sophus ochreata Carcasson, 1961. Forests of Mpanda and Kigoma.

Rather common in medium altitude forests of Wanzizi, less common further to

the north, e.g. at Sitwe, Kampisa, Lukandamira. In the sandstone formations it

occurs in low level forests. Not so in the basement area, except at Luntampa which

borders on the sandstone. Record: 1, 3a, 3b, 4a, Sa, 5b (900-2000 m).

Hamanamida daedalus Fabricius, 1775. Ubiquitous, throughout Mpanda, Kigoma

and Ufipa. In open woodland, savanna, forest and montane grassland. Record: wide-

spread (900-2200 m).

Aterica galene galene Brown, 1776. Very common in forest, bush and heavy wood-

land throughout Mpanda, Kigoma and Ufipa. Record: widespread (800-2200 m).

Pseudargynnis hegemone nyassae Bartel, 1905. Along rivers, forest margins and

areas flooded during the rainy season in Mpanda and Kigoma. Record: 1, 2a, 3a, 4a,

5a, 5b, 6, 8a, 8b, 9a (800-1500 m).

Catuna crithea Drury, 1773. Dense forests of Kigoma and possibly Mpanda in the

north, fluttering close to the ground and settling in spots of sun. Record: 1, 2a, 3a,

4a, Sa, 8a, 8b (800-1300 m, 1850 m in the Karobwa forest). The Karobwa forest

is high altitude forest, completely separated from lower forests by open grassland

and woodland further down. Also taken at Lukandamira, 1700 m.

Pseudacraea semire Cramer, 1779. Scarce. In forests of Mpanda and Kigoma. Rec-

ord: 1, 5a, 5b, 8a, 8b, Gombe (800-1500 m).

Pseudacraea boisduvali trimeni Butler, 1874. Forests and heavy woodland of Mpan-

da and Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b (800-1500 m).

Pseudacraea eurytus Linnaeus, 1758. Forests of Kigoma and Mpanda: Lubalizi,

Luntampa, Kasoge, Mihumu, Kefu, Ntakatta. Record: 1, 3a, 4a, Sa, 8a, 8b (800-

1500 m). The following forms occur: kunovoides Carp., 1930, conradti Obert., 1893,

opisthoxantha Carp., 1924, hobleyi Neave, 1904, bicolor Aur., 1898, ruhama Hew.,

1872, 2 f. terra Neave, 1904, 2 f. simulator B., 1873, 2 f. poggeoides Poult.,

1913, © f. fulvaria B., 1874, ® f. tirikensis Neave, 1904, f. terra ab. impleta Griinb.,

1910, È f. jacksoni Carp., 1949, f. grisea Carp., 1949.

Pseudacraea poggei Dewitz, 1879, forms poggei and carpenteri Poult., 1918. Wood-

land in Mpanda and Kigoma. Not common. Record: 6, 8a (800-1400 m).

Pseudacraea lucretia expansa Butler, 1887, with £ f. heliogenes Butler, 1896. Com-

mon in forests and heavy woodland throughout Mpanda and Kigoma. Record:

widespread (800-2000 m).

Pseudacraea deludens deludens Neave, 1912. Montane forests of Mahale in Kigoma

and Sitebi in Mpanda. Record: 1, 5b (1600-2000 m). Three specimens from Mahale

in December and June; eight specimens from Sitebi in September, 1970.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 181

Subfamily Neptidinae

Neptis saclava marpessa Hopffer, 1855. Common in forests, woodland and

savanna of Mpanda, Kigoma and Ufipa. Record: widespread, also at Gombe

(800-2000 m).

Neptis nemetes nemetes Hewitson, 1868 (text-fig. 12). Forests of Kigoma

and Mpanda. Very common in the sandstone formations, but much scarcer in the

basement area. Record: 4a, 5a, 6, 8a 8b, Biharamulo (800-1200 m, 1500 m

at Ntakatta).

Neptis serena Overlaet, 1955 (text-fig. 13). Woodland, savanna and forest

throughout Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b,

6, 7, 8a, 8b, 9a, 9b, Ufipa, Gombe (800-1800 m).

Neptis laeta Overlaet, 1955 (text-fig. 14). Same habitats as above. Common.

Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b, 9a, 9b, Ufipa,

Gombe (800-2200 m).

Neptis kiriakoffi Overlaet, 1955 (text-fig. 15). Woodland, savanna and riverine

forest of Mpanda and Kigoma. Very common. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b,

5a, 5b, 6, 7, 8a, 8b, 9a, 9b (800-2000 m).

Figs. 12—15. Male genitalia of Neptis species. 12, N. nemetes Hew., Mihumu, Kigoma, no. 480; a, valva,

lateral aspect; b, aedeagus. 13, N. serena Ovit., Sibweza, Mpanda, no. 182; a, valva, lateral aspect; b,

aedeagus; c, valva, dorsal aspect. 14, N. laeta Ovit., Sibweza, Mpanda, no. 271; a, valva, lateral aspect; b,

valva, dorsal aspect. 15, N. kiriakoffi Ovit.; a, typical valva, Sibweza, Mpanda, no. 179; b, atypical valva,

Morogoro, no. 347; c, atypical valva, Sibweza, Mpanda, no. 481; d, aedeagus.

182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Neptis gratiosa Overlaet, 1955 (text-fig. 16). Stunted woodland at rather high

altitude at Kampisa and at Ngondo and Mweze. Very local. Record: 3a, 5b

(1500-1700 m).

Neptis penningtoni van Son, 1953 (text-fig. 17). Brachystegia hills of Mpanda

and Kigoma. Not common. Record: 1, 3a, 4a, 5b, 6 (900-1700 m).

Neptis jordani Neave, 1910 (text-fig. 18). Riversides and marshy woodland.

Common. Record: 1, 2a, 2b, 4a, 4b, 5a, 8a, 8b (800-1200 m).

Neptis alta Overlaet, 1955 (text.-fig. 19). Brachystegia hills of Mpanda and

Kigoma. Common. Record: 1, 2b, 3a, 3b, Sa, 5b, 6, 8a, 8b (1000-1600 m,

1900 m at Mt. Sitebi).

Neptis metanira conspicua Neave, 1904 (= nysiades Hew. f. conspicua) (text-

fig. 20). Forests of Mihumu, Kefu, Luntampa, Lubalizi. Rare. Record: 3a, 4a, 5a, 8a,

8b (800-1200 m, 1500 m at Ntakatta).

Neptis puella Aurivillius, 1894 (text-fig. 21). Forests of Lubalizi, Kefu, and a forest

near Mt. Sitebi. Rare. Record: 3a, 5a, 5b, 8b (900-1700 m). General distribu-

tion: Ivory Coast, Cameroun to Zaire and Uganda.

Neptis trigonophora vansomereni Eltringham, 1929 (text-fig. 22). Rather common in

forests of Mpanda and Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b,

(800-1500 m).

Neptis strigata Aurivillius, 1894 (text-fig. 23). Not uncommon in forests of

Kigoma and Mpanda. Record: 1, 3a, 4a, 5a, 8a, 8b (800-1500 m).

Neptis nicoteles Hewitson, 1874 (text-fig. 24). Forests at Helembe, Mukuyu,

Luntampa, Lubalizi, Ntakatta. Rare. Record: 3a, 4a, 5a, 8a, 8b (800-1200 m, 1500 m

at Ntakatta). General distribution: Nigeria, Cameroun and Zaire to Uganda.

Neptis melicerta Drury, 1773 (text-figs. 25, 28). Mukuyu forest, Kefu forest,

Mugondozi river. Sandstone area only. Record: Sa, 8a, 8b (800-1400 m).

Neptis carcassoni van Son, 1959 (= melicerta Dr. ssp. carcassoni) (text-figs.

25, 28). Evergreen forests of Mpanda and Kigoma. Particularly in the sandstone

area. Record: 1, 3a, 4a, 5a, 5b, 6, 8a, 8b, 9b, Gombe (800-1500 m).

N. melicerta and N. carcassoni overlap considerably and should be recorded

as separate species. Except in area 5a (Ntakatta) N. goochi is only overlapped by car-

cassoni and could be considered a race of melicerta; however, the genitalia of all

three differ as shown in text-fig. 25.

Neptis goochi Trimen, 1879 (= melicerta Dr. ssp. goochi) (text-figs. 25, 28).

Forests of Mpanda and Kigoma. Common. Record: 1, 3a, 4a, 5a, 6, 8b (1000-

1500 m).

Neptis occidentalis occidentalis Rothschild, 1918 (text-fig. 26). Forests of Mweze

and Mahale in Kigoma. Not common and very local. Record: 1, 3a (1600-

2000 m).

Neptis ochracea reducta ssp. nov. (text-fig. 27, pl. 9 figs. 34-36). Montane

forests of Mpanda and Kigoma. Not uncommon. Record: 1, 3a, 5a, 5b (1500-2000 m).

Differs from the nominotypical race in reduction of the fore wing spots in space 1.

In all males examined the spots are reduced to a streak along the hind margin;

in one specimen hardly discernible. In ssp. ochracea there are two long spots in

space lb, usually conjoined and almost confluent with the spots 3 and 4, forming

a band. The yellow band of the hind wing is very variable, in most specimens

KIELLAND: Rhopalocera eastern side Lake Tanganyika 183

Pe DI

\ 27

SADE, Zar

21 b b a

Figs. 16—27. Male genitalia of Neptis species. 16, N. gratiosa Ovlt., Mweze, Mpanda, no. 613; valva,

lateral aspect. 17, N. penningtoni van Son, Sibweza, Mpanda, no. 186; valva; lateral aspect. 18, N. jordani

N., Sibweza, Mpanda, no. 183; valva, lateral aspect. 19, N. alta Ovit., Sibweza, Mpanda, no. 242; valva,

lateral aspect. 20, N. mentanira conspicua N., Mukuyu, Kigoma, no. 763; a, valva, dorsal aspect; b, val-

va, lateral aspect. 21, N. puella Aur., Lubalizi, Kigoma, no. 762; a, valva, dorsal aspect; b, valva, lateral

aspect. 22, N. triogonophora vansomereni Eltr., Sibweza, Mpanda, no. 475; a, valva, lateral aspect; b,

aedeagus. 23, N. strigata Aur., Kigoma, no. 486; a, valva, lateral aspect; b, aedeagus. 24, N. nicoteles

Hew., Ntakatta forest, Mpanda, 1500 m, no. 1464; a, gentitalia in lateral view with aedeagus and left

valva removed; b, valva in dorsal aspect. 25, N. melicerta melicerta Dr., a, valva, dorsal aspect; b, valva,

lateral aspect; N. carcassoni stat. n., c, valva, dorsal aspect; d, valva, lateral aspect; N. goochi Trim.; e,

valva, dorsal aspect; f, valva, lateral aspect. 26, N. occidentalis occidentalis Roth., Mweze forest, Kigoma,

no. 488; a, valva, lateral aspect; b, aedeagus. 27, N. ochracea reducta ssp. n., Lukandamira, Mahale,

Kigoma, no. 791; a, valva, lateral aspect; b, valva, dorsal aspect.

184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

much reduced and separated in space lb, in others wider and conjoined. The females

have much wider and paler hind wing bands and the fore wing pale spots less reduced.

Habitat. Montane forests and riverine forests in western Tanzania.

Holotype & : Tanzania, Mpanda, Mt. Sitebi, 2000 m, 2. v. 1970, J. Kielland. Allo-

type : Tanzania, Kigoma, Sitwe, 1700 m, 20. viii. 1960, J. Kielland. Paratypes:

Tanzania, Kigoma, Lukandamira, 1600 m, August 1970; 3 6 , 2 2, same, but October

1969; Mt. Sitebi, l.v. 1970, 16; Mpanda, Katuma, July 1972, 16 ; Sitwe, 29. ix.

63, 1 6. All. J. Kielland.

Holotype, allotype and paratypes in the National Museum, Nairobi; further para-

types in the British Museum (Nat. Hist.), London, and J. Kielland collection.

Neptis incongrua Butler, 1896. Flying sparingly in the Mbisi Forest, Ufipa (2300 m);

one female taken February 1978.

Subfamily Marpesiinae

Cyrestis camillus camillus Fabricius, 1781. Forests of Mpanda and Kigoma.

Particularly along streams. Record: 1, 2a, 3a, 4a, 5a, 5b, 8a, 8b (800-1500 m).

Subfamily Eunicinae

Sallya pechueli Dewitz, 1879. One male taken by T. H. E. Jackson’s collector

\ O melicerta

RER, Kasulu ‘ \

goma<_ I goochi

Ta PENARE | A ,

mn Uwinza = carcassoni

ee — —

= SN

5 ON \

\ À

Yo} \ A

fa LA \

e\_ a

)

Os f

N . 1

-. Kigoma’

N, 7

N DA er

È \Mpanda

EN à

“Ufipa 3

Lake

Tanganyika

\ Rukwa

Sumbawanga

Toll Zambia

Fig. 28. Distribution of Neptis species.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 185

Watulege in the Mpanda area. One male taken by me at Luegele in Kigoma,

1. xii. 1970, in open woodland. Record: 4a, Mpanda area (1100 m).

Sallya amulia rosa Hewitson, 1877. Woodland and swampy ground of Mpanda

and Kigoma. Record: 3a, 4a, 5a, 5b, 6, 8a, 8b (800-1300 m). Very common in certain

places of area 8a, 8b, particularly on hills near Helembe, in the dry season. Further

south less common and local.

Sallya occidentalium occidentalium Mabille, 1876. Very local in forests of Mpanda

and Kigoma. Record: 1, 4a, 5a, 5b, 8a, 8b (800-1500 m).

Sallya moranti dubiosa Strand, 1911. Forests and woodland of Mpanda and

Kigoma. Common. Record: 1, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-1500 m, 1900

m at Sitebi).

Sallya spec. Some males and females, closely related to moranti, were sent to the late

Mr. Peter Muteshi who was revising Sallya at the National Museum, Nairobi. Mr.

Muteshi was about to describe it as a new species. Forests of Luntampa, Lubalizi,

Kefu, Mukuyu, Ntakatta. Record: 3a, 4a, 5a, 8a, 8b (800-1200 m).

Sallya boisduvali boisduvali Wallengren, 1857. Forests and woodland of Mpanda

and Kigoma. Rather common. Record: 3a, 3b, 5a, 5b, 6, 8a, 8b, 9b, Gombe

(800-2000 m).

Sallya boisduvali omissa Rothschild, 1918. Record: K asoge forest at 1 (800 m).

Sallya garega garega Karsch, 1892. Forests of Mpanda and Kigoma. Not very

common. Record: 1, 3a, 3b, 4a, 5a, 5b, Gombe (1500 m).

Sallya umbrina Karsch, 1892. Common in forests and woodland of Kigoma

and Mpanda, Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 8a, 8b, Gombe (800-1600 m).

Subfamily Eurytelinae

Byblia acheloia acheloia Wallengren, 1857. Very common in woodland and savanna

of Mpanda, Kigoma and Ufipa. Record: widespread (800-2000 m).

Mesoxantha ethosea reducta Rothschild, 1918. Low level forests of Kigoma and

probably Mpanda in the north. Often occurring in small colonies. Local. Record:

1, 3a, 4a, 5a, 8a, 8b (800-1500 m).

Ariadne enotrea suffusa Joicey & Talbot, 1921. Forests and heavy woodland and

shrub in Mpanda and Kigoma. Common. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a,

8b, Biharamulo, Gombe (800-1500 m).

Ariadne enotrea archeri Carcasson, 1958. From Lubalizi, Sibweza, Kampisa and

other places. A doubtful subspecies since ssp. suffusa is found in the same areas.

Ariadne pagenstecheri Suffert, 1904. Forests of Mpanda and Kigoma. Common.

Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6 (1200-2300 m). A rather high altitude species. At

Kampisa, 1500 m, A. enotrea and A. pagenstecheri fly together. In most other places

they are not seen together. A. pagenstecheri more often flies at higher altitudes, while A.

enotrea frequents lower levels.

Neptidopsis ophione ophione Cramer, 1777. Very common in forests and woodland

of Mpanda, Kigoma and Ufipa. Record: widespread (800-2200 m).

Eurytela hiarbas hiarbas Drury, 1782. Forests throughout Mpanda, Kigoma and

Ufipa. Common. Record: widespread (800-2200 m).

Eurytela dryope angulata Aurivillius, 1898. Common in forests and heavy wood-

186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

land of Mpanda, Kigoma and Ufipa. Record: widespread (800-1700 m, 2000 m in

Ufipa).

Subfamily Vanessinae

Kallima rumia rattrayi Sharpe, 1904. Forests of Kigoma and Mpanda. Locally

common. Record: 1, 2a, 3a, 4a, Sa, 5b (800-1400 m, 1600 m at Lukandamira and

1700 m at Mt. Ipumba). Differs somewhat from ssp. rattrayi and may constitute an

undescribed race.

Kallima cymodoce Cramer, 1777, form lugens Schultze, 1912. Flying in low altitude

forests of Kigoma, in the sandstone area only. Local and uncommon. Mihumu,

K apalamasenga river, Mubondozi river, Kefu. Record: 8a, 8b (900-1100 m).

Kallima ansorgei Rothschild, 1899. The Ntakatta forest only. Very local. Record:

5a (1200-1500 m). K. ansorgei and K. rumia fly together, while neither of these have

been taken further inside the sandstone area where K. cymodoce occurs. Ntakatta is

situated on the border between sandstone and basement formations.

Apaturopsis cleocharis Hewitson, 1873. One specimen was observed by the author

in the Mukuyu forest in Kigoma. Record: 8a (800 m).

Hypolimnas misippus Linnaeus, 1764; © forms: misippus, inaria Cr., 1779, and

alcippoides B., 1883. Common in woodland and open habitats of Mpanda and

Kigoma. Record: widespread (800-1500 m).

Hypolimnas monteironis Druce, 1874. Luntampa and Ntakatta forests only. Very

rare and local. Record: 4a, 5a (1200-1500 m). This is probably an undescribed

race.

Hypolimnas dinarcha Hewitson, 1865. Kigoma area. Mukuyu forest, 16, Novem-

ber; Luntampa forest, May, June and March, a few specimens; Ntakatta forest Id.

Very rare and local. Record: 4a, 5a, 8a (800-1200 m). Intermediate between the

W. African and the Uganda race.

Hypolimnas dubius dubius Polisot de Beauvais, 1805. Common in forests of

Mpanda and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b (800-1700

m). Forms: dubius, damoclina T., 1869, anthedon Dbl., 1845.

Salamis temora Felder, 1867. Dense forests of Kigoma and the Ntakatta in

Mpanda. Record: 1, 4a, 5a, 8a, 8b, Gombe (800-1200 m, 1600 m at Lukandamira).

Salamis parhassus Drury, 1782; forms: parhassus and aethiops de Beauvais,

1805. Very common in forests and heavy woodland of Mpanda and Kigoma.

Record: widespread (800-2200 m).

Salamis anacardii Linnaeus, 1764. Common in woodland, savanna and forest

margins of Mpanda and Kigoma. Record: widespread (800-1600 m).

Salamis cacta cacta Fabricius, 1793. Not common in forests of Mpanda and

Kigoma, Mihumu, Kefu, Ntakatta, Lubalizi, Luntampa. Record: 3a, 4a, 5a, 8a, 8b

(800-1250 m).

Precis cloanthe Cramer, 1781. Very common in woodland and savanna of

Mpanda, Kigoma and Ufipa. Record: widespread (800-2200 m).

Precis touhilimasa Vuillot, 1892. Brachystegia woodland of Mpanda, Kigoma and

Ufipa. Preferring hills. Record: 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, Chala (1000-1800 m).

Precis artaxia Hewitson, 1864. Brachystegia woodland, savanna and grassland of

KIELLAND: Rhopalocera eastern side Lake Tanganyika 187

Mpanda, Kigoma and Ufipa. Common. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6,

7, 8a, 8b, Chala (800-1900 m). Forms: artaxia (d.s.f.) and nobilitata Thur., 1905

(w.s.f.). The latter is smaller than artaxia and occurs during the start of the rains, in

Nov. to Dec. Only tattered specimens of this form have been taken early January.

The form artaxia emerges late December and continues to emerge during the rainy

season, until May-June. From then on, during the dry season few specimens seem

to emerge. Specimens are increasingly tattered as the dry spell continues, until late

October.

Precis stygia gregorii Butler, 1895. Common in forests of SE and Kigoma.

Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe (800-1800 m).

Precis natalica natalica Felder, 1860. Common in woodland and savanna of

Mpanda, Kigoma and Ufipa. Record: widespread (800-1700 m).

Precis terea elgiva Hewitson, 1864. Very common in forests and woodland of

Mpanda, Kigoma and Ufipa. Record: widespread (800-2000 m).

Precis archesia Cramer, 1779; forms archesia (d.s.f.), intermediate f. chapunga

Hew., 1864, pelasgis Gt., 1819 (w.s.f.). Common in woodland and savanna of Mpan-

da, Kigoma and Ufipa. Record: widespread (800-1800 m).

Precis sinuata Plötz, 1880. Forests and woodland of Kigoma and Mpanda.

Rather common. Record: 1, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, 9a, 9b, Gombe

(800-1500 m).

Precis tugela Trimen, 1879. Forests and woodland of Mpanda and Kigoma. Rather

common. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6 (800-2000 m).

Precis rauana (?) kakamega Carcasson, 1961. Not uncommon in forests of Mpan-

da and Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b ( 1200-2000 m).

Precis actia Distant, 1880; forms: actia (d.s.f.), furcata R. & Jordan, 1903 (w.s.f.).

Woodland and savanna of Mpanda, Kigoma and Ufipa. Common. Record: wide-

spread (800-2000 m).

Precis pelarga Fabricius, 1775; forms: leodice Cr., 1777 (d.s.f.), harpyia F., 1781

(d.s.f.), pelarga (w.s.f.). Flying in woodland of Mpanda, Kigoma and Ufipa. Common.

Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6, 8b, Kishi in Ufipa, Gombe (800-1600 m).

Precis ceryne Boisduval, 1847; forms: ceryne (w.s.f.), tukuoa Wall., 1857 (d.s.f.).

Common in woodland and savanna of Mpanda, Kigoma and Ufipa. Record: wide-

spread (800-2000 m).

Precis antilope Feisthamel, 1850; forms: antilope (d.s.f.), simia Wall., 1857 (w.s.f.).

Common in woodland and savanna of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 3a,

3b, 4a, 5a, 5b, 6, 8b, Rukwa basin, Gombe (800-1700 m).

Precis cuama Hewitson, 1864. Taken in stunted woodland at Kampisa in Mpanda.

Very rare. Record: 5b (1500 m).

Precis octavia sesamus Trimen, 1883; forms: sesamus (d.s.f.), natalensis St., 1885

(w.s.f.), intermediate f. transiens Wich., 1918. Common in woodland and savanna

of Mpanda, Kigoma and Ufipa. Record: widespread (800-2100 m).

Precis sophia infracta Butler, 1888. Common in woodland and savanna of Mpanda,

Kigoma and Ufipa. Often flying in swampy areas and along river banks. Record:

1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Chala, Gombe (800-1800 m).

Precis westermanni westermanni Westwood, 1870. Two males taken Jan. 1973 in

the Ntakatta forest at a riverbed. Record: 5a (1400 m). General distribution: W. Afri-

188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

ca to Uganda and Kenya. Haldane (1969) recorded it from Ngara Distr. south of Buko-

ba, Tanzania.

Precis oenone oenone Linnaeus, 1764 (= clelia Cr., 1775). Common in open habitats _

and woodland of Mpanda, Kigoma and Ufipa. Record: widespread (800-2000 m).

Precis hierta hierta Fabricius, 1798 (= cebrene T., 1870). Open habitats and wood-

land of Mpanda, Kigoma and Ufipa. Common. Record: widespread (800-2000 m).

Precis orithya madagascariensis Guenee, 1864. Common in open habitats and

woodland of Mpanda, Kigoma and Ufipa. Record: widespread (800-2000 m).

Vanessula milca latifasciata Joicey & Talbot, 1928. Common in forests of Mpanda

and Kigoma. Record: 1, 2a, 3a, 4a, 5a, 5b, 6, 8a, 8b, 9b (800-1500 m).

Vanessa cardui Linnaeus, 1758. Ubiquitous in all open and wooded habitats of _

Mpanda, Kigoma and Ufipa. Record: widespread (800-2200 m).

Antanartia hippomene hippomene Hübner, 1806. Forests of Kigoma, Mpanda and

Ufipa. Not uncommon in highland. Record: 1, 3a, 5b, Mbizi Mt., Chala, Mbuzi Mt.

(1500-2300 m).

Antanartia schaenia dubia Howarth, 1966. Forests of Mpanda, Kigoma and Ufipa.

High altitude. Record: 1, 3a, 5b, Mbizi forest, Chala Mt., Mbuzi Mt. (1700-2300 m).

Antanartia delius Drury, 1782. Record: Gombe (900 m).

Subfamily Argynnidinae

Lachnoptera iole Fabricius, 1781. Ssp. intermediate between iole and ayresii Trimen,

1879. Common in forests of Mpanda and Kigoma. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a,

5b, 6, 8a, 8b, 9b (800-1500 m).

Phalanta columbina columbina Cramer, 1779. Common in forests of Mpanda,

Kigoma and Ufipa. Record: widespread (800-1800 m).

Phalanta phalantha aetiopica Rothschild & Jordan, 1903. Common in woodland,

savanna and open habitats of Mpanda, Kigoma and Ufipa. Record: widespread

(800-1600 m).

Issoria excelsior katangae Neave, 1910. Forests and forest margins surrounded

by open grassland in Ufipa. Record: Mbizi forest in Ufipa (2000-2300 m).

Family Acraeidae

Bematistes vestalis Felder, 1865. Forests of Kigoma in the sandstone area only.

Very local. Record: 8a, 8b (800-1100 m).

Bematistes alcinoe camerunica Aurivillius, 1893. A rare species occurring in the

Kasoge forest, Kefu, Luntampa and at Mihumu. September and January. Record:

1, 4a, 8a, 8b (800-950 m).

Bematistes umbra macarioides Aurivillius, 1893. A rare species, mainly in dense

forest of the sandstone area. Mihumu, Kefu, Ntakatta, Luntampa, Lubalizi. January,

February, April, September. Record: 3a, 4a, 5a, 8a, 8b (800-1400 m). General distri-

bution: From Ivory Coast to Nigeria, Cameroun and Zaire. Ssp. macarioides from

Central Zaire.

Bematistes aganice montana Butler, 1888. Rather common in forests and heavy

woodland of Mpanda and Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b (800-

KIELLAND: Rhopalocera eastern side Lake Tanganyika 189

1600 m). Females with very narrow fore wing white band occur at 2000 m on Mt.

Sitebi.

Bematistes scalivittata Butler, 1896. One female taken in the Mbisi Forest, Ufipa

(2200 m), February 1978.

Bematistes quadricolor itumbana Jordan, 1910. In montane forest of the Mahale

Mt. only, at Lukandamira. Very rare. Several males and one female taken August

1970 and August 1971. Record: 1 (1600-2000 m).

Bematistes poggei poggei Dewitz, 1879. Not uncommon in forests of Mpanda and

Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b (800-2000 m).

Bematistes macarista Sharpe, 1906. Locally common in forests of Mpanda and

Kigoma. Record: 1, 3a, 4a, 5a, 8a, 8b, Gombe (800-1500 m).

Bematistes persanguinea Rebel, 1914. Two males taken in the Mihumu forest,

Kigoma. September 1968. Very rare. Record: 8a (900 m). General distribution:

K ayonza forest, Uganda; Kasai in Zaire.

Bematistes epiprotea Butler, 1874. Dense forests of Kigoma in the sandstone

area only. Mukuyu, Mihumu, Kefu, Mugondozi, Mugombazi, northern Ntakatta.

Rather common. Record: 5a, 8a, 8b (800-1100 m, 1400 m at Ntakatta).

Bematistes epaea lutosa Suffert, 1904. Rather common in forests of Kigoma and

Mpanda. Record: 1, 3a, 4a, 5a, 8a, 8b (800-1200 m, 1500 m in the Ntakatta forest).

Differs slightly from typical ssp. lutosa. In the Kasekera forest at Gombe the author

has taken a male and a female of another race of epaea, differing from ssp. lutosa.

Acraea pentapolis epidica Oberthür, 1893. Very common in the Kasoge forest from

July to October, but not taken anywhere else in the study area. Its foodplant, a

Myrianthus species, is plentiful in this forest, but very scarce elsewhere. Record:

1 (800-1600 m). The only other place where a few Myrianthus trees have been

noticed, is at Mihumu and in a forest west of Mt. Kakungu. In February 1978,

larvae of pentapolis were observed on a Myrianthus tree at Mihumu (area 8a).

Acraea cerasa kiellandi Carcasson, 1964. Not common. In forests of Mpanda

and Kigoma. Record: 3a, 3b, 5a(1200-1500 m).

Acraea quirina quirina Fabricius, 1781. Forests of Kigoma and Ntakatta in

Mpanda. Record: 1, 5a, 8a, 8b, Gombe (800-1400 m).

Acraea kraka Aurivillius, 1893. Fifteen specimens were taken along a riverbed

in the Ntakatta forest, in half shade, September 1973, and a few more in 1974.

Although Ntakatta has been extensively collected by the writer for several years,

none had been seen before.

Acraea terpsichore neobule. Doubleday, 1847. Woodland of Mpanda, Kigoma and

Ufipa. Not uncommon. Record: 2a, 3a, 3b, 6, Rukwa basin (800-1200 m).

Acraea admata Hewitson, 1865. Forests of Kigoma and Mpanda at Ntakatta.

Common. Record: 1, 4a, 5a, 8a, 8b (800-1200 m, 1500 m at Ntakatta).

Acraea zetes zetes Linnaeus, 1758. Not very common. In woodland of Mpanda,

Kigoma and Ufipa. Record: 1, 3b, 4a, 5a, 5b, 6, 8a, 8b, Chala, Gombe (800-

2000 m).

Acraea anemosa Hewitson, 1865. Very common in savanna and woodland of

Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Chala

(800-1200 m, 1800 m at Chala).

Acraea pseudolycia astrigera Butler, 899. Not common in savanna of Mpanda

190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

and Kigoma, more common in Ufipa. Record: 3a, 3b, 4a, 6, 8a, 8b, Chala, Ruk- |

wa basin (800-1700 m). |

Acraea egina harrisoni Sharpe, 1904. Common in forest, open woodland and

grassland of Mpanda and Kigoma. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Ib,

Gombe (800-2000 m).

Acraea cepheus abdera Hewitson, 1852. Forests of Mpanda and Kigoma. |

Rare. Mukuyu, Kasoge, Wanzizi. Record: 1, 3b, 8a (800-1500 m). |

Acraea guillemei Oberthür, 1893. Savanna and woodland of Mpanda. Rare. Simbo, |

Kaliangkulukulu, Kabungu, Kapanda, Sibweza, Nkungwe. Record: Area 6 only (1000- |

1200 m). Wet season only. January to April. Not found on hills. |

Acraea chambezi Neave, 1910. Montane grassland of the Sitebi Mt. and Mt. Ipum-

ba at Mpanda. September 1970 and 1973. First East African records. Record:

5b (2000 m). General distribution: Zambia. |

Acraea periphanes Oberthür, 1893. Rather common. Locally very common in |

woodland and savanna of Mpanda and Kigoma. Record: 1, 3a, 3b, 4a, 4b, 5a, 6, 8a, 8b

(800-1300 m, 1700 m at Lukandamira). Forms with and without strong fore wing |

apical black patch. Also a form lacking discal spots on the fore wing particularly in |

the females. |

Acraea acrita taborensis Le Deux, 1932. Very common in woodland and savanna _

of Mpanda and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b, Gombe ©

(800-2000 m). |

Acraea chaeribula Oberthür, 1893. Wooded savanna of Mpanda at Sibweza,

Wanzizi and on flats south-west of Wanzizi. Rare and very local. August to Sep-

tember. Record: 2b, 3b, 6 (1100-1600 m).

Acraea leucopyga Aurivillius, 1904. One male taken at Chala in Ufipa, May 1972.

Open country. Record: Chala (1800 m). General distribution: Southern Tanzania

and Tunduma near Mbeya, Zambia and Malawi.

Acraea caldarena caldarena Hewitson, 1877. Common on savanna of Mpanda and

Ufipa. Also taken at Mt. Sitebi. Record: 3b, 5b, 6, Chala (1000-1300 m, 2000 m at

Sitebi).

Acraea pudorella pudorella Aurivillius, 1898. Not uncommon in woodland and sa-

vanna of Mpanda and Ufipa. Record: 6, Chala (ca. 1100 m, 1800 mat Chala).

Acraea rhodesiana Wichgraf, 1909. A single male taken in open woodland at

Sibweza. Record: 6 (1100 m). General distribution: Rhodesia. One specimen labelled

Dar es Salaam in the National Museum, Nairobi.

Acraea oncaea Hopffer, 1855. Woodland of Mpanda and Kigoma. Common in the

Lubalizi valley, rare elsewhere. One record from Ikola by T. H. E. Jackson.

Record: 1, 2b, 3a, 4a, Gombe (800-1700 m).

Acraea natalica natalica Boisduval, 1847. Common in woodland and savanna of

Mpanda, Kigoma and Ufipa. Record: widespread (800-1600 m).

Acraea anacreon bomba Grosse-Smith, 1898. Common in most habitats of Mpanda,

Kigoma and Ufipa. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b, Sumbawanga,

Chala, Mbuzi Mt., Gombe (800-1900 m).

Acraea rahira rahira Boisduval, 1833. Not uncommon in swampy plains and river-

sides of Mpanda and also at Lukandamira in Kigoma. Record: 1, 6, 7 (800-1100 m,

1600 mat Lukandamira).

KIELLAND: Rhopalocera eastern side Lake Tanganyika 191

Acraea encedon Linnaeus, 1758; forms daira G. & Salvin, 1884; infuscatoides Le

D., 1931; lycioides Le D., 1931. Very common in most habitats of Mpanda, Kigoma

and Ufipa. Record: 1, 2a, 2b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b, Chala, Gombe (800-

1700 m).

Acraea goetzi Thurau, 1903. Forest margins and grassland of Ufipa highland.

Common. Record: Mbizi forest, Chulwe Mt., Chala Mt., Mbuzi Mt. (1600-2000 m).

Acraea bonasia bonasia Fabricius, 1775. Forests of Mpanda and Kigoma. Not

common. Local. Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 5b (800-2000 m).

Acraea sotikensis sotikensis Sharpe, 1891. Common in forests of Mpanda, Kigoma

and Ufipa. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 8a, 8b, 9b, Chala, Mbizi Mt.

(800-2200 m).

Acraea cabira Hopffer, 1855. Heavy woodland and forest margins of Mpanda,

Kigoma and Ufipa. Common. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, 9a, Chala,

Gombe (800-2000 m).

Acraea acerata Hewitson, 1874. Common in woodland and riverine forests of

Mpanda, Kigoma and Ufipa. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, Chala, Mbuzi

Mt. (800-1700 m, 2000 m in Ufipa).

Acraea eponina Cramer, 1780. Very common in most habitats of Mpanda, Kigoma

and Ufipa. Record: widespread (800-2000 m).

Acraea ventura Hewitson, 1887. Not uncommon in forests, grassland along rivers

and open woodland of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, Sa, Sb, 6,

9b, Mbuzi Mt. (1000-1700 m, 2000 m at Mbuzi Mt.).

Acraea pharsalus pharsalus Ward, 1871. Common in forests and heavy woodland

of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, Kigoma,

8b, Mbuzi Mt. (800-2000 m).

Acraea perenna perenna Doubleday & Hewitson, 1847. Common in forests and

heavy woodland of Mpanda and Kigoma. Record: 1, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b,

Gombe (800-2000 m).

Acraea peneleos pelasgius Grosse-Smith, 1900. Forests of Kigoma and probably

Mpanda in the north. Not common. Record: 1, 8a, 8b (800-1000 m). General

distribution: Cameroun to Zaire, Uganda and W. Kenya.

Acraea penelope penelope Staudinger, 1896. Forests of Kigoma and probably Mpan-

da in the north. Rare. Record: 8a (900 m). General distribution: Zaire to Uganda

and W. Kenya.

Acraea ntebiae kigoma ssp. nov. (pl. 10 figs. 37-42). Differs from the nominate

subspecies in the very variable colour of the underside; in some males the colour is as in

the nominate subspecies, in others ochreous brown; upperside fore wing median spots

creamy, in some of the females hind wing ground colour orange red and the spots in the

fore wing space 2 and cell red. Females with ground colour invading the broad, black

marginal band in streaks nearly to the margin.

Habitat. Forests of the Mahale Mt. Mainly at high levels.

Holotype é . Tanzania, Kigoma, Lukandamira, 1600 m, 17. viii. 1970, J. Kielland.

Allotype 2 : as holotype. Paratypes: 4 d and 7 2 same data; 2 ó same, but 1.xi.

1972; 16 Kigoma, Ntondo, August 1971. Holotype, allotype and paratypes in the

National Museum, Nairobi; further paratypes in the British Museum (Nat. Hist.), Lon-

don, and J. Kielland collection.

192 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Acraea servona rhodina Rothschild & Jordan, 1905. Forests of Mpanda and Kigoma. |

Not very common. Record: 3a, 4a, 5a, 5b, 8a, 8b (800-1500 m). |

Acraea oreas oreas Sharpe, 1891. Forests of Mpanda, Kigoma and Ufipa. Rather -

common. Record: 1, 2a, 3a, 3b, 4a, Sa, 5b, 8a, 8b, Mbizi forest, Gombe (800- _

2000 m).

Acraea cinerea Neave, 1904. Collected by the Japanese expedition at Mahanga in

Kigoma, by T. H. E. Jackson’s African collector Watulege at Mahale. Record:

1, 8a. Altitude not specified.

Acraea quirinalis Grosse-Smith, 1900. A few specimens taken in the Ntakatta forest

flying along a riverbed and settling on wet sand. Very rare. Record: 5a (1400-1500 m).

General distribution: Forests from eastern Zaire to Uganda, W. Kenya and Tanzania

in the Usukuma area.

Acraea amicitiae amicitiae Heron, 1909. Montane forest of the Mahale Mt. only.

Record: 1 (1600-2000 m). General distribution: Burundi and Ruanda to W. Uganda.

Acraea alciope alciope Hewitson, 1852. Forests of Mpanda and Kigoma. Not

common. Record: 3a, 3b, 4a, 5a(1200-1500 m).

Acraea jodutta jodutta Fabricius, 1793. Forests of Mpanda and Kigoma. Not

common. Record: 1, 3a, 3b, 4a, 5a, 8a, 8b (800-1600 m).

Acraea esebria Hewitson, 1861. Forests of Kigoma and Mpanda. Common. Re- ©

cord: 1, 3a, 3b, 4a, 5a, 8a, 8b, Gombe (800-1600 m).

Acraea lycoa bukoba Eltringham, 1911. Rather common in forests of Mpanda and

Kigoma. Record: 1, 3a, 4a, Sa, 8a, 8b (800-1300 m).

Acraea johnstoni johnstoni Godman, 1885. Common in forests of Mpanda. Kigoma

and Ufipa. Record: 1, 3a, 3b, 4a, 4b, 5a, 5b, Chulwe Mt. (800-2000 m). Form

octobalia Karsch, 1894, at Lukandamira and Ntondo.

Pardopsis punctatissima Boisduval, 1833. In woodland at Sibweza, along the Uwin-

za road, Luntampa and at Mihumu. Very local and sporadic. Record: 4a, 6, 8a, 9a, 9b

(1060-1400 m).

Family Libytheidae

Libythea labdaca labdaca Westwood, 1851. Forests of Mpanda and Kigoma. Not

common. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-1800 m).

Family Riodinidae

Abisara neavei mahale ssp. nov. (text-fig. 30, pl. 14 figs. 54, 55). Resembling

the Cameroun race latifasciata Riley, 1931, in the wide, white discal band of both

wings, but in ssp. mahale the band is still wider, the hind wing ocellar yellow

ring is situated further from the margin, 0.8-1.1 mm in mahale, 0.5-0.6 mm in

other subspecies. On the underside of fore and hind wing subbasal and subtornal bands

narrower and fainter than those of the nominate race. The sexes are similar, but the

wings of the female are broader and more rounded than in the male. Length of fore

wing Ó 21 to 22 mm, £ 21.5 mm. Genitalia 4 same as ssp. neavei.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 193

Habitat. A somewhat weak, skipping flight amongst forest undergrowth, on which it

frequently settles.

Holotype 6 : Tanzania, Kigoma, Kasoge, 7.vi.1968, J. Kielland. Allotype OAs

holotype, 26.x.1967. Paratypes: 8 © and 1 ® of same data and collector. Holotype,

allotype and paratypes in the National Museum, Nairobi, further paratypes in the

British Museum (Nat. Hist.), London, and J. Kielland collection. This subspecies is

very common in the Kasoge forest on the western slope of Mt. Kungwe in the Mahale

Mt. range (800-1600 m) and has been taken at all times of the year. On the eastern

slope of the Mahale Mt. only a few specimens were collected at Lukandamira, 1600 m.

Away from Mahale, no trace has so far been found of this race.

Family Lycaenidae

Recent authors tend to divide this very large group of Rhopalocera into many

subfamilies; H. K. Clench (see Fox et al., 1965) even divides them into three separate

families, viz., Liptenidae, Liphyridae and Lycaenidae. There are still many uncertain-

ties and disagreements amongst specialists regarding the classification of the Lycaeni-

dae. In the following classification I have followed Eliot (1973), who divided the family

into 8 subfamilies of which 6, namely Lipteninae, Liphyrinae, Miletinae, Theclinae,

Lycaeninae and Polyommatinae are represented in Africa.

Subfamily Lipteninae

Alaena aurantiaca Butler, 1883. Occurring in the Ufipa area and also taken by the

Japanese Exp. at Mihumu and by the author near Mugombazi river. Very rare and

local. Record: 8a, 8b, Ufipa (in Kigoma 900-1200 m; in Ufipa not specified). General

distribution: Zambia.

Alaena kiellandi Carcasson, 1965. Very local, but sometimes rather common in

Mpanda and Ufipa. Mainly taken in December and a second generation sometimes

occurs in March. In open woodland, settling on rocks and grass. Sibweza, Sitwe,

Lugala Hill, Lungerengere, Chala. Record: 3a, 6, Chala in the ravine of Mtozi river

(1100-1600 m).

Alaena reticulata Butler, 1896. Montane grassland of Mt. Sitebi in Mpanda, 3 6

and 1 ® , 20.xii. 1970. Record: 5b (1950 m). General distribution: Mbeya in Tanzania

and Malawi.

Alaena bicolora Bethune Baker, 1924. Recorded by T. H. E. Jackson. Taken by his

collector Watulege in the Mpanda area. Record: no place or altitude specified.

Alaena spec. Two males and one female taken flying in a rocky place on a Brachyste-

gia clad hill close to Ntakatta. Record: 5a (1300 m).

Pentila inconspicua Hamilton H. Druce, 1910. Common in forests in Kigoma. Rec-

ord: 2a, 3a, 4a, 5a, 8a, 8b (1000-1500 m).

Pentila nyassana clarensis Neave, 1903. Common in heavy woodland, forest

margins and clearings of Mpanda and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a,

5b, 6, 8a, 8b, Gombe (800-1500 m).

Telipna sanguinea kigoma ssp. nov. (pl. 11 figs. 43-45). Closest to the nomino-

typical race from W. Africa and Zaire, but larger in both sexes; wings, upperside

194 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

ground colour paler; size of the subapical white patch of the fore wing inter-

mediate between ssp. sanguinea and ssp. depuncta Talb., 1937, from Uganda. The apic-

al black area more extended than in the other two races; inner margin of the black

area irregular and sharply angled basad, along vein 2, more so than in ssp. sanguinea.

Underside hind wing differs from the nominate race in the two long, costal transverse

streaks which are situated a little further apart. In the female the streaks are

Figs. 29—34. Male genitalia of Nymphalidae and Lycaenidae. 29, Pseudathyma plutonica expansa ssp.

n., holotype, Ntakatta forest, Mpanda, 1400 m, no. 1424; a, genitalia without aedeagus; b, aedeagus. 30,

Abisara neavei mahale ssp. n., holotype, Kasoge, Mahale, Kigoma, no. 510; a, tegumen-uncus; b, valva;

c, aedeagus. 31, Jolaphilus montana sp.n., paratype, Ujamba, Mahale, Kigoma, 2000 m, no. 406; a,

tegumen-uncus; b, valva. 32, Virachola diocles Hew., Chala, Ufipa, no. 1407; a, valva; b, subunci.

33, Virachola lorisona Hew., Uluguru Mt., Morogoro, no. 1620; a, valva; b, subunci. 34, Virachola ufipa

sp. n., paratype, Chala, Ufipa, no. 1408; a, valva; b, subunci.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 195

more extended towards the tornus, only 1.5 mm from the marginal, black border area;

in the females of the two other races this distance is 4 mm. Length of fore wing 4

25mm, 2 28mm.

Habitat. Riverine forest, particularly close to rivers, in half shade. Sitting for long

periods on the vegetation and flying up only when disturbed. The flight is rather weak

and reminds of the black and red Arctiid moths. Flight period from early rainy season

in November till the end of the rains in early May. Record: 3a, 4a, 5a, 8a, 8b (900-

1400 m).

Holotype 6 : Tanzania, Kigoma, Lubalizi, 6.iii. 1970, J. Kielland. Allotype® :

Same, but 11.iii.1970. Paratypes: Same, but 7.iii.1970, 2 6 ; same, but March 1970,

2 © ; Kefu, February 1972, 3 d , 1 2 ; same, but January 1972, 1 6 ; same, but April

1972, 19; same, but January 1973, 29; Ntakatta, 11.vi. 1974, 19 ; Mihumu, 18. xi.

1971, 1 © . Holotype, allotype and paratypes in the National Museum, Nairobi,

further paratypesin the British Museum (Nat. Hist.), London, and in J. Kielland col-

lection.

Ornipholidotos peucetia peucetia Hewitson, 1866. Local, but sometimes common in

heavy woodland and forest of Mpanda and Kigoma. Record: 2a, 6, 8a (800-1200 m).

Ornipholidotos overlaeti Stempffer, 1947. Very local and scarce in riverine forests

of Lubalizi, Ntakatta and Kefu. Rainy season only. Record: 3a, 5a, 8a, 8b (900-

1200 m). General distribution: Kenya and Uganda to Zaire.

Mimacraea krausei obsolescens Hawker Smith, 1926. Not uncommon, but local, in

riverine forests of Mpanda and Kigoma. Record: 2b, 3a, 4a, 6, 8a (900-1350 m).

Ovipositing on tree-trunks five feet above the ground. This species was listed as M. skop-

toles Ham. H. Druce by Carcasson (1966) and by Kielland (1968). According

to Stempffer it should be krausei.

Mimacraea marshalli marshalli Trimen, 1898. Not uncommon, but local in

Brachystegia woodland of Mpanda and Ufipa. Record: 3b, 5a, 6, 9b, Chala (1200-

1700 m).

Baliochila hildegarda Kirby, 1887 (= Teriomima aslauga Trim. f. hildegarda). Not

uncommon in woodland of Mpanda and Kigoma. Record: 2a, 3a, 3b, 4a, 4b, 5a, 5b,

8a, 8b (800-1500 m). Very variable and easily mistaken for other species. Reliable

identification only by dissection of the genitalia. The uncus of specimens from western

Tanzania differs somewhat from those of specimens from the eastern, central and

southern part of the country. It possibly indicates that several races are involved.

Baliochila lipara Stempffer & Bennet, 1953. One female taken by the Japanese Exp.,

December 1961, at Mukuyu. Record: 8a (ca. 900 m). General distribution: Rho-

desia, Zambia, Malawi, Kenya and East Tanzania.

Cnodontes vansomereni Stempffer & Bennet, 1953. In woodland and savanna of

Mpanda. Not common. Record: 6 (1050-1200 m).

Toxichitona gerda Kirby, 1890. In montane forest at Mt. Sitebi in Mpanda. One

specimen only, sent to Dr. Stempffer in Paris. Record: 5b (1900 m). General distri-

bution: Uganda.

Liptena xanthostola Holland, 1890. Common in riverine forests of Kigoma and

Mpanda. Record: 2a, 3a, 4a, 5a, 8a, 8b (900-1500 m). General distribution: W. Africa

to Zaire and Uganda.

Liptena ideoides Dewitz, 1886. Bushland along forest margins of Kigoma and into

196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Mpandaat Luegele river. Not uncommon. Record: 1, 2a, 3a, 4a, 5a, 8b (900-1400 m).

Liptena ilma ugandae Stempffer, 1964. In dark places of the Kasoge forest only.

Record: 1 (800-1000 m). General distribution: ssp. ugandae, Uganda; ssp. ilma, W.

Africa to Zaire. The Kasoge specimens differ somewhat from ssp. uganda.

Larinopoda tera Hewitson, 1873. Flying locally in unergrowth of forest at Mi-

humu, Kefu and Mugondozi rivers. Sandstone area only. Record: 8a, 8b (800-1000 m).

General distribution: W. Africa to Uganda.

Epitola miranda vidua Talbot, 1935. One female, June 1968, in riverine forest of

Lubalizi; one male Kefu forest, February 1971; one male, Ntakatta forest. Record:

3a, 5a, 8b (800-1600 m). General distribution: ssp. vidua, Uganda to Moyen Congo;

ssp. miranda, Ivory Coast.

Epitola pseudoconjuncta Jackson, 1962. A few specimens taken in the following

forests: Lubalizi, Kasha near Kefu, Ntakatta. Early morning and sometimes late

afternoon. Record: 3a, 5a, 8b (1000-1500 m).

Epitola kamengensis Jackson, 1962. January 1962 1 é taken in a Lubalizi forest,

other specimens observed; also males observed whirling over tops of bushes at forest

margin at Kapapa. Sent to Dr. Carcasson. Record: 3a, at 4a observed, 8b (1200 m).

General distribution: Uganda, W. Kenya. Early morning.

Epitola katera Jackson, 1962. One male taken February 1973 at the edge of the

Kefu forest, Kigoma and observed in the Ntakatta forest, early morning. High

flying species in tree tops and very difficult to catch. Record: 5a observed, 8b

(900 m, 1400 m at Ntakatta). As the specimen taken differs somewhat from katera,

it may represent anew race, or even an undescribed species.

Deloneura ochrascens littoralis Talbot, 1935 (acc. to Carcasson, pers. comm.). Not

common. In woodland of Mpanda and Kigoma; Nkungwe, Simbo, Ntakatta, Mihumu,

Mugombazi. Record: 5a, 6, 8a, 8b (900-1700 m).

Deloneura subfusca Hawker Smith, 1933 (= innesi van Son, 1949). Woodland at

Simbo in Mpanda. Very rare. Record: 6 (1200 m). General distribution: Katanga

and Malawi.

Hewitsonia similis ugandae Jackson, 1962. One female taken in a Mihumu forest.

Record: 8a (900 m). General distribution: Ivory Coast, Gabon, Ghana, to Zaire and

Uganda.

Subfamily Liphyrinae Doherty, 1889

Euliphyra species near leucyanea Hewitson, 1874. One female at riverine forest

margin, Lubalizi (3a), 1150 m, 30.xi.1970. The specimen flew before sunrise and

showed a peculiar behaviour. It soared up from the ground into the air and then

fluttered down like a dead leaf. It is larger than E. leucyanea, ground colour

upperside sooty brown with a pale area in the middle of the fore wing; colour and shape

of the hind wing as in E. leucyanea; underside similar to that of E. leucyanea, but bands

narrower and tending to be divided into round spots, while in E. leucyanea the bands

are more contiguous. E. leucyanea occurs from Sierra Leone to Cameroun.

Aslauga vininga Hewitson, 1875. One male May 1971, several males and two

females May 1972, taken in the Kefu forest. Settling on forest vegetation. Record:

8b (800-900 m). General distribution: West Africa to Uganda. The Tanzania speci-

KIELLAND: Rhopalocera eastern side Lake Tanganyika 197

mens are larger than specimens from other parts of Africa.

Aslauga lamborni Bethune Baker, 1913. One female taken in the Kasekera forest

at Gombe, December 1972; one female at Ntakatta. Record: 5a, Gombe (900 and

1400 m). General distribution: W. Africa to Uganda.

Aslauga purpurascens marshalli Butler, 1898. Not uncommon in woodland of

Mpanda and Kigoma. One female taken in riverine forest of Lubalizi. Record: 3a, 4a,

4b, 5b, 6, 8b (1000-2000 m).

Subfamily Miletinae Corbet, 1939

Spalgis lemolea H. H. Druce, 1890. Common in heavy woodland and forest of

Mpanda and Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Biharamulo

(800-1700 m).

Spalgis jacksoni Stempffer, 1967 (acc. to Stempffer). Flying along a stream in the

Kasha forest north of Kefu; settling on leaves of small trees and on wet sand; January

1972 and 1973. Record: 8b (1000 m). General distribution: Bwamba forest and Mon-

giro in Uganda.

Lachnocnema bibulus Fabricius, 1793. Common in Brachystegia woodland of

Mpanda and Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, 9a, 9b, Gombe

(800-1700 m).

Lachnocnema durbani Trimen, 1887. In woodland and riverine forests of Mpanda

and Kigoma. Not common. Record: 1, 3a, 4a, 5a, 5b, 8a, 8b (800-1600 m).

Lachnocnema spec., probably divergens Gaede, 1915. A male and a female of a

larger species taken at Simbo in Mpanda. Sent to Dr. Stempffer in Paris. Record:

6 (1100 m). General distribution of L. divergens: Uganda, Kenya, and Amani in Tan-

zania.

Subfamily Theclinae Swainson, 1831

Myrina silenus ficedula Trimen, 1879. Near Ficus trees at Sibweza and in open

grassland near Mweze. Record: 1, 3a, 4a, 5a, 5b, 6, Ufipa, Gombe (800-1750 m).

Oxylides faunus albata Aurivillius, 1895. Rather common in forests of Mpanda

and Kigoma. Particularly in the sandstone area. Record: 1, 2a, 4a, 5a, 5b, 8a, 8b (800-

1500 m). General distribution: Zaire to Uganda.

Aphnaeus orcas Drury, 1782, ssp. near hollandi Butler, 1902. Flying sparingly in

forests of Mpanda and Kigoma, Kasoge, Ntobo river near Mt. Sitebi, Lubalizi,

Mihumu, Ntakatta. Record: 1, 3a, 5a, 5b, 8a (800-2000 m).

Aphnaeus marshalli Neave, 1910. Taken by the Japanese Exp. at Mihumu and one

male at Kampisa by the writer. Extremely rare. Record: 5b, 8a (ca. 900 m). General

distribution: Rhodesia to Zambia and K atanga. È

Aphnaeus eriksoni kiellandi Stempffer, 1973. Mainly in open habitats of Mpanda

and Kigoma and one male at Chala in Ufipa. Mt. Sitebi, Mt. Kafulungu. Also taken

on Brachystegia clad hill at Nkungwe on flowering shrub. Not uncommon. Record:

3a, 4a, Sb, 6, Chala (1150 m at Nkungwe, 1900 m at K afulungu and Sitebi).

Aphnaeus flavescens flavescens Stempffer, 1954. One large female taken on wet sand

at Lubalizi river, September 1971. This is an extremely rare species. Record: 3a (1150

198 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

m). The specimen is larger than the types; length of forewing 23.5 mm.

Spindasis natalensis Westwood, 1852. Brachystegia woodland of Mpanda. Not un-

common. Record: 3b, 6 (1000-1500 m).

Spindasis nyassae Butler, 1884. Woodland of Mpanda at Nkungwe. Record: 6

(1100 m).

Spindasis victoriae Butler, 1884. Woodland of Mpanda and Kigoma. Common. Rec-

ord: 3a, 4a, 5b, 6, 8a, 8b, Gombe (900-2000 m).

Spindasis apelles Oberthür, 1878. Mainly open montane habitats, but also wood-

land of Mpanda, Kigoma and Ufipa. Common. Record: 1, 2b, 3a, 3b, 5a, 5b, 8a, 8b,

Mbuzi Mt., Chala (900-2000 m).

Spindasis nairobiensis Sharpe, 1904. Woodland of Mpanda and Kigoma. Uncom-

mon. Record: 2b, 3a, 3b, 5b, 6, 8a, 8b (900-2000 m).

Spindasis mozambica Bertoloni, 1851. Woodland and open grassland of Mpanda.

Kigoma and Ufipa. Common. Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Chala,

Gombe (900-2000 m).

Spindasis trimeni Neave, 1910. Common in woodland of Mpanda, Kigoma and

also taken at Muze in Ufipa. Record: 3a, 3b, 5b, 6, Muze (1000-2000 m).

Spindasis homeyeri Dewitz, 1886. Open habitats and woodland of Mpanda, Kigoma

and Ufipa. Rather common. Record: 1, 3a, 3b, 5b, 4a, 8b, Ufipa highland (900-

2000 m).

Spindasis ella Hewitson, 1865. Savanna and Brachystegia woodland of Mpanda

at Nkungwe. Rare. Record: 6 (1050 m).

Spindasis brunnea Jackson, 1965. (Carcasson, personal communication). Taken in

open habitat at Lubalizi river and at Sitebi Mt., 1 d and 1®. Record: 3a, 5b (1200

and 1700 m).

Spindasis menelas H. H. Druce, 1907. Riverine forest. One female taken on

vegetation at Katuma river near Mt. Sifuta. Extremely rare; known from Nigeria.

Male unknown. Record: 5b.

Spindasis crustaria Holland, 1890. Evergreen forests at Kasoge, Mihumu and

Gombe. Record: 1, 8a, Gombe (800-950 m).

Lipaphnaeus loxura Rebel, 1914. Common in the Kasoge forest of Kigoma, but

elsewhere scarce. Record: 1, 4a, 5a, Gombe (800-1700 m).

Lipaphnaeus aderna spindasoides Aurivillius, 1915. Forests of Mahale, Sitebi and

Mbuzi Mt. in Ufipa. Local. Record: 1, 5b, Mbuzi Mt. (1660-2000 m).

Chloroselas argentea Riley, 1932. Woodland of Mpanda. Uncommon. Record:

5b, 6, 7 (1000-1500 m). General distribution: Rhodesia.

Chloroselas minima Jackson, 1965. Woodland of Mpanda. Rare. Record: 6, 7 (900-

1100 m).

Chloroselas overlaeti Stempffer, 1956. Taken by T. H. E. Jackson’s collector Watu-

lege in Mpanda, by Zuzuki of the Japanese Exp. at Mihumu and by the writer near

Kigoma. Very rare. Record: 8a, Mpanda, Kigoma just east of town (altitude not

specified).

Chloroselas pseudozeritis tytleri Riley, 1932. This species occurs in Zambia and

northern and eastern parts of Tanzania and probably also in Mpanda and Kigoma

areas. Likely habitat: the Rukwa valley.

Axiocerses bambana Smith, 1900. Woodland and savanna of Mpanda, Kigoma and

KIELLAND: Rhopalocera eastern side Lake Tanganyika 199

Ufipa. Very common. Record: widespread (800-2000 m). Numerous specimens of

bambana have been dissected. A. bambana is very variable and the genitalia also vary

to some extent, but apparently not in accordance with external variation. It is therefore

not probable that this name covers more than one species. Specimens examined from

other parts of Tanzania, viz., Oldeani, Arusha, Singida, Morogoro, Mikumi, Ruaha

and Ngara, were all bambana.

Axiocerses amanga Westwood, 1881. Woodland and savanna of Mpanda, Kigoma

and Ufipa. Less common than A. bambana. Record: widespread (800-1700 m).

Aloeides conradsi angoniensis Tite & Dickson, 1973. Open habitats of Mpanda and

Kigoma. Particularly at high levels, in August to September. Very rarely taken in the

rainy season. Record: 2a, 3a, 3b, 5a, 5b, 6, 8a, at the ridge of Kararumpeta Mt. (1000-

2000 m).

Aloeides molomo kiellandi Carcasson, 1961. Open montane grassland of Mpanda

and Kigoma. Locally common. August to September only. Record: 2a, 3a, 3b, 4a, 5b

(1200-2000 m).

Tolaus bolissus aurora Clench, 1964. Riverine forests and forest margins of Mpanda

and Kigoma. Not uncommon. Record: 1, 3a, 4a, 5a, 5b, 8a, 8b (800-1700 m).

General distribution: Uganda to Zaire, W. Africa and Angola.

Stugeta bowkeri subinfuscata Grinberg, 1910. Scarce in woodland of Mpanda

and Kigoma. Record: 3a, 6, 8b (1000-1700 m).

Argiolaus crawshayi littoralis Stempffer, 1958. One female July 1970, Chala; one

female May 1971, Ntondo at Mahale. Record: 1, Chala(1700 m).

Argiolaus silas silarus H. H. Druce, 1885. Woodland and forest margins of Mpan-

da. Record: 3b, 5a, 6 (1050-1400 m).

Iolaphilus paneperata Druce, 1890. Riverine forests of Mugondozi, Mihumu and

Ntakatta. Rare. Record: 5a, 8a, 8b (1000-1400 m). General distribution: W. Africa to

Zaire.

Iolaphilus trimeni Wallengren, 1875. Brachystegia hills at Nkungwe in the Lubalizi

valley and at Mt. Sitebi. Rare. Record: 3a, 5b, 6(1150-2000 m).

Iolaphilus ndolae Stempffer, 1958. One male of this rare species was taken in a

montane forest of the Mbizi Mt., 10 miles east of Chala. The female has not yet been

described. Record: Mbuzi Mt. (2000 m), August 1973. General distribution: Zambia.

Iolaphilus montana spec. nov. (text-fig. 31, pl. 11 figs. 46-49). Male. Underside body

white; legs white; head black above, white below, one faint white spot on vertex top,

one faint white spot just above each eye; frons black, edged with white, finely dorsad,

„broadening ventrad; palpi second segment very long, first and second segment white,

last segment black, finely sprinkled with white near base laterally and ventrally; all seg-

ments narrowly black dorsally; antennae missing. Upperside fore wing iridescent

greenish blue, black border indented, at vein 2 two mm wide, in lb three mm,

broadening apically, 1.5 mm wide at basal half of costa. Inner margin lobed just

opposite sexual spot of hind wing; basal part of costa strongly convex.

Upperside hind wing iridescent greenish blue, costa and margins black, 1.5 mm wide;

a red, black bordered, tornal spot, a blue streak in 2, a brown androconial patch

partly in subcostal area partly in cell. Underside fore wing white, an ochreous

submarginal line slightly curving inwardly towards apex; margin finely bordered in

black; cilia white and sprinkled with black; an ochreous sexual hair tuft in space 1b.

200 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Underside hind wing white; a straight, ochreous line from red submarginal spot in 2 to

vein 8, at 2/3 from base to apex; a wavy, black subtornal line from vein 1b to near vein

2: a tornal red and black spot and a submarginal red spot in 2; a faint dusty

submarginal area, widest in 4 and 5. Length of fore wing 20 mm.

Genitalia. Uncus composed of two triangular lobes, separated by a gentle median

depression; in the middle of this, two small rounded knobs; subunci robust, gently

curved, bluntly tipped; vinculum broad, one to two small triangular dentations near

tegumen; valva ovaloid, tapering to a blunt apical tooth; two-third from base on the

ventral margin, an equally blunt, inwardly directed, strong tooth; aedeagus widely

open dorsally; vesica enclosing one long, spine-like cornutus; a short, thorny protuber-

ance apically on the right side; lower fultura very robust.

Female. Head and palpi as in the male; antennae black. Upperside fore wing

dark sky-blue basally up to middle of cell and along hind margin, invading 1b; a large

median, white area, invaded by blue scales, particularly along the veins; costa black;

apex and outer margin broadly black.

Upperside hind wing ground colour sky-blue, darker close to base and paler distally;

costa and inner half of area 6 whitish gray; apex and margin black; a large red

blotch in 1b, conjoining another in 2; anal lobe carmine red; discal black spots in 1b,

2, 3, 4 and 5 in a straight line; the area between the discal spots and the black mar-

gin, whitish blue; a long, black, white bordered tail at vein 1b. The specimen is very

worn along the edges and the tail at vein 3 is missing; the margin forms a tooth at

vein 3.

Underside wings as in the male; the wavy, black, subtornal line extended in area 2 as

a separate streak. Length of fore wing 23.5 mm.

Habitat. Frequents the margins of montane forests. Rare and local. Record: 1, 3b,

5b (1450-2200 m).

Holotype Ó : Tanzania, Mpanda, Wanzizi Hills, 1450 m, J. Kielland. Allotype 2:

Same, but Mt. Sitebi, 2000 m. Paratypes: Kigoma, Mahale, Ujamba, 2000 m, 16;

Kigoma, Sisaga Mt, 2200 m, November 1971, 12 ; Mpanda, Kampisa, 1500 m,

August 1973, 16 , same, but July 1972, 19 . All J. Kielland. Holotype and allotype

in the National Museum, Nairobi, paratype ó in the British Museum (Nat. Hist.),

London, and other paratypes in J. Kielland collection.

Philiolaus parasilanus parasilanus Rebel, 1914. Three males and one female taken

on bushes in woodland close to a riverine forest at Mihumu, November 1971. Record:

8a (ca. 900 m). General distribution: W. Africaand Zaire to Uganda and Kenya.

Aphniolaus pallene Wallengren, 1857. Woodland and savanna of Mpanda and Kigo-

ma. Not common. Record: 2b, 3a, 3b, 6, 8a, 8b ( 1000-1600 m).

Epamera violacea Riley, 1928. Riverine forest margins at Kampisa, Mt. Sitebi, Mt.

Ipumba. Very local. Record: 5b, 8b (900-2000 m). General distribution: Angola to

Zaire, Rhodesia and Malawi.

Epamera scintillans Aurivillius, 1905. Wooded hills of Nkungwe in Mpanda. Rare.

Record: 6 (1150 m). General distribution: W. Africa to Uganda and Sud Bahr el

Ghasal.

Epamera australis Stevenson, 1937. One female taken at forest margin of Wanzizi

Hills. Record: 3b (1500 m). General distribution: Rhodesia and Zambia to Zaire.

Epamera pseudopollux Stempffer, 1962. Forest margins of Kampisa, Sitebi and

KIELLAND: Rhopalocera eastern side Lake Tanganyika 201

Ipumba Mts. Very local. Record: 5b (1500-2000 m). General distribution: Kigezi in

Uganda.

Epamera pseudofrater Stempffer, 1962. Forests of Mihumu, Kefu, Ntakatta, K ampi-

sa, Mahale. Rare. Record: 1, 5a, 5b, 8a, 8b (900-1600 m). General distribution: Kigezi

and Kayonza forests in Uganda.

Epamera sidus Trimen, 1864. Woodland and open grassland of hills and mountains.

Record: 5a, 5b, 6, 8a, 8b (900-1900 m).

Epamera alienus alienus Trimen, 1898. Two males taken on summit of Mt. Sitebi,

in open short grassland with stunted and scattered vegetation, September 1970 and

September 1971. Record: 5b (2050 m). General distribution: From Transvaal to

Rhodesia, Zambia, Mozambique, Malawi and Songea in Tanzania; ssp. bicaudatus

Aur., 1905, from Cameroun, Nigeria and Upper Volta; ssp. uganda St., 1953, from

Uganda and Kenya.

Epamera aphnaeoides aphnaeoides Trimen, 1873. One female taken at Mtozi river

near Chala in Ufipa. Record: Chala (1700 m).

Epamera fontainei Stempffer, 1956. One male and one female taken in the Kefu

forest. February 1972. Extremely rare. Record: 8b (900-1000 m). General distribution:

W. Africa to Zaire.

Etesiolaus catori cottoni Bethune Baker, 1908. Forest margin in the Lubalizi valley,

16, 2.viii.1970; Ntakatta forest, 16, February 1972. Extremely rare. Record:

3a, 5a (1150-1500 m). General distribution: Zaire to Uganda and at Usambara in

Tanzania.

Hemiolaus caeculus caeculus Hopffer, 1855. Woodland of Mpanda and Kigoma.

Locally common. Record: 3a, 6, 8b (1000-1200 m).

Hypolycaena hatita ugandae Sharpe, 1904. Rather common in forests of Mpan-

da and Kigoma. Record: 1, 2a, a, 3b, 4a, 5a, 5b, 8a, 8b, 9b, Gombe (800-

1600 m, 2000 m at Mt. Sitebi).

Hypolycaena antifaunus latimacula Joicey & Talbot, 1921 (1924). Common in

forests and also in heavy woodland of Mpanda and Kigoma. Record: 1, 2a, 3a, 4a, 5a,

5b, 6, 8a, 8b, Gombe (800-1500 m). General distribution: W. Africa to Uganda.

Hypolycaena nigra Bethune Baker, 1914. Forests of the sandstone area. Mukuyu,

Kefu, Mugondozi, Ntakatta. Record: 4a, 5a, 8a, 8b (850-1000 m). General distribu-

tion: W. Africa to Uganda.

Hypolycaena dubia Aurivillius, 1895. Several males and females taken in riverine

forests at Mukuyu, Kefu, Kasoge, Ntakatta, Luntampa and Gombe. Not common.

Record: 1, 4a, 5a, 8a, 8b, Gombe (800-1300 m). General distribution: W. Africa to

Zaire and Uganda.

Hypolycaena liara plana Talbot, 1935. Common in forests and heavy woodland

of Mpanda and Kigoma. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe

(800-1600 m, 2000 m at Mt. Sitebi). Form obscura St., 1947, occurring at Kasoge,

Kefu, Luntampa and at Gombe. This form is nearly always smaller than the typical

form.

Hypolycaena auricostalis Butler, 1897. Common in woodland and open grassland of

Mpanda, Kigoma and Ufipa. Particularly at higher levels. Record: 1, 3a, 3b, 4a, 5a,

5b, 6, 8a, 8b, Chala, Mbuzi Mt. (1050-2000 m).

Hypolycaena philippus Fabricius, 1793. Very common in open habitats and wood-

202 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

land of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a,

8b, Ufipa, Gombe (800-1800 m).

Hypolycaena buxtoni buxtoni Hewitson, 1874. Common in open habitats and

woodland of Mpanda, Kigoma and Ufipa. Record: 1, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b,

Ufipa, Gombe (900-2000 m).

Hypomyrina nomenia nomenia Hewitson, 1874. Rather common in forests of Mpan-

da and Kigoma. Record: 1, 2a, 3a, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-1700 m).

Actis mimetica mimetica Karsch, 1895 (= perigrapha Karsch, 1895). In riverine

forests of Mpanda and Kigoma. Sitebi, Utobo river near Sitebi, Ntakatta and Lukan-

damira. Uncommon. Record: 1, 5a, 5b (1400-1700 m). General distribution of ssp.

mimetica: Uganda and Kenya.

Hypocopelates otraeda otraeda Hewitson, 1863. Not uncommon in forests of Mi-

humu, Mukuyu to Mugombazi and a few specimens taken in the Ntakatta forest.

Sandstone area only. Record: 5a, 8a, 8b (800-1100 m, 1500 m in the Ntakatta forest).

General distribution: Zaire to Uganda; ssp. modesta Talbot, 1935, Ivory Coast to

Cameroun.

Hypocopelates kafuensis Neave, 1910. Brachystegia woodland of Mpanda and Ki-

goma. Common. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b (800-1600 m, 2000 m

at Mt. Sitebi).

Pilodeudorix camerona Plötz, 1880. Not uncommon in woodland and forest margins

of Mpanda and Kigoma. Record: 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Gombe (900-

2000 m).

Pilodeudorix diyllus orientalis Stempffer, 1957. Rather uncommon in woodland and

forest margins of Mpanda and Kigoma. Record: 6, 8a, 8b (1000-1200 m). General

distribution: Nominotypical race from W. Africa; ssp. orientalis from Katanga to

Uganda.

Pilodeudorix zeloides Butler, 1901 (= P. zela zeloides Hewitson, 1869). Open

habitats of Sitwe, Sitebi and Mahale, on stunted trees and bushes. Uncommon. Record:

1, 3a, 5b, Gombe (900-1900 m). General distribution: Rhodesia to Malawi.

Pilodeudorix caerulea caerulea H. H. Druce, 1890. Not uncommon in woodland of

Mpanda and Kigoma. Record: 2a, 3a, 3b, 4a, 5a, 5b, 6, 8b (1000-1500 m, 2000 m

at Mt. Sitebi).

Pilodeudorix zelomina Rebel, 1914. Three females taken December 1970 in a forest

at Mt. Sitebi. Record: 5b (1900 m). General distribution: Burundi, Uganda.

Diopetes corruscans Aurivillius, 1897. Females rather common in the Kefu forest;

males rarely seen. Also taken at Mugondozi river and in the Ntakatta forest. Sand-

stone and bordering area only. Record: 5a, 8b (900-1300 m). General distribution:

Cameroun, Gabun.

Diopetes deritas Hewitson, 1874. Open habitats and forest margins of Mpanda and

Kigoma. Rare. Record: 1, 3b, 5b, 8a, 8b (900-1600 m). General distribution: Zaire

to Uganda.

Virachola odana H. H. Druce, 1887. Females locally common in forests of Kigoma;

males very rarely seen. One male taken near Sibweza, in woodland. Record: 1, 2a, 3a,

4a, 5a, 5b, 6, 8a, 8b (800-1000 m, 2000 m at Mt. Sitebi). General distribution: Kenya

and Uganda to W. Africa.

Virachola diocles Hewitson, 1869 (text-fig. 32). Rather common on Brachystegia

KIELLAND: Rhopalocera eastern side Lake Tanganyika 203

hills of Mpanda and Kigoma. Also Ufipa. Record: 3a, 3b, 6, 8a, Chala (1000-1200 m,

1800 m at Chala).

Virachola lorisona Hewitson, 1862 (text-fig. 33). Rather common on wooded hills

and forest margins of Mpanda and Kigoma. Record: 4a, 4b, 5a, 5b, 6, 8a, 8b (800-

1900 m).

Virachola ufipa spec. nov. (text-fig. 34, pl. 12 figs. 50, 51, pl. 13 figs. 52, 53). Some-

what between V. diocles Hew. and V. lorisona Hew., with forewing red area reaching

hind margin, but somewhat restricted; underside more like that of V. diocles than

that of V. lorisona. The latter has much cleared markings.

Description. Antenna brown, white ringed; club brown, underside white at base;

eyes hairy; frons dark brown with white lateral stripes which continue as almost a com-

plete circle around the eyes; palpi black dorsally, white ventrally and laterally, third

joint thin, as long as second joint, first joint as long as second and third combined.

Male. Tip of abdomen red; upperside fore wing black; red area hardly reaching cell

and vein 3, basally not reaching base of vein 2, reaching hind margin; the outer black

margin 3 mm wide at middle of area Ib, curved basad like a wedge in space la, leaving

a 2.5 mm gap of red area, which is extended as a fine line along the margin to near

tornus. Upperside hind wing red; lower part of cell black and gray, hairy; outer mar-

gin narrowly black, cilia black, a black androconial patch at base of vein 7; the tornal

lobe red, bordered black, hairy; a tail at vein 1b. Underside fore wing silvery grayish,

the red area of the upperside somewhat shining through; discal spots contiguous, same

shade as ground colour, outlined with dark gray and white on both sides; spots in 5 and

6 a little displaced towards margin, otherwise discal spots almost a straight line; there

are faint submarginal spots outlined in white; a white outlined patch at end of cell; the

hair pencil is dark brown. Underside hind wing ground colour as fore wing; discal spots

irregularly placed particularly those in area 2 and 3 which are placed on edge and a

little towards base; the discal spots are outlined in darker gray and white; the space

between the discal and submarginal spots is whitish; a narrow whitish stripe along

margin, then a very fine blackish stripe; cilia pale gray with a darker median stripe;

there is a red lunule with black center near margin in space 2; the lunule in 1b is red

with a small blue dot distad; also a red patch in la proximad of the black lobe; some

blue scales between the black lobe and the red patch, also a few blue scales at the tip of

the lobe; a rectangularly shaped and white bordered spot at the end of cell, one spot in

the cell, one basal spot in 7 and a faint spot just below the cell touching the cell spot.

Length of fore wing 14.8 to 15.8 mm.

Female. Fore wing brown with median, pale area as in lorisona; hind wing basal

part, costal fold and costa to vein 6 are brown; the rest of the wing up to distal part of

the cell pale violet blue with strongly marked, dark veins; a tail at vein 2; the spot of

the underside in area 2 shining faintly through; there is a brown submarginal line and

a blackish marginal line; cilia on both wings white.

Underside whitish gray; fore wing discal spots very faint, almost invisible, but this

may be an individual variation; there is an even, one mm wide submarginal band and

an equally wide marginal line, the outer border of this is finely black; hind wing

as in the male, but the lunules are yellowish, not red. Length of fore wing 17.5 mm.

Genitalia 6: Very similar to the genitalia of V. diocles; valva longer and more

slender as shown in the figures (text-fig. 34 vs. 32).

204 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Habitat. Open woodland. Record: Chala in Ufipa (1800 m).

Paratype: Same, but May 1972, 1 6. Holotype and allotype in the National Museum,

Nairobi, paratype in J. Kielland collection.

Virachola dinochares Smith, 1877. Locally common in woodland of Mpanda, Kigo-

ma and Ufipa. Record: 3a, 3b, 5a, 5b, 6, 7, 8a, 8b, Chala (800-1600 m, 2000 m at Mt.

Sitebi).

Virachola antalus antalus Hopffer, 1855. Very common in woodland and savanna

and open grassland of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b,

5a, 5b, 6, 7, 8a, 8b, 9a, 9b, Chala, Gombe (800-2200 m).

Virachola vansoni Pennington, 1948. Woodland and Brachystegia hills and montane

grassland of Mpanda, Kigoma and Ufipa. Not uncommon. Record: 2a, 3a, 3b, 4a, Sa,

5b, 6, 8a, 8b, Rungwa, Chala (900-2000 m). General distribution: described from

Zululand and recorded from Malawi.

Very difficult to separate from V. penningtoni van Son, 1939, and V. magdae Gif-

ford, 1953. Identified by Dr. H. Stempffer. This species was taken by the author

also in the Singida area, N. E. of Tabora and in Ruaha Nat. Park.

Virachola penningtoni van Someren, 1939. Among specimens sent to Dr. Stempffer

for determination, one female and one male appears to belong to this species. Taken in

woodland and montane grassland. Record: 5b at Sitebi, Rukwa basin (900-2000 m).

Capys catharus Riley, 1932. Open habitats at Sitwe, Mahale, Luntampa and Si-

tebi. Not common. Record: 1, 3a, 4a, 5a, 5b (1200-1900 m). General distribution:

Malawi and K atanga.

Capys disjunctus Trimen, 1895. Open grassland at Mt. Sitebi, Mahale and at

Gombe. Record: 1, 5b, Gombe (1500-2300 m).

Capys connexivus Butler, 1897. Open and wooded habitats of Mpanda, Ki-

goma and Ufipa. Rather common. Record: 1, 3a, 5b, Kishi, Chala (1200-2000 m).

Capys brunneus Aurivillius, 1915. One male taken in woodland at Lubalizi in Kigo-

ma, March 1971. Record: 3a(1200 m).

Subfamily Lycaeninae Leach, 1815

Lycaena abbotii Holland, 1892. Montane grassland of Ufipa. Here common. Record:

Sumbawanga area, Chala, Mbuzi Mt. (1800-2100 m). L. abbotii is by some authors

considered a race of L. phlaeas Linnaeus, 1761. The latter occurs at Ngara District,

north of Kigoma.

Subfamily Polyommatinae Swainson, 1827

Anthene (Cupidesthes) leonina Bethune Baker, 1903. One female from the Mihumu

forest, one female Kefu forest, one female K asoge and one male from a stream-bed near

Kasangatongwe N.W. of Ikola. Record: 1, 2b, 8a, 8b (800-900 m). General distribu-

tion: W. Africa to Zaire and Uganda.

Anthene (Cupidesthes) isobelae Jackson, 1965. Forests and woodland of Mpanda and

Kigoma. Very rare. Sibweza and Mihumu. Record: 6, 8a (900-1100 m). General

distribution: Uganda.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 205

Anthene (Cupidesthes) irumu Stempffer, 1948. On wet sand in the Kefu forest,

K asoge forest, on bushes at Luntampa and Ntakatta. Also on wet river sand at upper

Katuma. Record: 1, 4a, 5a, 5b, 8b (800-1600 m). General distribution: Central African

Rep. to Zaire.

Anthene (Cupidesthes) wilsoni Talbot, 1935. A very large species. A female was

taken in open vegetation near a riverine forest at Mihumu, November 1971. Record: 8a

(ca. 900 m). General distribution: Ethiopia, Sudan to Kenya.

Anthene rubricinctus jeanneli Stempffer, 1961. Woodland and riverine forests of

| Mpanda and Kigoma. Common. Record: 1, 2a, 3a, 3b, 4a, 4b, Sa, Sb, 6, 8a, 8b,

| Gombe (800-1600 m).

Anthene anadema H. H. Druce, 1890. Rather common in forests of Kigoma and

Mpanda. Record: 1, 4a, 5a, 5b, 8a, 8b, Gombe (800-1600 m).

Anthene ligures Hewitson, 1874. Forests and heavy woodland of Mpanda and Ki-

goma. Record: 1, 3a, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-1600 m).

Anthene hobleyi Neave, 1904. Montane forests on the Mahale Mt. Only place

recorded, but common here. Record: 1 (1700-2200 m).

Anthene barnesi Stevenson, 1940. Forest of Mbuzi Mt. in Ufipa. Rare. Record:

Mbuzi Mt. (2000 m).

Anthene definita Butler, 1899. Woodland and savanna of Mpanda, Kigoma and

Ufipa. Common. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Ufipa, Gombe (800-

2000 m).

Anthene indefinita Bethune Baker, 1910. Common in forests of Mpanda and Kigoma.

Record: 1, 2a, 3a, 3b, da, 4b, 5a, 5b, 6, 8a, 8b, Gombe (800-1600 m).

Anthene sylvanus Drury, 1773. Forests of Mpanda and Kigoma. Not uncommon.

Record: 1, 2a, 3a, 3b, 4a, 5a, 8b (800-1700 m).

Anthene alberta Bethune Baker, 1910. Common in forests of Mpanda and Kigoma.

Particularly in the sandstone area. Record: 1, 2a, 3a, 3b, 4a, Sa, 5b, 6, 8a, 8b (800-

1600 m).

Anthene schoutedeni Hulstaert, 1924. Forests of Mpanda and Kigoma. Rather

common. Record: 1, 2a, 3a, 3b, 4a, 5a, 8a, 8b (900-1600 m).

Anthene lemnos Hewitson, 1878. Forests of Mpanda and Kigoma. Record: 1, 2a, 3a,

3b, 4a, 5b (800-1800 m).

Anthene princeps Butler, 1876. Woodland and savanna of Mpanda, Kigoma and

Ufipa. Common. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe, Bihara-

mulo, Mbizi forest, Chala (800-2200 m).

Anthene liodes liodes Hewitson, 1874. Less common. In forests of Mpanda and Ki-

goma. Record: 1, 3a, 5a, 6, 8a, 8b (800-1300 m). Rather low altitude.

Anthene butleri livida Trimen, 1881. Two males only at Nkungwe in Mpanda.

Woodland. Record: 6 (1000 m).

Anthene nigropunctata Bethune Baker, 1910. Woodland of Mpanda and Kigoma.

Uncommon. Record: 3a, 4a, 5b, 6, 7, 8b, Biharamulo (800-1200 m).

Anthene otacilia Trimen, 1868. Woodland of Mpanda, Kigoma and Ufipa. Not

common. Record: 1, 3a, 3b, 6, 8b, Gombe, Chala, Mbuzi Mt. (800-1500 m, 1800 m at

Mt. Chala).

Anthene talboti Stempffer, 1936. Woodland of Mpanda and Kigoma. Not common.

Record: 3a, 5b, 6 (1000-1600 m). Pupae taken on species of Mimosaceae.

206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Anthene amarah Guerin, 1847. Very common in woodland of Mpanda, Kigoma and

Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Chala (800-1800 m).

Anthene lunulata Trimen, 1894. Common in woodland, savanna and open grass-

land of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a,

8b, 9a, Gombe, Ufipa (800-2000 m).

Anthene larydas Cramer, 1780. Very common in woodland and forests of Mpanda

and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b, 9a, 9b, Gombe

(800-1700 m).

Anthene crawshayi crawshayi Butler, 1899. Common in woodland and savanna of

Mpanda and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 5b, 6, 8a, 8b, Gombe (800-1500 m).

Anthene gemmifera Neave, 1910. Wooded hills of Mpanda and Kigoma. Very local.

Record: 3a, 5a, 5b, 6, 8b (1000-1700 m).

Anthene staudingeri Smith & Kirby, 1894. A few males and one female taken at a

riverbed inside the Ntakatta forest. Record: Sa (ca. 1500 m). General distribution:

W. Africa to Zaire and Uganda.

Anthene phoenicis Karsch, 1893. Forests and heavy woodland of Mpanda, Kigoma

and Ufipa. Not common. Record: 2a, 4a, 5a, 5b, 6, 8b, Chala (1000-1600 m). General

distribution: W. Africa to Uganda and Zambia.

Anthene hades Bethune Baker, 1910. Forests and woodland of Mpanda, Kigoma and

Ufipa. Common. Record: 1, 2a, 3a, 4a, 5a, 5b, 6, 8a, 8b, Gombe, Chala (800-

2000 m). General distribution: W. Africa to Zaire.

Anthene lamias Hewitson, 1878. Riverine forest near Mugombazi river and at

Kasoge in Kigoma, at Kampisa in Mpanda. Rare. Record: 1, 5b, 8b (800-1500 m).

General distribution: W. Africa to Zaire and Uganda.

Cupidopsis cissus Godart, 1824. Most open habitats of Mpanda, Kigoma and Ufipa.

Common. Record: widespread (800-2200 m).

Cupidopsis jobates Hopffer, 1855. Open habitats and woodland of Mpanda, Kigo-

ma and Ufipa. Less common than C. cissus. Record: 3a, 3b, 5a, 5b, 6, 8b, Ufipa

(1000-1800 m).

Pseudonacaduba sichela Wallengren, 1857. Very common in woodland and forest

of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b,

Chala, Gombe (800-2000 m).

Pseudonacaduba aethiops Mabille, 1877. One male taken on the Katuma-Mpanda

road; one male at Sibweza and several males in the Kasoge and Kefu forests.

Local and uncommon. Record: 1, 3a, 4a, 5a, 6, 8b (800-1400 m). General distri-

bution: W. Africa to Uganda and Kenya.

Lampides boeticus Linnaeus, 1767. Ubiquitous. Throughout Mpanda, Kigoma and

Ufipa (780-2200 m).

Uranothauma antinorii Oberthür, 1883. Common in woodland and forests of Mpan-

da, Kigoma and Ufipa. Particularly in the Mahale area. Record: 1, 4b, 5a, 5b, 8a,

8b, Gombe, Mbuzi Mt. (800-2000 m).

Uranothauma lunifer Rebel, 1914. Montane forest of the Mahale Mt. Male not

uncommon. June to November. Record: 1 (1600-2200 m).

Uranothauma poggei Dewitz, 1879. Woodland of Mpanda and Kigoma. Common.

The whole year. Record: 1, 2b, 3a, 5b, 6, 8a, 8b, Gombe (800-2000 m).

The genitalia of U. poggei and U. lunifer are identical. There is, however, no

KIELLAND: Rhopalocera eastern side Lake Tanganyika 207

doubt that they are separate species. U. lunifer lives in montane forest; U. poggei

is much more adapted to woodland and is generally found at lower levels, but it

flies together with U. lunifer over the whole range of the latter. The external characters

are constant in both species.

Uranothauma nubifer Trimen, 1895. Heavy woodland and forest of Mpanda, Kigo-

ma and Ufipa. Mostly at higher altitudes. The whole year. Common. Record:

1, 3a, 3b, 4b, 5b, 6, Sumbawanga, Chala, Mbuzi Mt. (1200-2000 m).

Uranothauma delatorum Heron, 1909. Forests of the Mahale Mt., Mt. Sitebi and

Mbuzi Mt. in Ufipa. Record: 1, 5b, Mbuzi Mt. (1600-2000 m).

Uranothauma falkensteini Dewitz, 1879. Forest of Kasoge and at Katuma river

and Gombe. Very local. The whole year. Record: 1, 6, Gombe (800-1200 m).

Uranothauma titei Cottrell, |1968]. Close to U. vansomereni Stempffer, 1951. Very

common in woodland and savanna of Mpanda, Kigoma and at Biharamulo. A female

taken at high altitude on Mt. Karobwa. Elsewhere at much lower levels. The whole

year. Record: 2b, 3a, 3b, 4b, 5b, 6, Biharamulo (1000-1200 m, 1700 m at Karobwa).

Dr. Cottrell wrote me in 1968 that he was naming this species U. titei. Its publication,

however, has not been confirmed.

Phlyaria cyara Hewitson, 1876. South of Kigoma only taken in the Kasoge forest;

north of Kigoma common in the Kasekera forest at Gombe. Record: 1, Gombe (800-

1700 m).

Phylaria heritsia intermedia Tite, 1958. Common in forest margins of Mpanda and

Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8b, Gombe (800-1700 m).

Cacyreus audeoudi Stempffer, 1936. Scarce in forests of the Mahale area, Ntakat-

ta and Luntampa forest. Record: 1, 4a, 5a (800-1600 m).

Cacyreus virilis Aurivillius, 1924. Forests and heavy woodland of Mpanda, Kigo-

ma and Ufipa. Common. Record: 1, 3a, 5a, 5b, 6, 9a, Mbizi forest, Chala Mt., Mbuzi

Mt. (1000-2200 m).

Cacyreus lingeus Cramer, 1781. Very common in woodland and forests of Mpanda

Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, Sb, 6, 7, 8a, 8b, Mbizi

forest, Chala, Mbuzi Mt., Gombe (800-2000 m).

Cacyreus palemon Cramer, 1782. Forests and forest margins of Ufipa. Common.

Record: Mbizi forest, Chulwe Mt., Chala, Mbuzi Mt. (2000 m).

Harpendyreus major Joicey & Talbot, 1924. Montane vegetation, forest margins

and clearings of the Mahale Mt. and Ufipa highland. Locally common. Record: 1,

Mbizi forest, Chulwe Mt. Chala Mt., Mbuzi Mt. (1600-2200 m).

Harpendyreus juno Butler, 1896. A few specimens taken on grassland near the

Mbisi Forest, Ufipa (2200 m), February 1978.

Syntarucus pirithous Linnaeus, 1767. Very common in all habitats of Mpanda, Ki-

goma and Ufipa. Record: widespread (800-2200 m).

Syntarucus jeanneli Stempffer, 1935. Very common in all habitats of Mpanda, Ki-

goma and Ufipa. Record: widespread (800-2200 m).

Syntarucus babaulti Stempffer, 1935. Common in most habitats of Mpanda and

Kigoma. Record: 1, 3a, 3b, 5a, 5b, 8a, 8b, Gombe (800-2000 m).

Syntarucus marginalis Aurivillius, 1925. Mpanda and Ufipa in open habitats. Very

rare. Record: 6, Sumbawanga (800-2000 m).

Syntarucus brevidentatus Tite, 1958. Woodland and open habitats of Mpanda and

208 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Kigoma. Not common. Record: 5b, 6, Gombe (1000-1500 m). |

Syntarucus pulcher Murray, 1874. Marshy habitats of Mpanda. Very rare inland, |

but more common along marshy shores of Lake Tanganyika during the dry season. |

Record: 2b, 6, 7(776-1100 m). |

Castalius hintza Trimen, 1864. Scarce in woodland of Mpanda, Kigoma and |

Ufipa. Record: 6, 8b, Chala (1000-1800 m).

Castalius calice calice Hopffer, 1855. Common in woodland and savanna of

Mpanda, Kigoma and Ufipa. Record: widespread (800-2000 m).

Castalius melaena Trimen, 1887. Common in woodland and savanna of Mpan-

da, Kigoma and Ufipa. Record: 3a, 3b, 5b, 6, Ufipa (800-1800 m).

Castalius margaritaceus Sharpe, 1891. Very common in forests of Mpanda, Ki-

goma and Ufipa. Both high and low level. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b

(800-2200 m). The specimens from this area are rather similar to C. carana Hew.,

1876, in external characters, but the genitalia correspond with that of margaritaceus.

It is very common in all forests south of Kigoma and also occurs at the Ngara

District, south of Bukoba.

Tarucus sybaris Hopffer, 1855. Brachystegia woodland and savanna of Mpanda,

Kigoma and Ufipa. Not common. Record: 1, 6, Chala (800-1800 m).

Zizeeria knysna Trimen, 1862. Very common in woodland and open grassland of

Mpanda, Kigoma and Ufipa. Record: widespread (800-2200 m).

Zizina antanossa Mabille, 1877 (= otis Fabr., 1787). Open grassland of Mpanda,

Kigoma and Ufipa. Less common. Record: 3a, Mbizi forest, in open grassland, Mbuzi

Mt., Gombe (1500-2100 m).

Zizula hylax Fabricius, 1775 (= gaika Trim., 1862). Very common in woodland and

open grassland of Mpanda, Kigoma and Ufipa. Record: widespread (800-1900 m).

Acticera lucida Trimen, 1883. Common in open habitats and woodland of Mpanda,

Kigoma and Ufipa. Record: widespread (800-2200 m).

Acticera stellata Trimen, 1883. Montane grassland of Ufipa. Record: Near Sumba-

wanga, Chala and Mbuzi Mt. (2000 m).

Azanus mirza Plôtz, 1880. Common in woodland and savanna of Mpanda, Kigoma

and Ufipa. Record: widespread (800-1900 m).

Azanus moriqua Wallengren, 1857. Common in woodland and savanna of Mpanda,

Kigoma and Ufipa. Record: widespread (800-1900 m).

Azanus natalensis Trimen, 1887. Common in woodland and savanna of Mpanda,

Kigoma and Ufipa. Record: widespread (800-1900 m).

Azanus jesous Guerin, 1847. Common in woodland and savanna of Mpanda,

Kigoma and Ufipa. Record: widespread (800-1900 m).

Azanus ubaldus Cramer, 1782. Woodland and savanna of Mpanda. Not common.

Record: 3b, 6 (1100-1600 m).

Eicochrysops mahallakoena Wallengren, 1857. Open montane grassland of

Mpanda, Kigoma and Ufipa. Common only in Ufipa. Record: 3a, 5b, Sumbawanga

(1700-2000 m).

Eicochrysops nandianus Bethune Baker, 1906. Montane grassland in Ufipa. Record:

Sumbawanga, Mt. Chulwe(1900 m).

Eicochrysops messapus Godart, 1819. Open habitats of Mpanda, Kigoma and

Ufipa. Common. Record: 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Ufipa, Gombe (900-2000 m).

KIELLAND: Rhopalocera eastern side Lake Tanganyika 209

Eicochrysops hippocrates Fabricius, 1793. Swampy land and riverine thickets of

Mpanda and Kigoma. Common. Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b,

Gombe (800-1700 m).

Lepidochrysops dollmani Bethune Baker, 1922. Brachystegia hills of Wanzizi. Ex-

tremely rare. November-December only. Record: 3b (1250-1400 m). General distribu-

tion: Rhodesia, Zambia.

Lepidochrysops cinerea cinerea Bethune Baker, 1922. Woodland and open grass-

land of Mpanda and Kigoma. Locally common. Record: 2a, 3a, 3b, 5a, 5b, 8a, 8b

(1000-2000 m). The species normally occurs in Brachystegia woodland, 1000-1400 m,

from late October to early December. In August and September specimens are found in

a quite different habitat, viz., montane grassland, between 1700 and 2000 m. At this

time of the year no specimens are found in the other habitat. It is, therefore, probable

that specimens from the two different habitats belong to two different species. But

they cannot be separated on the basis of external or genital characters. The geni-

talia of the Tanzania specimens differ somewhat from Kenya cinerea.

Lepidochrysops plebeja proclus Hulstaert, 1924. Brachystegia hills of Mpanda. Lo-

cal, in November-December. Record: 3b, 5b, 6 (1100-1600 m).

Lepidochrysops solwezii Bethune Baker, 1922. Brachystegia hills of Mpanda and

Kigoma. Common in November to December. Record: 2a, 3b, 4a, 5a, 5b, 6, 8a, 8b

(900-1700 m).

Lepidochrysops mpanda Tite, 1961. Open montane grassland of Mweze and at

Sitebi Mt. August to September. Record: 3a, 5b (1700-2000 m). Endemic to this

area.

Lepidochrysops intermedia cottrelli Stempffer, 1954. Common on a Brachystegia

hill near Ntakatta. A few specimens taken further north and at Lubalizi. Novem-

ber only. Record: 3a, 4a, 5a, 8b (1000 to 1400 m). General distribution:

Ndola in Zambia and Malawi.

Lepidochrysops anerius kiellandi Stempffer, 1973. Woodland of Mpanda and Ki-

goma. Sometimes common. November only. Record: 2a, 3b, 5a, 5b, 8a, 8b (900-

1400 m). General distribution: Nominotypical race described from Elisabethville in

Zaire. November 1971 this species was very abundant between Ntakatta and Mu-

gombazi river. Other years uncommon.

Lepidochrysops spec. Brachystegia woodland of Sibweza, Mabu, Wanzizi, Kampisa,

Ngondo. October to November. Not common. Record: 3a, 3b, 5b, 6 (1000-1600 m).

Lepidochrysops spec. Three males taken in open, montane grassland on Mt. Chala,

October 1971, and a series was taken at the same place in September 1973. Record:

Chala Mt., observed at Mbuzi Mt. (2000 m). Flying time: (August) September to early

October. Apparently a new species.

Lepidochrysops inyangae Pinhey, 1945. One male taken at Kabakaranga, Novem-

ber 1958; one female on a Brachystegia clad hill just north of Ntakatta. Novem-

ber only. Record: 2a, 5a (900-1400 m). General distribution: the species was only

known from Inyanga in Rhodesia.

Lepidochrysops spec. Open habitats of Chala in Ufipa. Record: Chala (1700-

1800 m). x

Lepidochrysops gigantea Trimen, 1898. Woodland in hilly country of Mpanda and

Kigoma. Rather common. October to January. Record: 2a, 3a, 3b, Sa, 5b, 6, 8b (1000-

1600 m).

210 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Lepidochrysops cupreus Neave, 1910. Montane grassland of Mpanda. Local. August

to September. Record: 3a, 5a, 5b (1600-2000 m).

Euchrysops barkeri Trimen, 1893. Common in woodland and savanna of Mpanda

and Kigoma. Record: 2a, 3a, 3b, 4a, 5a, 5b, 6, 7, 8a, 8b (800-1600 m).

Euchrysops dolorosa Trimen, 1887. Open habitats of Mpanda and Kigoma, Rather

common. Mainly in the dry season. Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6 (1000-

2000 m).

Euchrysops subpallida major Bethune Baker, 1922. Open, montane grassland of

Mpanda, Kigoma and Ufipa. Locally common. Record: 1, 3a, 5a, 5b, Mt. Chulwe,

Mbuzi Mt. (1600-2000 m).

Euchrysops mauensis Bethune Baker, 1923 (Det. by Dr. Stempffer). Montane grass-

land of Kigoma, Mpanda and Ufipa. Record: 1, 3a, 5a, 5b, Mbuzi Mt. (1500-

2000 m).

Euchrysops albistriatus koena Strand, 1911. On grassland with scattered shrubs and

trees at Mweze highland and at Lublizi. April and August. Record: 3a (1200-

1700 m). General distribution: W. Africa to Uganda.

Euchrysops malathana Boisduval, 1833. Very common in open habitats and wood-

land of Mpanda, Kigoma and Ufipa. Record: widespread (800-2200 m).

Euchrysops brunneus Bethune Baker, 1922. Taken at Masaba in Kigoma in open

habitats. Very rare. Record: 2a (1300 m).

Euchrysops osiris Hopffer, 1855. Very common in open and wooded habitats of

Mpanda, Kigoma and Ufipa. Record: widespread (800-2200 m).

Euchrysops spec. Small species with rather pointed fore wing and dark underside,

taken in open, stunted woodland at Kampisa in Mpanda. Record: 5b (1500 m).

Euchrysops spec. Small, light gray species with tails, taken in Brachystegia wood-

land at Wanzizi. Record: 3 b(1500 m).

Thermoniphas fontainei Stempffer, 1956. Kasoge forest only. Particularly along

water courses. Not common and very local. Dry season. Record: 1 (800-900 m).

General distribution: Moyen Congo to K atanga in Zaire.

Oboronia punctatus Dewitz, 1879. Dense forests of Kasoge, Mihumu, Kefu, Lubali-

zi. Local, but rather common. Record: 1, 3a, 8a, 8b (800-900 m). General

distribution: W. Africa to Uganda.

Oboronia gussfeldti Dewitz, 1879. Several specimens flying amongst vegetation

along Masale River, west of Mt. Karobwa. Record: 2a (1300 m); one female taken in

the riverine forest, January 1978.

Athysanota ornata vestalis Aurivillius, 1895. Forests of Kigoma and Mpanda.

Record: 1, 3a, 4a, 5a, 8a, 8b, Gombe (800-1400 m). General distribution: W. Africa

to Uganda.

Freyeria trochylus Freyer, 1845. Open habitats and woodland of Mpanda, Ki-

goma and. Ufipa. Record: 1, 2a, 3a, 3b, 4a, Sb, 6, 7, 8b, U fipam dl 000s

2000 m).

Family Hesperiidae

Subfamily Coeliadinae

Coeliades chalybe Westwood, 1852. Occurring in some evergreen forests of Kigoma

KIELLAND: Rhopalocera eastern side Lake Tanganyika 210

and extending into Mpanda. Mugondozi river, Kefu, Luntampa, Ntakatta and ob-

served at lower Ujamba, Mahale. Record: 1 (observed), 3a, 4a, 5a, 8b (1000-1400 m).

General distribution: W. Africa and Angola to Zaire, Uganda and N. W. Kenya.

Coeliades libeon Druce, 1875. Woodland and forest of Mpanda and Kigoma.

Common. Record: 1, 2a, 2b, 3a, 3b, 4a, Sa, 5b, 6, 8a, 8b, Gombe (800-1700 m).

All specimens collected south of the sandstone ridge are of the form /ibeon, while

all the ones collected and observed north of the ridge (Kefu Forest) belong to the

smaller form brussauxi Mabille, 1877, without a purple sheen and more distinct white

tornal border on the hind wing.

Coeliades anchises Gerstaecker, 1871. Taken on flowers on and near the beach of

Gombe Stream Nat. Park. Also in open woodland. Record: Gombe (800-1000 m).

‘ Coeliades forestan forestan Cramer, 1782. Very common in most habitats of Mpan-

da, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, Sa, 5b, 6, 7, 8a, 8b, Chala,

Gombe (800-2100 m).

Coeliades pisistratus Fabricius, 1793. Forest margins at Luntampa, Kampisa, Sib-

weza, Wanzizi and in Ufipa. Uncommon. Record: 1, 3a, 3b, 5b, 6, Mbuzi Mt.

(1000-1700 m).

Coeliades sejuncta Mabille & Vuillot, 1891. In the sandstone area and border

area to the basement formations. Rather uncommon. Record: 5b, 8a, 8b (800-

1500 m).

Coeliades hanno Plötz, 1879. Taken in riverine forest at Kampisa, Luntampa, Mt.

Ipumba and at Gombe, where it was rather common at the time of ale in De-

cember 1972. Record: 1, 4a, 5a, 5b, Gombe (800-1700 m).

Subfamily Pyrginae

Celaenorrhinus proxima proxima Mabille, 1877. Not common in forests of Mpan-

da and Kigoma. Record: 1, 3a, 3b, 4a, 5a, 8a, 8b (800-1700 m).

Celaenorrhinus homeyeri Plötz, 1880. Katuma riverine forest of Mpanda, forests

of Mugombazi, Mihumu, Kefu of Kigoma. Not common. Record: 5a, 5b, 6, 8a, 8b

(800-1000 m, 1700 m near Mt. Sitebi). General distribution: W. Africa to Zaire,

Uganda and Rhodesia.

Celaenorrhinus galenus Fabricius, 1793. Most specimens belong to the form gale-

nus. Very common in forests of Mpanda and Kigoma. Record: widespread (800-

2100 m).

Celaenorrhinus bettoni Butler, 1902. Rather common in forests of Mpanda and

Kigoma. Record: 1, 3a, 3b, 4a, 5a, 5b, 8a, 8b (800-1700 m).

Tagiades flesus Fabricius, 1781. Very common in woodland and forests of Mpanda,

Kigoma and Ufipa. Record: widespread (800-2000 m).

Eagris lucetia Hewitson, 1876. Common in forests of Mpanda, Kigoma and Ufipa.

Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 8b, Mbuzi Mt., Gombe (800-1600 m, 2000 m at

Mbuzi Mt.).

Eagris decastigma purpura Evans, 1937. Several males taken in the Kefu and Kasha

forests of Kigoma. November to February. Extremely local and rare. Record: 8b (900-

1000 m). General distribution: Uganda and Kenya; nominotypical race from W. Africa

to Zaire.

212 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Calleagris lacteus lacteus Mabille, 1877. Forests of Kasoge, the Mahale ridge, Kefu

and Mihumu. Very rare. Record: 1, 5a, 8a, 8b (800-2200 m).

Calleagris jamesoni Sharpe, 1890; forms: jamesoni and gava Evans, 1937. Com-

mon in woodland and savanna of Mpanda and Kigoma. Record: 2a, 3a, 3b, 5a, 5b, 6

(1000-1600 m).

Calleagris hollandi Butler, 1896. Uncommon and very local in woodland of Mpanda

and Kigoma. Dry season from May onwards. Record: 2a, 5b, 6, 8a (1000-1200 m).

General distribution: Angola to Katanga, Zambia, Malawi and Uganda.

Eretis umbra maculifera Mabille & Bouillet, 1916. Ufipa plateau in open grassland.

Common. Record: Chala Mt., Mbizi Mt. (2000 m).

Eretis lugens Rogenhofer, 1891. Common in forests of Mpanda, Kigoma and Ufipa

Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe, Mbuzi Mt., Kibondo

(800-2100 m).

Eretis melania Mabille, 1891. Common in woodland and forests of Mpanda, Kigo-

ma and Ufipa. Record: 1, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Chala, Mbuzi Mt. (1000-

2000 m).

Eretis mitiana Evans, 1937. Lukandamira forest at Mahale. From May to Ocotber.

Record: 1 (1600 m). General distribution: W. Africa to Uganda and Kenya.

Sarangesa laelius Mabille, 1877. Common in woodland of Mpanda, Kigoma and

Ufipa, Record: 1, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Chala (800-1700 m).

Sarangesa lucidella lucidella Mabille, 1891. Woodland of Mpanda at Mtikimu. Re-

cord: 6(1100 m).

Sarangesa phidyle Walker, 1870. Mpanda area in the south. Record: Rukwa basin

(800-1000 m).

Sarangesa seineri tanga Evans, 1937. Woodland of Mpanda and Kigoma. Rather

common. Record: 2b, 3b, 4a, 4b, 5a, 6, Gombe (800-1500 m).

Sarangesa motozi Wallengren, 1857; forms: motozi and pertusa Mabille, 1891.

Common in woodland and open habitats of Mpanda, Kigoma and Ufipa.

Record: 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Rukwa basin (900-1500 m).

Sarangesa ruona Evans, 1937. In forests of Wanzizi and woodland at Sibweza.

Also taken in the Rukwa basin and by Dr. Goodall at Gombe. Record: 3b,

6, Rukwa basin, Gombe (800-1500 m). General distribution: Ukerewe Island, Tan-

zania; Zambia and Malawi.

Sarangesa maxima Neave, 1910. Mpanda area and open habitats at Sumbawanga.

Record: Mpanda, Sumbawanga (1000-1960 m). General distribution: Katanga,

Zambia.

Sarangesa astrigera Butler, 1893; forms: astrigera and neavei Riley, 1921. Common

in woodland of Mpanda and Kigoma. Record: 2b, 3a, 3b, 4a, Sa, 5b, 6, 7, 8a, 8b (900-

1600 m).

Sarangesa haplopa Swinhoe, 1907. Woodland of Mpanda and Ufipa. Not common.

Record: 3a, 4a, 6, Chala (1060-1900 m).

Sarangesa tricerata Mabille, 1891. Woodland of Mpanda and Kigoma. Uncommon.

Record: 3b, 6 (1000-1500 m). General distribution: W. Africa to Zaire.

Sarangesa maculata Mabille, 1891. Common in forests of Mpanda and Kigoma.

Record: 1, 2a, 3a, 3b, 4a, 4b, Sa, 5b, 6, 8a, 8b, Gombe (800-1600 m).

Sarangesa brigida atra Evans, 1937. Dense woodland at Sibweza. Very rare.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 213

December 1970, January and April 1971. Record: 6 (1000 m).

Netrobalane canopus Trimen, 1864. Common in thickets and forest margins of

Mpanda and Kigoma. Record: 3a, 3b, 4a, 5a, 5b, 6, 8b, 9a, Gombe (1000-1600 m).

Caprona pillaana Wallengren, 1857. Common in woodland of Mpanda, Kigoma

and Ufipa. Record: 6, Chala, 8b (1000-1900 m).

Abantis tettensis Hopffer, 1855. Two males taken drinking water at Sibweza.

Extremely rare. Record: 6 (1060 m). Not seen for many years.

Abantis bismarcki arctomarginata Lathy, 1901. One specimen taken at forest

margin of a Lubalizi riverine forest. Record: 3a (1200 m). General distribution:

ssp. bismarcki, W. Africa to Kenya; ssp. arctomarginata, Zaire to Zambia and Ma-

lawi. Note: arctomarginata is a subspecies of bismarcki, according to Berger.

Abantis zambesiaca Westwood, 1874. Woodland of Mpanda and Kigoma. Com-

mon. Record: 2b, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b (800-1900 m).

Abantis paradisea paradisea Butler, 1870. Rather common in woodland of Mpan-

da and Kigoma. Record: 3a, 4a, 5a, 5b, 6, 8a, 8b (800-1500 m).

Abantis venosa venosa Trimen, 1889. Common in open woodland of Mpanda and

Kigoma. Record: 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-1700 m).

Spialia zebra bifida Higgins, 1925. Open grassland of Sitwe (Mweze). One record

only, October 1958. Record: 3a (1700 m). General distribution: Kenya to S. Sudan;

Ruaha in Tanzania.

Spialia depauperata Strand, 1911. Common in open habitats of Mpanda, Kigoma

and Ufipa. Record: 3a, 3b, 5b, 6, Chala, Mbuzi Mt. (1000-2000 m). General distri-

bution: Geita in Tanzania, Natal to Uganda.

Spialia colotes transvaaliae Trimen, 1889. Woodland of Mpanda at Sibweza only.

Rare. Record: 6 (1000-1200 m).

Spialia secessus Trimen, 1891. Common in woodland of Mpanda, Kigoma and

Ufipa. Record: 3a, 3b, 4b, 5b, 6, Lake Katawi, Mbizi forest, Mbuzi Mt. (1000-

2200 m).

Spialia dromus Plôtz, 1884. Common in woodland of Mpanda, Kigoma and Ufipa.

Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Biharamulo, Gombe, Mbuzi Mt. (800-

2000 m).

Spialia spio Linnaeus, 1767. Common in woodland and open habitats of Mpanda,

Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, Sa, 5b, 6, 7, 8a, 8b, Chala (800-

1700 m).

Spialia diomus ferax Wallengren, 1863. Open grassland in Ufipa. Record: Mbisi

Mt. (2200 m). One female taken February 1978. Underside markings correspond with ssp.

ferax and it is probably the first record of this race from Tanzania. General distribu-

tion: ssp. ferax from South Africa to Zambia, ssp. diomus Hopff., 1855, from Tanza-

nia, Kenya and Uganda to Ethiopia and Yemen.

Spialia mafa higginsi Evans, 1937. Open habitats of Kigoma, Mpanda and Ufipa.

Uncommon. Record: 1, 3a, 3b, 5b, Chulwe Mt., Chala Mt. (1200-2000 m).

Gomalia elma Trimen, 1862. Forests of Mpanda, Kigoma and Ufipa. Common.

Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8b, Chala Mt., Mbuzi Mt. (800-2200 m).

214 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Subfamily Hesperiinae

Metisella midas midas Butler, 1893. Very common in Mpanda, and Ufipa on

swampy grassland. Mainly in the wet season. Record: 3a, 3b, 4a, 4b, 5a, 5b, 6, Gombe,

Mbizi forest, Chulwe Mt., Chala Mt. (900-2200 m).

Metisella orientalis Aurivillius, 1925. Common in highlands of Mpanda, Kigoma

and Ufipa. Frequenting forest glades. Scarcer lower down. Mainly in the wet season.

Record: 1, 2a, 2b, 3a, 4a, 5a, 5b, Mbizi forest, Mbuzi Mt., Chala Mt. (1150-

2200 m).

Metisella trisignatus tanga Evans, 1937. Swampy forest glades of Mpanda and Ki-

goma. Locally common. Wet season. Record: 1, 3b, 5b, 6 (1000-2200 m).

Metisella kambove kambove Neave, 1910. Very common during the wet season in

Mpanda and also Kigoma. Record: 1, 2a, 3a, 3b, 4a, Sa, 5b, 6, Gombe (1000-1600 m).

General distribution: Zaire to Zambia.

Metisella carsoni Butler, 1898. Montane grassland of Kampisa, near Ntakatta and

the Mahale ridge. Local and rare. Wet season only. Record: 1, 5a, 5b (1500-2200 m).

General distribution: recorded from Fwambo in Tanzania.

Metisella formosus linda Evans, 1937. Woodland and savanna of Mpanda and Ki-

goma. Very common in the wet season. Record: 2a, 5b, 6, 8a, 8b (1000-1400 m).

Metisella willemi Wallengren, 1857. Common in woodland of Mpanda, Kigoma and

Ufipa. Record: 1, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8b, 9b, Gombe, Chala (1000-1800 m).

Tsitana wallacei Neave, 1910. Savanna and open woodland of Mpanda and Kigo-

ma. Rather rare and local. Record: 2b, 6, 8b, (1000-1450 m).

Isoteinon punctulata Butler, 1895. Common in the wet season in woodland and open

habitats of Mpanda and Kigoma. Record: 1, 2a, 2b, 3a, 4a, 6, 8b, 9a, 9b, Gombe

(1000-1700 m).

Ampittia capenas blanda Evans, 1946. Very common in woodland and savanna of

Mpanda, Kigoma and Ufipa. Record: widespread (800-1600 m).

Kedestes mohozutza Wallengren, 1857. One male taken at riverside in Mbisi

Forest, Ufipa(2200 m), February 1978.

Kedestes wallengrenii wallengrenii Trimen, 1883. Common in woodland and open

habitats of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8b, Gom-

be, Chala, Mbuzi Mt. (1000-2200 m).

Kedestes paola paola Plôtz, 1884. Two males taken settling on bare ground near a

riverine forest at Mt. Sitebi. Open grassland. July 1972. Record: 5b (1750 m). General

distribution: Angola and Zaire to Zambia; ssp. protensa Butler, 1901, W. Africa to S.

Sudan, Uganda, and East Tanzania.

Gorgyra aretina Hewitson, 1878. Forests of Kigoma from Lubalizi to Mihumu.

Rather common. Record: 1, 3a, 4a, 5a, 8a, 8b, Gombe (800-1300 m, 1700 m at

Ntondo). General distribution: W. Africa to Uganda and Kenya.

Gorgyra mocquerysii Holland, 1896. Taken in the Kasoge forest, Mkulya riverine

forest, Kefu, Mukuyu and Ntakatta forests and at riverine forest of Kampisa.

Rather rare. Record: 1, 3a, Sa, 5b, 8a, 8b (800-1500 m). General distribution: W. Afri-

ca and Angola to Uganda.

Gorgyra diva Evans, 1937. Woodland at Sibweza in Mpanda. Rare. Record: Sa, 6,

8b, Gombe (800-1500 m).

KIELLAND: Rhopalocera eastern side Lake Tanganyika 215

Gorgyra subfacatus Mabille, 1889. Not uncommon in forests of Mpanda and Ki-

goma at Gombe, Kefu and Ntakatta. Record: 5a, 8b, Gombe (900-1400 m). General

distribution: Usambara in Tanzania and W. Africa.

Gorgyra minima Holland, 1898. Woodland of Mpanda at Sibweza and Kapanda.

Rare. Record: 6 (ca. 1100 m). General distribution: W. Africa to Uganda and Kenya.

Gorgyra johnstoni Butler, 1893. Common in woodland and riverine forest of

Mpanda and Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b (800-1700 m).

Gorgyra bibulus Riley, 1929. Forests and forest margins of Mpanda and Kigoma.

Local. Record: 1, 3a, 5a, 5b (1500-2000 m). General distribution: W. Africa to

Uganda and Kenya.

Gorgyra kalinzu Evans, 1946. Forests of Kigoma and at Ntakatta and Mt. Sitebi

in Mpanda. Uncommon and very local. Record: 5a, 5b, 8a, 8b (900-1900 m). General

distribution: Uganda and Kasai in Zaire.

Note: According to De Jong (pers. comm.) the Tanzania specimens differ consider-

ably from the Uganda specimens and belong either to a new race or a new species.

Teniorhinus ignita Mabille, 1877. Open woodland and forests of Kigoma and

Mpanda. Not common except in certain forests of the sandstone formations as at

Mugondozi and Kasha forest. Record: 2a, 6, 8a, 8b (800-1100 m). General distribu-

tion: W. Africa to Uganda and Zambia.

Teniorhinus harona Westwood, 1881. Common in Brachystegia woodland of Mpan-

da and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe

(800-1700 m).

Teniorhinus herilus Hopffer, 1855. One record from Ugala Hills near Sibweza, a

few from the Rukwa valley and from Kasoge. Rare. Record: 1, 6, Kipenzi at the foot

of Ufipa escarpment (800-1250 m).

Pardaleodes incerta incerta Snellen, 1872. Common in forests of Mpanda and K igo-

ma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-1600 m).

Pardaleodes sator Westwood, 1852; form pusiella Mabille, 1877. The K asoge forest

only. Not uncommon. Record: 1 (800 m). General distribution: W. Africa to Uganda

and Kenya.

Pardaleodes bule Holland, 1896. Kasoge forest and Gombe Stream only. Rather

common at Kasoge. Record: 1, Gombe (800 m). General distribution: W. Africa to

Zaire, Uganda and Kenya. In Tanzania at Dodoma.

Xanthodisca vibius vibius Hewitson, 1878. Not uncommon in forests of Kigoma

from Lubalizi to Mihumu and at Ntondo near Mahale. Record: 1, 3a, 4a, 5a, 8a, 8b

(900-1700 m).

Acada biseriatus Mabille, 1898. Very common in Brachystegia woodland of Mpan-

da and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, Sa, 5b, 6, 7, 8a, 8b (800-

1700 m).

Osmodes thora Plôtz, 1884. Kasoge forest and at Gombe only. Not uncommon at

Kasoge. Record: 1, Gombe (800-900 m). General distribution: W. Africa to Zaire,

Uganda and Kenya.

Parosmodes morantii Trimen, 1873. Common in Brachystegia woodland and savan-

na of Mpanda and Kigoma. Record: 2a, 3a, 3b, 4a,4b, 5a, 5b, 6, Gombe (800-

1500 m).

Acleros placidus Plôtz, 1879. Kasoge forest and at Gombe only. Record: 1, Gombe

216 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

(800-1300 m). General distribution: W. Africa to Zaire, S. Sudan, Uganda and

Kenya.

Acleros ploetzi Mabille, 1879. Common in forests and forest margins of Mpanda and

Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 8a, 8b, Gombe (800-1600 m).

Acleros neavei Evans, 1937. In the Kasoge forest and forests of the sandstone area.

Record: 1, 5a, 8a, 8b (800-1400 m). General distribution: Uganda and Zaire.

Acleros mackenii Trimen, 1868. Very common in woodland and forest margins of

Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b,

Mbuzi Mt. (800-2000 m).

Acleros spec. Riverine forests of Mpanda and Kigoma. Rather common. Record:

1, 3a, 3b, 4a, 5a, 5b (1100-1500 m). Also known from Katanga by R. H. Carcasson

(pers. comm.).

Acleros substrigatus Evans, 1946. Riverine forest of Mweze highland. One record

only: 12 , October 1958. Record: 3a (1700 m).

Semalea pulvina Plôtz, 1879. Common in forests of Mpanda and Kigoma. Record:

1, 2a, 3a, 3b, 4a, 5a, 5b, 8a, 8b (800-2000 m).

Semalea sextilis Plotz, 1886. Forests of Mpanda at Wanzizi. Very rare. Record: 3b

(1500 m). General distribution: W. Africa to Zaire and Uganda.

Semalea arela Mabille, 1891. Rather common in forests of Mpanda and Kigoma.

Record: 1, 2a, 3a, 4a, 4b, 5a, 5b, 8a, 8b (800-2000 m).

Hypoleucis tripunctata draga Evans, 1937. Forests of Kasoge, Luntampa and

Ntakatta. Uncommon. Record: 1, 4a, 5a (800-1300 m). General distribution: Zaire

to Uganda and Kenya; ssp. tripunctata in W. Africa.

Hypoleucis ophiusa ophir Evans, 1937. Forests of Kigoma and Mpanda in the north-

west: Kasoge, Luntampa, Kefu, Mugondozi, Ntakatta, Mihumu. Not common. Rec-

ord: 1, 4a, 5a, 8a, 8b (800-1600 m). General distribution: Uganda; ssp. ophiusa in

W. Africa.

Meza larea Neave, 1910. Brachystegia woodland of Mpanda, Kigoma and Ufipa.

Not uncommon. Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6, 8b, Chala (1000-1700 m).

General distribution: Zaire, Zambia and Malawi.

Meza mabea Holland, 1894. Two males taken in the Kefu forest; several males

at Lubalizi and in the Ntakatta forest. Very rare and local. Record: 3a, 5a, 8b

(900-1500 m). General distribution: W. Africa to Cameroun. This is probably the

first record from eastern Africa.

Meza cybeutes pallida Holland, 1894. Forest margins at Lubalizi, Luntampa, Kam-

pisa and Ntakatta. Local and rare. Record: 3a, 4a, 5a, 5b (1200-1500 m). General

distribution: Angola and Zaire to Uganda; ssp. cybeutes occurs from W. Africa to

Zaire.

Andronymus neander neander Plôtz, 1884. Common in heavy woodland and forest

margins of Mpanda and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 7, 8a, 8b,

Gombe (800-1600 m).

Andronymus caesar philander Hopffer, 1855. Very common in heavy woodland

and forest margins of Mpanda and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b,

6, 8a, 8b, Gombe (800-1500 m).

Andronymus hero Evans, 1937. In riverine forests around Helembe, Mugondozi,

Zanza, Mihumu, Ntakatta, Lubalizi. Local and uncommon. Record: 3a, 5a, 8a,

KIELLAND: Rhopalocera eastern side Lake Tanganyika 217

8b (900-1400 m). General distribution: W. Africa to Angola and Zaire. Probably

first record from east Africa.

Andronymus helles Evans, 1937. From Kefu forest to Ntakatta. Very local and un-

common. Record: 5a, 8b (900-1400 m). General distribution: W. Africa to Zaire and

Malawi.

Andronymus marina Evans, 1937. Taken in the Ntakatta forest, Kefu and at Mi-

humu. Very local and rare (900-1600 m). General distribution: Toro and Mabira

forests in Uganda.

Andronymus fenestrella Bethune Baker, 1908. In dense thickets of riverine forests

of Mpanda and Kigoma. Not uncommon. Record: 2a, 3a, 3b, 4a, 5a, 5b, 8a, 8b (900-

1700 m). General distribution: W. Africa to Zaire, Uganda and Malawi.

Chondrolepis niveicornis Plôtz, 1883. Common in woodland and forests of Mpan-

da, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8b, Mbuzi Mt.

(800-2200 m). Specimens from western Tanzania are considerably larger than speci-

mens from Kilimanjaro, Arusha, Oldeani and east Tanzania. Genitalia identical.

Chondrolepis leggei Heron, 1909. A single male taken in the Lukandamira forest,

foot of Mahale. Record: 1 (1600 m). General distribution: Zaire to Uganda and

Kenya.

Zophopetes dysmephila Trimen, 1868. Near riverine forest at Kampisa. Flying at

dusk, after sundown. Males seen whirling in very rapid flight above a young, wild date

palm; females visiting flowers on herbs. April, July and September. Record: 5b

(1500 m). General distribution: S. Africa and Mozambique to Malawi, Uganda and

Senegal.

Zophopetes cerymica nobilior Holland, 1896. One male taken on flowering tree,

early morning at Gombe. Lush lake-shore vegetation. Record: Gombe (800 m).

General distribution: S. Africa, W. Africa, Mozambique, Malawi and Uganda.

Gamia shelleyi Sharpe, 1890. In the half-shade of thick forest at Luntampa, Lu-

balizi, Ntakatta and lower Ujamba at foot of Mahale Mt. Rare. Record: 1, 2a,

3a, 4a, 5a(1200-1300 m).

Artitropa reducta Aurivillius, 1925. Kasoge forest and Kefu forest. Very rare.

Record: 1, 8b (800-900 m). General distribution: Cameroun to Zaire, Kenya, Uganda

and Malawi. According to Berger reducta is a true species.

Artitropa cama Evans, 1937. Taken on bushes at a forest river in the Kefu forest.

One female March 1971; one male April 1972. Extremely rare and local. Record: 8b

(900 m). General distribution: Cameroun. Probably first record from East Africa.

Artitropa milleri milleri Riley, 1925. Taken June 1971 and 1972 at upper reaches of

Katuma river, between Mts. Sifuta and Sitebi; at Ntakatta February 1972. On

flowers in forest. Record: 5a, 5b (1400-1600 m).

Gretna carmen capra Evans, 1937. Rare in the Kefu and Kasoge forests. North

of Kigoma, at Gombe, it seems more common. Habit of settling on the bark of bare

tree trunks, where it blends perfectly. Record: 1, 4a, 5b, 8b, Gombe (800-900 m).

General distribution: the coast of Kenya and Tanzania.

Caenides leonora Plôtz, 1879. A single male taken in the Kasoge forest, February

1971, very early in the morning. Record: 1 (800 m). General distribution: W. Africa

to Zaire and Uganda; ssp. dux Evans, 1937, from Zaire and Malawi.

Caenides stoehri halma Evans, 1937. One specimen came to light at about 9 p.m.

218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

in a forest at Luntampa. Also taken at Lubalizi and Ntakatta in dense forest, in

daytime. April, September and November. Record: 3a, 4a, 5a (1200-1300 m). General

distribution: Uganda, Kenya and at Mpala west of Lake Tanganyika; nominotypical

race in W. Africa and Zaire.

Caenides dacela Hewitson, 1876. Forests of Kigoma and Mpanda. Not common.

Record: 1, 4a, 5a, 8a, 8b (800-1300 m). General distribution: W. Africa to Zaire and

Uganda.

Caenides hidarioides Aurivillius, 1896. Mukuyu forest in Kigoma and in the

Ntakatta forest in shady places. Rare. Record: 5a, 8a (800-1500 m). General distri-

bution: Sierra Leone and Ghana to Zaire. This is probably the first record from

East Africa.

Monza alberti Holland, 1896. Forests of Mpanda and Kigoma. Record: 1, 3b, 5b

(800-1500 m).

Monza cretacea crola Evans, 1937. Common in forests and heavy woodland of

Mpanda and Kigoma. Record: 1, 2a, 2b, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe

(800-1700 m).

Monza punctata Aurivillius, 1910. Forests of Mpanda and Kigoma. Record: 1, 5b

(1500-1700 m).

Fresna netopha Hewitson, 1878. Forest margins and forests of Mpanda and Ki-

goma. Not uncommon. Record: 1, 3a, 5a, 5b, 8b, Kibondo, Gombe (900-1500 m). Ge-

neral distribution: W. Africa to Uganda and Kenya.

Fresna nyassae nyassae Hewitson, 1878; forms: nyassae, ennuari Riley, 1921, and

plata Evans, 1937. Common in woodland of Mpanda and Kigoma. Record: 2a, 3a, 3b,

Figs. 35— 36, Male genitalia of Platylesches species. 35, P. ayresii Trim., Katuma river, Mpanda, 1600

m, no. 1090; a, tegumen-uncus; b, valva and saccus; c, aedeagus, 36. P. langa stat. n., Katuma river,

Mpanda, 1600 m, no. 1099; a, tegumen-uncus; b, valva; c, saccus; d, aedeagus.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 219

4a, 4b, 5a, 5b, 6, 8b, Gombe by Goodall (800-1500 m).

Fresna cojo Karsch, 1893. Two males taken in the Kefu forest February and April;

several males and a female taken at Lubalizi riverine forest on bean flowers and in the

Ntakatta forest, January 1974. Also August to September. A rare species. Record:

3a, 5a, 5b, 8b (900-1500 m). General distribution: Sierra Leone to Nigeria and

recorded from Kenya. A typical forest species.

Platylesches galesa Hewitson, 1877. Common in woodland and forests of Mpanda

and Kigoma. Record: 1, 2b, 3a, 3b, 4a, 5a, 5b, 6, 8b (800-1600 m).

Platylesches ayresii ayresii Trimen, 1889 (text-fig. 35). Sixteen males taken on wet

sand at upper Katuma river near Kampisa, June 1971 and June 1972. Record: 5b

(1600 m). This species, which Evans thought to be conspecific with his P. ayresii

langa, does in fact fly together with /anga in the recorded area. By dissecting the

genitalia of both species, it was found that the uncus in ayresii was much narrower

distally than in /anga and that in dorsal view the two distal protuberances were

much closer together; the valva with bipectinate dorso-distal process and less serrated

distad (figs. 35 and 36); the saccus is more sharply tapering in langa. In langa

the underside of the hind wing is only partly striated and the rest evenly sprinkled with

white. General distribution: According to Evans (1937) Mozambique to South Africa.

Platylesches langa stat. nov. (Platylesches ayresii langa Evans, 1937) (text-fig. 36).

Common in woodland and forests of Mpanda, Kigoma and Ufipa. Record: 3a, 4a,

4b, 5b, 6, Chulwe Mt., Mbuzi Mt. (1000-2000 m).

Platylesches robustus Neave, 1910. Forests and woodland of Mpanda, Sibweza,

Wanzizi, Kampisa, Lubalizi, Katuma. Locally common, particularly at Katuma. Rec-

ord: 3a, 3b, 5b, 6 (1060-1600 m). General distribution: S. Africa to Zaire, Zambia

and Malawi. One record from Geita in Tanzania and from Ngara.

Platylesches shona Evans, 1937. Forest at Kampisa in Mpanda. Very rare. Record:

5b (1500 m). General distribution: Zaire to Rhodesia and Zambia.

Platylesches moritili Wallengren, 1857. Rather common in woodland and riverine

forests of Mpanda and Kigoma. Record: 1, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe,

Kibondo (800-1700 m).

Platylesches fosta Evans, 1937. Taken at Kampisa, Ujamba at Mahale, Sitwe,

Ufipa. Highland forest margins. Uncommon. Record: 1, 3a, 5b, Mbuzi Mt. (1500-

2000 m). General distribution: known from Toro and Mpanga forests in Uganda.

Platylesches picanini Holland, 1894. Forests of Mpanda and Kigoma. Rather

common. Record: 1, 3a, 3b, 4a, 5a, 5b, 6, 8b, Gombe (800-1500 m, 2000 m at Mt.

Sitebi).

Platylesches rasta anka Evans, 1937. Montane forests of Mpanda, Kigoma and

Ufipa. Not uncommon. Record: 1, 5a, 5b, Mbizi forest, Mbuzi Mt. (1600-2100 m).

General distribution: Nominotypical race from Malawi and east Tanzania as at

Usambara and Mulundwe hill in Mikumi Nat. Park; ssp. anka described from W. An-

kole and Mpanga forest in Uganda.

Platylesches affinissima Strand, 1920. A small species closely resembling affinissima,

taken west of Mt. Karobwa, in the Kefu forest, at Mugombazi river, Ntakatta, Kam-

pisa and Lukandamira. Everywhere very rare. Record: 1, 2a, 5a, 5b, 8b (900-1700 m).

General distribution: Sierra Leone, Gambia, Central African Republic and Malawi.

This species has always been taken in forests.

220 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Platylesches tina Evans, 1937. Thirteen males taken on wet sand, in riverine forest

along the upper reaches of the Katuma river near Kampisa, July 1972. Record: 5b

(1600 m). General distribution: Malawi.

Brusa saxicola Neave, 1910. Woodland of Mpanda and Kigoma. Rather common

in the rainy season. Record: 1, 2a, 2b, 3a, 3b, 5a, 6, 8b, 9b (900-1700 m).

Brusa allardi Berger, 1967. Open rocky country of the Sitebi Mt. and near Mbizi

and at Chala Mts. Common in the dry season, but very local. According to Berger

also from Yampehu (Ugaraba) in Kigoma. All the specimens examined by the writer

from the Yampehu area were saxicola; also the habitat in that area differs consi-

derably (Brachystegia woodland) from the typical habitat of allardi (montane grass-

land with rocky outcrops). Record: (2a), 5b, Chala Mt., Mbizi Mt. (1800-2100 m,

Yampehu 1200 m). Always settling on rocks or bare ground. September 1970, 50

specimens were collected at Mt. Sitebi, all on rocks, one specimen observed at Mbizi

Mt. on rocks; several specimens taken on Mt. Chala on rocks. The flight is rapid, but

short and the insect difficult to detect. When settling, it seems to disappear due to its

cryptic markings and colour blending with the rocks.

Zenonia zeno Trimen, 1864. Very common in most habitats of Mpanda, Kigoma

and Ufipa. Record: widespread (800-2200 m).

Zenonia anax Evans, 1937. Woodland and forest margins of Mpanda and Kigoma.

Less common, but sometimes abundant at Mt. Sitebi. Record: 2a, 3a, 3b, 4a, 5a, 5b, 6

(1000-2000 m). General distribution: K atanga to Malawi.

Pelopidas mathias Fabricius, 1798. Common in woodland of Mpanda, Kigoma and

Ufipa. Record: 1, 3a, 3b, 6, 8b, Gombe, Chala (1000-2000 m).

Pelopidas thrax inconspicua Bertolini, 1850. Common in woodland of Mpanda and

Kigoma. Record: 1, 2a, 3a, 3b, 4a, 5a, 5b, 6, 7, 8b, Gombe (800-1500 m).

Borbo lugens Hopffer, 1855. Common in forests of Mpanda and Kigoma. Record:

1, 2a, 3a, 4a, 5a, 8a, 8b, Gombe (800-2000 m).

Borbo fatuellus Hopffer, 1855. One of the commonest skippers in Mpanda and Ki-

goma. In most habitats, but particularly woodland. Record: widespread (800-2000 m).

Borbo fallax Gaede, 1916. Woodland of Mpanda and Kigoma. Less common.

Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 6, 8b, Gombe (800-1700 m).

Borbo fanta Evans, 1937. Common in woodland of Mpanda and Kigoma. Also open

habitats at Mahale. Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 6, 8b, Gombe (800-2000 m).

General distribution: W. Africa to Zaire.

Borbo sirena Evans, 1937. Woodland of Mpanda and Kigoma, Sibweza and Ujom-

be. Rare. Record: 2b, 6 (1000-1200 m). General distribution: Zaire to Uganda, Kenya

and Zambia.

Borbo perobscura Hamilton Druce, 1912. Forests and forest margins of Mpanda,

Kigoma and Ufipa. Rather common. Record: 2a, 3a, 4a, Sa, 6, 8a, Muze (800-

1500 m).

Borbo detecta Trimen, 1893. Common in woodland of Mpanda and Kigoma. Rec-

ord: 2a, 3a, 3b, 4a, 5a, 5b, 6, 8b, Gombe (1000-1600 m).

Borbo micans Holland, 1898. Riverine forests and marshy habitats of Mpanda and

Kigoma. Record: 1, 2a, 3a, 3b, 4a, 4b, 5a, 5b, 6, 8a, 8b, Gombe (800-1700 m).

Borbo borbonica borbonica Boisduval, 1833. Common in woodland and open habi-

tats of Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6, 8b, Mbuzi

KIELLAND: Rhopalocera eastern side Lake Tanganyika 221

Mt., Gombe (800-2200 m).

Borbo gemella Mabille, 1884. Common in woodland of Mpanda, Kigoma and Ufipa.

Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6, Chala (800-1800 m).

Borbo holtzii Plôtz, 1883. Very common in woodland and savanna of Mpanda, Ki-

goma and Ufipa. Record: widespread (800-2200 m).

Borbo kaka Evans, 1946. A few specimens taken in the Kefu forest, Kasoge,

Luntampa, Ntakatta and at Lukandamira. Record: 1, 4a, 5a, 8b (900-1700 m). Gen-

eral distribution: K atera and Kayonza forests in Uganda; K akamega forest in Kenya.

Parnara naso monasi Trimen, 1889. Very common in forests and woodland of

Mpanda and Kigoma. Record: 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6, 8b, Gombe (800-1500 m).

Gegenes pumilio gambica Mabille, 1878. Common in woodland and savanna of

Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 5b, 6, 8b, Ufipa (800-

2000 m).

Gegenes niso brevicornis Plôtz, 1884. Very common in woodland and savanna of

Mpanda, Kigoma and Ufipa. Record: 1, 2a, 2b, 3a, 3b, 4a, 5a, 5b, 6, 7, 8b, Chala,

Gombe (800-1700 m).

Gegenes hottentota hottentota Latreille, 1823. In woodland and savanna at Sibweza

and Chala. Uncommon. Record: 6, 7, Chala (1060-1800 m).

SPECIES ERRONEOUSLY RECORDED FROM WESTERN TANZANIA

Liptena homeyeri Dewitz, 1884. Recorded by Stempffer, Bennet & May (1974)

from ”Tanzania, Mpala”. This locality is in Zaire, north-west of Lake Tanganyika.

Aphnaeus brahami Lathy, 1903. Recorded by Stempffer (1954) from “Tanganyika,

Mpala”, which is in Zaire, north-west of Lake Tanganyika.

SPECIES AND SUBSPECIES DESCRIBED FROM THE STUDY AREA OR ITS VICINITY

Carcasson (1966), in his account of the Japanese collection from the Kigoma area,

mentions 19 species and subspecies described from this area and its vicinity. Since

then the following taxa have been described (an asterisk indicates that the taxon is

described in the present paper):

*Graphium almansor wranghami Kielland, 1978

*G. porthaon tanganyikae Kielland, 1978

Charaxes cynthia mukuyu van Someren, 1969

C. lucretius maxima van Someren, 1971

C. ansorgei kungwensis van Someren, 1967

*C. ansorgei ufipa Kielland, 1978

C.jahlusa kigomana van Someren, 1971

C. numenes aequatorialis van Someren, 1972

C. imperialis ugandicus van Someren, 1972

C. bipunctatus ugandensis van Someren, 1972

C. chepalungu van Someren, 1969

C. grahamei van Someren, 1969

Palla publius kigoma van Someren, 1969

*Pseudathyma plutonica expansa Kielland, 1978

DDD TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

*Neptis ochracea reducta Kielland, 1978

*Belenois raffrayi similis Kielland, 1978

*Acraea ntebiae kigoma Kielland,1978

* 4 bisara neavei mahale Kielland, 1978

*Virachola ufipa Kielland, 1978

*Iolaphilus montana Kielland, 1978

Aphnaeus erikssoni kiellandi Stempffer, 1973

Lepidochrysops anerius kiellandi Stempffer, 1973

L. intermedia cottrelli Stempffer, 1954

*Telipna sanguinea kigoma Kielland, 1978

ACKNOWLEDGEMENTS

The writer wishes to express his thanks to the following persons: to Dr. R. H. Carcas-

son of the National Museum, Nairobi (until late 1968), for his willingness to supply in-

formation and help with identification; for his and the present entomologist Mr. M.

Clifton’s kindness to let me have free access to the Museum collection; to Dr. H.

Stempffer, Paris, for checking and correcting identifications of numerous Lycaenids;

to Dr. G. E. Tite of the Tring Museum, London, for advice regarding Lycaenids;

to Dr. C. B. Cottrell for correspondence concerning Neptis; to the late Dr. V.G. L. van

Someren for help with Charaxes; and to Dr. L. A. Berger, Tervuren, Belgium, and Dr.

M. Condamin, I. F. A. N., Dakar, for helpful correspondence. Finally, thanks are

due to Mr. M. Clifton for the tiresome task of correcting bad English, for

photographing specimens, for his and Dr. V. G. L. van Someren’s suggestions and

remarks, and to Mr. R. Mehl, Toyen Museum, Oslo, for additional photo’s.

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list of those recorded. — Journ. E. Afr. Nat. Hist. Soc. 27: 211—214.

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— Bull. B. M. (Nat. Hist.), Ent. Suppl. 9: 302.

Hübner, J., 1819. Verzeichniss Bekannter Schmetterlinge (7) 1:110.

Jackson, T. H. E., 1962. Notes on the Epitolinae with descriptions of a new genus and species and sub-

species (Lepidoptera: Lycaenidae). — Bull. B.M. (Nat. Hist.) (Ent.) 12: 125—162.

, 1969. A revision of the genus Telipna Aur. — Bull. B.M. (Nat. Hist.) (Ent.) 23 (1): 1—23.

Kielland, J., 1968. Rhopalocera from the eastern side of Lake Tanganyika, I. — Atalanta 2: 53—71.

, 1969. do. II. — Atalanta Norvegica, 3: 110—120.

Minig, A. 1976. Etude de la super espèce Charaxes manica Trimen et description de 2 espèces nou-

velles (Lepidopteres Nymphalidae). — Lambillionea 76 (1—2): 9—14.

Peters, W., 1952. Provisional check-list of the Butterflies of the Ethiopian Region. — E. W. Classey.

Stempffer, H., 1947. Note sur de Ornipholiditos du groupe de muhata Dewitz. — Rev. Zool. Bot.

Afr. 40 (2—3): 165—174.

—, 1951. Contribution à l’etude des Lycaenidae de la faune ethiopienne. — Bull. Soc. Ent. France

56: 119—128.

—, 1953. Contribution a l’etude des Lycaenidae de l’Afrique Equatoriale. — Ann. Mus. Roy.

Congo Belge 8 (27): 7—48.

— —, 1954. La réserve naturelle integrale du mont Nimba. II. Lepidopteres Lycaenides. — Bull.

I.F.A.N. 40: 347—354.

——, 1964. Révision of the genus Aphnaeus Hübner. — Trans. Roy. Ent. Soc. Lond. 105: 493—

516.

—, 1956. Contribution a l’etude des Lépidoptères Lycaenides de l’Afrique Equatoriale. — Ann.

Mus. Roy. Congo Belge 8 (49): 7—54.

——., 1957. Les Lepidopteres de l’ Afrique Noire francaise. 3, Lycaenides. I.F.A.N. 14.

224 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

—, 1957. Contribution a l’etude des Lycaenides de la faune ethiopienne. Bull. I.F.A.N. 19: 209—

DI:

—, 1963. La réserve naturelle integrale du mont Nimba. II, Lepidopteres Lycaenides. — I.F.A.N.

40: 423—427.

——, 1964. Contribution a l’etude des Lycaenides d’Afrique tropicale et equatoriale. — Bull. I.F.A.N.

(A) 26: 1226—1287.

—, 1965. Contribution a la faune du Congo (Brazzaville) Mission A. Villiers et A. Descarpen-

tries. — Bull. I.F.A.N. (A) 27: 1449—1465. .

——, 1966. Liste des Lepidoptéres Lycaenidae de Cote d’Ivoir actuellement connus. — Bull. I.F.A.N.

(A) 28: 1554— 1586.

——, 1967. Contribution a l’etude des Lycaenidae d’Afrique tropicale et équatoriale. — Bull. I.F.A.N.

(A) 29:978—1000.

——, 1967. The Genera of the African Lycaenidae. — Bull. B.M. (Nat. Hist.) (Ent.) Suppl. 10:

1—322.

—, 1972. Contribution a l’etude des Lycaenidae d'Afrique orientale (Lep.). — Bull. Soc. Ent.

France 77: 249—252.

Stempffer, H., & H. Bennet, 1953. A revision of the genus Teriomima Kirby. — Bull. B.M. (Nat.

Hist) (Ent.) 3: 77—104.

——, 1958. Revision des genres appartenant au groupe des Jolaus I. —Bull. I.F.A.N. 20: 1243—

1347.

, 1959. do. II. — Bull. I.F.A.N. 21: 226—325.

Tite, G. E., 1958. On three new species of the genus Syntarucus from the African region. — Entomologist

91:189— 191.

—, 1958. New African Lycaenidae (Lepidochrysops). — Entomologist 91: 115—118.

, 1961. New species of the genus Lepidochrysops. — Entomologist 94: 21—48.

Tite, G. E., & C. G. C. Dickson, 1973. The genus Aloeides and allied genera (Lepidoptera: Lycae-

nidae). — Bull. B.M. (Nat. Hist.) (Ent.) 29 (5): 227—280.

Van Someren, V. G. L., 1963. Revisional notes on African Charaxes I. — Bull. B.M. (Nat. Hist.).

(Ent.) 13 (7): 197—241.

——, 1964. do. II. — Bull. B.M. (Nat. Hist.) (Ent.) 15 (7): 183—235.

——, 1966. do. III. — Bull. B.M. (Nat. Hist.) (Ent.) 15 (7): 47—101.

——, 1967. do.IV.— Bull. B.M. (Nat. Hist.) (Ent.) 18 (9): 279—3 16.

——, 1969. do. V.— Bull. B.M. (Nat. Hist.) (Ent.) 23 (4): 77—166.

——, 1970. do. VI. — Bull. B.M. (Nat. Hist.) (Ent.) 25 (5): 199—250.

——, 1971.do. VII. — Bull. B.M. (Nat. Hist.) (Ent.) 26 (4): 183—226.

——, 1972. do. VIII. — Bull. B.M. (Nat. Hist.) (Ent.) 27 (4): 217—264.

—— , 1974. do.IX.— Bull. B.M. (Nat. Hist.) (Ent.) 29 (8): 417—487.

——, 1975.do. X.— Bull. B.M. (Nat. Hist.) (Ent.) 32 (3): 67—135.

225

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Plate 4. Figs. 13—16. Graphium almansor kigoma Carcasson, male, upper- and underside. 13—14, wet

season form; 15—16, dry season form.

Plate 5. Figs. 17—20. Graphium almansor wranghami ssp. n., male, upper- and underside. 17—18, wet

season form; 19—20, dry season form.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 229

24 Pl

Plate 6. Figs. 21—24. Graphium porthaon tanganyikae ssp.n., upper- and underisde. 21—22, male;

23—24, female.

230 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978 |

Plate 7. Figs. 25—26. Belenois raffrayi similis ssp. n., male, upper- and underside.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 231

Plate 8. Figs. 27 —30. Charaxes ansorgei ufipa ssp. n., upper- and underside. 27—28,

male; 29—30, female.

232 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

Plate 9. Figs. 31—33. Pseudathyma plutonica expansa ssp.n. 31—32, male, upper- and underside; 33,

female, upperside. Figs. 34— 36. Neptis ochracea reducta ssp. n. 34, female, upperside; 35—36, male,

upper- and underside.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 233

41 42

Plate 10. Figs. 37—42. Acraea ntebiae kigoma ssp.n., upper- and underside. 37—38, male; 39—40,

female; 42—42, female form.

234 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978

> -

Plate 11. Figs. 4345. Telipna sanguinea kigoma ssp.n. 43—44, male, upper- and underside; 45, female.

Figs. 46—49, Jolaphilus montana sp.n., upper- and underside. 46—47, male; 48—49, female.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 235

Plate 12. Figs. 50—51. Virachola ufipa sp.n., upperside. 50, male, holotype; 51, female, allotype.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL121, AFL. 4, 1978

236

Plate 13. Figs. 52—53. Virachola ufipa sp.n., underside. 52, male, holotype; 53, female, allotype.

KIELLAND: Rhopalocera eastern side Lake Tanganyika 237

Plate 14. Figs. 54—55. Abisara neavei mahale ssp. n., upperside. 54, male, holotype; 55, female, allotype.

(re

= 2

NOTEN

BDEEL 121 AFLEVERING 5 MUS. COMP. ZOOL 978

LIBRARY

FEBS 1979

HARVARD

TIIDBSCHRIFT 7

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

J. KLIMESCH. — Beitrag zur Kenntnis der Nepticulidenfauna von Anatolien und

der Insel Rhodos (Lepidoptera, Nepticulidae), p. 239—278, Fig. 1—82.

Tijdschrift voor Entomologie, deel 121, afl. 5 Gepubliceerd 31-XII-1978

BEITRAG ZUR KENNTNIS DER NEPTICULIDENFAUNA

VON ANATOLIEN UND DER INSEL RHODOS

(LEPIDOPTERA, NEPTICULIDAE)

von

J. KLIMESCH

Donatusgasse 4, 4020 - Linz, Osterreich

Mit 82 Figuren

ABSTRACT

In this paper the author gives a report of 30 Nepticulid species found in Anatolia and the Island of

Rhodes during several journeys from 1964—1977. Among the mentioned species there are 11 new ones:

Nepticula macrolepidella (on Quercus macrolepis), N. embonella (on Pyrus spinosa), N. azaroli (on

Crataegus azarolus), N. pyrellicola (on Rhamnus pyrellus), N. amygdali (on Prunus dulcis), N. muricatella

(on Sanguisorba muricata), N. styracicolella (on Styrax officinalis), Trifurcula (Ectoedemia) aegilopidella

(on Quercus macrolepis), T. (Fedalmia) trilobella (on Salvia triloba), T. (F.) albiflorella (on Nepeta nuda

albiflora) and T. (Fomoria) deschkai (on Hypericum hircinum and H. crispum). Detailed descriptions of

the new species are given together with drawings of the genitalia of the imagines and their mines.

EINLEITUNG

Die diesem Beitrag zugrunde liegenden Aufsammlungen und Beobachtungen

sind das Ergebnis mehrerer Reisen nach Anatolien (1964, 1966: Ende Mai bis

Mitte Juni; 1968: Anfang Juli; 1970: Ende Mai bis Anfang Juni) und Rhodos

(1971—1977, zwei- bis dreiwochige Aufenthalte, in der Hauptsache Ende August

bis Anfang Oktober, dreimal Mai bis Mitte Juni).

Wahrend auf den anatolischen Reisen der Lichtfang im Vordergrund der

Sammeltatigkeit stand, konzentrierte sich bei den Besuchen von Rhodos das

Interesse auf die Suche nach blattminierenden Lepidopteren. Von den dabei

festgestellten Arten sollen hier die Nepticuliden behandelt werden, Uber die aus

den besuchten Gebieten wie überhaupt aus dem kleinasiatischen und dem

ägäischen Raum nur sehr spärliche Nachweise vorliegen. Die Aufsammlungen

enthalten insgesamt 30 Arten, darunter 11 noch unbeschriebene.

In den meisten Fällen konnte die Bestimmung der Arten auf Grund von

Zuchten, nur bei wenigen Arten mit Hilfe von Minen allein durchgeführt werden.

Soweit bei den erzielten Imagines beide Geschlechter oder nur & vorlagen,

wurden Genitaluntersuchungen, dazu oft auch von der nächstverwandten Art,

vorgenommen und bei den Beschreibungen ausgewertet. Genitaluntersuchungen

unterblieben in den Fällen, in denen nur weibliches Material zur Verfügung stand,

da dieses bekanntlich bei vielen Arten noch keine befriedigenden Aus-

239

240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

wertungsmôglichkeiten bietet. So fehlt derzeit u.a. noch die Moglichkeit der

Sichtbarmachung feinster Details der Strukturen der Bursa copulatrix, die

Unterscheidungsmerkmale bieten konnten.

Einige der bemerkenswertesten Ergebnisse der Aufsammlungen mogen hier

erwahnt werden. So vor allem die Auffindung einer Nepticula-Art an Styrax

officinalis L., offenbar der ersten an einer Styracacee. Die systematische Stellung

der Art bleibt vorläufig noch unklar, da nur zwei 9 vorliegen.

Interessant war auch die Feststellung, daß offenbar Quercus macrolepis Kotschy

— aegilops L.) eine eigene Lepidopterenfauna zu haben scheint, wie man auf

Grund der Blattminierer und der Bewohner der charakteristischen großen

Eichelbecher annehmen kann. So wurden u.a. in den Blättern zwei neue

Nepticuliden (N. macrolepidella und T. aegilopidella) gefunden, daneben konnten

aber keine weiteren, von anderen weichblätterigen Eichen bereits bekannte

Nepticuliden festgestellt werden. Quercus ehrenbergi Kotschy hat dagegen drei

Nepticula (s.l.) -Arten mit europäischen Eichen gemeinsam.

Eine an Pistacia lentiscus L. gefundene Weberina-Art wird vorläufig noch zu der

aus dem Mittelmeergebiet bekannt gewordenen Weberina minimella (Rebel) (=

lentiscella Groschke) gestellt — trotz einiger anatomischer Unterschiede. Zur

Klärung der Frage, ob es sich nur um zwei differenzierte Formen oder distincte

Arten handelt, sind weitere Beobachtungen und Untersuchungen notwendig. In

der Nomenklatur werden hier im allgemeinen die von Johansson (1971)

gemachten Vorschläge berücksichtigt, die eine Teilung der Familie Nepticulidae

in zwei große Genera (Nepticula und Trifurcula) mit einer Unterteilung des

letzteren vorsieht. Diese scheint wenigstens vorläufig die beste Lösung zu Gunsten

einer leichten Übersicht zu sein.

In Dankbarkeit gedenkt Verfasser der unermüdlichen Mithilfe seiner

unvergeßlichen Frau, der tatkräftigen Unterstützung in botanischen Belangen

durch Frau Dr. F. Sorger und Herrn H. Metlesics, Wien, und der

kameradschaftlichen Unterstützung und Hilfe seiner Freunde E. Arenberger, der

Ehegatten Ing. W. und G. Glaser, der Herren G. Deschka und R. Johansson.

SYSTEMATISCHER TEIL

Es werden zuerst die bereits bekannten Arten behandelt, darauf folgen die

Neubeschreibungen.

Nepticula aurella (Fabricius, 1775)

(Fig. 1,2)

Tinea aurella Fabricius, 1775: 666.

Nepticula aurella; Staudinger & Rebel, 1901: 224. Beirne, 1945: 203. Johansson, 1971: 243.

Stigmella aurella; Hering, 1957: 909.

Rhodos, Mt. Smith und Straße nach Kalithea, 6. vi.1974, verlassene Minen an

Rubus fruticosus agg. zwischen Gebüsch. Die beobachteten Minen weisen teilweise

KLIMESCH: Nepticulidenfauna von Anatolien 241

Fig. 1—2. Nepticula aurella F. la, b: Minen an Rubus fruticosus agg., Rhodos, Kalithea, 6 vi. 1974; 2: 3

Kopulationsapparat, GU/Kli-324, Italia, Liguria, Bordighera, ex 1. 13.x1.1938, Rubus fruticosus agg. Fig.

3—4. N. mespilicola Frey, Minen an (3) Amelanchier parviflora Boiss. und (4) Cotoneaster racemiflora C.

Koch, beide Anatolia, NW-Ende des Beysehir gölü, 18.vi.1966. Fig. 5. N. trimaculella Hw., Mine an

Populus italica Moench, Rhodos, Paradissi, 28.viii.1973. Fig. 6. Minen an Ulmus glabra Huds. (?),

Rhodos, Rodini, 21.x.1971, von (6a) N. u/mivora Fologne und (6b) N. viscerella Stt. Fig. 7. N. basiguttella

Hein., Mine an Quercus ehrenbergi Kotschy, Anatolia, Mut, 8.vii.1968.

242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

eine kräftige, teils eine aufgelockerte, den Gang bis auf einen schmalen Rand

erfüllende Kotspur auf. Eiablage oberseitig (Fig. | a, b). Aus derartigen Minen

wurden aus verschiedenen Gebieten des Mittelmeerraumes Imagines erzielt, die

als Nepticula aurella (F.) im Sinne Beirnes (1945) anzusehen sind. Charakteristisch

für diese Art ist im männlichen Genitale die Bewehrung des Aedoeagus, die u.a.

aus einer schmalen, fuchsschwanzartigen Gruppe von Cornuti besteht (Fig. 2).

Verbreitung: Spanien, Frankreich, England, Irland, teilw. in den Skand.

Ländern, Mitteleuropa, SO-Europa und Nordafrika, Italien.

Nepticula mespilicola Frey, 1856

(Fig. 3, 4)

Nepticula mespilicola Frey, 1856: 392. Staudinger & Rebel, 1901: 224.

Stigmella mespilicola; Hering, 1957: 77.

Anatolia, NW-Ende des Beysehir gölü, ca 1100 m, 18.vi.1966, zahlreiche

verlassene und eine besetzte Mine an Amelanchier parviflora Boiss. Daraus

schlüpfte am 4.vii.1966 eine weibliche Imago, die sich von mitteleuropäischen

Stücken dieser Art nicht unterscheidet. Die Mine (Fig. 3) erscheint fast platzartig,

sie verläuft stark gewunden am Blattrand und weist eine kräftige Kotlinie auf.

Eiablage auf der Blattoberseite.

Am selben Standort wurden gleichzeitig an Cotoneaster racemiflora C. Koch elf

verlassene Gangminen (Fig. 4) gefunden, die mit Vorbehalt zu N. mespilicola

gezogen werden. Auch bei diesen ist die Eiablage oberseitig, der Verlauf der

Minen ist aber weniger stark gewunden, auch sind sie schmäler. Diese Merkmale

dürften aber auf die verschiedene Konsistenz der Substrate zurückzuführen sein.

Nepticula paliurella (Gerasimov, 1937)

Stigmella paliurella Gerasimov, 1937: 285; 1952: 251. Hering, 1957: 737 (=? zizyphi Walsingham, 1911:

190).

Nepticula paliurella; Klimesch, 1940: 177—179, 188—189.

Anatolia, Mut (Mersin), 8.vi.1966. Verlassene Minen an Paliurus spina-christi

Mill. Eine ostmediterrane Art.

Nepticula trimaculella (Haworth, 1829)

(Fig. 5)

Tinea trimaculella Haworth, 1829: 583.

Nepticula trimaculella; Staudinger & Rebel, 1901: 228.

Stigmella trimaculella; Hering, 1957: 810.

Rhodos, Paradissi, 28.viii.1973. Nur drei verlassene Minen an Populus italica

Moench; darunter eine stark gewundene, in sich verschlungene (Fig. 5). Die

Minen entsprechen mitteleuropaischen dieser Art.

Verbreitung: Europa bis in die Skandinavischen Lander. England.

KLIMESCH: Nepticulidenfauna von Anatolien 243

Nepticula ulmivora Fologne, 1860

(Fig. 6a)

Nepticula ulmivora Fologne, 1860: 92. Klimesch, 1975a: 6—8 (= ulmifoliae Hering, = ulmicola Hering).

Nepticula ulmifoliae Hering, 1931: 531—532.

Nepticula ulmicola Hering, 1932a: 568.

Stigmella ulmivora; Beirne, 1945: 199. Hering, 1957: 1089. Gustafsson, 1975: 3—4.

Stigmella ulmifoliae; Hering, 1957: 1089.

Stigmella ulmicola; Hering, 1957: 1089.

Rhodos, Rodini. Im Juni 1976 und im September 1972 und 1973 sowie im

Oktober 1971 verlassene Minen an Ulmus glabra Huds. (?) sowohl im schattigen

Parkgelände als auch außerhalb desselben an Straßenrändern nicht selten (Fig.

6a). In stark besonnten Blattern wurden Minen festgestellt, die die Merkmale der

Hering’schen Minenform ulmicola aufwiesen: an einer Blattrippe angelehnt,

gerade verlaufend, mit kompakter Kotablage. In Schattenblättern kommen

dagegen langere Minen mit aufgelockertem, halbkreisförmig abgelagerten Kot

und stark gewundene Minen zarterer Kotlinie vor (f. ulmifoliae Her.). Imagines

wurden nicht gezüchtet.

Nepticula viscerella Stainton, 1853

(Fig. 6b)

Nepticula viscerella Stainton, 1853: 3958. Klimesch, 1975a: 2—4 (= tauromeniella Groschke).

Nepticula tauromeniella Groschke, 1944: 117—118.

Stigmella viscerella; Beirne, 1945: 199. Hering, 1957: 1090.

Stigmella tauromeniella; Hering, 1957: 1090.

Rhodos, Rodini, 21.x.1971 (verlassene Minen), 22.ix.1972 besetzte Minen,

Imagines daraus 10.x.1972 und 13.iv.1973; 31.vin.1973 besetzte Mine, Imago

daraus 24.ix.1973; 1.vi.1973 besetzte Mine, 24.ix.1973 Imago. Die Minen an Ulmus

glabra Huds.(?)

Die Minen (Fig. 6b) entsprechen vollständig jenen, die Groschke (1944)

veranlaßte, eine neue Art — tauromeniella — anzunehmen. Sie sind infolge des

dickblätterigen Substrates auffallend kurz, in ihren Gangwindungen derart

zusammengedrängt, daß Einzelheiten nicht mehr wahrgenommen werden können.

Aus derartigen Minen wurden Imagines erzielt, die sich von mitteleuropäischen N.

viscerella Stt. nicht unterscheiden lassen (Klimesch, 1975).

Verbreitung: Lokal in ganz Europa, einschl. England. Fehlt in den Skand.

Ländern. Die Art scheint in südlichen Ländern häufiger aufzutreten.

Nepticula basiguttella Heinemann, 1862

(Fig. 7)

Nepticula basiguttella Heinemann, 1862: 258. Staudinger & Rebel, 1901: 221. Johansson, 1971: 256—258

(= cerricolella Klim.).

Stigmella basiguttella; Hering, 1957: 869.

Stigmella cerricolella Klimesch, 1946: 160— 162. Hering, 1957: 869.

Anatolia, Mut (Mersin), 8.vii.1968. Zwei verlassene Minen an Quercus

244 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

Fig. 8—9. N. samiatella Z. 8: Mine an Quercus ehrenbergi Kotschy, Anatolia, Kizilcahamam, l.vii.1968;

9: & Kopulationsapparat, GU/Kli-4109, dieselben Daten, ex |. 1.vii. 1968. Fig. 10—13. N. rhamnophila

Ams. 10: ¢ Kopulationsapparat, Holotypus, GU/Kli-721 (Coll. Amsel), Palaestina, Tabgha, See

722, dieselben Daten, ex |. 29.11.1933; 12: ¢ Kopulationsapparat, GU/Kli-1251, Rhodos, Lindos, ex I.

8.xi.1971, Rhamnus oleoides L.; 13: Mine an Rhamnus oleoides L., Rhodos, Lindos, 23.x.1971. Fig. 14. N.

freyella Heyd., Minen an Convolvulus althaeoides L., Rhodos, Kalithea, 1.1x.1973.

KLIMESCH: Nepticulidenfauna von Anatolien 245

ehrenbergi Kotschy. Die Minen (Fig. 7) weisen die für basiguttella

charakteristischen Merkmale auf: Gangminen, in denen der dunkelgrüne, spater

dunkelbraune Kot den Fraßgang vollständig ausfüllt, ohne Ränder freizulassen.

Eiablage oberseitig.

Eine an verschiedenen Eichenarten durch ganz Europa verbreitete Art.

Nepticula samiatella (Zeller, 1839)

(Fig. 8, 9)

Lyonetia samiatella Zeller, 1839: 215.

Nepticula samiatella; Johansson, 1971: 256—258.

Es unterbleibt die Anführung weiterer Zitate, da samtliche Autoren (Herrich-

Schaffer, Heinemann-Wocke, Stainton u.a.) die Art nicht richtig erkannt, sie

vielmehr mit den nahestehenden, besonders mit atricapitella Hw. und ruficapitella

Hw. vermischt haben. Erst Johansson (1971) gelang es, die Art einwandfrei

festzulegen.

Anatolia, Kizilcahamam, im ausgedehnten, aus Quercus ehrenbergi Kotschy

bestehendem Eichenbuschwald sowohl Minen (1.vii.1968) als auch Imagines am

Licht. Die Imagines wurden durch GU 4109 und 4110/Kli als zu samiatella Z.

gehorig bestatigt.

Außerdem liegen vom gleichen Fundort 4 von Holzschuh, Wien, am Licht

erbeutete & vor (GU 1314), die sich ebenfalls als N. samiatella Z. erwiesen.

Die Mine ist hinsichtlich der Dichte und Breite der Kotablage etwas

veränderlich. Meist handelt es sich um Gangminen mit kräftiger, aufgelockerter

Kotlinie, die breite Ränder des Ganges freiläßt (Fig. 8). Eiablage oberseitig.

Raupe hellgelb, Kopf hellbraun.

Die männlichen Imagines besitzen keine Duftschuppen auf den Hinterflügeln.

Das männliche Genitale (Fig. 9) ist ausgezeichnet durch einen langgestreckten

Aedoeagus, der eine schmale Reihe kräftiger Cornuti besitzt.

Nepticula rhamnophila Amsel, 1935

(Fig. 10— 13)

Nepticula rhamnella ssp. rhamnophila Amsel, 1935: 317.

Stigmella rhamnophila; Hering, 1957: 890.

Rhodos, Lindos, 23.x.1971, zahlreiche verlassene und bezetzte Minen an

Rhamnus oleoides L. Sträuchern an felsigen Stellen. Imagines (11 &, 14 9) daraus

5—14.xi.1971. |

Anatolia, Kizilcahamam, 29.v.1970, einige besetzte Minen an Rhamnus spec. im

Merkmalen mit N. rhamnophila übereinstimmen.

Die Minen (Fig. 13) verlaufen meist am Blattrand, sie sind stark gewunden und

enden oft platzartig. Kotlinie kräftig, breit, anfangs den Gang fast gänzlich

ausfüllend, später breite Ränder aufweisend. In kleinen Blättern nimmt die Mine

246 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

das ganze Blatt ein. Das Blatt wird durch einen Schlitz in der Oberseite verlassen.

Eiablage meist unterseitig. Kokon weiß, zart.

Die Imagines von N. rhamnophila weisen gegenüber der sehr nahestehenden N.

rhamnella H.-S. in beiden Geschlechtern schwarzbraune, leicht violett glänzende

Vorderflügel auf, während bei rhamnella das 3° etwas heller getönt erscheint als

Genitalunterschiede (Fig. 10, 11, 12) sind geringfügig. Sie liegen in der Hauptsache

in der flacheren Einbuchtung des Uncus und der Bewehrung des Aedoeagus, der

bei rhamnella außer zahlreichen verstreuten Cornuti noch eine Cornutalplatte

besitzt.

Zu N. rhamnophila Ams. dürften auch zwei am NW-Ende des Beysehir gölü

(Anatolien, Isparta) an Rhamnus rhodopaeus Velenovsky gefundene, verlassene

Minen gehören (14.vi.1966).

Nepticula freyella Heyden, 1858

(Fig. 14— 16)

Nepticula freyella Heyden, 1858: 175. Staudinger & Rebel, 1901: 225.

Stigmella freyella; Hering, 1957: 321.

Rhodos, Kremasti und Kalithea, an Straßenrändern und Brachen, zahlreiche

besetzte Minen Ende August und anfangs September 1973. Die Imagines

erschienen vom 9. bis 14.1x.1973 und stimmen völlig mit solchen aus Mitteleuropa

überein. Das männliche Genitale ist ausgezeichnet durch einen bogenförmigen,

gezähnten Uncus, zwei freie Gnathosäste, ventral grob gezähnte Valven, die in

einen nach innen gerichteten Fortsatz enden. Aedoeagus mit zahlreichen

verstreuten Cornuti bewehrt (Fig. 15). Die Bursa copulatrix des weiblichen

Genitale besitzt kein Signum; die Bursawand weist keine Struktur auf (Fig. 16).

Die Mine (Fig. 14) ist stark gewunden mit meist zarter Kotlinie. Eiablage

unterseitig. Raupe gelb mit brauner Kopfkapsel. Futterpflanzen: Convolvulus

arvensis L. und C. althaeoides L.

Verbreitung: Mittel- und Osteuropa, Corsica, Nordafrika.

Nepticula aceris Frey, 1856

(Fig. 17, 18)

Nepticula aceris Frey, 1856: 386. Staudinger & Rebel, 1901: 222.

Nepticula szoecsi Klimesch, 1955: 423—424.

Stigmella aceris; Hering, 1957: 20—21.

Anatolia, Kizilcahamam, Buschwaldgebiet, 3.vi.1970, zahlreiche besetzte

Minen an Acer tataricum L., aus denen vom 15—17.vi.1970 19 Imagines schlupften.

Während ein Teil der Tiere in ihren äußeren Merkmalen mitteleuropäischen

Stücken gleichen, weisen 8 Exemplare die für N. szoecsi Klim. (1955)

charakteristischen bindenlosen Vorderflügel auf. Angesichts der völligen

Übereinstimmung der Genitalien (Fig. 18) beider Formen besteht kein Zweifel,

KLIMESCH: Nepticulidenfauna von Anatolien 247

Fig. 15—16. Nepticula freyella Heyd. 15: & Kopulationsapparat, GU/Kli-1323, Rhodos, Kalithea, ex 1.

10.1x.1973, Convolvulus althaeoides L.; 16: 9 Kopulationsapparat, dieselben Daten. Fig. 17—18. N.

aceris Frey. 17: Mine an Acer tataricum L., Anatolia, Kizilcahamam, 3.vi.1970; 18: &

Kopulationsapparat, GU/Kli-4100, dieselben Daten, ex |. 16.vi.1970. Fig. 19—20. N. nivenburgensis

Preiß. 19: Minen an Salix alba L. (?), Rhodos, Kalithea, 1.ix.1973; 20: 4 Kopulationsapparat, GU/Kli-

4129, dieselben Daten, ex I. 15.ix.1977.

248 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

daß wir es nur mit einer Art zu tun haben. Diese scheint jedoch in Ungarn und in

Anatolien zeichnungslose Formen zu entwickeln.

Die Mine (Fig. 17) beginnt aus einem auf der Blattunterseite abgelegten Ei als

zarter sowohl in der Blattspreite als auch am Blattrand verlaufender Gang. Dieser

erweitert sich rasch und weist eine breite, nur sehr schmale Rander freilassende, in

Querbogen abgelagerte Kotspur auf. Der Kot ist ursprünglich dunkelgrün, von der

Farbe des Parenchyms, dunkelt aber bald bis schwarzlichbraun. Das Blatt wird

von der Raupe durch einen Schlitz in der Oberseite verlassen. Raupe lebhaft grün.

Kokon weiß, zart.

Nepticula nivenburgensis Preißecker, 1942

(Fig. 19—21)

Nepticula nivenburgensis Preißecker, 1942: 208— 211. Hering, 1943: 273—278.

Stigmella nivenburgensis; Hering, 1957: 928.

Rhodos, Straße nach Kalithea, Mine an Salix alba L. (?), ex 1. 19.ix.1973; weitere

Minen am gleichen Fundort: eine am 3.vi.1974 (Imago daraus 23.vi.1974),

zahlreiche Minen 1—S.ix.1977 (Imagines 15—24.1x.1977).

Gangmine (Fig. 19), meist unterseitig beginnend, zart, vielfach an eine Rippe

angelegt, daher wenig auffallend, mit scharfer, unterbrochener, den Gang anfangs

ganz ausfüllender, ca % der Gangbreite ausmachender Kotlinie. Während die

Mine anfangs im Schwammparenchym verläuft, nimmt sie im zweiten Abschnitt

beide Parenchyme in Anspruch, nicht selten verläuft sie jedoch durchwegs

unterseitig. Preißecker (1942) beschreibt die Mine sehr eingehend, er erwähnt

jedoch nicht, wo die Raupe die Mine zur Verpuppung verläßt. Bei den Zuchten

von Rhodos wurde bisher nur ein unterseitiges Verlassen des Blattes beobachtet.

Ei unterseitig, Raupe hellgelb.

Die rhodischen, ausschließlich durch Zuchten erhaltenen Exemplare stimmen

in allen Merkmalen mit mitteleuropäischen überein.

N. nivenburgensis ist auf Grund der männlichen Genitalien leicht zu erkennen.

Sie gehört in die Verwandtschaft der N. /uteella und N. glutinosae, die im

männlichen Genitale durch einen verbreiterten, apical flachen Uncus und einen

kurzen, mit gruppenspezifischen Cornuti bewehrten Aedoeagus ausgezeichnet

sind (Fig. 20). Am weiblichen Genitale (Fig. 21) fällt die Bursa copulatrix durch

feine Querwülste skulpturierte Bursawand auf. Es ist ein Signum vorhanden, das

aus einer langgestreckten, ovalen, randgekerbten Platte besteht, die an die Signa

der trifurculoiden Gruppe erinnert, aber keine wabenförmige Struktur aufweist.

Bei einem anderen Präparat (GU 4131/Kli) wurde neben der einen Platte noch

eine zweite festgestellt.

Auf Grund der Minenfunde und der Zuchtergebnisse sind zweifellos zumindest

zwei Generationen anzunehmen, dabei sind die im Frühjahr auftretenden Raupen

der ersten Generation weitaus seltener als die der Herbstgeneration.

Verbreitung: die Art wurde aus dem östl. Nieder-ôsterreich (Umgeb. v. Wien)

beschrieben, sie wurde auch in Ungarn (Szöcs) gefunden und in Oberitalien

(Klimesch) festgestellt. Es scheint sich demnach um eine südöstlich verbreitete

Art zu handeln.

KLIMESCH: Nepticulidenfauna von Anatolien : 249

Fig. 21. Nepticula nivenburgensis Preiß., 9 Kopulationsapparat, GU/Kli-4130, Rhodos, Kalithea, ex |.

15.1x.1977, Salix alba L. (?). Fig. 22a, b. N. promissa Stgr., Minen an Rhus coriaria L., Rhodos, Mt.

Smith, 25.v.1974. Fig. 23—24. Trifurcula (Ectoedemia) caradjai (Hering). 23: Mine an Quercus ehrenbergi

Kotschy, Anatolia, Mut, 8.vii.1968; 24: Mine an Quercus sp. (pubescens?), Rhodos, Salakos, 29.viii.1973.

Fig. 25. 7. (E.) spec. pr. suberis (Stt.), Mine an Quercus coccifera L., Rhodos, Trianta, 2.ix.1973.

250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

Nepticula promissa Staudinger, 1870

(Fig. 22)

Nepticula promissa Staudinger, 1870: 325. Staudinger & Rebel, 1901: 228. Klimesch, 1942: 395—396.

Stigmella promissa; Hering, 1957: 781.

Rhodos, Mt. Smith, 10.v.1975, besetzte Minen an Rhus coriaria L., aus denen

vom 25—29.v.1977 drei weibliche Imagines erzielt wurden.

Die Mine ist in ihrem Verlauf und in der Kotspur sehr variabel, bald dem

Blattrand folgend, bald in der Blattspreite (Fig. 22a, b). Die Kotablage kann breit,

aufgelockert, in Querbogen oder zarter, kompakt sein und kann in ein und

derselben Mine abandern. Demnach kann der Kot die Mine fast ganz erfullen

oder breite Rander freilassen. Hering (1957) ist geneigt, in den Minen mit zarter

Kotlinie eine noch unbeschriebene Art zu sehen.

Ei unterseitig, Raupe grun mit brauner Kopfkapsel, Kokon hellbraun.

An Pistacia atlantica Desf. wurden am gleichen Standort sowohl im Mai als auch

im September-Oktober nicht selten verlassene und nur einmal zwei besetzte

Minen (mit grunen Raupen) mit zarter Kotspur gefunden, die nach Ansicht des

Verfassers zu N. promissa Stgr. gehoren durften. Eine mediterrane Art, die bis ins

sudliche Mittel-Europa und Sud-Ungarn reicht.

Trifurcula (Ectoedemia ) caradjai (Hering, 1932)

(Fig. 23, 24)

Nepticula caradjai Hering, 1932b: 16. Groschke, 1944: 118—119.

Stigmella caradjai; Hering, 1957: 867.

Anatolia, Mut (Mersin), 8.vii.1968, besetzte Minen an Quercus ehrenbergi

Kotschy; Imagines daraus vom 5—11.viii.1968, 3 9 die mit solchen aus dem

Mittelmeergebiet (aus Quercus pubescens agg.) übereinstimmen.

Die charakteristische Mine (Fig. 23) beginnt mit kurzen, ganz mit Kot erfüllten

Windungen, die alsbald in einen ovalen bis kreisrunden Platz übergehen, in dem

der Kot im unteren Teil bald wolkig aufgelockert, bald kompakt

zusammenhängend an den Seiten abgelagert wird. Ei auf der Blattunterseite.

Rhodos, Salakos, 26.viii.1973, einige verlassene und wenig besetzte Minen an

Quercus pubescens agg. (?) (Fig. 24) mit oberseitiger Eiablage. Imagines daraus (2

Q) am 2.v.1974.

Die Art ist wie auch die übrigen Vertreter dieser Gruppe einbrütig. Die

anatolischen Funde lassen aber auf eine — wenigstens teilweise — zweite

Generation schließen.

Eine mediterrane Art, die am Alpenostrand bis über Wien hinaus nach Norden

vordringt.

Trifurcula (Ectoedemia) species

(Fig. 25)

Rhodos, Rodini, 1.ix.1973 und Trianta, 30.viii.1973 und 1977. Neben

KLIMESCH: Nepticulidenfauna von Anatolien 251

zahlreichen, vor mehr oder minder kurzer Zeit von den Raupen verlassenen

Platzminen an Quercus coccifera L. wurden auch einige besetzte gefunden. Daraus

entwickelten sich eine Imago am 16.ix.1973 sowie weitere am 24.iv und 4.v.1974,

an weichblätterigen Eichen lebenden T. caradjai Hering, an welche Art auch die

Minen erinnern.

Die Mine (Fig. 25) beginnt als gewundener, ganz mit Kot erfüllter Gang,

erweitert sich aber später zu einem Platz, in dem die Kotablage an dessen Basis

erfolgt. Eiablage oberseitig.

Die Mine ähnelt auch sehr jener der einbrütigen, im Februar an Quercus suber

letzteren ist aber wesentlich robuster als jene der rhodischen Art. Erst bei

Vorliegen eines zahlreicheren Zuchtmaterials in beiden Geschlechtern kann die

an Quercus coccifera L. gebundene Species geklärt werden.

Trifurcula (Ectoedemia) terebinthivora Klimesch, 1975

(Fig. 26—28)

Trifurcula (Ectoedemia) terebinthivora Klimesch, 1975b: 19—23.

Anatolia, Mut (Mersin), 8.vii.1968, zahlreiche verlassene Minen an Pistacia

terebinthus L. Anatolia, Kanlidivane, in einem Buschwaldgebiet an der Straße

Silifke-Mersin, 31.v.1970, mehrere, meist erst halbentwickelte Minen an dem

gleichen Substrat. Imagines daraus 24—30.vi.1970. Rhodos, Lindos, 20.x.1971,

zahlreiche verlassene alte Minen an Pistacia terebinthus L.

Mine (Fig. 26) anfangs stark gewundener Gang zwischen zwei Nebenrippen, von

schwarzem Kot fast ganz erfüllt. Später Platz mit dicht wolkig abgelagertem Kot,

der nur im letzten Abschnitt breitere Ränder freiläßt. Raupe fast farblos, glasig

erscheinend, mit schwarzbrauner Kopfkapsel. Kokon dunkelgrau. Eiablage auf

der Blattunterseite, meist an der Mittelrippe.

Männlicher Kopulationsapparat (Fig. 27). Vinculum schmal, flach

ausgeschnitten, Ventralarme der Transtillae relativ lang. Valven mit leicht

konkavem Ventralrand; sie enden in einen einwärts gerichteten Fortsatz.

Pseuduncus schwach sklerotisiert, plattenförmig. Uncus schwach gewölbt, apical

gezahnt. Gnathosäste apical verschmolzen. Aedoeagus lang, mit einer

Cornutalplatte und zwei dornartigen Cornuti.

Weiblicher Kopulationsapparat (Fig. 28). Das Signum der Bursa copulatrix

besteht aus zwei langgestreckten Platten mit unregelmäßig sechseckiger

wabenähnlicher Struktur.

Trifurcula (Ectoedemia) groschkei (Skala, 1943) comb. nov.

(Fig. 29— 34)

Nepticula groschkei Skala, 1943: 86.

Stigmella groschkei; Hering, 1957: 1117. Klimesch, 1948: 77—78.

Rhodos; Feraklos, Lindos, Kalathos, Massari, Archangelos, Salakos und

252 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

Fig. 26—28. Trifurcula (Ectoedemia) terebinthivora Klim. 26: Minen an Pistacia terebinthus L.; Anatolia,

Kanlidivane (Mersin), 31.v.1970; 27: & Kopulationsapparat, GU/Kli-1137, dieselben Daten, ex |.

30.vi.1970; 28: 9 Kopulationsapparat, GU/Kli-1138, dieselben Daten. Fig. 29—30. 7. (E.) groschkei

(Skala). 29: Minen an Vitex agnus-castus L., Rhodos, Kremasti, 3.ix.1973; 30; G Imago, Italia, Liguria,

Quiliano (Savona), ex I. 5.v.1945, Vitex agnus-castus L.

KLIMESCH: Nepticulidenfauna von Anatolien 253

Kremasti, an StraRenrändern, in trockenen Bachbetten und an Ufern sowie in der

Nahe des Meeresstrandes an Vitex agnus-castus, Ende August bis Oktober, meist

verlassene Minen. Aus den wahllos eingetragenen Blattern, die feucht gehalten

wurden, entwickelten sich in der Folge im Laufe von 8—10 Tagen einzelne Minen.

Ein Teil der Imagines entwickelte sich noch im September, wahrend die Mehrzahl

der Kokons überwinterte.

Die Mine (Fig. 29) beginnt als zarter Gang aus einem wahrscheinlich unterseits

abgelegtem Ei; sie zeigt eine unterbrochene schwarze Kotlinie, die die Rander

freiläßt. In der zweiten Phase entwickelt sie sich zu einem rundlichen Platz, meist

zwischen Mittelrippe und Blattrand, seltener in der Blattspitze. Die Kotablage ist

hier zentral, entweder wolkig aufgelockert oder kompakt. In dickeren Blättern ist

die Mine häufig unterseitig, sonst beiderseitig. Die blaßgrünliche Raupe mit

hellbrauner Kopfkapsel verläßt die Mine stets durch einen Schlitz auf der

Blattunterseite. Kokon rund bis schwach oval, olivbräunlich. Mehrbrütig.

Die Imagines sind etwas kleiner als solche von Italien (Ligurien, Quiliano-

Savona) (Fig. 30, 31) und etwas weniger deutlich gezeichnet. In den Genitalien

(Fig. 32, 33, 34) stimmen jedoch beide Populationen überein. Die systematische

wabenförmigen Struktur der ovalen, randgekerbten Sigma (Fig. 34).

Verbreitung: Bisher wurde T. groschkei in Sizilien (Locus classicus), Ligurien,

Dalmatien, Albanien, Griech. Mazedonien und Peloponnes festgestellt.

Vergeblich suchte nach ihr Verfasser in der Schlucht des Torrente de Parreis auf

Mallorca (Balearen) und bei Port Bou; an beiden Stellen kommt die Futterpflanze

häufig vor. Es könnte sich demnach um eine ostmediterrane Art handeln.

Trifurcula (Weberina) platani (Müller-Rutz, 1934) comb. nov.

(Fig. 35)

Weberina platani Müller-Rutz, 1934: 122—123; 1936: 672. Klimesch, 1940: 191—192.

Niepeltia platani; Hering, 1957: 789.

Rhodos, Stadtgebiet, einzelne verlassene Minen an Platanus orientalis L.,

Oktober 1971. Petaloudes, 28.viii.1973, verlassene Minen, darunter nicht selten

auch solche mit zarter Kotlinie. Funtukli, 19.x.1971, einzelne leere Minen, alle an

Platanus orientalis L.

Aus dem unterseitig abgelegten Ei entwickelt sich eine meist sehr stark

gewundene Gangmine. Kotablage in Halbkreisbogen, zuerst dunkelgrun, dann

dunkelbraun nachdunkelnd, den Gang ganz erfüllend (Fig. 35a), an gleiche

Erscheinungen wie bei N. aceris Frey an Acer-Arten erinnernd. Nicht selten

wurden aber auch Minen mit sehr zarter Kotlinie, sich oft in ihren stark

gewundenen Verlauf kreuzend, gefunden (Fig. 35b). Auch hier befindet sich die

Eischale auf der Blattunterseite. Selten wurde eine vollstandige Entwicklung der

Mine im Schwammparenchym festgestellt. In allen Fallen verläßt aber die Raupe

das Blatt durch einen Schlitz in der Blattoberseite. Hering (1957) zieht die Mine

mit zarter Kotspur zu einer noch unbeschriebenen Art. Zur Bestätigung dieser

Ansicht wären Imagines aus Zuchten notwendig.

254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

Fig. 31—34. Trifurcula (Ectoedemia) groschkei (Skala). 31: 9 Imago, Italia, Liguria, Quiliano (Savona),

5—20.v.1945; 32: & Kopulationsapparat, GU/Kli-211, dieselben Daten, ex |. v.1945; 33: &

Kli-1317, Italia, Liguria, Quiliano (Savona), ex I. 16.1x.1944; alle auf Vitex agnus-castus L.

Eine mediterrane Art, die auch noch im südlichen Mitteleuropa (Tessin und

Trentino) vorkommt.

KLIMESCH: Nepticulidenfauna von Anatolien 255

Fig. 35a, b. Trifurcula (Weberina) platani (M.R.), Minen an Platanus orientalis L., Rhodos, Petaloudes,

28.viii.1973. Fig. 36—39. 7. (W.) minimella (Rbl.). 36: & Kopulationsapparat, GU/Kli-4108, Rhodos,

Rodini, ex |. 8.xi.1972; 37: & Kopulationsapparat, Mallorca, Palma Nova, ex |. 7.ix.1968; 38: 9

Kopulationsapparat, GU/Kli-1312, Rhodos, Rodini, ex |. 27.x.1972: 39b: Minen, Rhodos, Rodini,

22.1x.1972; alle auf Pistacia lentiscus L.

256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

Trifurcula (Weberina) minimella (Rebel, 1926) comb. nov.

(Fig. 36—39)

Nepticula minimella Rebel, 1926: (106)—(110).

Nepticula species; Klimesch, 1942: 396.

Weberina lentiscella Groschke, 1944: 115—117.

Niepeltia lentiscella; Hering, 1957: 781.

Rhodos, Rodini, zahlreiche Minen, darunter mehrere besetzte, an Pistacia

lentiscus L.-Büschen an durch Trockenmauern geschützten Stellen, 22.ix.1972:

Imagines daraus 27.x—8.xi.1972 (5 3,6 9).

Aus dem oberseitig abgelegten Ei entwickelt sich die sehr charakteristische

Mine als fadendünner Gang, meist am Blattrand oder diesem zustrebend, mit

mehr oder wenigen Windungen. Später erfolgt die Kotablage in einer sehr breiten,

kompakten Spur, die nur geringe Ränder freiläßt (Fig. 39a und 39b).

Die Imagines von Rhodos sind durchwegs kleiner als solche von den Balearen

(Mallorca): 4.25 mm zu 5 mm Flügelspannung; auch sind sie dunkler getönt,

besonders die 9. Dies gilt auch für die Duftschuppen auf den Hinterflügeln des &,

die bei den rhodischen Tieren dunkler braun gegen hell rostfarben bei den

mallorquinischen erscheinen. Noch auffallender sind Verschiedenheiten in den

Genitalien (bes. beim &) von Tieren beider Gebiete: männliche rhodische Stücke

weisen am Ventralrand der Valven bei % (Fig. 36) einen kräftigen Vorsprung auf,

bei mallorquinischen Exemplaren ist dagegen bei % ein fingerförmiger, gezähnter

Fortsatz vorhanden (Fig. 37). Auch die Bewehrung des Aedoeagus ist bei beiden

Formen einigermaßen verschieden: bei der rhodischen sind zahlreiche kleine

höckerartige Cornuti und zwei grössere, stabförmige, etwas gebogene sichtbar,

während die mallorquinische in der Hauptsache zwei Cornutalplatten besitzt. Der

weibliche Kopulationsapparat der rhodischen Form (Fig. 38) weist in der Struktur

der Signa reticulata der Bursa copulatrix kleinere und gestrecktere Wabenzellen

auf als bei der mallorquinischen Form.

Auf Grund dieser Feststellungen dürfte man bei der Form von Rhodos

zumindest eine Subspecies der 7. minimella Rbl. annehmen können. Trifurcula

(Weberina) minimella wurde von Rebel (1926) auf Grund eines von Nowak bei

Sucurac (Dalmatien) gefangenen ¢ beschrieben. Die Etikette des betr. Tieres

lautet “Sucurac, 9.23, Nowak”. Dieses Stück (Holotypus) wurde anatomisch

untersucht (GU/Kli-446a in Coll. Mus. Vind.) und völlige Übereinstimmung mit

der Form von Mallorca festgestellt. Bedauerlicherweise ist das Typenmaterial der

Weberina lentiscella Groschke nach einer Mitteilung des Museums für Naturkunde

in Stuttgart, wo Groschkes Sammlung aufbewahrt war, verschollen. Groschke hat

aber Minen, die vom Verfasser bei Zaton-Gruz (Dalmatien) an Pistacia lentiscus L.

gesammelt wurden (Klimesch, 1942) als zu seiner Weberina lentiscella gehörig

anerkannt (Groschke, 1944). Aus mit diesen übereinstimmenden, auf Mallorca

gesammelten Minen erhielt Verfasser Imagines, auf die die Beschreibung

Groschkes (1944) in allen Punkten paßt, die genitaliter auch der W. minimella Rbl.

entsprechen und daher nichts mit jenen von Rhodos zu tun haben. Es kann somit

daraus mit Sicherheit die Synonymie der Weberina lentiscella Groschke mit W.

minimella Rebel gefolgert werden.

KLIMESCH: Nepticulidenfauna von Anatolien 257

Daher wurde aus dem mallorquinischen Material ein Neotypus (G) und ein

Paratypus (9) ausgewählt. Das betreffende Typenmaterial trägt folgende

Bezettelung: “& Ins. Baleares, Mallorca, Palma Nova, ex 1. 26.6.1968, Mine an

aber mit Schlüpftag der Imago “25.8.1968”.

Über die Generationen der Rhodos-Form läßt sich mangels entsprechender

Beobachtungen nichts aussagen. Das durch Zucht erzielte Imagines-Material

entwickelte sich im Verlaufe von 4—5 Wochen, während die am 10—12.iii.1968

auf Mallorca eingesammelten Raupen die Falter von Ende April bis Ende August

1968 ergaben.

NEUBESCHREIBUNGEN

Nepticula macrolepidella spec. nov.

(Fig. 40—45)

In die ruficapitella Hw.-Gruppe gehörig, von allen europäischen Arten aber

äußerlich auffällig verschieden.

Fühler beim ¢ bis fast 3/4 der Vorderflügellänge reichend, gelblichgrau bis

graubraun, die einzelnen Glieder nicht deutlich hervortretend, beim 9 kürzer,

etwas über 1/2. Augendeckel groß, weiß.

Kopfhaare bei beiden Geschlechtern im Gesicht hellgelblich bis bräunlich, im

Nacken dunkelbraun bis schwarz. Nackenschöpfe gelblichweiß, beim 9 zuweilen

dunkelbraun. Maxillarpalpen lang, hellgelblich; Labialpalpen kürzer. Thorax und

Abdomen gelblichgrau, die Tegulae gelblich, mehr oder weniger stark von

schwarzen Schuppen bedeckt. Beine hellgelb, beim g die Tarsen des 3.

Beinpaares grau gefleckt.

Feinschuppig. Expansion 4—4.5 mm. Vorderflügel-Grundfarbe hellgelb,

stellenweise dicht mit schwarzbraunen Schuppen bedeckt. Die Grundfarbe tritt an

der Basis, am Vorderrand und in einem Streifen am Innenrand nach der Mitte

ungetrübt hervor. Die restlichen Teile der Vorderflügel sind mehr oder minder

stark schwarzbraun beschuppt (Fig. 40—41). Seltener kommen stärker

verdunkelte Stücke — bei beiden Geschlechtern — vor, stets aber bleibt an der

Basis, am Vorder- und Innenrand eine gelbliche Aufhellung. Vorderflügelfransen

gelblichgrau, nicht scharf vom Flügelgrund abgesetzt, um den Tornus am

Innenrand dunkler grau. Hinterflügel grau mit hellgrauen Fransen.

Kopulationsapparat. Das männliche Genitale zeigt die typische Bauart der

ruficapitella-Gruppe (Fig. 42). Vinculum anal flach ausgeschnitten. Uncus doppel-

zackig, median flach gekerbt. Gnathosäste voneinander getrennt. Valven mit

gerade verlaufendem Ventralrand, distal mit zwei einwärts gekrümmten, spitzen

Vorsprüngen, von denen der untere kräftig ausgebildet ist. Aedoeagus sehr kräftig,

mit zahlreichen Cornuti, die lateral von kürzeren, kleineren begleitet werden.

Charakteristisch sind vier sehr große, bis zur halben Länge des Aedoeagus

reichende Cornuti im unteren Teil.

Beim Weibchen (Fig. 43) sind die vorderen Gonapophysen gestreckt, am

258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

Fig. 40—45a. Nepticula macrolepidella spec. nov. 40: g Imago, Rhodos, Kremasti, ex |. 24.ix.1973,

Quercus macrolepis Kotschy; 41: 9 Imago, dieselben Daten, ex 1. 17.1x.1973; 42: & Kopulationsapparat,

Rodini, ex I. 11.x.1972, Quercus macrolepis Kotschy; 44: Minen an Q. macrolepis Kotschy, Rhodos,

macrolepis Kotschy.

KLIMESCH: Nepticulidenfauna von Anatolien 259

oberen Ende verbreitert, die hinteren stabformig. Das Signum der Bursa

copulatrix besteht aus einer nach innen gerichteten, im oberen Abschnitt

gebogenen Reihe kräftiger Stacheln. Lateral befinden sich außerdem kleinere

Felder von kurzen Stacheln und Höckern.

Die neue Art weist keine näheren Beziehungen zu den übrigen Arten der

ruficapitella-Gruppe auf.

Futterpflanze: Quercus macrolepis Kotschy (= aegilops L.).

Mine. Aus dem oberseitig abgelegten Ei entwickelt sich eine beiderseitige

Gangmine, die zuerst mit kompakt abgelagertem Kot ganz erfüllt ist. Später

lockert sich die Kotspur auf und läßt mehr oder minder breite Ränder in der Mine

frei. Die Mine erinnert (Fig. 44) sehr an die der ruficapitella-Gruppe. Meist e

twickeln sich nur einzelne Minen im Blatt. An einem völlig verstaubten Quercus

macrolepis-Strauch an einem Straßenrand bei Kremasti (Rhodos) konnte ein

Massenvorkommen der Mine anfangs September 1973 beobachtet werden. In den

einzelnen, ganz vom Straßenstaub bedeckten Blättern konnten mindestens 5—10

Minen festgestellt werden. Bei entsprechender Feuchthaltung entwickelten sich in

den folgenden Tagen noch weitere Minen!

Raupe hellgelb, Kopfkapsel braun. Kokon sehr zart, durchscheinend, hellgelb,

oval.

Puppe (Fig. 45) der Bauart der Pupa libera, also mit frei beweglichen

Gliedmaßen angehörend; sie ist flach mit deutlich sichtbaren Maxillarpalpen und

Coxen. Stigmen vom |. Hinterleibssegment bis zum 5. Dorsal ab dem 3. Abdo-

minalsegment je 16 kurze, kleine, in Reihen angeordnete Dornen.

Der Beschreibung liegen 111 Imagines zu Grunde, die folgende Bezettelung

tragen: 68 4, 41 9 = “Mine an Quercus macrolepis, 3.9.1973, Zucht Nr. 1084,

Rhodos, Kremasti, ex 1. 15.—26.9.1973, J. Klimesch”. Der Holo- und ein

Paratypus tragen folgende Etiketten: “Mine an Quercus macrolepis, 22.9.1972,

Zucht Nr. 1054, Rhodos, Rodini, ex 1. 15.10.1972”.

Nepticula embonella spec. nov.

(Fig. 46, 47, 49)

In die oxyacanthella-Gruppe gehörig, der N. pyri Glitz zunächst verwandt.

Fühler schwarzbraun, beim & bis % der Vorderflügellänge reichend, beim 9

etwas kürzer. Maxillarpalpen gelblich, ebenso die kürzeren Labialpalpen.

Kopfhaare bei beiden Geschlechtern von rostfarben bis dunkelbraun und

schwarz variierend, Nackenschöpfe hellbraun. Augendeckel gelblichweiß. Thorax

und Abdomen dunkelgrau, unterseits heller. Beine hellgrau.

Eine kleine Art: Expansion 2.5—3 mm. Glattschuppig. Vorderflügel einfarbig

dunkelgraubraun mit mattem Glanz. Fransen dunkelgrau, um den Apex heller mit

licht glänzenden Enden. Hinterflügel grau mit helleren Fransen, die am Innenrand

dunkler erscheinen. Unterseite aller Flügel grau.

Kopulationsapparat. Männchen (Fig. 46): Vinculum relativ schmal, flach

ausgeschnitten. Valven mit bauchig verlaufendem Ventralrand und einem

260 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

O.1mm

0.1mm 0.1 mm

Fig. 45b. Nepticula macrolepidella spec. nov, 3 Puppe in Dorsalansicht, aus Zucht, Rhodos, Kremasti,

ix.1973, Q. macrolepis Kotschy. Fig. 46—47, 49. N. embonella spec. nov. 46: ¢ Kopulationsapparat,

GU/Kli-1307, Rhodos, Embonas, ex I. 13.ix.1973, Pyrus spinosa Forsk.; 47: 9 Kopulationsapparat, GU/

Kli-4103, dieselben Daten, ex I. 20.iii.1974; 49: Minen an Pyrus spinosa Forsk., selben Fundort,

29.viii.1973. Fig. 48. N. pyri Glitz. G Kopulationsapparat, GU/Kli-1249, Austria sup., Umg. v. Linz, ex I.

14.ix.1935, Pyrus communis agg. Fig. 50—51. N. azaroli spec. nov. 50: G Kopulationsapparat, GU/Kli-

1262, Anatolia, NW-Ende d. Beysehir gölü, 14.vi.1966; 51: 9 Kopulationsapparat, GU/Kli-4102,

Rhodos, Mt. Smith, ex I. 18.ix.1973; beide an Crataegus azarolus L.

KLIMESCH: Nepticulidenfauna von Anatolien 261

zahnartigen Vorsprung im oberen Teil sowie einem fingerartigen Fortsatz im

apicalen Bereich. Gnathos mit zwei distalen Âsten. Aedoeagus mit einer Gruppe

verstreuter kleiner Cornuti und einem Cornutalfeld.

Weibchen (Fig. 47). Gonapophysen von der üblichen Bauart: die vorderen

flachig verbreitert, die hinteren stabformig. Die schwach sklerotisierte Bursa

copulatrix weist kein Signum auf.

Die zunächst stehende Nepticula pyri Glitz ist größer (4.5—5 mm) und viel

starker, kupferig, glanzend.

Im mannlichen Kopulationsapparat zeigt N. pyri vor allem Unterschiede im

Gnathos, deren Aste hier kürzer und in der Basis breiter sind (Fig. 48). Im

Aedoeagus ist die Zahl der verstreuten Cornuti groRer, das Cornutalfeld kleiner.

Die Richtigkeit der Bestimmung der hier zum Vergleich mit der neuen Art

herangezogenen Nepticula pyri Glitz-Stucke wurde von Herrn Roland Johansson

durch Überprüfung und Vergleich mit pyri-Originalen in dankenswerter Weise

bestatigt.

Futterpflanze: Pyrus spinosa Forsk. (= amygdaliformis Vill.). Aus dem meist

unterseitig, seltener oberseitig abgelegten Ei entwickelt sich eine wenig

gewundene, eine unterbrochene Kotspur aufweisende Gangmine, die sich in der

zweiten Phase platzartig verbreitert und nun eine breite, aufgelockerte Kotlinie

zeigt. Raupe grun mit hellbrauner Kopfkapsel. Kokon oval, von fester Konsistenz,

dunkelbraun.

Bei der Zucht der Ende August 1973 eingetragenen Raupen schlüpften die

Imagines bereits nach 8—10 Tagen, ein kleiner Teil entwickelte sich jedoch erst

nach der Uberwinterung im April 1974. Die Art ist sicher mehrbritig. Fundorte:

Buschwaldgebiete und Straßenränder bei Embonas, Lindos, Trianta auf Rhodos.

Bereits fruher wurden anderwarts von der neuen Art meist leere Minen, nur

einzeln aber besetzte gefunden. Es wurden keine Imagines erzielt. Hering hielt die

Minen als zu Nepticula minusculella H.-S. gehorig. Folgende Fundorte wurden

registriert: Mazedonien (Drenovo-Kavadarci, Stari Dojran, Ohrid-Sv. Stephan),

Anatolien (Manisa). Der Beschreibung liegen 60 Exemplare zugrunde; sie tragen

folgende Bezettelung: “Mine an Pyrus spinosa, 29.8.1973, Zucht Nr. 1078, ex 1.

10.—20.9.1973 und ex 1. 20.2.—20.3.1974, Rhodos, Embonas, J. Klimesch’’. Der

Holotypus und die Paratypen wurden als solche unter diesem Material

gekennzeichnet.

Nepticula azaroli spec. nov.

(Fig. 50, 51, 53)

In die pomella-Gruppe gehörig, der N. pygmaeella Hw. zunächst stehend.

Fühler dunkelgrau, beim g etwas über 1/2 der Vorderflügellänge reichend,

Kopfhaare bei beiden Geschlechtern hell rostfarben, Nackenschôpfe lichter.

Augendeckel gelblichweiß.

Thorax und Hinterleib dunkelgrau, unterseits hellgrau, glänzend. Beine grau,

unterseits gelblichgrau.

Expansion 3—4 mm. Glattschuppig. Vorderflügel einfarbig dunkel aschgrau,

262 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

Fig. 52. Nepticula pygmaeella Hw., 3 Kopulationsapparat, GU/Kli-1261, Austria inf., Dürnstein, ex I.

25.iii.1936, Crataegus oxyacantha L. Fig. 53. N. azaroli spec. nov. Mine an Crataegus azarolus L.,

Rhodos, Rodini, 4.ix.1973. Fig. 54—55. N. pyrellicola spec. nov. 54: Q Imago, Anatolia, Mut (Mersin),

ex 1. 19.vi.1966, Rhamnus pyrellus O. Schwarz; 55: Mine an R. pyrellus O.S., dieselben Daten, 6.v1.1966.

Fig. 56—57. N. amygdali spec. nov. 56: g Imago, Rhodos, Lindos, Prunus dulcis D. A. Webb, ex. |.

28.vi.1976; 57: & Kopulationsapparat, GU/Kli-4121, dieselben Daten, ex I. 14.vii.1976.

KLIMESCH: Nepticulidenfauna von Anatolien 263

matt glänzend. Fransen dunkelgrau, nicht vom Flügelgrund abgesetzt.

Hinterflügel grau mit ebensolchen Fransen. Unterseite aller Flügel einfarbig grau.

Kopulationsapparat. Männchen (Fig. 50). Vinculum tief ausgeschnitten.

Ventralrand der Valven fast gerade verlaufend, apical mit kurzem, nach innen

gerichteten fingerformigen Fortsatz. Uncus median ausgeschnitten, mit zwei

lateralen Kerben. Aedoeagus mit drei Reihen kraftiger Cornuti und einer

Cornutalplatte bewehrt.

Weibchen (Fig. 51). Die vorderen Gonapophysen flächig verbreitert, die

hinteren stabförmig. Das Vestibulum bauchig erweitert. Die Bursa copulatrix

verhältnismäßig klein, schwach sklerotisiert, ohne Signum oder sonstige

erkennbare Struktur der Bursawand.

Nepticula azaroli steht der an Crataegus oxyacantha und C. monogyna lebenden N.

pygmaeella Hw. sehr nahe, N. pygmaeella ist aber größer und robuster (Expansion

4.5—5.5 mm), hat dunkelbraune, matt violett schimmernde Vorderflügel und

heller rostfarbene Kopfhaare.

Im männlichen Genitale fallen vor allem Unterschiede im lateralen Bereich des

Uncus auf; dieser ist bei azaroli durch gerundete Höcker ausgezeichnet, bei

pygmaeella (Fig. 52) sind diese deutlich mehr zugespitzt. Die Cornuti des

Aedoeagus scheinen bei pygmaeella zahlreicher vorhanden zu sein als bei azaroli.

breiter, das Vestibulum erscheint stärker ausgebaucht als bei azaroli. Beiden Arten

fehlt ein Signum, auch zeigt die Bursawand keine erkennbaren Strukturen.

Futterpflanze: Crataegus azarolus L. Die Mine beginnt häufig an einer Rippe aus

einem unterseitig abgelegten Ei, meist an der Mittelrippe, zuerst als zarter mit

schwarzem Kot ganz erfüllter Gang, dann platzartig verbreitert mit zentraler,

aufgelockerter Kotlinie. Der Kot füllt hier die Mine nur zu 1/3—1/4 aus (Fig. 53).

Raupe hellgelb, Kopfkapsel braun, Kokon oval, flach, glatt, braun.

Fundorte. Anatolia, Mut (Mersin), einzelne besetzte Minen, 6.vi.1966, Imago

daraus 20.vi.1966; NW-Ende des Beysehir gölü (Isparta), 1100 m, Minen

14.vi.1966, ex 1. 29.vi.1966. Rhodos, Rodini, und Mt. Smith, nahe der Akropolis,

Mitte Mai 1974 einzelne Minen, häufig dagegen im September 1972 und 1973. Aus

letzteren wurden zahlreiche Imagines erzielt, die sich nach einer ca 10-tagigen

Puppenruhe ausnahmslos noch im Herbst entwickelten, zu einer Zeit in der die

Futterpflanze bereits ihr Laub abgeworfen hatte. Dies legt die Vermutung nahe,

vollziehen dürften. Eine Überwinterung der Imagines erscheint wohl wenig

wahrscheinlich. Ein Hochzeitsflug der Art wurde am 18.vi.1966 am NW-Ende des

Beysehir gölü (Anatolien) bei Sonnenaufgang beobachtet: zahlreiche Z

schwärmten bei völliger Windstille knapp über dem Crataegus azarolus-Gebüsch.

Beim Lichtfang, der in nächster Nähe dieser Büsche ausgeführt wurde, erschienen

niemals Imagines an der Leinwand.

Der Holotypus sowie der Paratypus und weitere Paratypen wurden unter den 88

Exemplaren der Zuchten ausgewählt. Die Tiere tragen folgende Bezettelung:

“Mine an Crataegus azarolus, Zucht Nr. 1052, 22.9.1972, Rhodos, Rodini, ex 1.

10.—11.10.1972 (4 &, 4 9)”; sowie “Zucht 1085, Mine 4.9.1973, Crataegus

azarolus, Rhodos, Akropolis, ex 1. 18.—30.9.1973 (34 8,44 9), J. Klimesch.”

264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

Nepticula pyrellicola spec. nov.

(Fig. 54, 55)

Eine kleine, in die Verwandtschaft der Nepticula crenulatae Klim. gehorige Art.

Da nur 2 9 vorliegen, ist die Beschreibung der Art als vorläufig zu werten.

Fühler bis 1/2 der Vorderflügellänge reichend, dunkelgrau, schwach geringt.

Kopfhaare hell rostgelb, Nackenschopfe weißlich. Augendeckel weiß. Maxillar-

palpen weißlich wie auch die kleineren Labialpalpen.

Thorax und Hinterleib grau, unterseits etwas heller. Die gleiche Färbung weisen

auch die Beine auf; die Hinterschienen und die Hintertarsen weißlich geringt.

Expansion 3 mm. Grobschuppig. Grundfarbe der Vorderflügel grau, dicht mit

schwarzgrauen Schuppen bedeckt, nach 1/2 der Vorderflügellänge mit einer

breiten, weißen median etwas eingeengten Querbinde. Fransen hellgrau, vom

Flügelgrunde scharf abgesetzt, im Apex weißlichgrau, im Tornus am Innenrand

dunkelgrau. Unterseite der Vorderflügel einfarbig dunkelgrau, am Innenrand

etwas aufgehellt (Fig. 54). Die ähnliche N. crenulatae Klim. ist wesentlich dunkler

und besitzt eine schmälere, matt silbern glänzende Querbinde.

Nepticula pyrellicola gehört in die Gruppe der an Rhamnus-Arten lebenden

Species (N. catharticella Stt., N. rhamnella H.-S., alaternella Le Md.). Die

Feststellung ihrer nächsten Verwandtschaft wird erst nach anatomischer

Untersuchung des noch unbekannten & möglich sein.

Futterpflanze: Rhamnus pyrellus O. Schwarz.

Mine. Aus dem unterseits abgelegten Blatt entwickelt sich die Mine zuerst

oberseitig, anfangs sehr schwer erkennbar, meist am Blattrand, seltener in der

Blattspreite, in grosseren Blattern erst im Endabschnitt stark gewunden. In

kleineren Blattern nimmt die beiderseitige Mine die Blattflache in mehreren

Windungen fast ganz ein (Fig. 55). Kokon zart, weißlich.

Die Minen wurden zahlreich, doch nur zwei von Raupen besetzt, in der Ebene

von Mut (Mersin, Anatolien) am 6.vi.1966 auf fast sterilem Boden, auf dem die

Futterpflanze verstreut vorkommt, gefunden. Aus der Zucht (Nr. 890) schlüpften

zugrunde gelegt wurden.

Nepticula amygdali spec. nov.

(Fig. 56—59)

In die N. anomalella Goeze-Gruppe gehörig.

Fühler schwarz, beim g etwas über 1/2 der Vorderflügellänge reichend, beim 9

bis 1/2. Maxillarpalpen gelblichweiß. Kopfhaare bei beiden Geschlechtern hell

rostfarben. Nackenschöpfe weiß, ebenso die Augendeckel.

Thorax und Abdomen schwarz, Analende bei beiden Geschlechtern gelblich.

Beine hellgrau, unterseits gelblichgrau, die Tarsen des 3. Beinpaares dunkel

gefleckt.

Expansion: 3.5—4 mm. Grobschuppig. Vorderflügel-Grundfarbe bei beiden

Geschlechtern dunkelgrau, dicht mit schwarzen Schuppen bedeckt, lediglich im

Außenfeld und am Tornus ist die Grundfarbe noch schwach erkennbar. Fransen

KLIMESCH: Nepticulidenfauna von Anatolien 265

10mm

Fig. 58—59. Nepticula amygdali spec. nov. 58: 9 Kopulationsapparat, GU/Kli-4128, Rhodos, Treas,

Prunus dulcis D.A. Webb, ex 1. 4.x.1977; 59: Minen an P. dulcis D.A.W., Rhodos, Lindos, 6. vi.1976. Fig.

60, 62. N. muricatella spec. nov. 60: & Kopulationsapparat, GU/Kli-1321, Anatolia, Meram b. Konya,

Sanguisorba muricata (Spach) Focke, ex 1. 2.vii.1966: 62: Minen an S. muricata (S.) F., selben Fundort,

12.vi.1966. Fig. 61. N. thuringiaca Petry, & Kopulationsapparat, GU/Kli-1246, Austria inf., Dürnstein,

Ilias, ex I. 25.iii.1973, Styrax officinalis L.

266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

hellgrau, scharf vom Flugelgrund abgesetzt, im Tornus dunkelgrau. Hinterflügel

bei beiden Geschlechtern dunkelgrau; beim ¢ treten am Vorder- und Innenrand

bis ca. 1/2 der Flügellänge schwarze Duftschuppen auf. Fransen dunkelgrau,

ebenso die Unterseite aller Flügel (Fig. 56).

Kopulationsapparat. Männchen (Fig. 57). Uncus flach ausgeschnitten, Gnathos-

äste von einander entfernt inseriert. Valven mit vortretendem Ventralrand.

Vinculum schmal, flach ausgeschnitten. Aedoeagus mit einer dichten Reihe

kräftiger Cornuti und einer Cornutalplatte bewehrt.

Weibchen (Fig. 58). Die Gonapophysen von der üblichen Bauart. An der

Bursawand fällt eine körnige Struktur auf. Ein Signum fehlt.

Die systematische Stellung dieser isoliert stehenden Art ist derzeit noch unklar.

Futterpflanze: Prunus dulcis (Mill.) D. A. Webb (= Amygdalus communis L.).

Mine. Eiablage ober- oder unterseitig. Gangmine, vielfach unterseitig

beginnend, fast immer an eine Blattader angelehnt verlaufend, Kotlinie

unterbrochen, schwarz, den Gang fast vollständig ausfüllend. Die Mine ist daher

in diesem Stadium schwer sichtbar. Im weiteren Verlauf ist die Mine vielfach

gewunden mit aufgelockerter Kotlinie, die meist schmale Ränder freiläßt. Erst in

diesem Stadium fällt die relativ kurze Mine auf. Die Raupe verläßt das Blatt auf

der Blattoberseite (Fig. 59).

Raupe hellgelb mit deutlich grün durchscheinendem Rückengefäß; Kopfkapsel

hellbraun. Kokon dünnhäutig, gelblichweiß.

Es können zumindest zwei Generationen angenommen werden. Besetzte Minen

wurden im Juni und im September gefunden. Fundorte: Rhodos, Lindos, Treas,

Salakos, Filerimos. Auf trockenen Steilhängen und an Rändern von Kulturzonen,

meist an verwilderten Mandelbäumen.

20.5.1976, ex 1. 24.6.1976”. Aus dem Material von Lindos wurden der Holotypus

und Paratypen ausgewählt.

Nepticula muricatella spec. nov.

(Fig. 60, 62)

In die Verwandtschaft von N. thuringiaca Petry gehörend.

Fühler beim & etwas über 1/2 der Vorderflügellänge reichend, beim 9 bis 1/2,

grau, schwach heller geringt erscheinend. Kopfhaare gelblichbraun, beim g etwas

dunkler, grau getönt. Nackenschöpfe gelblich. Augendeckel weiß. Maxillarpalpen

Thorax ockerbraun, Hinterleib graubraun, unterseits aufgehellt. Beine hellgrau,

die Tarsenglieder des dritten Beinpaares leicht dunkler gefleckt.

Expansion: 4—4.75 mm. Glattschuppig. Vorderflügel bei beiden Geschlechtern

einfarbig ockerbraun, beim & manchmal etwas dunkler, grau getönt. Fransen

hellgrau, nicht vom Flügelgrund abgesetzt. Hinterflügel grau, Fransen lichter.

Männlicher Kopulationsapparat (Fig. 60). Uncus gekerbt mit lateralen

KLIMESCH: Nepticulidenfauna von Anatolien 267

Vorsprüngen. Gnathos mit zwei distanzierten Asten. Verlauf des Ventralrandes der

Valven gerade, apical mit einem einwarts gekrümmten Fortsatz. Transtillae kurz.

Vinculum schmal, basal bogig ausgeschnitten. Aedoeagus mit zahlreichen kleinen

Cornuti bewehrt.

Futterpflanze: Sanguisorba muricata (Spach) Focke.

Mine. Eiablage unterseitig. Gangmine, zuerst zart mit schwarzer, den Gang fast

ganz ausfüllender Kotlinie, später stark verbreitert, dem Blattrand folgend und

schlieRlich einen Teil der Blattspreite einnehmend mit kräftiger, etwas

aufgelockerter Kotspur. Kleine Blatter werden vollstandig ausgeweidet und lassen

dann charakteristische Merkmale kaum mehr erkennen (Fig. 62). Zur

Verwandlung verläßt die Raupe das Blatt auf der Oberseite. Raupe hellgelb,

Kopfkapsel ebenso. Kokon zarthäutig, oval, hellbraun.

Die Minen wurden am 12.vi.1966 auf einem felsigen Standort an sehr

geschützten Stellen zwischen vorspringenden Felsblöcken gefunden; die meisten

Minen waren bereits verlassen, ebenso am 9.vii.1968 bei einem weiteren Besuch

der Fundstelle: Meram bei Konya (Anatolien). Es wurden nach zweiwöchiger

Die nahestehende Nepticula thuringiaca Petry ist etwas größer, robuster,

dunkler, besonders im weiblichen Geschlecht, bei dem die Vorderflügel

dunkelgrau-braun getönt sind. Das männliche Genitale der N. thuringiaca weist im

apicalen Teil des flacher eingekerbten Uncus beiderseits je zwei zapfenartige

Vorsprünge auf. Der kräftigere Aedoeagus besitzt dicht gestellte größere,

dornenförmige Cornuti und eine Cornutalplatte (Fig. 61).

Das zur Beschreibung verwendete Material, aus dem der Holotypus und

Paratypen ausgewählt wurden, weist folgende Bezettelung auf: 2 g “ex |.

2.7.1966”, 5 Q “ex 1. 30.6., 1.7., 2.7.1966, Anatolia, Meram b. Konya, Mine

12.6.1966 an Sanguisorba muricata, Zucht Nr. 893”.

Nepticula styracicolella spec. nov.

(Fig. 63, 64)

Da nur zwei 9 vorliegen, kann über die systematische Stellung der neuen Art

derzeit noch nichts ausgesagt werden.

Fühler nicht 1/2 der Vorderflügellänge erreichend, schwarz mit schwach

hervortretenden Gliederenden. Kopfhaare hell rostfarbig, Nackenschöpfe

hellgelb. Augendeckel gelblichweiß. Maxillarpalpen und Labialpalpen hellgelb.

Thorax und Hinterleib schwarz, metallisch glänzend, unterseits heller grau.

Beine oberseits schwarz, unterseits heller, bräunlich.

Expansion: 4 mm. Grobschuppig. Vorderflügel schwarz, im basalen Teil mit

metallischem, violetten Glanz, bei 3/4 der Vorderflügellänge eine schwach

gebogene, stark metallisch glänzende, messingfarbene Querbinde. Fransen

hellgrau, scharf vom Flügelgrund abgesetzt, um den Apex heller, weißlich, im

Tornus dunkelgrau. Unterseite dunkelgrau. Hinterflügel grau mit ebensolchen

Fransen.

Futterpflanze: Styrax officinalis L.

268 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

10mm

Fig. 64 a-c. Nepticula styracicolella spec. nov. Minen an Styrax officinalis L., Rhodos, Mt. Profitis Ilias,

19.x.1976. Fig. 65—67. Trifurcula (Ectoedemia) aegilopidella spec. nov. 65: g Imago, Rhodos, Rodini,

Kopulationsapparat, GU/Kli-4107, dieselben Daten, ex |. 17.iv.1973.

Mine. Eiablage oberseitig, häufig an einem Blattnerv als schwarzer,

punktartiger Höcker leicht erkennbar. Mine sehr variabel. In eünnen

Schattenblättern ist sie sehr lang, stark gewunden (Fig. 64a, c), oft sich kreuzend,

vielfach am Blattrand verlaufend, mit zarter, breite Ränder freilassender, öfters

kurz unterbrochener schwarzer Kotlinie. In dickeren Blättern ist die Mine

KLIMESCH: Nepticulidenfauna von Anatolien 269

merklich kürzer, die Kotlinie kräftiger, haufig in Halbkreisbogen abgelagert und

nur einen schmalen Rand des Ganges freilassend (Fig. 64b). Manchmal konnen

beide Arten der Kotablage in ein und derselben Mine kombiniert auftreten,

ahnlich wie z.B. bei Nepticula tiliae Frey. Fast immer findet sich nur eine Mine in

einem Blatt, selten sind es zwei. Die Mine dieser Art ist mit ihrer Variabilitat ein

gutes Beispiel dafür, wie sehr Standort und Sonneneinstrahlung die Lange der

Mine und die Art der Kotablagerung beeinflussen. Zur Verwandlung verläßt die

Raupe die Mine durch einen halbkreisförmigen Schlitz auf der Blattoberseite. Die

Raupe ist hellgelb mit hellbrauner Kopfkapsel. Kokon oval, hellbraun, von

lockerem Gespinst umgeben.

Auf Rhodos wurde die Art an folgenden Orten festgestellt: im Petaloudes-Tal,

das durch das massenhafte Vorkommen von Callimorpha quadripunctaria Poda

Berühmtheit erlangt hat, Gräben bei Treas und Trianta, Funtukli und Mt. Profitis

Ilias bei ca. 800 m. Die Standorte der Art bezw. ihrer Futterpflanze sind Gebüsche

an Gräben und Bachläufen, wo sich Styrax in Gesellschaft von Platanus orientalis,

Myrthus italica und Laurus nobilis findet; im Unterholz von Waldungen aus Pinus

halepensis und Cupressus. Sie ist demnach in Höhenlagen von 20 bis ca 800 m

verbreitet. Über die Generationsfolge (es sind wohl mindestens zwei anzunehmen)

kann noch nichts ausgesagt werden. Besetzte Minen können von Ende Mai bis

Ende September — stets sehr einzeln — gefunden werden.

“Rhodos, Petaloudes, ex 1. 25.3.1973, aus Mine an Styrax officinalis, Zucht Nr.

1054 A, 25.9.1973”; gleicher Fundort, “11.6.1976”, auf einem Styrax-Blatt bei der

Eiablage gefangen. Die betr. Stücke wurden als Holo- und Paratypus gekenn-

zeichnet.

Trifurcula (Ectoedemia) aegilopidella spec. nov.

(Fig. 65—69)

In die weitere Verwandtschaft der Trifurcula (Ectoedemia) caradjai Hering

gehörige Art.

Fühler dunkelgrau, beim ¢ bis etwas über 1/2 der Vorderflügellänge reichend,

gelblich. Augendeckel gelblichweiß. Maxillarpalpen und Labialpalpen hellgelb.

Thorax und Abdomen schwarzbraun, ebenso die Beine. Unterseits sind alle

diese Teile heller, gelblich.

Expansion 4—4.25 mm. Grobschuppig. Vorderflügel gestreckt, Grundfarbe

Diese lassen die Grundfarbe — vor allem beim 4 — etwas hervortreten,

besonders im Bereich des Tornus. Bei 1/2 des Vorderflügels eine schwach schräg

gestellte, beim g häufig durch einzelne schwarze Schuppen getrübte, schmale,

gelblichweiße Querbinde, die am Vorder- und am Innenrand etwas verbreitert ist.

Beim Q tritt diese Binde, da dort die Deckschuppen dichter sind als beim &,

schärfer hervor. Die hellgrauen, gelblich glänzenden Fransen des Außenrandes

sind vom Flügelgrund scharf abgesetzt, sie sind im Tornus dunkler grau.

Hinterflügel grau mit helleren Fransen; beim g ist die Flügelfläche von der Basis

270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

01 mm 10mm 0.1mm

Rhodos, Rodini, Quercus macrolepis Kotschy, ex |. 17.iv.1973; 69: Minen an Q. macrolepis Kotschy,

derselben Ort, 22.ix.1972. Fig. 70—72. T. (Fedalmia) trilobella spec. nov. 70: 3 Kopulationsapparat,

4096, dieselben Daten, ex |. 25.ix.1975; 72: Minen an S. triloba L., ex 1. 22.v.1974.

bis etwa 1/2 von schmalen, dunkelgrauen Duftschuppen bedeckt; sie sind am

dichtesten am Vorderrand inseriert, fehlen jedoch am Innenrand. Unterseite der

Vorderflügel grau, beim 4 median schwärzlich verdunkelt, an der Basis ein

Buschel langer, gelblicher Dufthaare.

KLIMESCH: Nepticulidenfauna von Anatolien 271

Männlicher Kopulationsapparat (Fig. 67). Uncus zungenformig. Ventralrand

der Valven flach ausgeschnitten. Gnathosäste von einander entfernt inseriert.

Vinculum schmal, flach ausgeschnitten. Aedoeagus mit zahlreichen kleinen

Cornuti und einer Cornutalplatte bewehrt.

Weiblicher Kopulationsapparat (Fig. 68). Vordere Gonapophysen flachig

verbreitert, hintere stabformig. Die dorsal auf dem Vestibulum zum Ductus bursae

gelegene Platte ist schwach sklerotisiert, die beiden Fortsatze dagegen starker. An

der Wand der Bursa copulatrix befindet sich sowohl dorsal als auch ventral eine

ovale Platte mit relativ großer wabenformiger Struktur. Gegenüber den

(entfernter) verwandten, ebenfalls an weichblätterigen Eichen lebenden Arten der

T. caradjai Her. und subbimaculella Hw.-Gruppe erscheint aegilopidella viel

kleiner, zarter und heller, beim Z durch Duftschuppen und im Genitale durch die

weit voneinander entfernten Gnathosäste ausgezeichnet.

Futterpflanze: Quercus macrolepis Kotschy (= aegilops L.).

Mine. Eiablage oberseitig. Anfangsgang der Mine knäuelartig gewunden,

Kotspur schwarz, zuerst kompakt, später mehr oder minder aufgelockert. In

Schattenblättern ist die Mine gedehnt, im breiten, oft platzartig erscheinenden

Gang läßt die aufgelockerte Kotlinie breite Ränder frei (Fig. 69). In den der Sonne

stärker ausgesetzten, dickeren Blättern ist die Mine oft auffallend kurz, die

Windungen der Kotlinie können dann kaum wahrgenommen werden; sie füllen

die Mine völlig aus, ohne Ränder freizulassen. Raupe smaragdgrün mit brauner

Kopfkapsel. Kokon oval, flach, braun. Besetzte Minen wurden nur im September

(1972, 1977) gefunden. Einbrütigkeit ist daher anzunehmen.

Fundorte: Rhodos: Rodini, Kremasti, Treas.

Das durch Zucht erzielte Material (34 &, 71 9) trägt folgende Bezettelung:

“Mine 22.9.1974 an Quercus macrolepis, Zucht Nr. 1054, ex 1. 17.—30.4.1973,

Rhodos, Rodini; J. Klimesch.” Aus diesem Material wurden der Holotypus, der

Allotypus und weitere Paratypen ausgewählt.

Trifurcula (Fedalmia) trilobella spec. nov.

(Fig. 70— 72)

In die Verwandtschaft der 7. rosmarinella Chret. gehörig.

kürzer; dunkelbraun. Augendeckel gelblichweiß. Maxillarpalpen hellgelb.

Kopfhaare hell rostfarben, ebenso die Nackenschöpfe.

hervortretende Legeröhre zugespitzt erscheinend. Beine gelblichbraun, unterseits

lichter.

Expansion 5—5.5 mm. Grobschuppig. Vorderflügel gestreckt, Grundfarbe

hellgelb, dicht mit dunkelbraunen Schuppen bedeckt, die nur im Tornus etwas

spärlicher auftreten. Kein Sexualdimorphismus. Der Gesamteindruck der

Vorderflügel ist — mit unbewaffnetem Auge betrachtet — dunkel lederbraun. Die

hellgelben Fransen scharf abgesetzt, im Tornus etwas dunkler. Hinterflügel

‚hellgrau mit hellgraugelblichen Fransen, die im Apex etwas dunkler sind.

272 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

Unterseite der Vorderflugel hellgrau, gegen den Apex dunkler, am Innenrand

gelblich aufgehellt.

Männlicher Kopulationsapparat (Fig. 70). Pseuduncus schwach sklerotisiert,

den spatelformigen Uncus etwas uberragend. Ventralrand der Valven schrag

verlaufend mit einem nach innen gerichteten apicalen, fingerformigen Fortsatz.

Lateralarme der Transtillae kurz. Gnasthosaste apical verschmolzen. Aedoeagus

kraftig, mit einem langen, stabformigen, schwach gebogenen, zugespitzten

Cornutus und einer Cornutalplatte bewehrt.

Weiblicher Kopulationsapparat (Fig. 71). Ausgezeichnet durch die langen

Gonapophysen, die ein weites Herausstrecken des Ovipositors ermoglichen.

Dadurch kann die Eiablage durch den dichten Haarfilz hindurch auf die

Epidermis der Futterpflanze (Salvia triloba L.) erfolgen. Die Bursa copulatrix weist

zwei langgestreckte Platten mit wabenförmiger Struktur auf, wie sie für die

trifurculoiden Arten charakteristisch sind.

Die Bauart der Genitalien weist die neue Art in die Verwandtschaft der T. (F.)

rosmarinella Chret., hamirella Chret. und stoechadella Klim. Unterschiede

bestehen beim Z im Vinculum, im Verlauf des Innenrandes der Valven und in der

der vorderen Gonapophysen von denen der genannten Arten abweichen.

Äußerlich ist trilobella durch die dunklere, lederbraune Vorderflügel-Färbung

von den genannten, bedeutend heller gelbbraunen Arten zu unterscheiden.

Futterpflanze: Salvia triloba L.

Mine. Gangmine (Fig. 72), lang, stark gewunden, sowohl am Blattrand als auch

in der Blattspreite verlaufend. Die unregelmäßig kurz unterbrochene, schwarze

Kotlinie nimmt bei der erwachsenen Raupe etwa 1/3 der Minenbreite ein.

Eiablage meist oberseitig. Raupe gestreckt, hellgelb, mit brauner Kopfkapsel.

Kokon oval, von fester Konsistenz, marginal mit einzelnen abstehenden

Gespinstfäden.

Fundorte: Rhodos; Rodini, oberes Petaloudes-Tal bei der Kapelle Kalopetra,

Mt. Smith bei Rhodos-Stadt auf gebüschreichen, schattigen Standorten.

Besetzte Minen wurden am 5. und 6. Mai 1975 gefunden, die Imagines

schlüpften im Laufe des September 1975. Die Art ist sicher einbrütig. Die Minen

entwickeln sich zu Beginn des Frühjahres in den jungen Blättern der beginnenden

Vegetationsperiode, daher konnten anfangs Mai meist nur mehr verlassene Minen

gefunden werden.

Hering führt in seinen Bestimmungstabellen (1957 : 933, Nr. 4558) an Salvia

spec. eine Nepticuliden-Mine an; sie wird als Gangmine beschrieben mit einer

schwarzen Kotlinie, die mehr als die halbe Gangbreite einnimmt. Ob es sich hier

um die neue Art handelt, können nur Imagines aus Zuchten entscheiden.

folgende Bezettelung: “Mine 6.—8.5.1975 an Salvia triloba, ex |. 16.9.— 16.10.1975,

Zucht Nr. 1133, Rhodos, Rodini, J. Klimesch”. Aus diesem Material wurde der

Holotypus, ein Allotypus und weitere Paratypen ausgewählt.

KLIMESCH: Nepticulidenfauna von Anatolien 273

NIS

N N

ISIS

= a

Fig. 73, 75. Trifurcula (Fedalmia) albiflorella spec. nov. 73: & Kopulationsapparat, GU/Kli-1322,

Anatolia, Meram p. Konya, ex I. 10.vii.1966, Nepeta nuda albiflora Gams; 75: Mine an N. nuda albiflora

Gams, derselbe Ort, 12.vi.1966. Fig. 74. 7. (F.) saturejae (Parenti), g Kopulationsapparat, GU/Kli-1318,

Italia, Piemonte, Susa, ex I. 8.vi.1966, Calamintha nepeta agg. Fig. 76—78. T. (Fomoria) deschkai spec.

nov. 76: 2 Imago, Rhodos, Mt. Smith, ex I. 3.x.1973, Hypericum crispum L.; 77: & Kopulationsapparat,

GU/Kli-1306, Rhodos, Etonas, ex |. 15.ix.1973, Hypericum spec.: 78: 9 Kopulationsapparat, GU/KIi-

4106, Rhodos, Rodini, ex 1. 3.vi.1974, Hypericum spec.

274 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

Trifurcula (Fedalmia) albiflorella spec. nov.

(Fig. 73, 75)

In die Verwandtschaft der 7. (F.) saturejae (Parenti) gehörig. Es liegen 3 ¢ vor.

Fühler etwas über 1/2 der Vorderflügellänge reichend, gelblichgrau, schwach

geringt. Kopfhaare rostfarbig, Nackenschopfe gelb. Augendeckel groß, weiß.

Maxillarpalpen wie auch die Labialpalpen gelblich.

Thorax und Hinterleib graubraun, das Analende gelblich. Beine oberseitig

gelblichgrau, die Tarsen des 3. Beinpaares dunkler gefleckt, unterseits wie auch

das Abdomen gelblich.

Expansion: 4.25—5 mm. Grobschuppig. Vorderflügel-Grundfarbe hellgelb,

dicht mit graubraunen Schuppen bedeckt. Am Innenrand tritt aber die

Grundfarbe fleckartig hervor. Fransen weißlichgrau, vom Flügelgrund scharf

abgesetzt. Unterseite hellgrau, am Vorderrand dunkler. Hinterflügel hellgrau mit

lichteren Fransen.

Die nahe verwandte Trifurcula (Fedalmia) saturejae (Parenti) ist viel dunkler,

schwarzgrau. Anatomisch sind beide Arten leicht durch die Struktur des

männlichen Kopulationsapparates auseinanderzuhalten. Bei T. albiflorella (Fig.

73) verläuft der Ventralrand der Valven schräg, der apicale Fortsatz ist gerade,

fingerförmig. Bei T. (F.) saturejae (Fig. 74) springt der Ventralrand bauchig vor,

der apicale Fortsatz ist breiter und krallenartig gekrümmt.

Futterpflanze: Nepeta nuda albiflora Gams.

Mine. Die Stelle der Eiablage konnte nicht ermittelt werden. Die Mine (Fig. 75)

stellt einen breiten Gang dar mit einer zentralen, aufgelockerten Kotlinie, die

breite Ränder freiläßt; sie verläuft meist am Blattrand. Es wurde nicht untersucht,

ob ähnlich wie bei T. (F.) saturejae auch Stengelminen oder kombinierte Stengel-

Blattminen vorkommen.

Fundort. Felsiges Gelände bei Meram-Konya (Anatolien) auf einer durch

vorspringende Felsblocke geschützten Stelle. Der Beschreibung liegen 3 & mit

folgender Bezettelung zugrunde: “Mine 12.6.1966 an Nepeta nuda albiflora, Zucht

Nr. 892, ex 1. 10.—11.7.1966, Anatolia, Meram p. Konya, J. Klimesch”.

Trifurcula (Fomoria) deschkai spec. nov.

(Fig. 76—78)

In die Verwandtschaft der 7. (F.) septembrella Stt. gehörend.

Fühler dunkel graubraun, kaum merklich dunkler geringt. Bei beiden

Geschlechtern bis ca. 3/4 der Vorderflügellänge reichend. Kopfhaare bei beiden

Geschlechtern rostbräunlich, Nackenschöpfe gelblichweiß. Augendeckel gelb-

lichweiß, apical breit schwarzbraun gerandet. Maxillarpalpen und Labialpalpen

weißlichgrau. Abdomen schwarzlichgrau, unterseits etwas lichter. Analende beim

cd hellgrau. Beine schwarzgrau, unterseits lichter; die letzten zwei Tarsenglieder

des 3. Beinpaares auch oberseits hellgrau.

Expansion je nach Substrat der Raupen: 3.75—4.25 mm (aus Hypericum crispum

und Hypericum spec.), 4—4.75 mm (aus Hypericum hircinum) (Fig. 76). Kein

KLIMESCH: Nepticulidenfauna von Anatolien DIS

Fig. 79—80. Trifurcula (Fomoria) septembrella (Stt.). 79: & Kopulationsapparat, GU/Kli-1310, Austria

dieselben Daten. Fig. 81—82. T. (F.) deschkai spec. nov. 81: Minen an Hypericum crispum L., Rhodos,

Mt. Smith, 5.1x.1973; 82: Minen an Hypericum spec., Rhodos, Etonas, 31.viii.1973.

Sexualdimorphismus. Grobschuppig. Vorderflügel-Grundfarbe schiefergrau, dicht

mit schwarzen Schuppen überdeckt; die Grundfarbe tritt nur im Außenfeld etwas

zu Tage, besonders beim &. Im Tornus ein weißer Fleck am Innenrand. Fransen

weißlich, glänzend, scharf vom Flügelgrund abgesetzt, im Tornus und am

276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

Innenrand dunkelgrau. Hinterflügel dunkelgrau mit ebensolchen Fransen.

Unterseite aller Flügel einfarbig dunkelgrau.

Männlicher Kopulationsapparat (Fig. 77). Pseuduncus schwach sklerotisiert,

den Uncus überragend. Uncus gestutzt. Gnathosäste verschmolzen. Valven mit

schräg verlaufendem Ventralrand. Arme der Transtillae lang ausgezogen, den

Außenrand des Vinculums erreichend. Aedoeagus mit einer Gruppe von Cornuti

und einer Cornutalplatte bewehrt.

Weiblicher Kopulationsapparat (Fig. 78). Die Signa der Bursa copulatrix

bestehen aus zwei langgestreckten, an den Enden gerundeten Platten mit der fur

trifurculoide Arten charakteristischen Wabenstruktur.

Die nahestehende 7. (F.) septembrella (Stt.) weist im männlichen Genitale (fig.

79) nur geringe Unterschiede auf. Sie bestehen hauptsachlich in der Bewehrung

Wabenstruktur der Signa reticulata kleiner als bei deschkai.

Auch die Unterschiede der äußeren Merkmale sind bei beiden Arten gering. T.

(F.) septembrella (Stt.) ist größer (Expansion 5—6 mm), der weiße Fleck im Tornus

ist meist deutlicher ausgeprägt, die Fransen am Außenrand sind matter, hellgrau,

nicht glänzend weiß wie bei T. deschkai. Die Augendeckel sind meist nur wenig

dunkel gerandet.

Futterpflanzen: Hypericum hircinum L., Hypericum spec. und Hypericum crispum

Mine. Eiablage unterseitig. Die meist stark gewundene, mit Kot fast ganz

erfüllte Jugend-Gangmine geht häufig fast vollständig im späteren, pustelartigen

Platz auf, in dem die Kotablage zentral erfolgt. An Hypericum hircinum-Blättern

konnten nicht selten mehrere Minen in einem Blatt festgestellt werden. Bei dem

schmalblätterigen Hypericum crispum werden nur die grösseren, im Innern einer

Pflanze schattig gelegenen Hochblätter befallen (Fig. 81). Raupe hellgelb,

Kopfkapsel braun. Kokon gestreckt, in einem mit Gespinst ausgekleidetem Teil

der Mine, von der ein von der Raupe vorbereiteter, mit Gespinstfäden

ausgekleideter Gang zur Schlupfstelle auf der Blattunterseite führt. Mehrere

Generationen sind anzunehmen.

Fundorte: Rhodos: Ethonas b. Malona, Archangelos (an A. hircinum und spec.),

Rodini (A. spec.), Rhodos, Stadtgebiet (7. crispum und H. spec.), Mt. Smith (4.

crispum), Kremasti, Paradissi (H. crispum). Die Standorte der T. deschkai sind

Brachfelder (H. crispum), schattige Gebüschränder (H. hircinum) und

Straßenböschungen (H. spec.); sie besiedelt also ein durch extreme Verhältnisse

gekennzeichnetes Verbreitungsgebiet, während die europäische 7. septembrella

Waldgebiete bevorzugt.

Die neue Art wurde von dem erfolgreichen Entomologen Gerfried Deschka

durch den fast gleichzeitigen Fund von Minen an den drei Substraten entdeckt.

Ihm sei deshalb die Art zugeeignet.

Der Beschreibung dienten 47 Exemplare mit folgender Bezettelung: “Mine an

Rhodos, Ethonas-Malona”. Aus derselben Futterpflanze und vom gleichen

KLIMESCH: Nepticulidenfauna von Anatolien 277

gleichen Fundort: “Mine an Hypericum spec. 31.8.1973, Zucht 1081, ex 1.

crispum, Zucht 1081A, ex 1. 29.9.—8.10.1973”, 2 &, 2 9. Das Typenpaar (Holo-

und Allotypus) wurde aus den aus Hypericum hircinum erzielten Züchtlingen von

Ethonas 1973 ausgewählt; diese und alle übrigen Paratypen wurden entsprechend

gekennzeichnet.

LITERATUR

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der Familie Nepticulidae. — Polskie Pismo ent. 42: 689 — 709.

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Fologne, E., 1860. Nepticula ulmivora. — Ent. Wkly Intell. 8: 92.

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tien). — Mitt. munch. ent. Ges. 32: 395—397.

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——, 1948. Zur Frage der verwandtschaftlichen Beziehungen einiger Stigmella-Arten auf Grund des

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——, 1953. Die europäischen Trifurcula- und Ectoedemia-Arten. — Z. wien. ent. Ges. 38: 160—195.

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278 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978

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BEN-7 5682

È DEEL 121 AFLEVERING 6 MUS. COMP. AB.

LIBRARY

FEBS 1979

HARVARD

| TIJDSCHRIFT UNIVERSITY

| VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

WILLIAM RAMIREZ B. — Evolution of mechanisms to carry pollen in Agaonidea

(Hymenoptera Chalcidoidea), p. 279— 293, text-figs. |—21.

Tijdschrift voor Entomologie, deel 121, afl. 6 Gepubliceerd 31-XII-1978 .

EVOLUTION OF MECHANISMS TO CARRY POLLEN IN

AGAONIDAE (HYMENOPTERA CHALCIDOIDEA)

WILLIAM RAMIREZ B.

Facultad de Agronomia, Universidad de Costa Rica

With 21 text-figures

ABSTRACT

It is possible that the pre-agaonid wasps visited the pre-Ficus inflorescense in order to eat pollen.

Later in the evolution of the pre-fig plant, the pre-agaonid started using the ovules for oviposition.

Once the pre-fig evolved a closed inflorescense with very tight entrances, only the pre-agaonids which

were more successful in penetrating the pre-figs, and those which carried more pollen to the interior of

the pre-fig, were positively selected. Most of the pollen carried on the body and wings of the ancestral

agaonids was probably rubbed off from the insect body. Those areas of the insect body which entered

in least contact with the ostiolar scales must have carried some pollen to the interior of the receptacles.

The areas of the wasps which probably were least cleaned off by ostiolar scales were the inner or medial

sides of the coxae, the sternum, the neck, petiolar areas, and undoubtedly the buccal cavity and the

digestive tract. Selection would favor those agaonids with more bristles on the mesosternum and coxae

in addition to those which were efficient in carrying pollen in the buccal cavity or the digestive tract, if

they regurgitated the pollen on the stigmata of the flowers. Four main systems of pollen transport seem

to have evolved in the Agaonidae: 1) improved capacity to carry pollen in the buccal cavity and other

hidden areas of the body, 2) hairiness of the mesosternum as well as the coxae, 3) mesosternal

concavities (corbiculae or pollen pockets), and 4) corbiculae on the front coxae which are used to

facilitate the transfer of pollen from the anthers to the sternal corbiculae or pockets and to carry

pollen.

Concerning the trends in the evolution of the corbiculae, it is possible that the first structures, other

than hairs or bristles, that the agaonids evolved to carry pollen were the sternal corbiculae. This idea is

based on the fact that there are many agaonids without corbiculae, with only sternal corbiculae, and

with sternal and coxal corbiculae. There is only one report of two agaonids with only coxal corbiculae.

The external corbiculae have apparently evolved in three different lines of the agaonids, whereas the

coxal corbiculae seem to have evolved independently in four lines. It is not surprising to find such a

high degree of convergence in organisms, such as the agaonids, which spend much of their lives in very

similar environments (the interior of the syconium) and are apparently exposed to similar forces of

selection.

INTRODUCTION

Man has wondered for many centuries about the function of the small insects

(the agaonids) which penetrate and develop inside the fruits (syconia) of some

varieties of the edible fig (Ficus carica) and in other species. Aristotle, circa 340

B.C., described a small creature called ‘“psen’ which pierced unripe figs and thus

caused them to remain on the tree until mature. Theophrastus, a pupil of Aristotle,

gave a clear account of caprification (pollination of the figs by the agaonids), but

279

280 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 6, 1978

like his teacher he believed that the fig wasps nibbled at the cultivated figs and

caused them to develop (Condit, 1947). During the twenty centuries following the

time of Theophrastus, little was added to the store of information about fig

pollination and fig wasps.

The prevailing view until 1969, which was based primarily on observations of

Blastophaga psenes, the pollinator of the edible fig, was that pollination is effected

by pollen grains adhering to the wasp’s body surface and perhaps to its appendages

(Müller, 1883; Eisen, 1896; Knuth, 1906; Baker, 1913; Pemberton, 1921;

Buscalioni & Grandi, 1938). Other views generally supported the transport of

pollen on the body surface (Wiebes, 1963; Hill, 1967; Galil & Eisikowitch, 1968).

Recently it has been found that in many other fig species the situation is not as

simple as in the common fig. Independently and almost simultaneously, Ramirez

(1969), in Costa Rica, and Galil & Eisikowitch (1969), in East Africa, found

specific organs in some agaonids which serve for pollen transport from the mature

figs to the young ones of several species of Ficus.

The object of this work is to try to explain the evolution, as well as the sequences

and convergences, of the mechanisms to carry pollen in the Agaonidae (table 1).

MEANS OF POLLEN TRANSPORT

It is possible that most of the pollen carried on the body and wings of the

ancestral agaonids was rubbed off when the wasps squeezed between the ostiolar

scales. Those areas of the insect’s body, however, which entered in less contact:

with the ostiolar scales must have carried some pollen to the interior of the

receptacles. The areas of the wasps which probably were least cleaned off by

ostiolar scales were: the inner or medial sides of the coxae, the sternum, the neck

and petiolar areas, the wings, and undoubtedly the buccal cavity and digestive |

tract. On a population basis and presumably on an individual basis, selection

should have favored those wasps that carried more pollen on the areas mentioned;

as a consequence more efficient pollination was accomplished after the wasp

entered the young inflorescences. Selection would therefore favor wasps with

more bristles on the mesosternum and coxae in addition to those which were

efficient in carrying pollen in the buccal cavity, other hidden areas of the body and

digestive tract, if they regurgitated pollen on the stigmata of the flowers when they:

were ovipositing.

Four main systems of pollen transport apparently evolved in the Agaonidae: 1)

Improved capacity to carry pollen in the buccal cavity, as well as in the digestive

tract!). This system is probably derived from the pollen eating activity of the pre-

agaonid wasps (Ramirez, 1976). To this group of wasps belong Tetrapus and

') According to Wiebes (1977b) “in many Agaonid species, pollen can sometines be found in small

quantities on several places of the body, e.g., between the mouth parts or under the hypopygium

...” Galil & Neeman (1977) showed for Blastophaga psenes (the pollinator of the edible fig) that the

females carry pollen into intersegmental and pleural invaginations which form in the shrunken body

following water loss.

RAMIREZ B: Mechanisms to carry pollen in Agaonidae

Table 1. List of fig species groups with a list of the agaonid pollinators (modified from Hill, 1967) of

Subgenus Section Subsection Agaonidae Corbiculae

Absent Sternal Coxal

Urostigma Urostigma Blastophaga + +

(Group E)

(=Platyscapa)

Leucogyne Maniella + +

Conosycea Conosycea Blastophaga + +?

Deilagaon +

Eupristina + +

Waterstoniella +

Dictyoneuron Waterstoniella +

Eupristina + +

Benjamina Parapristina + +

Stilpnophylum Blastophaga + +?

clavigera

(=Blastophaga

Group G)

Malvanthera Pleistodontes +

Pleistodontes') + de 19

Galoglychia Agaon +

Alfonsiella + +

Allotriozoon +

Elisabethiella + +

Paragaon +

Americana Pegoscapus + +

Pharmacosycea Oreosycea Blastophaga + +

Group F

Dolichoris + +

Pharmacosycea Tetrapus +

Ficus Ficus Blastophaga +

Group A

Rhizocladus Blastophaga +

Group A

Kalosyce Blastophaga +

Group A

Sinosycidium?)

Eriosycea Blastophaga +

Group B

Sycidium Scabrae Blastophaga +

Group B

Varinga Blastophaga +

Group B

Phaeopilosae Blastophaga +

Group C

Paleomorphe Paleomorphe Liporrhopalum +

Copiosae Blastophaga +

Group D

Sycomorus Adenosperma Ceratosolen +

Neomorphe Ceratosolen +

Prostratae Ceratosolen +

Pungentes Ceratosolen +

Pseudopalmeae Ceratosolen +

Rivulares?) A

Sycocarpus Ceratosolen +

Sycomorus Ceratosolen +

each group, and the presence or absence of corbiculae

1) Wiebes (1977b) reports that two species of Pleistodontes have only coxal corbiculae

2) Probably pollinated by a wasp of Blastophaga Group A

3) Probably pollinated by a Ceratosolen wasp

281

282 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 6, 1978

Figs. 1—4. Mesosterna of: 1, Blastophaga pumilae, without corbiculae; 2, B. (Waterstoniella) sundaica,

without corbiculae but with abundant sternal bristles; 3, Deilagaon megarhopalus, with incipient

corbiculae (right corbiculae with some pollen); 4, B. (Waterstoniella) masii, with more developed

corbiculae and less bristles than Deilagaon megarhopalus (fig. 3).

RAMIREZ B: Mechanisms to carry pollen in Agaonidae 283

probably those Blastophaga that inhabit the figs of sections Ficus, Kalosyce and

Rhizocladus. Section Sinosycidum is probably also pollinated by this type of wasp.

‘ 2) Hairiness of the mesosternum as well as of the coxae. This system of

transportation of pollen is apparently found in Blastophaga (Waterstoniella)

sundaica (fig. 2) and Pleistodontes blandus which possess mesosterna with abundant

long bristles; however, much pollen was also found in their digestive tracts. 3)

Sternal corbiculae, i.e., pollen carrying concavities in the mesosternum (figs.

3—10). 4) Corbiculae on the front coxae (figs. 16—20) which are used to facilitate

the transfer of pollen from the anthers to the sternal corbiculae, as well as a means

to carry pollen (Galil et al., 1973). In the forms with well-developed corbiculae,

pollen is not found in the digestive tract.

Sternal corbiculae!)

It isprobable that in some of the agaonids with hairy mesosterna (as in fig. 2) a

small flange or tegulum evolved by mesal growth of the most lateral angle of the

“pleural suture” (as in fig. 3). This tegulum formed a small cavity in which some

pollen could be transported (as in fig. 3). This flange is hereafter referred to as the

corbicular tegulum. A small tegulum which forms a small corbicula is still found in

Deilagaon megarhopalus (fig. 3), Blastophaga errata and Blastophaga (Waterstoniella)

masii (fig. 4). However, parts of the mesosterna of the latter two are covered with

long bristles as in Blastophaga (Waterstoniella) sundaica (fig. 2) and Pleistodontes

blandus, two wasps without sternal corbiculae.

Once the mesosternal tegulum formed a functional cavity to carry pollen, its

enlargement was positively selected. Larger concavities were formed by

progressive mesal growth of the tegulum. A small sternal corbicula accompanied

by sparse sternal hairiness is found in Blastophaga (Waterstoniella) masii (fig. 4).

The mesal extension of the corbicular tegulum to form a larger cavity was

accompanied by the loss of more and more sternal bristles, as suggested by

Blastophaga (Pegoscapus) cumanensis (fig. 6), Blastophaga javana (fig. 5) and

Blastophaga (Eupristina) adempta. However, some of the sternal hairs were retained

close to the sternal corbiculae and became specialized to rub the pollen from the

arolia of the font legs and/or to hold the pollen in place in the corbiculae (fig. 6).

In other agaonids the corbicular tegulum continued to grow mesally until a

much bigger cavity was formed. A wide, mesal corbicular opening is found in the

corbiculae of wasps such as Blastophaga gestroi, B. javana (fig. 5) and B. sumatrana;

in the subgenera Pegoscapus (fig. 6), Eupristina and Parapristina; in the genera

Maniella, Alfonsiella, Elisabethiella, in the Blastophaga wasps (=Platyscapa of

Wiebes, 1977a) inhabiting Old World section Urostigma for example, in

Blastophaga constabularis (= B. coronata; Wiebes, 1963); in Dolichoris vasculosae;

and probably in all of the Blastophaga wasps which inhabit Old World figs of

section Oreosycea, subgenus Pharmacosycea.

In other wasps the corbicular tegulum grew mesally and ventrally until the

corbiculae became almost closed. This condition is found in wasps such as Agaon

hamiferum modestum (fig. 7) and Allotriozoon prodigiosum.

1) Mesothoracal pollen pockets of Galil & Eisikowitch (1969).

284 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 6, 1978

Figs. 5—9. Mesosterna of: 5, Blastophaga javana, with large, open sternal corbiculae and few

mesosternal bristles; 6, Blastophaga (Pegoscapus) cumanensis, right side, showing corbicula with

corbicular bristles and some pollen inside; 7, Agaon hamiferum modestum, with almost closed sternal

corbiculae; 8, Blastophaga jacobsi, with closed sternal corbiculae; 9, Ceratosolen pilipis, with closed

corbiculae (sternal pockets of Galil & Eisikowitch, 1969).

RAMIREZ B: Mechanisms to carry pollen in Agaonidae 285

The maximum mesal and ventral growth of the corbicular tegulum occurs in

Blastophaga jacobsi (fig. 8), in all wasps of the genus Ceratosolen (fig. 9), in

Liporrhopalum (fig. 10), Platyscapa quadraticeps and probably B. tentacularis.

Galil & Snitzer-Pasternak (1970) report that the pollen pockets of Platyscapa

quadraticeps are complex and highly specialized structures rather than mere

Fig. 10. Mesosternum of Liporrhopalum mindanaensis, with closed corbiculae. Figs. 11—13. Front legs

of: 11, Blastophaga psenes (the pollinator of the edible fig), without coxal combs; 12, Deilagaon

megarhopalus, with incipient sternal corbiculae (note the aggregation of coxal bristles); 13, Blastophaga

jacobsi, with large, closed, sternal corbiculae (note the presence of coxal combs of bristles, which are

probably used to move the pollen from the sternum to the corbiculae).

286 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 6, 1978

depressions covered by a fold of the body wall. Each pocket is a small saccular

organ with well-defined openings. The two marginal exits are very narrow. The

third opening is the largest and gaps widely when the adjacent sternum is touched

Figs. 14—15. Front legs of: 14, Ceratosolen pilipis, with closed, sternal corbiculae (note the coxal comb);

15, Liporrhopalum mindanaensis, with only sternal corbiculae. Figs. 16—17. Font coxae of: 16,

Blastophaga malabarensis, with coxal corbiculae surrounded by abundant bristles; 17, B. nervosae, with

coxal corbiculae and combs.

RAMIREZ B: Mechanisms to carry pollen in Agaonidae 287

parallel to the inner border of the pocket and closes shortly afterwards. These

observations have not been verified, but I have observed that in Ceratosolen

arabicus and other Ceratosolen wasps the corbicular tegulum extends ventrally and

touches the mesosternum. However, the corbicular tegulum is apparently not

fused to it; thus a narrow slit is found along the entire mesal side of the structure.

When the corbicula of a Ceratosolen arabicus was carefully pressed, pollen came

out along the entire mesal slit or entrance of the structure. According to Wiebes

(personal communication) there is some misunderstanding about the pocket of

Platyscapa quadraticeps, there being not more than one opening for each pocket.

Coxal corbiculae

The front coxae of Sycophaga sycomori (a wasp that penetrates the fig in order to

lay eggs, but which is not a pollinator) possess almost no hairs. The same condition

is also found in Blastophaga (Waterstoniella) sundaica, although some thick bristles

are found on the mesal side. The rest of the front legs of both species are also

sparsely hairy. All the species of Tetrapus, in addition to Pleistodontes plebejus, also

possess very smooth coxae.

From the supposedly condition of smooth legs, hairiness became more

consipicuous in some agaonids. In Blastophaga psenes (fig. 11) the entire leg is

hairier than those of the previously mentioned wasps; the hairs of the coxa of

Blastophaga psenes are apparently randomly distributed over its entire surface (fig.

11). It is probable that hairiness of the mesal side of the coxa of some agaonids was

positively selected because it favored pollen transport.

Although the fore coxal hairs of Deilagaon megarhopalus (fig. 12) are

concentrated toward the mesal side, they occur without any noticeable pattern.

However, in Blastophaga jacobsi (fig. 13), Ceratosolen pilipes (fig. 14) and most

other Ceratosolen wasps, the fore coxal hairs are denser on the mesal side of the

coxa, forming a well-defined mesal area of strong bristles which Galil &

Eisikowitch (1969) call coxal combs. In other agaonids the coxal hairs also became

concentrated, but instead of forming a random grouping they form well-defined

rows. A similar condition is found in Liporrhopalum mindanaensis (fig. 15) and

Allotriozoon prodigiosum. Grandi (1917: fig. IV, 3) also reports and illustrates the

front coxa of Blastophaga valentinae as possessing a row of hairs similar to the one

found on Liporrhopalum mindanaensis.

Another tendency in the evolution of the coxal structures to carry pollen was

the formation of a mesal cavity. A primitive coxal corbicula completely

surrounded and covered with hairs is found in Blastophaga malabarensis (fig. 16).

The corbiculae and surrounding hairs of this species were found to be completely

full of and covered with pollen. Blastophaga nervosae (fig. 17) has an incipient coxal

corbicula, but possesses far fewer hairs than Blastophaga malabarensis. The coxal

bristles of Blastophaga nervosae are organized in a more definite row (fig. 17); a

similar condition is also found in Platyscapa quadraticeps. Pleistodontes imperialis

(fig. 18) possesses a well-defined row of hairs located in a more dorsal position

than those of Blastophaga nervosae (fig. 17). It is not sure if Pleistodontes imperialis

has coxal corbiculae.

288 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 6, 1978

Figs. 18—19. Front legs of: 18, Pleistodontes imperialis, with open, sternal corbiculae and apparently

incipient coxal corbiculae; 19, Elisabethiella stueckenbergi, with well developed coxal corbiculae and an

outer coxal comb of bristles. Fig. 20. Front coxa of Blastophaga boschmai, with well developed coxal

corbiculae and an inner comb of bristles.

289

RAMIREZ B: Mechanisms to carry pollen in Agaonidae

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Once the coxal corbiculae evolved, there was a tendency to lose most of the

bristles located inside the cavities and to organize the peripheral bristles into more

definite rows or combs (figs. 17—20). Two tendencies in the formation of these

rows of bristles are found. In some wasps, the corbicular cavity is fenced by a

medial row of bristles, as in Blastophaga nervosae (fig. 17). In Maniella delhiensis the

coxal cavity has a well-defined row of bristles on its mesal side, but some hairs are

still present inside the corbicula. This condition attains its maximum expression in

Blastophaga boschmai (fig. 20), in which the coxal cavity is very deep and its long

bristles form a good barrier to hold the pollen in the corbicula. In other agaonids

the corbicular bristles form a more ventral row of bristles. This feature is found in

an incipient condition in Blastophaga constabularis (= Platyscapa coronata). A well-

defined mesal row of ventral bristles is found in most New World wasps of the

subgenus Pegoscapus and in the Old World Blastophaga (Eupristina) adempta,

Blastophaga (Parapristina) verticillata, Alfonsiella longiscapa and Elisabethiella

stueckenbergi (fig. 19).

TRENDS IN THE EVOLUTION OF THE CORBICULAE IN THE AGAONIDS

It is very well possible that the first structures, other than hairs or bristles, that

the agaonids evolved to carry pollen were the sternal corbiculae. This idea is based

on the fact that there are many agaonids without corbiculae, with only sternal

corbiculae, and with sternal and coxal corbiculae; although Wiebes (1977b)

reports that Pleistodontes blandus and mandibularis have only coxal corbiculae.

However, it is possible that the ancestors of some agaonids evolved both kinds of

corbiculae and that the coxal corbiculae were later lost in some; this is a very

unlikely possibility. The agaonids which could have lost the coxal corbiculae

would be the Blastophaga that inhabit the figs of sections Eriosycea, Scabrae,

Varinga, Phaepilosae and Copiosae (sensu Ramirez, 1974) and also the wasps of the

genera Ceratosolen and Liporrhopalum. Other agaonids which could have lost the

coxal corbiculae and inhabit the figs of subgenus Urostigma are: Agaon,

Allotriozoon, Paragaon, Deilagaon megarhopalus, the subgenus Waterstoniella, and

the two wasps (Blastophaga carlosi and B. mariae) that pollinate F. tuerckheimii in

Central America and Mexico. All of the wasps mentioned immediately above

possess only sternal corbiculae. The question then arises: Did each of these

structures evolve once or more than once in the history of the Agaonidae? This

question is difficult to answer. However, based upon the morphology of some of

the structures of each group of agaonids and the correlation of those structures

with the presence or absence of corbiculae, it is here concluded that the sternal

corbiculae, as well as the coxal ones, seem to have developed at least three or four

times during the evolution of the agaonids. It is not surprising to find such a high

degree of convergence in organisms, such as the agaonids, which spend much of

their lives in very similar environments (the interior of the syconium) and are

apparently exposed to similar forces of selection. Hill (1967: 53) notes that,

presumably because of the uniformity. of the micro-habitat, there is considerable

convergence in various characters in the family Agaonidae.

RAMIREZ B: Mechanisms to carry pollen in Agaonidae 291

Sternal corbiculae

These structures have apparently evolved in three different lines of the

agaonids: 1) In the ancestor (fig. 21: 18 — 19) of the Blastophaga that live in the

dioecious figs of sections Copiosae, Eriosycea, Phaeopilosae and Sycidium (all sensu

Ramirez, 1974). This type of corbicula was inherited by Ceratosolen (fig. 9) and

Liporrhopalum (fig. 10). In these two groups of wasps, as well as in Blastophaga

group C (i.e., Blastophaga jacobsi) (fig. 21, S), the corbiculae are closed rather than

open (fig. 5). 2) In the ancestor (fig. 20: 3 — 4) of Dolochoris vasculosae and

Blastophaga group F (i.e. Blastophaga boschmai) which inhabit the figs of Old

World section Oreosycea (subgenus Pharmacosycea). 3) In the ancestors of several

of the agaonid groups which inhabit the figs of subgenus Urostigma.

First evolutionary line of sternal corbicüae

The evolution from Blastophaga without corbiculae to Blastophaga with open

corbiculae and to Blastophaga, Ceratosolen and Liporrhopalum with closed sternal

corbiculae agrees with the line of evolution postulated by Wiebes (1963: 105) for

Ceratosolen. Wiebes states that the genus Ceratosolen appears to be a derivative of

Blastophaga and seems to be still connected with it through the pollinators of

subsection Sycidum (sensu Corner, 1965).

Second evolutionary line of sternal corbiculae

The second evolutionary line occurred in the ancestor (fig. 21: 3 — 4) of the

agaonids which inhabit the Old World section Oreosycea of subgenus

Pharmacosycea. This section of figs is inhabited by Blastophaga group F and one

species of Dolichoris (D. vasculosae). These wasps are characterized by their open

sternal corbiculae and the presence of coxal corbiculae.

Third evolutionary line of sternal corbiculae

The ancestors of some of the agaonid groups (fig. 21: 5 — 6) which inhabit the

figs of subgenus Urostigma reflect a third evolutionary line. Considering the

presence or absence of sternal corbiculae in the pollinators of subgenus Urostigma,

we find several main groups of wasps. These are: 1) In Pleistodontes, Pleistodontes

imperialis has sternal corbiculae and apparently coxal corbiculae (Ramirez, 1974).

Wiebes (1977b) reports that P. froggatti and regalis have sternal corbiculae, while

P. imperialis has sternal and coxal corbiculae. P. blandus and mandibularis have

only sternal corbiculae, in P. rennellensis he also finds a very vague trace of sternal

corbiculae. P. longicaudus, plebejus, immaturus and rieki have no coxal and sternal

corbiculae (Wiebes, 1977b); 2) Two Waterstoniella species (W. jacobsoni and W.

sundaica) (fig. 2), which do not have sternal corbiculae!); 3) Another Waterstoniella

(i.e., Blastophaga (W.) masii) (fig. 4), as well as Deilagaon megarhopalus (fig. 3),

which possess very small sternal corbiculae; 4) Agaon (fig. 7), Allotriozoon and

Paragaon, which have only large sternal corbiculae; 5) Alfonsiella and

Elizabethiella, which have large sternal as well as coxal corbiculae (fig. 19); and 6)

1) According to Wiebes (1976), ‘In some of the Blastophaginae, however, the sternal pollen-carying

structures seem to be secondarily lost (e.g., in some Waterstoniella) or are in the process of becoming

lost (as in Waterstoniella masii).”

292 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 6, 1978

The subgenus Pegoscapus (except Blastophaga (P.) carlosi and B. (P.) mariae),

Blastophaga clavigera (Blastophaga group G), Maniella delhiensis, Blastophaga group

E (= Platyscapa of Wiebes, 1977a) and Eupristina and Parapristina, which have

coxal and sternal corbiculae.

Coxal corbiculae

These structures are found in some of the agaonid groups which inhabit the

subgenus Urostigma and in those which inhabit the monoecious figs of section

Oreosycea (fig. 21: P and Q). The groups of wasps which have coxal corbiculae and

develop in the receptacles of subgenus Urostigma are: 1) Alfonsiella and

Elisabethiella (fig. 19) found in section Galoglychia; 2) Pegoscapus (as in fig. 19)

except Blastophaga (P.) carlosi and B. (P.) mariae, found in section Americana; 3)

the Blastophaga found in sections Stilpnophyllum and Urostigma; 4) Maniella found

in section Leucogyne; 5) Eupristina and Parapristina found in section Conosycea;

and 6) Blastophaga and Dolichoris vasculosae which inhabit the figs of sections

Oreosycea of subgenus Pharmacosycea (fig. 20). Pleistodontes imperialis also seems

to have coxal corbiculae (fig. 18).

It seems that the coxal corbiculae have evolved four times in the symbionts of

Urostigma as follows: 1) in the ancestor of Alfonsiella and Elisabethiella (fig. 21: 8 —

11); 2) in the ancestor (fig. 21: 7 — 12) of the agaonids which pollinate sections

Americana, Leucogyne, Stilpnophyllum and Urostigma, i.e., wasps of the subgenus

Blastophaga, the genus Maniella and the subgenus Pegoscapus; 3) in the

ancestor (fig. 21: 16 — 17) of Eupristina and Parapristina; 4) in the ancestor

(fig. 20: 3 — 4) of the wasps that pollinate the section Oreosycea of subgenus

Pharmacosycea and apparently in Pleistodontes imperialis (fig. 18). Wiebes (1977b)

reports that P. blandus and mandibularis have only coxal corbiculae.

Interrelation between sternal and coxal corbiculae

It seems that in most Blastophaga, as well as Ceratosolen and Liporrhopalum, that

possess only sternal corbiculae, groups of ventral bristles are present on the coxae

(figs. 13—15). Galil & Eisikowitch (1969) call these bristles ‘* coxal combs” and

state that during pollen loading the pollen is first deposited on the underside of the

thorax and afterwards is pushed by the coxal combs into the sternal pockets. The

coxal combs of the wasps mentioned could have evolved at the same time as the

sternal corbiculae as a means to move the pollen from the venter of the thorax to

the corbiculae. In 1971, Dr. Galil and the author further observed that several

species of Pegoscapus used the coxal corbiculae bristles to move the pollen from

the thorax to the sternal corbiculae although the coxal corbiculae are also used to

carry pollen (Galil et al., 1973).

From these observations it seems likely that the sternal corbiculae and coxal

bristles arose simultaneously, the latter to fill the former, and that coxal corbiculae

arose later to increase the amount of pollen carried; although Wiebes (1977b)

reports that “Pleistodontes blandus and P. mandibularis have only coxal

corbiculae”.

RAMIREZ B: Mechanisms to carry pollen in Agaonidae 293

ACKNOWLEDGMENTS

I wish to express my appreciation to those who have helped me in this study;

especially to Dr. Rafael L. Rodriguez, Universidad de Costa Rica, for reading the

manuscript and suggesting various changes; the University of Kansas; Universidad

de Oriente, Venezuela, for providing a predoctoral scholarship; the Smithsonian

Institution for providing a predoctoral research appointment and facilities at

Balboa, Canal Zone.

BIBLIOGRAPHY

Baker, C. F., 1913. A study of caprification of Ficus nota. — Philip. Sci. 8: 63—84.

Buscalioni, L., & G. Grandi, 1938. Ficus carica L., la sua biologia, la sua coltivazione e i suoi rapporti

con l’insetto pronubo. — Boll. Ist. Ent. Univ. Bologna 10: 223—280.

Condit, I., 1947. The Fig. Chron. Bot., 222 p. — Waltham, Mass.

Corner, E. J. H., 1965. Check-list of Ficus in Asia and Australasia with keys to identification. — Gdns.

Bull. Singapore 21: 1—186.

Eisen, 1896. Biological studies on figs, caprifigs and caprification. — Proc. Calif. Acad. Sci. 1: 11—51.

Galil, J., & D. Eisikowitch, 1968. On the pollination ecology of Ficus sycomorus L. in East Africa. —

Ecology 49: 259—269.

—., 1969. Further studies on the pollination ecology of Ficus sycomorus L. (Hymenoptera Chalci-

doidea, Agaonidae). — Tijdschr. Ent. 112: 1—13.

Galil, J., & Y. Snitzer-Pasternak, 1970. Pollination in Ficus religiosa L. as connected with the structure

and mode of action of pollen pockets of Blastophaga quadraticeps Mayr. — New Phytol. 69:

775—784.

Galil, J., W. Ramirez B. & D. Eisikowitch, 1973. Pollination of Ficus costaricana and F. hemsleyana by

Blastophaga estherae and B. tonduzi in Costa Rica (Hymenoptera: Chalcidoidea, Agaonidae).

— Tijdschr. Ent. 116: 175—183.

Galil, J., & G. Neeman, 1977. Pollen transfer and pollination in the common fig (Ficus carica L.). —

New Phytol. 79: 163—171.

Grandi, G., 1917. Contributo alla conoscenza degli Agaonini (Hymenoptera, Chalcididae) di Giava. —

Boll. Lab. Zool. Gen. Agr. Portici 12: 3—60.

Hill, D. S., 1967. Figs (Ficus spp.) and fig-wasps (Chalcidoidea). — J. Nat. Hist. 1: 413—434.

Knuth, P., 1906. Hand-book of Flower Pollination 1: 380 p. — Clarendon Press, Oxford.

Muller, H., 1883. The Fertilization of Flowers: 669 p. — Macmillan, London.

Pemberton, C. E., 1921. The fig wasp in its relation to the development of fertile seed in the Moreton

Bay Fig. — Hawaiian Plant. Rec. 24: 297— 319.

Ramirez B. W., 1969. Fig wasps: mechanisms of pollen transport. — Science 163 (3867): 580—581.

, 1974. Coevolution of Ficus and Agaonidae. — Ann. Missouri Bot. Gard. 61: 770—780.

, 1976. Evolution of Blastophagy. — Brenesia 9: 1—13.

Wiebes, J. T., 1963. Taxonomy and host preferences of Indo-Australian fig wasps of the genus Ceratoso-

len (Agaonidae).—Tijdschr. Ent. 106: 1—112.

——., 1976. A short history of fig wasp research.— Gdns. Bull. Singapore 29: 207—232.

—, 1977a. Agaonid fig wasps from Ficus salicifolia Vahl and some related species of the genus Pla-

tyscapa Motschoulsky (Hym. Chalc.). — Neth. J. Zool. 27(2): 209—223. |

———, 1977b. Indo-Malayan and papuan fig wasps (Hymenoptera, Chalcidoidea) 7. Agaonidae, main-

ly caught at light. — Zool. Med. 52(11): 137—159.

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