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Library of the Museum of Comparative Zoology 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 REGISTER VAN DEEL 121 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) by 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. Ld (9°) u OQ; Oy BB — SAN — BH UN — w +O Oy +O 40 +0 40 40 40 40 40 40 wo Os 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 9 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. 12 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 Or 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 nr ON THE STIGMELLA-NEPTICULA CONTROVERSY (LEPIDOPTERA) by 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. 13 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- stems: xx + 178 pp. — Leipzig. 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. INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE, 1964. Adopted by the XVth International Con- ress of Zoology, 2nd ed.: XX + 176 pp. — London. Also amendments of 1974 in Bull. zool. 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 group. — Ent. scand. 2: 241—262, 52 figs. KARSHOLT, O. & E. SCHMIDT NIELSEN, 1976. Systematisk fortegnelse over Danmarks sommerfugle. Ca- talogue ofthe Lepidoptera of Denmark: 128 pp. KirBy, W. F., 1897. Lloyds Natural History. A Handbook to the Order Lepidoptera 5 (3): 332 pp. , 1923. Superfamily Nepticuloidea Family 5 Nepticulidae. In: W. T. M. Forbes (ed.), Lepidopte- | WILKINSON: Stigmella-Nepticula 21 KLoET, G. S. & W. D. Hincks, 1972. A check list of British Insects 2nd ed. (revised) 2, Lepidoptera. — 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. Ges. 36: 4—9. LEMPKE, B. J., 1976. Naamlijst van de Nederlandse Macrolepidoptera. Uitgave Bibliotheek Koninklijke Nederlandse Natuurhistorische Vereniging, 21: 97 pp. McDuNNOUGH, J., 1939. Check List of the Lepidoptera of Canada and the United States of America. II. Microlepidoptera. Mem. sth. Calif. Acad. Sci. 2(1). MEYRICK, E., 1895. Handbook of British Lepidoptera. VI. + 843 pp. — London. OKEN, L., 1815. Lehrbuch Naturg. 3. PETERSEN, W., 1930. Die Blattminierer-Gattungen Lithocolletis und Nepticula (Lep). Teil Il: Nepticula Z. — Stettin. ent. Ztg. 91: 1—82, 3 pls., 16 figs. SATTLER, K., 1973. A catalogue of the family-group and genus-group names of the Gelechiidae, Holco- pogonidae, Lecithoceridae and Symmocidae (Lepidoptera). — Bull. Br. Mus. nat. Hist. (Ent.) B Lond.: 153— 282. 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. , in: GRAY, J. E., 1854b. List of British Animals in the collection of the British Museum 16: Lepi- doptera: 199 pp. — London. 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”. mee ae VINTI Pik (nec CUS FR Tat Ak \ | mec Por n eh REP vf u R 3 pean ; Pa ER M 7 ME 14 n da PAIA, ti È Fa bf È ; A ES hi it > A d À x: 1 x 1 Wy È agito} | i : N 29 ate n Me a Lie sr ess mae Re N i ; IATA held FAT Be Ae Ù A A OD a DES, FN \ si i fl | | N R È È ne i # ui i i mi lie 4 AR eal MUS, CGMP. ZOOL 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) by 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 [Introduction vi... ee RP RITI I Co ete ete eee eee 23 DE Kevitoltheispecies and checklist 2 2... 2.2200 ee Ceci lone 28 de her Saale Goss TOO OOO III: 32 bl, JA — o osints.5 NOI DDD 0 0.0.6.0 a 0.0'0:0:0/8.0)0.0.0.00 0.0.0.0:0,9 000.010 105 Se CEYLON KOR 2000680224 00 00 08 00/0000 600 op o0bIb010 00.10 de db 0.0.0 © OOG000G050 125 6. General outline of the evolution of Spialia .................................... 133 UBRIACO NEN = rene la à pen le 135 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 23 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; Je 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. Dr 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 ue 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 SE 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 7 | | | 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). se ) i 2 H ia ! i Ri e / ( sl N À j i Eee PEN À i i ¢ EEn \ ER jer I } GES N £ x FRS nl 4 & if ; i fi PA) È Alf we PS À n 5 BAE EN > ¢ à a IE Al à e C- i- ? SU { Di = nn pee f i 1 EE 4 5 ! ZA = > fu di < i } ; era) \ a | : ; I N I Lo) VV Ma + = par 3 i à / Neu ee Map 7 er O Us: 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 07 OO ij 7 ge} LD GR LE 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 do 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 ou 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 bh 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 er 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 Le 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 94 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 a 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 = ? s SS Sn Map 21 Spialia depauperata ES 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 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 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 x x ex x 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 © 2 3 fe) 5 © ro) O = O 3 u ep) ep) © O has u © © o © © x lo) © I) a ic © © = © © u en (8) Vv) ke) T 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 5 kn © Cle ® Le 3 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 © N e) Q O 2 O) © m 3 | es © = © © © © © 5 © = = 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. 7. LITERATURE Aurivillius, C., 1882. Recensio critica lepidopterorum Musei Ludoviciae Ulricae quae descripsit Carolus a Linne. — Kong]. Svenska Vet.-Akad. Handl. 19 (5): 1—188, pl. 1. ——, 1891, Verzeichniss einer vom Herrn Fritz Theorin aus Gabun und dem Gebiete des Camerun- flusses heimgebrachten Schmetterlingssammlung. — Ent. Tidskr. 12: 193—228. ——, 1910. Lepidoptera. In: Y. Sjöstedt, Wissenschaftliche Ergebnisse der schwedischen zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch- Ostafrikas, 1905—1906. — P. Palmquists Aktiebolag, Stockholm. ——, 1925. Hesperiidae. In: A. Seitz, Die Grossschmetterlinge der Erde 13: 505—588. — Lehmann Verlag, Stuttgart. Beadle, L. C., 1974. The inland waters of tropical Africa. — Longman, London and New York. Berger, L.-A., 1962. Lepidopteres Hesperiidae de Guinée et Côte d’Ivoire. — Bull. PIFAN 24 (A): 447—463. ——. 1967. Contribution a la faune du Congo (Brazzaville). Mission A. Villiers et A. Descarpentries. 65. Lepidopteres Hesperiidae. — Bull ’IFAN 29 (A); 1763—1781. ——, 1968. Contribution à l’étude de la faune de la basse Casamance (Sénégal). 20. Lepidoptera Hes- periidae. — Bull. ’IFAN 30 (A): 205—220. Booth, A. H., 1958. The zoogeography of West African Primates: a review. — Bull. ’IFAN 20 (A): 587—622. Butler, A. G., 1888. An account of three series of Lepidoptera collected in North-West India by Major Yerbury. — Ann. Mag. Nat. Hist. (6)1: 196—209. ——, 1898. On a collection of Lepidoptera made in British East Africa by Mr. C. S. Betton. — Proc. zool. Soc. London 1898: 395—444. ——, 1899. On two small collections of butterflies made by Mr. Richard Crawshay during 1898 in British East Africa. — Proc. zool. Soc. London 1899: 417—427. ——, 1901. On a small collection of butterflies from the Kikuyu Country and Mombasa made by the Rev. K. St. Aubyn Rogers. — Ann. Mag. Nat. Hist. (7)8; 111—113. 136 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978 Carcasson, R. H., 1964. A preliminary survey of the zoogeography of African butterflies. — E. Afr. Wildl. Journ. 2: 122—157. Carpenter, G. D. H., 1935. The Rhopalocera of Abyssinia: a faunistic study. — Trans. R. ent. Soc. London 83: 313—447. Condamin, M., 1960. Lépidoptères Hesperiidae récoltés au Cameroun par C. de Raemy. — Bull. PIFAN 22 (A): 919—923. ———, 1969. Lepidoptera Rhopalocera. In: Le Parc National du Niokolo-Koba 3. — Mem. l’IFAN 84: 373—392. ——, 1973. Monographie du genre Bicyclus. — Mem. l’IFAN no. 88: 1—324. Cramer, P., 1782. Uitlandsche Kapellen, 4. — S. J. Baalde, Amsterdam; B. Wild, Utrecht. Druce, H., 1875. A list of the collection of diurnal Lepidoptera made by J. J. Monteiro, Esq., in Angola, with descriptions of some new species. — Proc. zool. Soc. London 1875: 406—417. Evans, W. H., 1937. A catalogue of the African Hesperiidae in the British Museum. — British Museum (Nat. Hist.), London. ——, 1951. Revisional notes on African Hesperiidae. — Ann. Mag. Nat. Hist. (12) 4: 1268—1272. Fabricius, J. C., 1793. Entomologia Systematica, 3. — C. G. Proft, Copenhagen. Gabriel, A. G., 1954. Expedition to South-west Arabia 1937-8. Vol. 1: 351—391. — British Museum (Nat. Hist.), London. Gifford, D., 1965. Butterflies of Malawi. — Society of Malawi, Blantyre. Hamilton, A. C., 1974. The history of the vegetation. In: Lind & Morrison, East African Vegetation. — Dengan London. Hennig, W., 1960. Die Dipteren-Fauna von Neuseeland als systematisches und tiergeographisches Problem. — Beitr. Entom. 10: 221—329. ——, 1966. Phylogenetic systematics. — Univ. of Illinois Press, Chicago. Heron, F. A., 1909. Ruwenzori Expedition Reports. 12. Lepidoptera Rhopalocera. — Trans. zool. Soc. London 19: 141—178. Higgins, L. G., 1924. On the Ethiopian species of the genus Hesperia (Lep. Rhop.), with descriptions of two new species. — Trans. ent. Soc. London 1924: 64—120. Holland, W. J., 1896. A preliminary revision and synonymic catalogue of the Hesperiidae of Africa and the adjacent islands, with descriptions of some apparently new species. — Proc. zool. Soc. London 1896: 2—107. ——., 1920. Lepidoptera of the Congo. — Bull. Amer. Mus. Nat. Hist. 43: 109—369. Hopffer, C., 1855. Monatsber. k. Akad. Wiss. Berlin 1855: 643. , 1862. In: W. C. H. Peters, Naturwissenschaftliche Reise nach Mossambique. Zoologie V, pl. 27 figs. 9, 10. Jong, R. de, 1972. Systematics and geographic history of the genus Pyrgus in the Palaearctic region (Lepidoptera Hesperiidae). — Tijdschr. Ent. 115: 1—120. ——., 1974. Systematics and evolution of the Palaearctic Spialia species (Lepidoptera, Hesperiidae). — Tijdschr. Ent. 117: 225—271. ——, 1976. Affinities between the West Palaearctic and Ethiopian butterfly faunas. — Tijdschr. Ent. 119: 165—215. ——, 1977. Problems about Spialia ploetzi (Aurivillius) (Lepidoptera, Hesperiidae). — Ent. Ber., Amst., 37: 41—44,. Karsch, F., 1896. Aethiopische Hesperiiden. — Ent. Nachr. 22: 372—378. Kauffmann, G., 1955. Spialia sertoria Hoffmansegg en Corse. — Rev. fr. Lep. 15: 38—40. Larsen, T. B., 1977. The butterflies of Eastern Oman and their zoogeographic composition. In: The Scientific results of the Oman Flora and Fauna Survey 1975: 179—208. — Journal of Oman Studies, Special Report. Lattin, G. de, 1967. Grundriss der Zoogeographie. — Gustav Fischer Verlag, Stuttgart. Lindsey, A. W., & L. D. Miller, 1965. Superfamily Hesperioidea. In: R. M. Fox e.a., The Butterflies of Liberia. — Mem. Amer. ent. Soc. 19: 1—438. Linnaeus, 1767. Systema Naturae, Ed. XII. Stockholm. Livingstone, D. A., 1976. The Nile — Palaeolimnology of headwaters. In: J. Race (ed.), The Nile, biology of an ancient river. — W. Junk, The Hague. Moore, F., 1865. On the lepidopterous insects of Bengal. — Proc. zool. Soc. London 1965: 755—823. Moreau, R. E., 1952. Africa since the Mesozoic. — Proc. zool. Soc. London 121: 869—913. , 1955. Ecological changes in the Palaearctic Region since the Pliocene. — Proc. zool. Soc. London 125: 253—295. —, 1966. The bird faunas of Africa and its islands. — Academic Press, London and New York. DE JONG: The genus Spialia 137 Murray, D., 1959. The genitalia and food-plants of the South African Hesperiidae (Lepidoptera: Rho- palocera). — Durban Mus. Novit. 5: 219—230. Neave, S. A., 1910. Zoological collections from Northern Rhodesia and adjacent territories: Lepidoptera Rhopalocera. — Proc. zool. Soc. London 1910: 2—86. Niceville, L. de, 1889. On new and little-known butterflies from the Indian region. — Journ. Bombay Nat. Hist. Soc. 4: 177. Oberthiir, Ch., 1912. Observations sur les Hesperidae du genre Syrichthus. — Et. Lep. Comp. 6: 47—120. Ogata, M., Y. Okada, H. Okagaki, & A. Sibatani, 1957. Male genitalia of Lepidoptera: morphology and nomenclature. III. Appendages pertaining to the tenth somite. — Ann. ent. Soc. Am. 50: 237—244. Picard, J., 1947. Notes sur les Hesperiidae des regions palearctiques. Tribus des Erynnidi, Carcharodidi et Pyrgidi. — Bull. Soc. ent. Fr. 52: 129—134. —— 1950. Hesperiidae du Liberia collectés par P. L. Dekeyser et B. Holas. — Bull. PIFAN 12 (A): 625—628. Plôtz, C., 1884. Analytische Tabellen der Hesperiinen-Gattungen Pyrgus und Carcharodus. — Mitt. naturw. Ver. Neu-Vorpomm. 15: 1—24. Rebel, H., 1899. — Sber. Akad. Wiss. Wien (math.-naturw. Kl.) 1899: 360. , 1907. Lepidopteren aus Südarabien und von der Insel Sokotra. —Denkschr. math.-naturw. Kl. kais. Akad. Wiss. Wien 71: 1—100. ——, 1914. Wissenschaftliche Ergebnisse der Expedition R. Graver nach Zentralafrika. Lepidoptera. — Ann. k.k. naturhist. Hofmus. Wien 28: 219—294. Reverdin, J. L., 1933. Hesperia galba Fab. ab. minuscula nova. — Bull. Soc. lép. Genève 7: 67. Rzoska, J., e.a., 1976. Palaeo-ecology. In: J. Rzoska (ed.), The Nile, biology of an ancient river. — W. Junk, The Hague. Rzoska, J., & G. E. Wickens, 1976. Vegetation of the Nile Basin. In: J. Rzoska (ed.), The Nile, biology of an ancient river. — W. Junk, The Hague. Sevastopulo, D. G., 1974. The butterflies of the Shimba Hills. — Entomologist’s Rec. J. Var. 86: 131—136. Shephard, H. H., 1935. Lepidopterorum Catalogus, Pars 69. Hesperidae, Subfamilia Pyrginae III. — W. Junk, The Hague. Someren, V. G. L. van, 1939. Butterflies of the Chyulu Range. — J. E. Afr. Ug. nat. Hist. Soc. 14: 130—151. Strand, E., 1911. Zoologische Ergebnisse der Expedition des Herrn Hauptmann a. D. Fromm 1908/09 nach Deutsch-Ostafrika. 1. Lepidoptera. — Mitt. zool. Mus. Berlin 5: 277— 304. Swanepoel, D. A., 1953. Butterflies of South Africa. — Maskew Miller Ltd, Cape Town. Swinhoe, C., 1908. On the species of Hesperidae from the Indo-Malayan and African Regions, des- cribed by Herr Plôtz, with descriptions of some new species. — Trans. ent. Soc. London 1908: 1—36. , 1913. Lepidoptera Indica, X. — L. Reeve & Co., London. Talbot, G., 1932. Lepidoptera from British Somaliland. — Bull. Hill Mus. Witley 4: 182—188. Trimen, R., 1864. Descriptions of some new species of butterflies found in Southern Africa. — Trans. ent. Soc. London 1864: 175—180. —, 1866. Rhopalocera Africae Australis, 2. — W. F. Mathew, Cape Town. —, 1870. Notes on butterflies collected by J. H. Bowker, Esq., in Basuto-land, South Africa; with descriptions of some new species. — Trans. ent. Soc. London 1870: 341—390. ——, 1889. South-African butterflies, 3. — Trübner & Co., London. ——, 1891. On butterflies collected in tropical South-Western Africa by Mr. A. W. Eriksson. — Proc. zool. Soc. London 1891: 59—107. —, 1898. On some new or little known species of African butterflies. — Trans. ent. Soc. London 1898: 1—15. Vari, L., 1976. South African Lepidoptera, 7. Descriptions and notes on new taxa of Rhopalocera, Ann. Transvaal Mus. 30: 121—141. Vavilov, N. I., 1922. The law of homologous series in variation. — J. Genet. 12:47— 89. Wallengren, H. D. J., 1863. Lepidopterologische Mittheilungen. — Wien. ent. Monatschr. 7: 137—151. Warren, B. C. S., 1926. Monograph of the tribe Hesperiidi (European species) with revised classification of the subfamily Hesperiinae (Palaearctic species) based on the genital armature of the males. — Trans. ent. Soc. London 74: 1—170. ——, 1957. Hitherto overlooked anatomical data concerning the genital structures in the Rhopalocera. — Trans. ent. Soc. London 109: 361—377. 138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 3, 1978 West, R. G., 1968. Pleistocene geology and biology. — Longman, London. 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 L = ee Ÿ 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). rs PAR tl di X FREE dr A Kl i BED- SE MUS. COMP. ZOOL. 1978 | DEEL 121 AFLEVERING 4 LIBRARY FEB8 1979 HARVARD | = > TIIDSCHRIFT 5% | VOOR ENTOMOLOGIE UITGEGEVEN DOOR SR EE 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 | te On Oi A PROVISIONAL CHECKLIST OF THE RHOPALOCERA OF THE EASTERN SIDE OF LAKE TANGANYIKA by 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 ER Toll Zambia 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 au À © D — TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 4, 1978 39 9, 25 a È 2022,20 29 VE © EN i Be -Kigëmax7 CE 31 2 0.-- “Mpandaz 33 Lake Tanganyika Sumbawanga IES Sed 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 ZA Kigoma N I = KE Ve ore ee = —— LK: + i = LS Kätavi > Nr G SE - 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 DI 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. 5 ae (re IJ = 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 Genezareth, ex 1. 24.iii.1933, Rhamnus palaestina Boiss.; 11: © Kopulationsapparat, Allotypus, GU/Kli- 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 Buschwaldgebiet, Imagines daraus (3 ©) 10.vi.1970, die in den äußeren 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 das ©. Die Kopfhaare der rhamnophila sind heller als die der rhamnella. Die 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 16 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, sämtliche ©. Diese entsprechen in ihren äußeren Merkmalen und im Habitus der 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 L. und ©. ilex L. vorkommenden Trifurcula (Ectoedemia) suberis Stt. Die Imago der 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 Stellung der Art ist noch etwas unsicher. Auffällig ist beim © das Fehlen einer 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: & Kopulationsapparat, GU/Kli-4132, Rhodos, Lindos, ex |. 22.ix.1977; 34: © Kopulationsapparat, GU/ 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 Pistacia lentiscus, 10.3.1968, Zucht No. 972, J. Klimesch”. © die gleichen Daten, 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, GU/Kli-1304, dieselben Daten, ex I. 20.ix.1973; 43: © Kopulationsapparat, GU/Kli-1300, Rhodos, Rodini, ex I. 11.x.1972, Quercus macrolepis Kotschy; 44: Minen an Q. macrolepis Kotschy, Rhodos, Rodini, 22.ix.1972; 45a: & Puppe in Ventralansicht, aus Zucht, Rhodos, Kremasti, ix.1973, ©. 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 51 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, beim © bis 1/2. Maxillarpalpen gelblich, ebenso die kürzeren Labialpalpen. 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. Beim © (Fig. 51) sind nach Johansson bei azaroli die vorderen Gonapophysen 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, daß die © der Herbstgeneration die Eiablage an den Knospen der Futterpflanze 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 am 19. und 20.vi.1966 je ein ©, die als Holo- und Paratypus der Beschreibung 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, ex I. 17.iv.1936, Potentilla verna agg. Fig. 63. N. styracicolella spec. nov., © Imago, Rhodos, Mt. Profitis 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. Der Beschreibung liegen 55 Imagines (24 3, 31 ©) zu Grunde; sie tragen folgende Daten: 22 &, 30 © “Lindos, Minen 10.6.1976, Zucht Nr. 1144; 1 & Lindos, Mine 5.6.1974, Zucht Nr. 1103, ex 1. 28.6.1974; 1 © Filerimos, Mine 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 beim & grau, beim © gelblichweiß. 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 Puppenruhe 2 & und 5 © erzielt. 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, Quercus macrolepis Kotschy, ex |. 22.iv.1973; 66: © Imago, dieselben Daten, ex |. 27.iv.1973; 67: & 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. Das zur Beschreibung verwendete Material (2 ©) trägt folgende Bezettelung: “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, beim © bis 1/2. Kopfhaare bei beiden Geschlechtern hell rostgelb, Nackenschöpfe 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 gelblichweiß, dicht — besonders beim © — mit schwarzen Schuppen bedeckt. 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 Fig. 68—69. Trifurcula (Ectoedemia) aegilopidella spec. nov. 68: © Kopulationsapparat, GU/Kli-1309, 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, GU/KIi-4065, Rhodos, Rodini, Salvia triloba L., ex |. 30.ix.1975; 71: © Kopulationsapparat, GU/Kli- 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. Fühler beim ¢ ca. 3/4 der Vorderflügellänge erreichend, beim © kaum merklich kürzer; dunkelbraun. Augendeckel gelblichweiß. Maxillarpalpen hellgelb. Kopfhaare hell rostfarben, ebenso die Nackenschöpfe. Thorax lederfarben, Abdomen hellgelb, beim ¢ anal gestutzt, beim © durch die 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 Bewehrung des Aedoeagus, während beim © die löffelartig verbreiterten Enden 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. Der Beschreibung liegen 44 Imagines (29 g, 15 ©) zu Grunde. Diese tragen 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 RS N N N N N N NN DO Co OR TX ISIS RM = 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 sup., Umg. v. Linz, ex I. 17.111.1932, Hypericum perforatum L.; 80: © Kopulationsapparat, GU/Kli-4105, 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 des Aedoeagus und in den Armen der Transtillae. Beim © erscheint die 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 L. 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 Hypericum hircinum, 31.8.1973, ex 1. 9.—13.9.1973”, 4 &,5 ©, “Zucht Nr. 1080, Rhodos, Ethonas-Malona”. Aus derselben Futterpflanze und vom gleichen Fundort: “Mine 22.5.1975, Zucht 1134A, ex 1. 11.— 20.6.1975”, 10 &, 5 ©. Vom KLIMESCH: Nepticulidenfauna von Anatolien 277 gleichen Fundort: “Mine an Hypericum spec. 31.8.1973, Zucht 1081, ex 1. 10.—18.9.1973”, 10 3,9 ©. “Rhodos, Mt. Smith, Mine 5.9.1973 an Hypericum 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 Amsel, H. G., 1933. Die Lepidopteren Palästinas. — Zoogeographica 2: 1—146. , 1935. Neue palästinensische Lepidopteren. — Mitt. zool. Mus. Berl. 20: 317. Beirne, B. P., 1945. The male genitalia of the British Stigmellidae. — Proc. R. Ir. Acad., Sect. B, 50: 190— 218. Borkowski, A., 1972. Studien an Nepticuliden, Teil IV. Bemerkungen zur Nomenklatur und Systematik der Familie Nepticulidae. — Polskie Pismo ent. 42: 689 — 709. Fabricius, J. C., 1775. Systema Entomologiae. — Kortius, Flensburg. Fologne, E., 1860. Nepticula ulmivora. — Ent. Wkly Intell. 8: 92. Frey, H., 1856. Die Tineen und Pterophoren der Schweiz. — Meyer und Zeller, Zürich. Gerasimov, A. M., 1937. Minierende Motten, V. — Mitt. zool. Mus. Berl. 22: 285. , 1952. Fauna USSR, Insects, Lepidoptera 1/2: 251. Groschke, F., 1944. Neues uber Minierer aus dem Mittelmeergebiet. — Mitt. munch. ent. Ges. 34: 115—124. Gustafsson, B., 1975. Anmarkningar om Nepticula ulmivora gruppen. — Ent. Tidskr. 96: 3—4. Haworth, A. H., 1803—1829. Lepidoptera Brittanica. — London. Heinemann, H., 1862. Einige Bemerkungen uber die Arten der Gattung Nepticula. — Wien. ent. Mo- natschr. 1862: 258. Hering, M., 1931. Minenstudien 12. — Z. Pflkrankh. PflPath. PflSchutz 41: 531—532. , 1932a. Minenstudien 13. — Z. Pflkrankh. PflPath. PflSchutz 42: 568. —, 1932b. Die Minenfauna von Tighina (Bender) in Bessarabien. — Bull. Acad. Roum. Sect. Scient. 15: 16. —, 1943. Untersuchungen über die Weiden-Nepticulen, I. — Z. wien. ent. Ges. 28: 273—278. , 1957. Bestimmungstabellen der Blattminen von Europa. — Verlag Junk, ’s-Gravenhage. Heyden, C. von, 1858. Nepticula of the Convolvulus. — Ent. Wkly Intell. 4: 175. Johansson, R., 1971. Notes on Nepticulidae I. A revision of the Nepticula ruficapitella group. — Ent. Scand. 2: 241—262. Klimesch. J., 1940. Uber eine Nepticuliden-Ausbeute von Triest. — Z. wien. Ent.-Ver. 25: 176—179. , 1942. Uber Microlepidopteren-Ausbeuten aus der Gegend von Zaton bei Gravosa (Süddalma- tien). — Mitt. munch. ent. Ges. 32: 395—397. —, 1946. Neue Stigmella-Arten. — Z. wien. ent. Ges. 31: 160—172. ——, 1948. Zur Frage der verwandtschaftlichen Beziehungen einiger Stigmella-Arten auf Grund des Baues des männlichen Kopulationsapparates. — Z. wien ent. Ges. 33: 49—82. ——, 1953. Die europäischen Trifurcula- und Ectoedemia-Arten. — Z. wien. ent. Ges. 38: 160—195. —, 1955. Nepticula szöcsi spec. nov. — Annls hist.-nat. Mus. natn. hung. 7: 423—424. — _, 1968. Die Lepidopteren-Fauna Mazedoniens, IV. Microlepidoptera. — Prirodonauten muzej, - Skopje. —, 1975a. Die an Ulme lebenden europäischen Nepticula-Arten. — Opuscula Zoologica 135: 1—15. —, 1975b. Uber neue mediterrane und kanarische Nepticuliden. — Mitt. münch. ent. Ges. 65: 1—28. —.,, 1976. Zur Genitalmorphologie und Lebensweise der Fedalmia (Stigmella) saturejae (Parenti, 1963). — Z. Arb.-Gem. ost. Ent. 28: 1—3. Muller-Rutz, J., 1934. Uber Microlepidopteren. — Mitt. schweiz. ent. Ges. 16: 122—123. 278 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 5, 1978 Preißecker, F., 1942. Zwei neue Nepticula-Arten aus dem Gebiet des Reichsgaues Wien. — Z. wien. Ent.-Ver. 27: 208—211. Rebel, H., 1926. Beitrag zur Mikrolepidopteren-fauna Dalmatiens. — Verh. zool.-bot. Ges. Wien 74—75: 106—110. Skala, H., 1939. Miner in deutschen Landen. — Ztschr. ost. Ent. Ver. 24: 27—30, 138—146, 149—152. , 1943. Nepticula groschkei spec. nov. — Z. wien. ent. Ges. 28: 86. Stainton, H. T., 1853. Nepticula viscerella Stainton. — Zoologist 1853: 3958. , 1967. The Tineina of Syria and Asia Minor. — Van Voorst, London. Staudinger, O., 1870. Beschreibung neuer Lepidopteren des europäischen Faunen-Gebiets. — Berl. ent. Z. 1870: 325. Staudinger, O., & H. Rebel, 1901. Catalog der Lepidopteren des palaarktischen Faunengebiets. — Ver- lag Friedlander & Sohn, Berlin. Szôcs, J., 1977. Hyponomia et Cecidia Lepidopterorum Fauna Hungariae. — Akad. Kiad. Budapest. Walsingham, Lord, 1911. Algerian Microlepidoptera. — Entomologist’s mon. Mag. 47: 190. Zeller, P. C., 1839. Versuch einer naturgemässen Eintheilung der Schaben. — Isis, Leipzig 1839: 215. 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 . À 4 EVOLUTION OF MECHANISMS TO CARRY POLLEN IN AGAONIDAE (HYMENOPTERA CHALCIDOIDEA) by 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 + 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 “v dno13 vspydojsyjg ‘7 :Q dno18 Dänydorsojg ‘ À :wunjbdoyssodi] ‘X ‘1ousodrao BuO] YIM u2/05010429 ‘A :10JISOdIAO JOYS YUM 493705010495) ‘N ‘10JISOdIAO JOYS YUM v2/09010499 ‘JL ‘> dno18 pävydoisvjg ‘Ò :s140y91J0q ‘d ‘S21U0p0]s1]d ‘O ‘U0DSY "N :UODBDADJ ‘IN :U00201410]]Y ‘7 ‘D]]2141990S1]F “A :vjjaisuofp *f ‘(odvasdinjg =) A dno13 v8vydorsvyg ‘] ‘ojjaiuopy ‘H ‘O dno18 v8vydoisvjg ‘D !sndp3sodag ‘A :vjjaiuojssajoy “A :punsiadvaog ‘Q :vuustadng ‘D *dno18 uovspjiaq ‘g ‘sndvajay “y :2epluoe8vy 10} weiSOpejo v ‘17 ‘314 Pek Yen NES 1582 Mes DAN MAN el HEI AS AS JAN 290 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 121, AFL. 6, 1978 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. 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