QL SA ENT. NOTA LEPIDOPTEROLOGICA Published by Societas Europaea Lepidopteroiogica (SEL) Volume 80 - Number 2 - 2007 SOCIETAS EUROPAEA LEPIDOPTEROLOGICA e.V. http://www.soceurlep.org HONORARY MEMBERS Günter Ebert (D), Pamela Gilbert (GB), Barry Goater (GB), Peter Hattenschwiler (CH), Prof. Dr Niels P. Kristensen (DK), Prof. Dr Vladimir Kuznetzov (RU), Dr Yuri P. Nekrutenko (UA) COUNCIL President: Dr Gerhard Tarmann (A) Vice-President: Prof. Dr Joaquin Baixeras (E) General Secretary: Dr David Agassiz (UK) Treasurer: Dr Robert Trusch (D) Membership Secretary: Willy De Prins (B) Ordinary Council Members: David Demerges (F), Prof. Dr Konstantin A. Efetov (UA), Dr Jörg Gelbrecht (D), Dr Andras Kun (H), Dr Nils Ryrholm (S) Editors: Dr Bernard Landry (CH), Dr Matthias Nuss (D) © Societas Europaea Lepidopterologica (SEL) ISSN 0342-7536 Type setting: blattwerk | dd Printed by Druckhaus Dresden GmbH All rights reserved. No part of this journal may be reproduced or transmitted in any form or by any means, electronic or mechanical including photocopying, recording or any other information storage and retrieval system, without written permission from the publisher. Authors are responsible for the contents of their papers. NFO 11 2007 N OTA LIBRARIES LEPIDOPTEROLOGICA A journal focussed on Palaearctic and General Lepidopterology Published by the Societas Europaea Lepidopterologica e.V. http://www.soceurlep.org Editors Dr Bernard Landry (Genève, CH), e-mail: bernard.landry @ ville-ge.ch Dr Matthias Nuss (Dresden, D), e-mail: matthias.nuss@snsd.smwk.sachsen.de Editoriai Board Dr Enrique Garcia-Barros (Madrid, E), Prof. Dr Roger L. H. Dennis (Wilmslow, UK), Dr Thomas Fartmann (Münster, D), Dr Axel Hausmann (Munich, D), Dr Peter Huemer (Innsbruck, A), Ole Karsholt (Copenhagen, DK), Dr Yuri P. Nekrutenko (Kiev, UA), Dr Erik van Nieukerken (Leiden, NL), Dr Thomas Schmitt (Trier, D), Dr Wolfgang Speidel (Bonn, D), Dr Carlos Lopez-Vaamonde (F), Dr Niklas Wahlberg (Turku, FIN) Volume 30 No. 2 - Dresden. 15.11.2007 - ISSN 0342-7536 Contents Andrew Wakeham-Dawson, Otakar Kudrna & Roger L. H. Dennis. Description of androconia in the Palaearctic Asian Pseudochazara baldiva (Moore, 1865) butterfly species-group (Nymphalidae: Satyrinae) with designation of two lectotypes and reference to type and other material in the Natural History Museum, Eon QT RE eek 211-223 Katalin Pecsenye, Judit Bereczki, Mariann Szilägyi & Zoltän Varga. High level of genetic variation in Aricia artaxerxes issekutzi (Lycaenidae) populations Er \oschera RO ARTE en ee are 225-234 H. Christof Zeller-Lukashort, Marion E. Kurz, David C. Lees & Michael A. Kurz. A review of Micropterix Hübner, 1825 from northern andieenttal Europe (Mieropterisidae) uses eseen naar 235-298 Pasi Sihvonen. Mating behaviour and copulation mechanisms in the genus ScOnuia Geomeltidae»Sterrhinae) aa. 299-313 Wolfgang A. Nässig & Rolf G. Oberprieler. The nomenclature of the family-group names of Eupterotidae (Bombycoidea) ..............uuusssesssennnenessnnnnneeneeennnnnnneeennn nn 315-327 210 Contents Frans Cupedo. Geographical variation and Pleistocene history of the Erebia pandrose — sthennyo complex (Nymphalidae; Satyrinae)..................... 329-353 Jérôme Pages. Une nouvelle espèce d’Hyponephele du Pakistan (Nymphalidae: Satyrinae) a... needs EEE 355-359 Jérôme Pages. Une nouvelle espèce de Pseudochazara du Pakistan (Nymphalidae, Satyrinae) keine ER 361-365 Thomas J. Witt, Vasiliy D. Kravchenko, Wolfgang Speidel, Josef Mooser, Amy Junnila & Günter C. Müller. A new Amata species from Israel (Aretiidae, SYNOMINAS) use reed en on Cee 367-373 Josef J. de Freina. Creatonotos omanirana sp. n. aus dem Oman und dem Iran (Artie: Are tna) 22258200 at RUN ee EE RS EEE 375-386 Axel Kallies. A revision of the clearwing moth species described by Zukowsky from China with additional notes on Sesiidae species from the Mell collection (SeSitdae) sms en a eee ae ee 387-396 Peter Hättenschwiler, Paul Sammut & Michael Zerafa. Rediscovery of Sciopetris melitensis Rebel, 1919 and description of its morphology and life history (Psychidaey Aue riesen ee ee 397-406 David Agassiz. Prays peregrina sp. n. (Yponomeutidae) a presumed adventive species 1n Greater London... nee ar es 407-410 Roman V. Yakovlev & Thomas J. Witt. Dyspessa aphrodite sp. n. from Greece (Cossidae) rise nd en IE 411-414 Roman V. Yakovlev. Taxonomic notes on Acossus Dyar and Parahypopta Daniel (Cossidas) nassen dere EEE 415-421 Instructions Tor authors vn ee ee 0 SEHE 422-424 Book reviews ee a ecke ee ER 224, 360, 374 Nota lepid. 30 (2): 211-223 211 Description of androconia in the Palaearctic Asian Pseudochazara baldiva (Moore, 1865) butterfly species-group (Nymphalidae: Satyrinae) with designation of two lectotypes and reference to type and other material in the Natural History Museum, London ANDREW WAKEHAM-DAWSON !, OTAKAR KUDRNA * & ROGER L. H. DENNIS ° Mill Laine Farm, Offham, Lewes, East Sussex, BN7 30B, UK; e-mail: andrew @ wakeham-dawson.orangehome.co.uk Naturmuseum Siidtirol, Bindergasse |, 39100 Bozen, Italy; e-mail: Kudrna.MEB @t-online.de Institute for Environment, Sustainability and Regeneration, Room s122, Mellor Building, Staffordshire University, College Road, Stoke on Trent, ST4 2DE, UK; e-mail: RLHDennis@aol.com D w Abstract. Sakai’s (1981) hypothesis (made in relation to material from Afghanistan) that the Pseudo- chazara baldiva-complex includes three species: P. baldiva (Moore, 1865), P. lehana (Moore, 1878), and P. droshica (Tytler, 1926) is tested by visual and statistical comparison of androconia scales from type and other specimens. The hypothesis is modified for a wider Central Asian study area to include P. bal- diva, P. droshica, and P. gilgitica (Tytler, 1926) as the names that have priority to represent three spe- cies-groups, each of which exhibits distinctively shaped androconia. Evans’ (1932a) misidentifications are resolved by designation of lectotypes for Eumenis mniszechii balucha Evans, 1932a and Eumenis mniszechii balucha f. pallida Evans, 1932a, and these two names are sunk in new synonymy as junior sub- jective synonyms of P. gilgitica and P. baldiva respectively. The findings of this study present a hypothesis of relationships for future testing against molecular and ecological data. Introduction The Pseudochazara de Lesse, 1951 butterflies of Afghanistan have been reviewed by Sakai (1981) as have those of Pamir, Turkmenistan, Uzbekistan, Tajikistan, Ladak and Kyrgyzstan (Tshikolovets 1997, 1998, 2000, 2003, 2005a, 2005b). Within this genus and endemic to Central Asia is a closely related group of mountain Pseudochazara taxa allied to P. baldiva (Moore, 1865). This is probably a natural complex of spe- cies artificially complicated by lepidopterists’ enthusiasm for naming taxa on the basis of location and differences in wing colour and pattern. Wing colour, in particular, is an unreliable taxonomic character in this genus (Wakeham-Dawson & Dennis 2001). Building on the work of Gross (1978) and Sakai (1981), we are presenting a rigorous approach to taxon determination by looking at the androconia: male scent scales used in courtship of satyrine butterflies (Tinbergen 1972) and thus probably more closely linked to reproductive isolation between biological species than wing colour. The present study is based on type and other specimens held in the Natural History Museum, London (BMNH). Androconia shape is compared visually and by statistical analysis of measurements made on the scales. We test Sakai’s (1981) hypothesis (made in relation to material from Afghanistan) that the P. baldiva-complex includes three spe- cies: P. baldiva, P. lehana, and P. droshica and propose a modified hypothesis based on a wider range of material from Central Asia. Synonymies are suggested based on these Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 DA Os WAKEHAM-Dawson et al.: Androconia in Palaearctic Asian Pseudochazara butterflies results and a taxonomic/nomenclatural complication is resolved by designation of two lectotypes. The findings of this study present a hypothesis of relationships for fu- ture testing against molecular and ecological data. Methods Biometrics: androconia were prepared and measured us- ing the methods described in Wakeham-Dawson (2006), Wakeham-Dawson & Kudrna (2000, 2005, 2006) and Wakeham-Dawson et al. (2003). In summary, androco- nia are removed from the forewing sex brands of male specimens and fixed on microscope slides. These slides are deposited in the BMNH and cross-referenced to specimens by authors’ and museum reference numbers. Measurements of androconium length (AL) and breadth (AB) are made using an eyepiece graticule (Fig. 1) from between five and ten androconia per specimen (where possible). The shape of the androconia is expressed as a ratio A (length/breadth) and the androconia are drawn to scale. Where possible type or topotype material has been Fig. 1. Measurements of andro- used and specimens determined by comparison to origi- conia from Pseudochazara type na] taxonomic descriptions. specimens held in the Natural BR | History Museum, London. (a) Statistical analysis: to test Sakai’s (1981) three-group en length (AL) from hypothesis measurements and ratios were subjected to asal stalk (bs) to terminal points ; above apex (tp); (b) androconium k-means clustering (Legendre & Legendre 1998). The breadth (AB) across the widest analysis was ‘seeded’ with three ‘type’ specimens as clus- part of lamina (1). Terminology a : after Kudrna (1977). ter centre points. These were specimens AWD628 to rep- resent P. baldiva (a syntype), 631 for P. droshica (a syn- type), and 618 for P. lehana (a topotype). The k-means clustering technique starts with k random clusters and then moves objects between clusters until (1) variability is minimized within clusters and (2) variability is maxi- mized between clusters. The method 1s related to Analysis of Variance (ANOVA), and the success of the operation is determined from the F statistics associated with each variable (or dimension). It is a suitable technique for situations where a certain number of groups is suspected to exist in a dataset and allows testing a priori classifications by ‘seeding’ the analysis with ‘type’ specimens (see Wakeham-Dawson et al. 2004 for another example of the use of this technique). The k-means clustering was carried out on standardized variables (each value taken from the mean and divided by the standard deviation to give zero mean, unit variance) to ensure comparability among the three variables. As it was not possible to normalise variable AB, results from this technique were compared with those using Kruskal-Wallis ANOVA by ranks. These results were then compared with an ANOVA test on more limited material (n = 15) where all three variables were standardized (each normal) on wing lengths. Nota lepid. 30 (2): 211-223 215 Material studied Unless otherwise stated, all specimens are male. B. M. Rhopalocera Type Nos can be compared against Riley & Gabriel’s (1924) catalogue. AWD = A. Wakeham-Dawson; OK = Otakar Kudrna. These names are written out in full on data labels, but are ab- breviated in the following list. IR Pseudochazara baldiva (Moore, 1865: 499, pl. 30, fig. 4). Originally named as ‘Lasiommata baldiva’. Sakai (1981) figures P. baldiva syntypes (male: pl. 24 figs 15-18; female: pl. 24 figs 19-20). Type locality: ‘Spiti and Tibet’. Syntype in Type Collection drawer 1-58: ‘[N. W. Himalaya], Upper Kuna- wur, Spiti, Lang 84’, ‘B. M. Type | No. Rh. 3724’, ‘Syntype | Lasiommata | baldıva Moore | det[ermined by] P. Ackery 1977’, ‘Coll. Moore 94-67’, ‘AWD | Androconia | preparation | 649°. Specimen 649 is accompanied by a female syntype (‘B. M. Type | No. Rh. 3725’) with similar data. Syntype in Main Collection drawer 3-22A: same data but ‘AWD Androconia slide 628’, ‘BM(N.H.) | Rhopalocera | Slide No. | 30610’. Non type specimen in Main Collection drawer 3-22A: ‘Skardo, July, 2000 ft’, ‘Leech Coll. 1901-173’, ‘AWD Androconia slide 629’, ‘ BM(N.H.) | Rhopalocera | Slide No. | 30608’. Specimen 629 is probably P. gilgitica (see below in Results). Pseudochazara lehana (Moore, 1878: 227). Originally named as ‘Hipparchia lehana’. Syntype figured by Tshikolovets (2005a, pl. 23 fig.8). Type locality: “Leh [34.10N, 77.35E], Kharbu [34.33N, 75.58E], 13000 ft, Ladakh’. Topotype in Main Collection drawer 3-23: ‘Himalayas, K[h]ardong [34.16N, 77.38E], 14000 ft, Aug. 1889’, ‘McArthur Coll. (Leech Coll. 1901-173)’, ‘AWD Androconia slide 618’, "BM(N.H.) | Rhopalocera | Slide No. | 30604’. Pseudochazara turkestana (Grum-Grshimailo, 1893: 384). Originally named as ‘Satyrus lehana turkes- tana’. Type locality: ‘Mts of Turkestan and Thian Schan (west)’. Topotype (syntype?) in Main Collec- tion drawer 3-23: ‘Boro Chozo’ , 1.vii.[18]89’, ‘Elwes Coll.’, ‘Coll. Gr[um]- Gr[shimailo]’, “AWD Androconia slide 620’, ‘BM(N.H.) | Rhopalocera | Slide No. | 30606’. Pseudochazara clarissima (Seitz, 1908: 128). Originally named as ‘Saryrus mniszechii clarissima’. Seitz (1908, pl. 43 figs f & g) figures this taxon. Type locality: ‘West China’. Topotype? (see Tshiko- lovets, 2005a: 105) in Main Collection drawer 3-23: ‘N. Kashmir, Hunza, 9450 ft, 21.viii.[19]13’, ‘No. 60’, ‘R. W. Hingston 1914-161’, “AWD Androconia slide 619’, ‘BM(N.H.) | Rhopalocera | Slide No. | 30605’. Pseudochazara esquilinus (Fruhstorfer, 1911: 308). Originally named as ‘Eumenis mniszechi [sic] esquilinus’. Type locality: ‘Alai’. ‘Type’ in Type Collection drawer 1-17: ‘Alai Mont[,] 1905[,] Korb’, ‘Fruhstorfer Coll. B.M. 1937-285’, ‘AWD | Androconia | preparation | 644’. Specimen 644 is accom- panied by a female with the same data. Pseudochazara droshica (Tytler, 1926: 256). Originally named as ‘Eumenis lehana droshica’. Sakai (1981) figures P. droshica syntypes (male: pl. 26 figs 5-6; female: pl. 26 figs 7-8). Type locality: ‘Drosh and Shandur, Chitral’. Cotype in Type Collection drawer 1-58: ‘Chitral, Jhela Drosh’, ‘B. M. Type | No. Rh. 10762’, ‘Capt. S. W. Harris 98-182’, ‘AWD | Androconia | preparation | 654’. The distinctively torn female syntype illustrated by Sakai (1981) appears to be no longer present in drawer 1-58. Syntype in Main Collection drawer 3—22A: ‘Chitral, 20.vi.[19]10, [on] wall’, ‘Syntype (det. P. Ackery 1979)’, ‘H. C. Tytler Coll. Brit. Mus. 1941-92’, ‘AWD Androconia slide 631’, “BM (N.H.) | Rhopalocera | Slide No. | 30612’. Pseudochazara gilgitica (Tytler, 1926: 256). Originally named as ‘Eumenis lehana gilgitica’. Type locality: ‘Ghizer, Gilgit’. Syntype in Type Collection drawer 1-58: ‘Chitral | Shandur Pass | vii. 1901 | el[evation] 11000 ft | Coll. G. A. Leslie | & W. H. Evans | 1903-284’, ‘B. M. Type | No. Rh. 110764’, ‘AWD | Androconia | preparation | 653’. Specimen 653 is accompanied by a female syntype (“B. M. Type | No. Rh. 10765’) with the following data: ‘Chitral | Shandur Pass | 11—14000 ft | 10.viii.[19]03 | A. R. C. Saunders | 1904-237’. Syntype in Main Collection drawer 3-22A: same data as specimen 214 WAKEHAM-DAWSON et al.: Androconia in Palaearctic Asian Pseudochazara butterflies 10. u: 653, but ‘AWD Androconia slide 630a’, ‘BM(N.H.) | Rhopalocera | Slide No. | 30613’. Syntype in Main Collection drawer 3—22A: ‘Gilgit, Ghizer’, ‘9. 23 [= ix.1923?]’, ‘H. C. Tytler Coll. Brit. Mus. 1941-92’, “AWD Androconia slide 630b’, "BM(N.H.) | Rhopalocera | Slide No. | 30614’. Pseudochazara balucha (Evans, 1932a: 113). Originally named as ‘Eumenis mniszechii balucha’. Type locality: ‘Urak, Baluchistan’. Evans’ (1932b) inclusion of ‘Ziarat, Kojak and Bogra’ in the type locality is no longer relevant with the designation of a lectotype for this taxon. Lectotype in Type Collection drawer 1-58: ‘Baluchistan | Urak | 10.vi.[19]28’, ‘W. H. Evans | B.M. 1932-274’, , ‘AWD | Syntype | det[ermined] 24.xi.2006’, ‘AWD & OK | Lec- totype | [designated] 24.x1.2006’ (see below in Results and Discussion), ‘AWD | Androconia | prepara- tion | 655°. Specimen 655 is accompanied by a female specimen with the following data: ‘Baluchis- tan | Ziarat | 17.v1.[19]28°, ‘W. H. Evans | B.M. 1932-274’. Three specimens (formerly topotypes) in Main Collection drawer 3-23: 1- ‘Baluchistan | Ziarat | 26.vi.1928’, ‘W. H. Evans | B.M. 1929-98’, ‘AWD | Androconia | preparation | 657”; 2- same data, but ‘21.vi.1928’, “AWD Androconia slide 624a’, ‘BM(N.H.) | Rhopalocera | Slide No. | 30619’; 3- same data, but ‘9.vii.1931’, ‘W. H. Evans | BM 1935-7’, ‘AWD Androconia slide 624b’, ‘BM(N.H.) | Rhopalocera | Slide No. | 30620’. Pseudochazara balucha f. pallida (Evans, 1932a: 113). Originally named as ‘Eumenis mniszechii balucha f. pallida’. Type locality: ‘Ziarat, Baluchistan’. Lectotype in Main Collection drawer 3-23: ‘Baluchistan | Ziarat | 17.vi.[19]28’, ‘W. H. Evans | BM 1932-274’, , ‘Syntype male | Eumenis mniszechii | balucha f. pallida | Evans | det[ermined by] P. Ackery 1979’, ‘ AWD & OK | Lectotype | [designated] 24.x1.2006’ (see below in Results and Discussion), ‘AWD | Androconia | slide 625 | 10.xi1.2004’, "BM(N.H.) | Rhopalocera | Slide No. | 30621’. Specimen 625 has been misidentified and unnecessarily named by Evans. It is probably P. baldiva or P. lehana (see below in Results and Discussion). Sakai (1981) figures specimen 625 (pl. 28 figs 17-18) and a female syntype (pl. 28 figs 19-20). Topotype: same data, but ‘1.vi.1930’, “W. H. Evans | B.M. 1934-491’, ‘AWD | Androconia | preparation | 656’. Specimen 656, like specimen 625, is probably P. baldiva or P. lehana (see below in Results and Discussion). Mislabeled specimen: same data, but 30.v11.1930°, ‘W. H. Evans | B.M. 1934-491’, ‘AWD | [slide] 643 | 6.v.2005 | Androconia’, “BM (N.H.) | Rhopalocera | Slide No. | 30624’. Specimen 643 is most probably a worn and faded specimen of P. balucha (see below in Results and Discussion). Pseudochazara watsoni Clench & Shoumatoff (1956: 148). Originally named as ‘Pseudochazara mnis- zechii watsoni’. Type locality: ‘Kotal Pass, 3800 m, Afghanistan’. Paratype in Main Collection drawer 3-23: ‘Afghanistan, Kotal Pass, August [17, 19]17’, ‘Paratype [Series] No. 279, Carn. Mus. Ent. Avinoff Coll. CM Acc. 14608’, ‘3 Danish Exp. Cent. Asia 148, N. Harlov, #127’, ‘Brit. Mus. 1974-415’, ‘AWD Androconia slide 621’, "BM(N.H.) | Rhopalocera | Slide No. | 30607’. Pseudochazara atambegi Wyatt & Omoto (1966: 161). Originally named as ‘Satyrus (Pseudochazara) baldiva atambegi’. Type locality: “Bala Quran, Anjuman, Badachshan’. Topotype in Main Collection drawer 3-22A: ‘Hindu Kush, Bala Quran, 17.v11.1971, Y. Ishikawa’, ‘AWD Androconia slide 627’, “BM(N.H.) | Rhopalocera | Slide No. | 30609’. . Pseudochazara wakhilkhani Wyatt & Omoto (1966: 164). Originally named as ‘Satyrus (Pseudocha- zara) watsoni wakhilkhani’. Type locality: ‘Bala Quran, Anjuman Valley, Badachshan’. Topotype in Main Collection drawer 3-23: ‘Afghanistan, NE Hindu Kush Mts, Bala-Quran, 3000-3400 m, 7-22.v11.1971, Anjuman V{alley], S. Sakai leg.’, ‘Brit. Mus. 1977-372’, ‘AWD Androconia slide 622’, ‘BM(N.H.) | Rhopalocera | Slide No. | 30617’. Non type specimen in Main Collection drawer 3-23: ‘Afghanistan, Kho-i-baba Mts, Mt Shah Fuladi, 3300 m, 8-12.v111.1974’, ‘Brit.-Mus. 1977-415’, ‘AWD Androconia slide 623’, "BM(N.H.) | Rhopalocera | Slide No. | 30618’. Results Measurements from the androconia are given in Table | (together with the nomenclatu- ral type status of each specimen) and representatives of androconia are illustrated in Figs 2-5. These neomorphic (see Warren 1963) androconia can be grouped by visual Nota lepid. 30 (2): 211-223 25 comparison into three main shapes: (1) very bulbous at the base (P. droshica and P. atambegi; Fig. 2), (2) rela- tively tall and thin with lam- ina widest near base giving a triangular appearance (P. gilgitica and P. balucha; Fig. 3), and (3) relatively shorter and thinner with lamina wid- est further up from base than in (2) giving a cigar-shaped appearance (P. baldiva, P. le- hana, etc.; Figs 4 & 5). Group (1) also includes aberrant scales from specimen 657, which have similar length to normal P. balucha scales, group (2) above, but are ex- tremely bulbous at the base like P. droshica scales. The k-means clustering analysis produced three Fig. 2. Androconia (drawn to scale, with scale bar representing 0.25 significantly differentiated mm) from specimens in Cluster 1: P. atambegi (AWD627), P. dro- Ver shica (631 & 654), and aberrant P. balucha scale (657b). clusters based on significant differences in the measure- ments (Androconium length, AL: F,,, = 23.0, P < 0.001; Androconium breadth, AB: F,, = 46.5, P < 0.001; Shape ratio, fe F,, ,= 41.2, P< 0.001). However, two of the ‘type’ specimens: 618 (P. lehana) and 628 (P. Baldıva) are placed in the same cluster. This is not a surprising outcome when we consider our findings from visual comparison of androconia in Figs 2-3. There are certainly three androconium shapes here as indicated by the three clusters, but from this analysis we have to reject Sakai’s (1981) hypothesis that P. lehana, P. dro- shica, and P. baldiva are the three names that have nomenclatural priority to represent the clusters of type specimens before us. From our analysis we can see that P. lehana (Moore, 1878) is a junior subjective synonym of P. baldiva (Moore, 1865). Re-visit- ing our androconia drawings in Figs 2-5 indicates that P. gilgitica (Tytler, 1926) is the name that has priority to represent the cluster of relatively tall, thin androconia in our data set. So we reran the k-means analysis ‘seeding’ with 628 to represent P. baldiva, 631 for P. droshica, and 653 for P. gilgitica (all these specimens are syntypes). This second analysis produces the same Euclidean distances between clusters and similar F-statistics (Androconium length, AL: F,,, = 21.8, P < 0.001; Androconium breadth, AB: F’,,, = 44.2, P < 0.001; Shape ratio, A: F,,, = 40.2, P < 0.001) as the first run and is represented in Fig. 6 as a non-metric multidimensional (NMMS) plot 22M 216 WAKEHAM-Dawson et al.: Androconia in Palaearctic Asian Pseudochazara butterflies 653 624a 624b 657a 629 Fig. 3. Androconia (drawn to scale, with scale bar representing 0.25 mm) from specimens in Cluster 2: P. balucha (AWD655, 624a, 624b & 657a), a specimen (643) labeled as P. balucha f. pallida, which is probably P. balucha, and a specimen (629) labeled as P. baldiva, which is probably P. gilgitica. Specimen 653 is P. gilgitica. of Euclidean distances. In Cluster 1 (four specimens represented by circles in Fig. 6): P. droshica (631 & 654) clusters with P. atambegi (627) and specimen 657b (aberrant scales from a P. balucha specimen) also clusters with this group. In Cluster 2 (seven specimens represented by triangles in Fig. 6): P. gilgitica (653) clusters with P. balucha (624a, 624b, 655 & 657a (normal scales)) and a specimen (643) labeled as P. balucha f. pallida. Specimen 629 (labeled as P. baldiva) also clusters with P. gilgitica, but 629 is not a type specimen and may therefore be P. gilgitica. In Cluster 3 (13 specimens represented by squares in Fig. 6): P. baldiva (628 & 649) clusters with P. watsoni (621), P. wakhilkhani (622 & 623), P. esquilinus (644), P. clarissima (619), P. lehana (618) and P. turkestana (620). Specimens 625 (syntype) and 656 that are labeled as P. bal- ucha f. pallida in the BMNH collection also cluster with P. baldiva. Specimens 630a and 630b (syntypes) labeled as P. gilgitica in the BMNH collection also cluster with P. baldiva, but are at the edge of the cluster closest to the P. gilgitica cluster. The an- droconia from specimens 630a and 630b are not cigar-shaped and are widest relatively close to their base. Nota lepid. 30 (2): 211-223 217 Fig. 4. Androconia (drawn to scale, with scale bar representing 0.25 mm) from specimens in Cluster 3: P. clarissima (AWD619), P. balucha f. pallida (656 & 625), P. lehana (618), P. baldiva (628), P. wa- khilkhani (622), and P. turkestana (620). Kruskal-Wallis ANOVA by ranks supported these contrasts (H, | _,,): AL = 13.2, P = 0.0013; AB = 16.1, P = 0.0003; A = 9.7, P = 0.008); all groups were separated using multiple comparisons: Clusters 1 and 2 by variables AB and A, Clusters 2 and 3 by variables AL and AB, and Clusters | and 3 by variable A. Using data standard- ized on wing lengths for a limited sample of 15 individuals (for which wing measure- ments were available), a one-way ANOVA confirmed inter-group (cluster) hetero- geneity (7, = 16.0, P < 0.001) despite small sample size, to which AB and A contri- 6,20 buted to heterogeneity. Discussion The use of visual and quantitative comparison of specimens in the current study sug- gests that Sakai’s (1981) nomenclature for the three androconia-shape groups based on material from Afghanistan should be replaced with the synonymy given below. This synonymy is based on material that includes more type specimens and covers a wider 218 WAKEHAM-DaAwson et al.: Androconia in Palaearctic Asian Pseudochazara butterflies Fig. 5. Androconia (drawn to scale, with scale bar represen- ting 0.25 mm) from specimens in Cluster 3: P. esquilinus (AWD 644), P. wakhilkhani (623), P. wat- soni (621), P. gilgitica (630a & 630b), and P. baldiva (649). See 630a 630b 649 text in relation to the placing of specimens 630a and 630b. Central Asian study area. Additional probable synonymies of nominal taxa, for which material was not available to us in this study (but whose androconia have been reliably illustrated in other studies), are given in square brackets with references. These names have been used in so many combinations in the literature (e.g. see Gross 1978; Tuzov 1997; Tshikolovets 2005a, b) that, with two exceptions (see text below), we do not at- tempt to identify which of these are new synonymies. Pseudochazara baldiva (Moore, 1865: 499) Hipparchia lehana Moore (1878: 227) Satyrus lehana var. turkestana Grum-Grshimailo (1893: 384) [Satyrus lehana var. sagina Heyne (1894: 542) (ref: Gross, 1978)] [Satyrus baldiva var. tarbagata Staudinger (1901: 57) (ref: Wakeham-Dawson & Kudrna, 2005) |] Satyrus mniszechii clarissima Seitz (1908: 128) Eumenis mniszechi [sic] esquilinus Fruhstorfer (1911: 308) Eumenis mniszechii balucha f. pallida Evans (1932a: 113), syn. n. Pseudochazara mniszechii watsoni Clench & Shoumatoff (1956: 148) Satyrus (Pseudochazara) watsoni wakhilkhani Wyatt & Omoto (1966: 164) [Satyrus (Pseudochazara) watsoni dargaga Wyatt & Omoto (1966: 165) (ref: Gross, 1978)] [Pseudochazara pseudobaldiva Gross (1978: 62) (ref: Gross, 1978)] [Pseudochazara pakistana Gross (1978: 63) (ref: Gross, 1978)] Nota lepid. 30 (2): 211-223 219 Pseudochazara droshica (Tytler, 1926: 256) [Pseudochazara porphyritica Clench & Shoumatoff (1956: 150) (refs: Gross, 1978; Sakai, 1981)] [Satyrus (Pseudochazara) baldiva panjshira Wyatt & Omoto (1966: 160) (ref: Gross, 1978)] Satyrus (Pseudochazara) baldiva atambegi Wyatt & Omoto (1966: 161) [Satyrus (Pseudochazara) turkestana badachshana Wyatt & Omoto (1966: 162) (ref: Sakai, 1981)] [Pseudochazara kopetdaghi Dubatolov, 1989: 138 (ref: Dubatolov, 1989)] Pseudochazara gilgitica (Tytler, 1926: 256) Eumenis mniszechii balucha Evans (1932a: 113), syn. n. 12 63 1.0 619 A ale 656 O 627 0.6 625 O oO 628 654 0.4 618 EH 644 O m O0 D 4623 630b 0.2 622 O O649 WN > SE? 621 < ae on 657b 04 653 Se O A À 624b -0.6 655 A =. 624a A -1.0 629 A 12 643 A 14 1,5 1.0 -0.5 0.0 0.5 1.0 15 2.0 2.5 3.0 AXIS 1 Fig. 6. A non-metric multidimensional (NMMS) plot of Euclidean distances between specimens (Aliena- tion = 0.005 and Stress = 0.0003 indicating that the clusters are accurately represented); see text for expla- nation. Cluster 1 (circles): P. droshica (631 & 654), P. atambegi (627), and specimen 657b (aberrant scales from a P. balucha specimen). Cluster 2 (triangles): P. gilgitica (653), P. balucha (624a, 624b, 655 & 657a (normal scales)), specimen 643 labeled as P. balucha f. pallida and specimen 629 labeled as P. baldiva. Cluster 3 (squares): P. baldiva (628 & 649), P. watsoni (621), P. wakhilkhani (622 & 623), P. esquilinus (644), P. clarissima (619), P. lehana (618), P. turkestana (620), P. balucha f. pallida (625 & 656) and P. gilgitica (630a & 630b, see text in relation to the placing of these specimens). The three ‘type’ speci- mens used to seed the k-means analysis are indicated by filled symbols. 220 WAKEHAM-Dawson et al.: Androconia in Palaearctic Asian Pseudochazara butterflies Tab. 1. Androconium length (AL) from basal stem to terminal points and androconium breadth (AB) across the widest point of the lamina (mm) (means + sd), and ratio A (AL/AB; no units) (mean + sd) that represents overall shape of androconia from 23 specimens among 12 nominal taxa of Pseudochazara butterflies. Spec. No. = Authors’ numbers allocated to specimens and androconia slides. BMNH No. = BMNH Rhopalocera Slide Number; * = No. not yet allocated at time of publication. N = number of androconia measured. Status: C = cotype, P = paratype, S = syntype, L = lectotype, T = type, TT = topo- type, TT* = topotype status removed by designation of lectotype, NOT = not type material, + = mislabeled in BMNH (this is probably P. gilgitica), ++ = mislabeled in BMNH (this is probably P. balucha). Status BMNH No. A 0.344 0.066 5.19 TT AWD627 30609 +0.004 | +0.002 Taxon P. atambegi P. baldiva NOT+ | AWD629 30608 vn 11 eo nee 0.332 0.032 10.45 S AWD628 30610 OO | one P. baldiva P. baldiva S # NOT++ 30624 m | oath — © P. f. pallida P. f. pallida P. f. pallida P. balucha P. balucha * (aberrant 0.082 349 IT AWD657b scales) + 0.005 * (normal 0.406 0.049 8.34 Tr AWD657a | scales) +0.009 | +0.002 me [anno [ao Ten, | 28% [14% me [avr [mn Tem, | 208, [28% me [ans [as [026, | 203, [102% 0.358 0.056 6.36 € AWD654 |* +0.000 | + 0.000 0.317 0.056 6.00 S AWD631 30612 +0.011 | +0.011 0.339 0.034 10.15 T AWD644 d +0.015 | +0.003 0.371 0.036 10.38 S AWD630a 30613 +0.011 + 0.004 S AWD630b 30614 0.356 0.042 8.67 +0.009 | +0.004 | + 1.04 S AWD653 3 0.396 0.039 10:35 +0.011 | +0.003 | +0.90 TT AWD618 30604 0.331 0.029 11.67 + 0.011 +0.004 | +1.81 Ss? AWD620 30606 0.351 0.029 12.20 +0.008 | +0.001 + 0.34 NOT AWD623 30618 0.345 0.035 9.78 +0.008 | +0.002 | +0.52 LE AWD622 30617 0.338 0.032 10.56 +0.013 | +0.002 | +0.74 AWD621 30607 0.368 0.036 10.38 +0.008 | +0.003 | + 0.98 P. balucha P. balucha P. balucha P. clarissima P. droshica © P. droshica P. esquilinus P. gilgitica P. gilgitica P. gilgitica P. lehana P. turkestana P. wakhilkhani P. wakhilkhani P. watsoni = Nota lepid. 30 (2): 211-223 | Our comparison of androconia shows that specimen AWD625 (which has been identi- fied as a syntype of Eumenis mniszechii balucha f. pallida Evans 1932 by P. R. Ackery) is clearly a different taxon from Eumenis mniszechii balucha and not just a pale form. This difference has been noted previously by Sakai (1981) and Wakeham-Dawson (2006). Evans (1932a: 113) describes his taxa as follows within a key of satyrid taxa that he includes in the genus Eumenis: ‘WSF [Wet Season Form] band dark; on upf [up- per-surface of fore-wing] decreasing to dorsum, lower ocellus always present and usu- ally 2 white dots between the ocelli: unf [under-surface of fore-wing] discal line clear, submarginal line zigzag. DSF [Dry Season Form] band very pale, on upf of even width, lower ocellus rarely present and white dots absent; unf discal line clear, submarginal line sinuous. [Eumenis] mniszechii balucha, nov. (58-65). The Tawny Rockbrown. Baluchistan. NR. (DSF pallida, nov).’ Evans (1932b: 201-202) gives the following additional descriptions for his two taxa: ‘45. ‘Eumenis mniszechii balucha’, Evans and ‘pallida’, Evans. The Tawny Rockbrown. A large dark brown species with a broad tawny band across both wings. It flies from the Caucasus and Asia Minor to Central Asia, the N.-W. Frontier to Ladak and W. China. It is common from June to August at Urak, Ziarat, the Kojak and Bogra, and at Ziarat in May and June (perhaps elsewhere) there is to be met a very pale form. The ordinary local form differs from its allies in the following respects: on the forewing above the band decreases posteriorly, there is always a lower ocellus and usually two white dots between the ocelli: on the underside of the forewing the central line 1s clearly marked and the submarginal line is zigzag. The form pallida, probably an early brood, has the band very pale brown; forewing below the central line is distinct and the submarginal line is sinuous.’ In addition (p. 197) he states: ‘All the types will be placed in the British Museum’. Evans had a mixed series of specimens before him and misidentified his ‘f. pallida’ specimens. Androconia shape indicates that specimen 625 (labeled as balucha f. palli- da) is probably a member of our P. baldiva group. Specimen 656 (placed in the balucha f. pallida series) is probably also a member of our P. baldiva group. However, specimen 643 (also placed in the balucha f. pallida series) has P. balucha-shaped androconia (when compared with syntype specimen 655; see Fig. 3) and has been mis-placed as P. balucha f. pallida in the BMNH collection because it is worn and faded. In addition, the shape of the orange band on the forewings is like that of syntype P. balucha rather than syntype P. balucha f. pallida. Furthermore specimen 643 is labeled as having been captured in late July, while true P. balucha f. pallida specimens are labeled as having been captured in early to mid June. To resolve any confusion of identity or nomenclature identified above, we hereby designate specimen AWD625 as Lectotype of Eumenis mniszechii balucha f. pallida Evans 1932a and synonymize this name with Lasiommata baldiva Moore 1865 (New Synonymy). In addition, we designate specimen AWD655 as Lectotype of Eumenis mniszechii balucha Evans 1932a and synonymize this name with Eumenis lehana gilg- itica Tytler, 1926 (New Synonymy). Our three-group synonymy is fairly bold and there is a danger that our nomenclature fails to acknowledge genetic variation between isolated populations of these butterflies. 222 WAKEHAM-DAWsoN et al.: Androconia in Palaearctic Asian Pseudochazara butterflies The relationships between these taxa may be much more subtle with clines of varia- tion and/or the effects of periodic isolation and recombination of mountain populations during ice ages. So the next stage in this research will be to test our hypothesis with additional material, molecular data and ecological field observations. Currently we do not know to what extent or even whether different androconia shape is associated with different pheromone production and how in turn this may be related to isolation be- tween taxa. One of our P. balucha specimens (AWD657) merits special discussion in this respect. Two shapes of androconia are present on this specimen. The majority are typical P. balucha scales, but three scales on the microscope slide are shaped like large P. droshica scales. We are confident that these scales are not contamination from another specimen as colour, location and size all indicate that they belong to specimen 657. What has caused this ‘mutation’, and is it the same mechanism that has given rise to androconia shape differences between populations and taxa? Can we assume that both shapes of scale on this specimen produce the same chemicals? If so, this again raises the question as to whether shape differences correspond to chemical differences between taxa. Or is the presence of two scale types the result of hybridization between taxa? Acknowledgements We thank Geoff Martin, Jim Reynolds and Phil Ackery (Department of Entomology, The Natural History Museum, London at Wandsworth), Goulven Keineg (Entomology Library, Wandsworth) and Alison Harding (Entomology Library, Tring) for their assistance in this study. Comments by Yuri Nekrutenko, an anonymous referee, and the editors improved an earlier draft of this paper. References Clench, H. K. & N. Shoumatoff 1956. Lepidoptera Rhopalocera (Insecta) from Afghanistan. The 3" Da- nish Expedition to Central Asia. — Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 118: 141-191. Dubatolov, V. V. 1989. [New species of butterflies (Lepidoptera, Lycaenidae, Satyridae) from Middle Asia]. — Proceedings of the Zoological Institute, Leningrad 200: 136-140. Evans, W. H. 1932a. The Identification of Indian Butterflies. Second Edition Revised. — Bombay Natural History Society, Madras. 454 pp., 32 pls. Evans, W. H. 1932b. The butterflies of Baluchistan. — Journal of the Bombay Natural History Society 36(1): 196-209. Fruhstorfer, H. 1911. Satyridae. Pp. 285-401. — In: Seitz, A. (ed.), Die Gross-Schmetterlinge der Erde, vol. 9. Die Indo-Australischen Tagfalter. — Alfred Kernen Verlag, Stuttgart. Gross, F. J. 1978. Beitrag zur Systematik von Pseudochazara-Arten (Lep., Satyridae). — Atalanta, Wurzburg 9: 41-103. Grum-Grshimailo, G. E. 1893. Lepidoptera palaearctica nova. Il. Horae Societatis — Entomologicae Rossicae 27: 379-386. Heyne, A. 1894. Pp. 529-576. — In: Rühl, F. & A. Heyne, Die palaearktischen Grossschmetterlinge und ihre Naturgeschichte. Vol. 1, Lief 11.- E. Heyne, Tagfalter. Leipzig. Kudrna, O. 1977. A Revision of the Genus Hipparchia Fabricius. — Classey, Faringdon. 300 pp. Legendre, P. & L. Legendre 1998. Numerical Ecology. Second Edition. — Elsevier, Oxford. xv, 853 pp. Moore, F. 1865. List of diurnal Lepidoptera collected by Capt. A. M. Lang in the N. W. Himalayas. — Proceedings of the Zoological Society of London, 1865: 486-509. Nota lepid. 30 (2): 211-223 DES Moore, F. 1878. Descriptions of new species of Lepidoptera collected by the late Dr F. Stoliczka dur- ing the Indian-Government Mission to Yarkund in 1873. — Annals and Magazine of Natural History (ser. 5) 1: 227-237. Riley, N. D. & A. G. Gabriel 1924. Catalogue of the Type Specimens of Lepidoptera Rhopalocera in the British Museum. — BMNH, London. 62 pp. Sakai, S. 1981. [Butterflies of Afghanistan]. — Kodansha, Tokyo. 271 pp., 48 pls. Seitz, A. 1908. 4. Familie: Satyridae. Pp. 79-153, pl. 43. — In: Die Grossschmetterlinge de Erde..., 1. Abteilung: Die Grossschmetterlinge des Palaearktischen Faunengebietes. 1. Band: Die Palaearkti- schen Tagfalter. — Fritz Lehmann Verlag, Stuttgart. Tinbergen, N. 1972. The courtship of the grayling Eumenis (= Satyrus) semele (L.). Pp. 197-249. — In: Tinbergen, N. (ed.), The Animal in its World: explorations of an ethologist. 1, Field Studies 1932-1972. — Allen & Unwin, London. Tshikolovets, V. V. 1997. The Butterflies of Pamir. — Slamka, Bratislava. 282 pp., 46 pls. Tshikolovets, V. V. 1998. The Butterflies of Turkmenistan (Lepidoptera, Rhopalocera). — Tshikolovets, Brno- Kyiv. 237 pp., 34 pls. Tshikolovets, V. V. 2000. The Butterflies of Uzbekistan (Lepidoptera, Rhopalocera). — Tshikolovets, Brno- Kyiv. 400 pp., 49 pls. Tshikolovets, V. V. 2003. The Butterflies of Palaearctic Asia. IV. The Butterflies of Tajikistan (Lepidoptera, Rhopalocera). — Tshikolovets, Brno-Kyiv. 500 pp., 77 pls. Tshikolovets, V. V. 2005a. The Butterflies of Palaearctic Asia. V. The Butterflies of Ladak (N.-W. India) (Lepidoptera, Rhopalocera). — Tshikolovets, Brno-Kyiv. 176 pp., 30 pls. Tshikolovets, V. V. 2005b. The Butterflies of Palaearctic Asia. VI. The Butterflies of Kyrgyzstan (Lepi- doptera, Rhopalocera). — Tshikolovets, Brno-Kyiv. 511 pp., 108 pls. Tuzov, V. K. (Ed.) 1997. Guide to the Butterflies of Russia and Adjacent Territories. Vol. 1. Hesperiidae, Papilionidae, Pieridae, Satyridae. — Pensoft, Sofia, Bulgaria. 480 pp., 78 pls. Tytler, H. C. 1926. Notes on some new interesting butterflies from India and Burma, pt. 1. — Journal of the Bombay Natural History Society 31(2): 248-260. Wakeham-Dawson, A. 2006. Descriptions of wing androconia from some Pseudochazara de Lesse, 1951 (Lepidoptera: Nymphalidae, Satyrinae) type specimens in The Natural History Museum, London. — Entomologist’s Gazette 57: 99-107. Wakeham-Dawson, A. & R. L. H. Dennis 2001. A quantitative description of the male genitalia of 23 taxa of Pseudochazara de Lesse, 1951 (Lepidoptera: Nymphalidae, Satyrinae). — Entomologist’s Gazette 52: 227-250. Wakeham-Dawson, A. & O. Kudrna 2000. A quantitative description of androconia from Staudinger’s Pseudochazara de Lesse, 1951 (Lepidoptera: Nymphalidae, Satyrinae) type specimens in the Zoolo- gical Museum of the Humboldt University of Berlin. — Entomologist’s Gazette 51: 75-81. Wakeham-Dawson, A. & O. Kudrna 2005. Further descriptions of androconia from Staudinger’s Pseu- dochazara de Lesse, 1951 (Lepidoptera: Nymphalidae, Satyrinae) type specimens in the Zoologisches Museum der Humboldt-Universitat zu Berlin. — Entomologist’s Gazette 56: 139-146. Wakeham-Dawson, A. & O. Kudrna 2006. Description of wing androconia from the lectotype of Pseudo- chazara caucasica (Lederer, 1864) (Lepidoptera: Nymphalidae, Satyrinae), with notes on the topotype wing androconia of related taxa. — Entomologist’s Gazette 57: 137-141. Wakeham-Dawson, A., P. Jaksic, J. D. Holloway & R. L. H. Dennis 2004. Multivariate analysis of male genital structures in the Hipparchia semele-muelleri-delattini complex (Lepidoptera: Nymphalidae, Satyrinae) from the Balkans: how many taxa? — Nota lepidopterologica 27(2/3): 103-124. Wakeham-Dawson, A., R. Parker, E. John & R. L. H. Dennis 2003. Comparison of the male genitalia and androconia of Pseudochazara anthelea acamanthis (Rebel, 1916) from Cyprus, Pseudochazara anthelea anthelea (Hiibner, 1924) from mainland Turkey and Pseudochazara anthelea amalthea (Frivaldsky, 1845) from mainland Greece (Lepidoptera: Nymphalidae, Satyrinae). — Nota lepidoptero- logica 25(4): 251-263. Warren, B. C. S. 1963. Monograph of the Genus Erebia. - BMNH, London. vii + 407 pp., 104 pls. Wyatt, C. & K. Omoto 1966. New Lepidoptera from Afghanistan [1.]. — Entomops 5: 138-167. 224 Book reviev Huemer, P. & S. Erlebach 2007. Schmetterlinge Innsbrucks. Artenvielfalt einst und heute. — Universitätsverlag Wagner, Innsbruck. 319 pp. ISBN 978-3-7030-0432-2. 25.00 €. (in German) With 118,000 inhabitants Innsbruck is the fifth largest city of Austria. Situated in the Inn river valley, it is protected by high mountain chains to the north as well as to the south. It comprises an area of 105 square kilometers and altitudes range from 565 to 2,641 meters. During the last decades, urbanisation increased seriously, causing an alarming decline of butterflies and moths, a characteristic situation in all of Central Europe. In spring 2004 a team of lepidopterists started to record the current butterfly and moth fauna of Innsbruck. Until 2006, recording took place from March to October every year using nets, artificial lights, sugar baits, and pheromones. An astonishing number of nearly 1,200 species was recorded. However, analyzing the historical data dating back 150 years, 920 species for- merly known from Innsbruck could not be found again. Though there are several species difficult to record, like Nepticulidae, the decline is also obvious in butterflies. Their number of species decreased from 138 to 87, or a third the species, since lepidopterological research started in Innsbruck 150 years ago. The largest decline happened in the period of 1950 to 1979. Altogether, 437 species have been recorded for the last time during that period. More details are given and analyzed in the excellent book on the butterflies and moths of Innsbruck by Huemer and Erlebach. Written in German and in a popular science style it will be understandable for entomologists as well as amateurs in this field. It is illustrated with 255 color figures, mostly photographs of Lepidoptera and their habitats, but also maps and graphs. The book is divided into nine chapters, an (1) introduction, (2) a chapter on nomenclature, characters, the metamorphosis and lifespan of Lepidoptera, (3) diversity of species in Innsbruck and their systematics, (4) history of lepidopterological research in Innsbruck (5) a description of the areas of unspoiled nature and the habitats of Lepidoptera, (6) threats, local losses of Lepidoptera and their conservation, (7) experiencing nature, (8) literature, and (9) an appendix with the list of species known from Innsbruck for the periods before 1878, 1879-1914, 1915-1949, 1950-1979, 1980-1999, and since 1999. Altogether, the reader can learn about Lepidoptera in general, the history of their study in Innsbruck, changes to their habitats and reasons for their decline, and he can get an introduction to the literature and an idea of the current composition of the lepidopteran fauna of Innsbruck. Thus, the book summarizes the knowledge on Lepidoptera in Innsbruck for professionals, amateurs, and those who are just interested in nature, and it might be a starting point to study or just to enjoy butterflies and moths. This book is not just written by lepidopterists for lepidopterists. It has the potential to interest non-lepidopterists to enjoy butterflies and moths, to respect them, and to care for them in the future. MATTHIAS Nuss Nota lepid. 30 (2): 225-234 225 High level of genetic variation in Aricia artaxerxes issekutzi (Lycaenidae) populations in Northern Hungary KATALIN PECSENYE !, JuDIT BERECZKI !, MARIANN SZILAGYI ! & ZOLTAN VARGA !:? ! Department of Evolutionary Zoology and Human Biology, University of Debrecen, Debrecen, Egyetem tér 1., Hungary; e-mail: pecskati@tigris.unideb.hu > HAS-DU Research Group for Evolutionary Genetics and Conservation Biology Abstract. Aricia artaxerxes issekutzi imagos were collected from eight localities in two regions in Northern Hungary (Bükk Mountains and Aggtelek Karst region) in 1999 and 2000. Enzyme polymorphism was analysed at 16 enzyme loci using polyacrylamide gel electrophoresis. In the analysis of the data, F-sta- tistics was computed and the total genetic variation Was partitioned into within and between population components. Nei’s genetic distances were calculated and UPGMA dendrogram was constructed on the basis of the distance matrix. Hierarchical F-statistics and AMOVA were computed to study the pattern of genetic differentiation among the samples. PCA analysis was also carried out using the allele frequencies of the samples. The Hungarian populations of A. artaxerxes issekutzi exhibited a high level of enzyme polymorphism. Both the dendrogram and the results of PCA indicated a clear differentiation between the Bükk and Karst regions. Most parameters of polymorphism and also the level of differentiation among the local populations were similar in the two regions. Nevertheless the average number of alleles per locus was significantly lower in the samples of the Bükk Mts than those of the Karst ones. In addition, the Bükk populations possessed a significantly lower portion of the species allele pool compared to the Karst ones. It was mostly the consequence of genetic drift due to the small population sizes in the Bükk Mts. Introduction Survival of natural populations in fragmented landscapes is an increasingly impor- tant topics in conservation biology (Ricketts 2001; Ries & Debinski 2001; Baguette & Schtickzelle 2001). Habitat fragmentation results in population subdivision and isola- tion. Consequently, it has grave influence on the genetic structure of populations result- ing in decreasing effective population size and loss of genetic variation (Thomas et al. 1998). Nevertheless, different species may experience the same fragmented habitat in a different way (Thomas & Harrison 1992). Thus, surveys of genetic differentiation among local populations within a species have become more and more embedded in conservation studies (Schmitt & Hewitt 2004). Butterflies have been considered as sensitive indicators of changes in cultural landscapes. As a consequence, studies ap- plying population genetic techniques to butterfly conservation surveys have assumed increasing importance. Aricia artaxerxes (Fabricius, 1793) is a widespread Eurasiatic species. It occurs from Northern Europe to Central Asia and East Siberia (Obraztsov 1935, 1936). In the north- western periphery of the range and also in the mountainous parts of southern Europe, it is subdivided into several subspecies (Beuret 1954; Kaaber & Hoegh-Guldberg 1961; Urbahn 1964; Hoegh-Guldberg 1966, 1968; Hoegh-Guldberg & Jarvis 1969; Kames 1969). One of these subspecies occurs in calcareous areas of the Carpathian basin such as the Bükk Mountains and the Aggtelek Karst region in Northern Hungary, de- scribed as A. artaxerxes issekutzi Balogh, 1956 (Varga 1961, 1968). The documented food plants of A. artaxerxes issekutzi are Helianthemum ovatum and Geranium san- guineum. However, the caterpillars of A. artaxerxes (different subspecies) have been Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 226 PECSENYE et al.: Genetic variation in Aricia artaxerxes in Hungary reared under artificial conditions on different Geranium species and Erodium cicu- tarium (Hoegh-Guldberg & Jarvis 1969; Varga pers. obs.). The favoured habitats of this subspecies are dry or semi-dry swards rich in flowers. Aricia artaxerxes issekutzi is a monovoltine species. Females lay their eggs usually singly on the lower surface of the leaves or on the flower buds of the food plant (Malicky 1969, Varga 1968). They have about 100 eggs but only 3-5 eggs will be layed on a plant. In this way, the eggs of a female are distributed fairly evenly within the habitat. Larvae develop first on the food plant. Later they are taken to ant nests. Aricia artaxerxes, however, is only a facultative myrmecophilous species (Malicky 1969). The Brown Argus butterflies are known to be highly variable Lycaenid species (Aagard et al. 2002). They are, therefore, appropriate to analyze the genetic consequences of habitat degradation and fragmentation. The aim of the present study was to compare the level and structure of genetic variation in strong and declining populations of A. artaxerxes issekutzi ın Northern Hungary. Materials and Methods Samples. Aricia artaxerxes issekutzi samples were collected from two subregions of northern Hungary: Aggteleki Karst region (strong populations) and Bükk Mountains (declining populations). Altogether 12 samples were collected from 8 populations in 1999 and 2000 (Fig. 1). In this way, the samples exhibited two types of hierarchy: geo- graphic regions and populations within the regions (spatial pattern); years and popula- tions collected in the same year (temporal pattern). Imagos were collected mostly in July, after the main egg-laying period and stored at —80°C until electrophoresis. Sample sizes varied between 12 and 40, according to the size of the populations. Enzyme studies. Allozyme polymorphism was studied at 16 different loci by vertical polyacrylamide gel electrophoresis: aconitase (Acon), alcohol dehydrogenase (Adh), aldehyde oxidase (Aox), esterase (Est), glutamate dehydrogenase (Gdh), glutamate oxalacetate transaminase (Gor), glucose-6-phosphate dehydrogenase (G6pdh), a-glyc- erophosphate dehydrogenase (aGpdh), hexokinase (Hk), isocitrate dehydrogenase (Idh), lactate dehydrogenase (Ldh), malate dehydrogenase (Mdh), malic enzyme (Me), phosphoglucose isomerase (Pgi), phosphoglucomutase (Pgm), and superoxid dis- mutase (Sod). Thoraxes were homogenized in 300y1 extraction buffer and these sam- ples were used to study Got, aGpdh, G6pgdh, Hk, Idh, Ldh, Mdh, Me, Pgi, Pgm, and Sod. Abdomens were homogenized in 1501] extraction buffer and these extracts were used to analyse Acon, Adh, Aox, Est, and Gdh. The extraction buffer, the electrophore- sis buffer systems and running conditions, together with the staining solutions used for each enzyme are described in Bereczki et al. (2005). Genotypes of the different indi- viduals were scored according to their enzyme pattern. Statistical analyses. Genotype and allele frequencies were calculated on the basis of banding patterns. Measures of genetic variation (average number of alleles, % propor- tion of polymorphic loci, average observed heterozygosity) were calculated for each sample. The distribution of alleles with different frequencies between the two geo- graphic regions was compared using generalised linear models. The computation was Nota lepid. 30 (2): 225-234 2 carried out with GLIM 4 (Francis et al. 1994). Genetic differentiation among the populations was first analysed by Wright’s F-statistics (Wright 1978; Weir I ON. Szelce 1996). In this analysıs, the total genetic 7 variation of the samples (F,,.) was parti- tioned into within (F.) and between pop- ulation components (F,,.). The analyses were conducted by FSTAT version 1.2 (Goudet 1995). An exact test for popula- tion differentiation (Raymond & Rousset 1995a) was also conducted to test for in- Kesskeäbre 77 dependence of the allelic composition of the populations. Genepop, version 1.0 (Raymond & Rousset 1995b) was used to perform this test. Allele frequencies were used to estimate Nei’s genetic distances (Nei 1972) and an UPGMA dendrogram (Sneath & Sokal 1973) was constructed on the basis of these data. The computa- tion of genetic distances was performed by Biosys-I, Release 1.7 (Swofford & Selander 1981). The distribution of the total genetic variation at various lev- els of the hierarchy was also studied by AMOVA (Excoffier et al. 1992; Weir 1996). In this analysis, the total genetic variation is partitioned into three components: among groups, among populations within a group and within population. AMOVA was carried out by Arlequin, version 2.000 (Schneider et al. 2000). In the last part of the study, we carried out a principal component analysis (PCA) using the genotypic com- position of the individuals to show the size of overlap in the genetic variation of the populations in a reduced space of variables. PCA analyses were performed running R Package Version 4.0 (Casgrain & Legendre 2001). Tohonya "no... Szölöhegy er Zsidérét — @ Fig. 1. Sample sites of A. artaxerxes issekutzi in northern Hungary. Results Level of enzyme polymorphism. Three of the 16 loci analysed (Hk, aGpdh, Sod) did not have an alternative allele in any of the investigated sample. The samples exhibited a very high level of polymorphism. As a whole, the portion of polymorphic loci was 57.3% and the average frequency of heterozygotes was 24.6% (Tab. 1). Heterozygote deficiency was observed in all samples, which proved to be significant in 8 cases out of the total 12 (Tab. 1). The total number of alleles was 71 at the 16 loci investigated. We have calculated the portion of alleles the samples actually possessed of the total 71 (Tab. 1: A). In general, populations contained 64% of the species gene pool. The 228 PECSENYE et al.: Genetic variation in Aricia artaxerxes in Hungary Tab. 1. Parameters of enzyme polymorphism in the Aricia artaxerxes issekutzi samples. N: sample size; n,: average number of alleles per locus; A: average portion of the allele pool of the species; P: portion of the polymorphic loci; F,,: index representing the within population component of genetic variation. *: sig- nificant at 0.05 level; **: significant at 0.01 level; ***: significant at 0.001 level. po NS in, AE EP aa 0.214 | 6.3 Dre | | Subtotal Hacava 0 0 Menes i, 2 Subtotal Hacava | Haragistya | 35.1 | 3.06 | 690 | 563: | 0.249 | 0029 | | Ménes | 218 | 288 | 648 | 500 | 0239 | 0.132" |__| Szélôh | 25.8 | 288 | 690 | 563 | 0.255 00930 | | Tohonya | 394 | 288 | 648 | 625 | 0242 5) ona foe 0 4 Subtotal | 30.9 | 3.00 | 69.0 | 58.8 | 0.249 | 0.103** _ | Total | | Ss | 642 |573 | 0246 Tome er) ES | | | Szeke | 120 | 244 | 549 | 500 | 0.234. Room RTE RS Tab. 2. Results of F-statistics for the A. artax- samples were grouped in two ways. First, erxes issekutzi samples. F,.: index indicating the the samples collected in 1999 were split ac- total genetic variation of the samples; F_.: index representing the within population component; COrding to their geographic origin (Bükk vs. F.: fixation index. *: significant at 0.05 level; Karst populations). Except for the param- *#%: significant at 0.01 level. eters indicating allelic richness, there was no significant difference in the level of poly- oo Bsn morphisms between the samples collected in the two geographic regions. Both the aver- age number of alleles per locus (F, ,,=8.09; 0.05>P>0.01) and the portion of the species gene pool the population possessed (4° =7.73 0.01>P>0.001) were, however, significantly higher in the populations collected in the Karst region compared to those of the Bükk eee next step bes to sort i ae of ek SE the Karst regıon accordıng to the years (gen- erations) they were collected. Although the samples collected in 2000 exhibited a slightly - | higher polymorphism these differences were not significant for any of the parameters (Tab. 1). We also compared the distribution of alleles with different frequencies between Nota lepid. 30 (2): 225-234 229 0.050 0.000 9Hac 9Toh OHac 9Har OMen Karst OToh 0Szöh 9Men OHar 9Szel 0.05

P>0.01). The Bükk populations carried less rare alleles but were fixed for more alleles than the Karst populations (Fig. 2). At the same time, the two samples collected in the Karst region proved to be similar (4°,=4.04; P>0.05). Structure of the genetic variation. The results of F-statistics indicated that a substan- tial portion of this variation was observed within the samples. The average F,, value suggested significant heterozygote deficiency within the samples (Tab. 2), which was attributable to 7 of the 13 loci. At the same time, the samples exhibited a relatively low level of genetic differentiation (Tab. 2: F,,). Yet, 8 of the total 13 loci proved to be sig- nificantly differentiating (Tab. 2). Nei’s (1972) genetic distances were calculated and a UPGMA dendrogram was con- structed on the basis of the distance matrix. The dendrogram showed an obvious geo- graphic pattern (Fig. 3). The samples originating from the two regions were clustered in two well differentiated branches. Moreover, the two samples collected in Hacava, which is situated in the Slovakian part of the Karst region were separated from the other Karst samples. At the same time, the samples of the other Karst populations (except for the ‘99 sample from the Tohonya ridge) collected in the two consecutive years (gen- erations) were scattered randomly in the middle branch (Fig. 3). This indicates a fairly high level of random changes in their allele frequencies. In the analysis of the structure of genetic differentiation, we contrasted the spatial and temporal components of the between sample variation. Accordingly, we computed AMOVA using Arlequin. In this analysis, the total genetic variation can be analysed in- volving the within sample component as well. However, we could not include all levels of the hierarchy above the sample in a single run. We, therefore, conducted the analy- 230 Di fferentiation among the populations within the regions |; Different ation among the samples within the populations ll Temporal variation Differentiation between the regions Spatial variation Fig. 4. Distribution of the total between sample variation in A. artaxerxes issekutzi. Fig. 5. Results of the PCA analysis for the A. artaxerxes is- sekutzi samples. A: Bükk samples; @: Karst samples col- lected in 1999; O: Karst samples collected in 2000. ein [Yer Tr, [ Baio Ro Ron on ne (6) (6) (6) PECSENYE et al.: Genetic variation in Aricia artaxerxes in Hungary ses in two series. In the first run, the levels of the hierarchy were: regions (group of samples) and samples. In the second run, the fol- lowing levels of the hierarchy were considered: populations (group of samples) and samples. The results suggested that most of the total genetic Variation was attributable to the within sample component (95.08%). The distribution of the between sample variation revealed that a much higher portion of vari- ation was explained by the differ- ences among the populations (Fig. 4: spatial variation) than by those within them, i.e. between the sam- ples collected in different gen- erations from the same population (Fig. 4: temporal variation). The indices of differentiation also in- dicated that the spatial component of variation was far larger than the temporal one. F,,, the index repre- senting the level of differentiation among the populations was highly significant (F,,= 0.034, P<0.001) while F,,, the index representing the level of differentiation among the samples within the populations was not (F,,= 0.011, P>0.05). Sımilarly to the dendrogram, the results of the PCA analysis also in- dicated a clear geographic pattern of differentiation. The samples ex- hibited two large clouds of points Tab. 3. F,, values and the results of exact test at different levels of the hierarhy in the A. artaxerxes issekutzi samples. The num- bers in brackets indicate the significantly differentiating loci. *: significant at 0.05 level; **: significant at 0.01 level; ***: sig- nificant at 0.001 level. Nota lepid. 30 (2): 225-234 25 Tab. 4. Results of AMOVA for the A. artaxerxes issekutzi samples. BR: between region variation, WRBP: between population variation within a region; WPBS: between sample (year/generation) variation within a population; WS: within sample (year/generation) variation. Indices of differentiation were determined in two analyses with different levels of hierarchy: Analysis 1 — region and sample; Analysis 2 — popula- tion and sample. F,,,: index of differentiation among the regions; F,,,: index of differentiation among the populations; F,,„: index of differentiation among the samples (year/generation) within a region; F,,: index of differentiation among the samples (year/generation) within a population; F,_: index of total differentia- tion among the samples (year/generation). Indices of differentiation 0.059%** 0.035*** in the reduced space of variables (Fig. 5). The two axes explained 56.7% of the genetic variation of the samples. The 95% ellipses drawn according to the geographic regions and the years the samples were collected indicated an evident separation between the Karst and Bükk populations along the first axis. The allele frequency distribution at the Got and Aox loci contributed most to this axis. In accordance with the results of the hierarchcal F-statistics the samples collected in two consecutive years/generations in the Karst populations comprised largely overlapping clouds. Discussion In line with other European lycaenid butterflies studied we found a high level of poly- morphism in Aricia artaxerxes issekutzi populations. Polyommatus coridon, P. bellar- gus and P. icarus has been surveyed in many Western and Central European populations (Schmitt & Seitz 2001a, b; Schmitt et al. 2003). Those results revealed a high level of polymorphism e.g. the proportion of polymorphic loci ranged from 0.42 to 0.85. The mean number of alleles per locus has been especially high (3.0-3.5) in the southern European P. coridon populations (Schmitt & Seitz 2001a; Schmitt et al. 2002). Aagaard et al. (2002) have reported a high level of genetic variation in the northern European populations of Aricia artaxerxes and A. agestis. They found 1.84 alleles per locus on average with some loci having 6 or even 8 alleles. Our data indicated an even higher polymorphism than those of Aagaard et al. (2002). In the Hungarian Aricia popula- tions, the average number of alleles ranges between 2.2 and 3.3. Moreover, we detected 17, 11 and 9 alleles at the Est, Pgi and Pgm loci respectively. As a consequence of the high number of alleles per locus, we also observed an exceptionally high frequency of heterozygotes (average H =0.246). As far as known, in butterfly populations enzyme studies has only revealed such a high level of heterozygosity in P. hispana populations (Schmitt et al. 2005). Although most parameters of polymorphism were similar in the two regions studied, the average number of alleles per locus and the average portion of the species gene 232 PECSENYE et al.: Genetic variation in Aricia artaxerxes in Hungary pool each population possessed were significantly lower in the Bükk samples than in the Karst ones. Moreover, the distribution of alleles among the three frequency catego- ries (rare, common and fixed alleles) was also different in the two regions. Namely, the fixed alleles were more frequent in the Bükk populations, whereas the rare alleles were more common in the Karst ones. It thus appears that populations living in the Bükk plateau have a lower allelic richness than those of the Karst region. A possible explanation of this situation is that Aricia populations are smaller and more isolated in the Bükk plateu than in the Karst region. Aricia artaxerxes prefers short-grass habitats at moderately high altıtudes, which are rich in flowering dicotyledonous plants. The main nectar sources of this species are small Fabaceae species with yellow flowers like Lotus corniculatus, Melilotus spp. (Varga pers. obs.). Moreover, females should find the small larval food plants in the lower vegetation. These circumstances can only be maintained under suitable edaphic-microclimatic conditions, e.g. at the rupicolous margin of karstic dolinas and/or by appropriate man- agement. In general, mountain grasslands, however, are prone to succession process without appropriate management. The structure will change due to overgrowth by tall grasses and by the extension of shrubby vegetation. This process has been quite rapid on the higher (about 800-850 m) and more humid (over 800 mm precipitation pro year) Biikk plateau as a consequence of abandoned mowing. Accordingly, Aricia artaxerxes issekutzi, which was fairly common in the sixties (Varga pers. obs.) became rare and more localised during the last two decades. Management has started in the last 2—3 years in order to save these populations. The situation is quite different in the Aggtelek Karst region. Lower precipitation (about 600 mm per year) and the shallow, karstic substrate is associated with an essentially slower succession. As a consequence, a more favourable structure and diversity of vegetation has been maintained in the Karst region than on the Bükk plateau. Both short-grass swards with abundant Helianthemum ovatum and xerothermic forest-steppe fringes with Geranium sanguineum remained widely distributed. Moreover, these habi- tat patches compose a kind of network on the Karst plateaus facilitating migration. Thus, Aricia populations have remained rather strong there. In contrast, the suitable habitat patches have bocome much smaller and relatively isolated in the Bükk Mts. The consequence of small population size and isolation is the enhanced effect of genetic drift, which results in an increased probability of allele fixation and loss of rare alleles (Frankham et al. 2002; Allendorf & Luikart 2006 and references therein). We detected both of these symptoms in the Bükk populations. The level of differentiation was significant among the Hungarian A. artaxerxes issekutzi samples. Aagard et al. (2002) also detected significant inhomogeneity among the A. ar- taxerxes populations in the UK and Scandinavia. Assuming the relatively strong effect of genetic drift in the Bükk populations we expected a higher level of differentiation in this region than in the Karst one. Contrary to our presumption, the F, values indicated a fairly similar level of genetic differentiation in the two regions. This suggests that the size of the Bükk populations has decreased recently, 1.e. they have been exposed to the effect of genetic drift for a short period. Though they have lost several rare alleles and a number of loci have become fixed this time period has not been long enough to enhance genetic differentiation among the local populations. Nota lepid. 30 (2): 225-234 253 Acknowledgements The study was supported by the OTKA T046612 and NKFP-3 B/023/2004 projects. The technical assistance of V. Mester in the electrophoretic work is very much respected. The support of the Nature Conservation Authorities of Hungary and especially the cooperation of the Aggtelek National Park Direction is highly appreciated. References Aagaard, K.,K. Hindar, A. S. Pullin, C. H. James, O. Hammarstedt, T. Balstad & O. Hanssen 2002. Phylo- genetic relationship in brown argus butterflies (Lepidoptera: Lycaenidae: Aricia) from north-western Europe. — Biological Journal of the Linnean Society 75: 27-37. Allendorf, F. W. & G. Luikart 2006. Conservation and the Genetics of Populations. — Blackwell Publi- shing, Malden, USA. Baguette, M. & N. Schtickzelle 2001. Local population dynamics are important to the conservation of metapopulations in highly fragmented landscapes. — Journal of Animal Ecology 40: 404-412. Bereczki, J., K. Pecsenye, L. Peregovits & Z. Varga 2005. Pattern of genetic differentiation in the Macu- linea alcon species group (Lepidoptera, Lycaenidae) in Central Europe. — Journal of Systematics and Evolutionary Research 43: 157-176. Beuret, H. 1954. Die Lycaeniden der Schweiz. Teil II. — Mitteilungen der Entomologische Gesellschaft Basel (N.F.) 4, suppl.: 204-222. Casgrain, P. & P. Legendre 2001. The R package for multivariate and spatial analysis, version 4.0 d5 User’s manual. Excoffier, L., P. Smouse & J. Quattro 1992. Analysis of molecular variance inferred from metric distances among DNA haplotypes: Application to human mitochondrial DNA restriction data. — Genetics 131: 479-491. Francis, B., M. Green & C. Payne 1994. GLIM 4. The statistical system for generalised linear interactive modelling. —- New York University Press Inc. Frankham, R., J. D. Ballou & D. A. Briscoe 2002. Introduction to Conservation Genetics. — Cambridge University Press, Cambridge. Goudet, J. 1995. Fstat version 1.2: a computer program to calculate F-statistics. — Journal of Heredity 86: 485. Hoegh-Guldberg, O. 1966. North European groups of Aricia allous G-Hb. Their variability and Relation- ship to A. agestis Schiff. — Natura Jutlandica 13: 1-184. Hoegh-Guldberg, O. 1968. Evolutionary trends in the genus Aricia. — Natura Jutlandica 14: 1-76. Hoegh-Guldberg, O. & F. V. L. Jarvis 1969. Central and North European Ariciae (Lep.). Relationships, Heredity, Evolution. — Natura Jutlandica 15: 1-106. Kaaber, S. & O. Hoegh-Guldberg 1961. Aricia allous Hb. ssp. vandalica nov. — Flora og Fauna. Meddelse fra Naturhistorisk Museum Aarhus 67: 122-128. Kames, P. 1969. Aricia artaxerxes spp. nova hercynica. — Natura Jutlandica 15: 107-119. Malicky, H. 1969. Übersicht über Präimaginalstadien, Bionomie und Ökologie der mitteleuropäischen Lycaenidae (Lepidoptera). — Mitteilungen der entomologischen Gesellschaft Basel 19: 25-91. Nei, M. 1972. Genetic distance between populations. — The American Naturalist 106: 283-292. Obraztsov, S. 1935. Lycaena (Aricia) allous Hb. und ihre östlichen Rassen. — Folia Zoologica et Hydrobiologica Riga 8: 141-150. Obraztsov, S. 1936. Weiteres tiber Lycaena (Aricia) allous Hb. — Folia Zoologica et Hydrobiologica Riga 9: 58-61. Raymond, M. & F Rousset 1995a. An exact test for population differentation. — Evolution 49: 1280-1283. Raymond, M. & F Rousset 1995b. GENEPOP ver.1.2, a population genetics software for exact tests and ecumenicism. — Journal of Heredity 86: 246-249. Ricketts, T. H. 2001. The Matrix Matters: Effective Isolation in Fragmented Landscapes. — The American Naturalist 158: 88-99. Ries, L. & D. M. Debinski 2001. Butterfly responses in habitat edges in the highly fragmented prairies of Central Iowa. — Journal of Animal Ecology 70: 840-852. 234 PECSENYE et al.: Genetic variation in Aricia artaxerxes in Hungary Schmitt, Th. & G. M. Hewitt 2004. The genetic pattern of populationthreat and loss: a case study of but- terflies. — Molecular Ecology 13: 21-31. Schmitt, Th. & A. Seitz 2001a. Allozyme variation in Polyommatus coridon (Lepidoptera, Lycaenidae): - identification of ice-age refugia and reconstruction of postglacial expansion. — Journal of Biogeogra- . phy 28: 1129-1136. Schmitt, Th. & A. Seitz 2001b. Intraspecific allozymatic differentiation reveals the glacial refugia and the postglacial expansions of European Erebia medusa (Lepidoptera: Nymphalidae). — Biological Journal of the Linnean Society 74: 429-458. Schmitt, Th., A. Giessl & A. Seitz 2002. Postglacial colonisation of western Central Europe by Polyomma- tus coridon (Poda, 1761) (Lepidoptera, Lycaenidae): evidence from population genetics. — Heredity 88: 26-34. Schmitt, Th., A. Giessl & A. Seitz 2003. Did Polyommatus icarus (Lepidoptera, Lycaenidae) have distinct glacial refugia in southern Europe? Evidence from population genetics. — Biological Journal of the Linnean Society 80: 524-538. Schneider, S., D. Roessli & L. Excoffier 2000. Arlequin ver. 2.000: A software for population genetic data analysis. — Genetics and Biometry Laboratory, University of Geneva, Switzerland. Sneath, P. H. & R. R. Sokal 1973. Numerical Taxonomy. — W. H. Freeman, San Francisco. Swofford, D. L. & R. B. Selander 1981. Biosys-1: A FORTRAN program for the comprehensive analysis of electrophoretic data in population genetics and systematics. — Journal of Heredity 72: 281-283. Thomas, C. D. & S. Harrison 1992. Spatial dynamics of a patchily distributed butterfly species. — Journal of Animal Ecology 61: 437-446. Thomas, C. D., J. K. Hill & O. T. Lewis 1998. Evolutionary consequences of habitat fragmentation in a localised butterfly. — Journal of Animal Ecology 67: 485-497. Urbahn, E. 1964. Neuere Auffassungen in der Maculinea alcon-rebeli-Frage (Lepidoptera, Lycaeni- dae). — Entomologische Abhandlungen, Staatliches Museum fiir Tierkunde, Dresden 32: 69-77. Varga, Z. 1961. Zoogeographical survey of the Macrolepidoptera-fauna of the N-Hungarian Karst area (in Hung. with German summary). — Folia entomologica hungarica 14: 345-386. Varga, Z. 1968. Bemerkungen und Ergänzungen zur taxonomischen Beurteilung und Ökologie der im Karpatenbecken vorkommenden Populationen von Aricia artaxerxes Fabr. (= A. allous G.-Hb., A. montensis Vrty., Lepidoptera, Lycaenidae) — Acta Biologica Debrecina 6: 171-185. Weir, B. S. 1996. Genetic Data Analysis II: Methods for Discrete Population Genetic Data. — Sunderland, Sinauer Associates, Inc. Wright, S. 1978. Evolution and the Genetics of Populations. vol. 4. Variability Within and Among Natural Populatons. — University of Chicago Press, Chicago. Nota lepid. 30 (2): 235-298 235 A review of Micropterix Hübner, 1825 from northern and central Europe (Micropterigidae) H. CHRISTOF ZELLER-LUKASHORT !, MARION E. Kurz’, Davin C. LEES * & MICHAEL A. KURZ * ! Forsthubfeld 14, 5303 Thalgau, Austria; e-mail: christof.zeller@gmx.net > Sportplatzstraße 23, 5303 Thalgau, Austria; e-mail: marion.kurz@ gmx.at > Department of Entomology, Natural History Museum, Cromwell Road, London SW7 5BD, U.K.; e-mail: del@nhm.ac.uk * Reischenbachweg 2, 5400 Hallein-Rif, Austria; e-mail: michael.kurz@gmx.at Abstract. All known species of the Palaearctic genus Micropterix Hübner, 1825 (Micropterigidae) from northern, north-western, central and north-eastern Europe including the Alps are reviewed. The purpose of this review is to permit reliable identification of the genus in this region. Adults, male and female genitalia are illustrated for eighteen species. For these, taxonomy, adult life history (preferred biotopes, phenology and pollination records) and variability is detailed. Distribution is summarized for each species. Treated species are fully diagnosed and also compared with extralimital taxa. The presently known species diver- sity of Micropterix is assessed and a brief synopsis of diagnostic characters for the genus is given in the context of what is known for the family worldwide. A checklist is provided and a key is presented based on adult wing pattern and characters of the male and female genitalia. Advice on genitalic preparation and information about suitable collection sites is also provided. Introduction The genus Micropterix Hübner, 1825 is distributed in the temperate and subtropical zones of Eurasia, as well as of Northern Africa (Kristensen 1984a). Up to now, 83 species are known (72 have been described), taking into account probable synony- my, of which 49 occur in Europe, 16 additional species in Asia (very few of which reach the eastern Palaearctic), and about 17 additional in northern Africa. This makes Micropterix by far the most species rich genus of Micropterigidae, a family numbering about 132 described and about 93 undescribed species up to end of 2006 (Kristensen 1999; Hashimoto 2006; Gibbs in litt.). Most of the described species occur in the Mediterranean region, and several undescribed species are to be expected in the moun- tains of Asia. It is the purpose of this paper to review eighteen species of the Palaearctic genus Micropterix. We treat all species known from northern, north-western, central and north- eastern Europe, including Fennoscandia, Great Britain and Ireland, France (including the northern slopes of the Pyrenees), the entire Alps, Hungary, Romania, Slovakia, Poland, Belarus and the Baltic States (Fig. 122). All species are figured in colour, and their genitalia are illustrated. Our focus in this paper is on the aspects of the species most useful for identification, namely wing colour patterns and abdominal morphol- ogy, for which we present a key. We also summarise what is known of adult life history and emphasise how little is known of the evolutionary relationships even of European species. We present for now a largely phenetic arrangement of species. In this paper we build on the previous important identification treatments by Heath (1987), Kozlov (1989, 1990a, b) and Whitebread (1992). Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 236 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Material and methods For many years the authors have collected specimens of Micropterix throughout the western Palaearctic region. These specimens form the basis of the presented study. Furthermore many other specimens, including type specimens from the following mu- seum and private collections have been investigated: BMNH British Museum of Natural History, London CZ Collection Christof Zeller HNSA Museum Haus der Natur, Salzburg IEUP Istituto di Entomologia, Università degli Studi, Pavia, Italy LSUK Linnean Society, London MK Collection Michael Kurz MNHN Museum National Histoire Naturel, Paris MSNM Museo Civico di Storia Naturale, Milano MSNV Museo Civico di Storia Naturale, Verona NHMK Landesmuseum für Kärnten, Klagenfurt NHRS Naturhistoriska Riksmuseet, Stockholm NMW Naturhistorisches Museum, Wien OLML Oberösterreichisches Landesmuseum, Biologiezentrum, Linz SMNK Staatliches Museum für Naturkunde, Karlsruhe TLMF Tiroler Landesmuseum Ferdinandeum, Innsbruck USNM Smithsonian National Museum of Natural History, Washington ZMHB Museum für Naturkunde, Humboldt-Universität zu Berlin ZMUC Zoological Museum, Copenhagen ZSM Zoologische Sammlung des Bayrischen Staates, München Collection sites. Best collection sites are boundaries of shrub vegetation and wood- land, where the adults gather on flowering vegetation often very locally, but often also in great abundance. The specimens fly only short distances, but can frequently be found crawling around pollinating many different blossoms. Some species prefer herbaceous plants, others shrubs or trees. In the southern latitude or lower elevation areas Micropterix were more usually found in shaded areas than areas exposed to full sunshine; in more northern latitudes, they often occur in open sunny areas. Ata given locality, adults aggregate usually on a limited range or even only a single plant species. At higher elevations some species can often be found around elfin woodland, feed- ing on the pollen of Pinus mugo Turra. Ecologically more specialized species inhabit marshes, dwarf vegetation, open grassland or alpine meadows. In the Mediterranean, specimens often rest on the underside of leaves or in shady places during the hottest hours of the day, being active mainly in the morning and in the evening. In woodland, specimens may be active throughout daytime. The adults are attracted to flowers of the following gymnosperm and angiosperm (monocot and dicot) plant families and genera or species (Adler et al. 1994; Kozlov 2006; Pankhurst 1999: Schwartz-Tzachor 2004; Zeller-Lukashort, Kurz & Kurz 2006; Helmut Deutsch, Markus Fluri, Peter Huemer, Utsugi Jinbo, all pers. comm .; this work): Pinaceae (Pinus mugo Turra), Ranunculaceae (Caltha palustris L., Actaea spicata L., Ranunculus spp.), Caryophyllaceae (Lychnis flos-cuculi L., Stellaria holostea L.), Fagaceae (Fagus sylvatica L. (pollen on leaves, not flowers), Quercus spp.), Urticaceae Nota lepid. 30 (2): 235-298 237 (Urtica dioica L.), Brassicaceae (Cardamine pratensis L.), Sapindaceae (Acer pseudo- platanus L.), Rosaceae (Aruncus dioicus (Walter) Fernald, Crataegus sp., Filipendula ulmaria (L.) Maxim., Rosa pendulina L., Rosa canina L. agg), Fabaceae (Spartium junceum L., Vicia sp.), Euphorbiaceae (Mercurialis perennis L.), Apiaceae (Heracleum sp.), Cistaceae (Helianthemum sp., Cistus salvifolius L.), Ericaceae (Rhododendron ferrugineum L., Vaccinium sp.), Pyrolaceae (Pyrola sp.), Primulaceae (Cyclamen per- sicum Mill.), Rubiaceae (Galium sp.), Scrophulariaceae (Veronica chamaedrys L.), Plantaginaceae (Plantago media L.), Oleaceae (Fraxinus ornus L., Olea europaea L., Syringa vulgaris L.), Caprifoliaceae (Sambucus nigra L., Lonicera xylosteum L.), Liliaceae (Paris quadrifolia L.), Orchidaceae (Dactylorhiza romana (Sebast & Mauri) Soö), Cyperaceae (Carex spp.) and Poaceae. External characters used for identification. Forewing markings and the colour of the scales of the head are the external characters used for the identification of the Micropterix species dealt within this paper. The colouration of the wings ıs solely due to the physical properties of the wing scales (which are of the ‘fused’ type and on the upper surface as in other lower grade Lepidoptera). Wing pattern is dominated by a combination of metallic golden, silvery and purplish elements. The basic marking scheme on the forewing seems to be a purplish ground colour with three transverse golden fasciae. This basic scheme is modified in many ways, such as by additional spots and fasciae, or by shifting the purplish ground colour to a golden one and the colour of the fasciae varying to a whitish or silvery one. Although of great importance for identification, forewing markings do vary to a significant degree. This variation seems to be not only genetically determined, but may be influenced by local habitat conditions, as discussed below under Life history, and on a geographic basis according to metapopulation structuring (Kozlov 1995). We include several instances here of lo- cal, geographically distinct colour morphs (see e.g. Micropterix paykullella (Fabricius, 1794), Micropterix aruncella (Scopoli, 1763)). Schematic wing drawings used in the identification key are encoded as given in Fig. 1. brownish golden - purple - bluish violet Es | reddish golden-coppery bronzy golden - greenish golden TT | | whitish golden - silvery Anatomical characters used for identification. Several anatomical details, present- ing valuable characters in some other lepidopterous families, like wing venation or number and size of spurs on the legs, are not useful for species identification in the ge- nus Micropterix. Male genitalia are very characteristic in this genus and always present sufficient details for a reliable specific identification. Female genitalia, although less useful, sometimes yield useful exoskeletal characters on segments IX and X, and to some extent from the shape and length of the receptaculum seminis. Nevertheless, in Fig. 1. Code of schematic wing drawings. 238 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe some cases, the geographical distribution has also to be considered, since many similar species are allopatrıc or endemic to a relatively small area. The main features of male and female abdominal characters are explained in Figs 2 and 3. Preparation techniques (genitalia preparation). The whole of the abdomen is re- moved and then transferred to a 10%-solution of caustic potash. By gentle shaking, the solution is heated until the abdomen becomes transparent and the ‘soft tissues’ have been completely dissolved. After rinsing the abdomen with water, it can be transferred to a drop of a 7:3 mixture of water and glycerol on a micro slide. The preparation should be as clean and devoid of scales as possible when inspecting it under 20-30 fold magnification. Most drawings and many photos of the female genitalia have been made in this stage of preparation, since the receptaculum seminis may be recognized very well now. Details of sclerotized parts have been added to the drawings using permanent preparations. For determination purposes only, it is normally not necessary to remove the sclerotized parts or the phallus from the male genitalia. Also it is not necessary to stain the prepara- tion, since all diagnostically important characters are strongly sclerotized. Therefore the preparation can be transferred to 96% ethanol for at least 15 minutes and afterwards mounted in Euparal in lateral position. For this purpose, the genitalia are first fixed in a drop of Euparal overnight before the cover slip is brought into position with a second drop of Euparal. With this technique, squeezing the genitalia can be avoided best. For taxonomical studies it may be necessary to remove the sclerotized parts and the phallus, to choose another position than the lateral one or to squeeze the genitalia (especially when investigating the spinoid (thickened) setae of the accessory claspers). When preparing female genitalia, it is important to know that the inner structures of the abdomen are not sclerotized at all. Removing the pregenital exoskeleton and staining the preparation is therefore not advisable, because dehydration during the process strongly deforms the receptaculum seminis. Furthermore sternum IX may be damaged, which is important for some species identifications. After rinsing the preparation with water, the following procedure can therefore be used for the mounting of the female genitalia. Firstly, the last two segments of the abdomen, forming a short, retractable ovipositor, are expanded carefully. These segments must not be damaged. A micro-pipette, that has been drawn above a flame to a very fine tip is connected to a rubber tube with a mouth-part and filled with a dispersion of chlorazol black in water. Then the tip of the pipette is inserted into the abdomen up to segment VI or VII, beginning at the proximal end. A tiny amount of the stain suspension is now injected by gentle pressure. In this way the receptaculum seminis may be stained blue without opening the exoskeletion. This also helps to avoid distortion of the membranous receptaculum seminis when dehydrating the preparation with 96% ethanol. Finally the preparation is transferred to Euparal and mounted in the usual way. A lateral position fits the natural flattening of the abdomen, although for inves- tigation of sternite IX a dorso-ventral mounting is to be preferred. Data archives. The data of all investigated specimens have been archived in a da- tabase, which is accessible via internet: http://www.nkis.info/nkis/auscollabfrage. cgi?uid=guest&lang=e (Kurz et al. 2000-2007). To create a list, just input “Micropterix” into field “Scientific name” and click on Button “Search”. Each specimen has been la- belled with a unique identification (ID) number representing the record number of the Nota lepid. 30 (2): 235-298 239 database. With this ID-number, the data of each specimen can be checked individually on www.nkis.info, menu “DATA ANALYSIS”. Furthermore, these records, together with original life observations, contribute to dynamically generated distribution maps. Analyses concerning phenology, vertical distribution, biotope preferences, interactions with other organisms and many others can be generated in a similar way. All these statistics are generated from original data and do not present literature compilations, although reliable literature records may to a small extent contribute to the original data. In any case, data are tagged as to whether they originate from investigated specimens, from living observations or from literature records. Results Micropterix Hübner, 1825 Micropterix Hübner, 1825 b: 426. Type species: Tinea podevinella Hübner, 1813, by original designation. Micropteryx Zeller, 1839: 185 (unjustified emendation). Eriocephala Curtis, 1839: pl. 751. Type species: Phalaena (Tinea) calthella Linnaeus, 1761, by mono- typy. Microptericina Zagulajev, 1983: 113. Type species: Micropteryx amasiella Staudinger, 1880, by original designation. The genus Micropterix has been suggested to represent the sister group of all other mem- bers of the basalmost group of extant Lepidoptera (e.g. Kristensen 1984a; Kristensen & Nielsen 1979), the Zeugloptera [= Micropterigidae Herrich-Schaffer, 1855] notably on the basis of its complete lack of a hindwing vein R and retention of minute remnants of the eighth abdominal sternite in males (Kristensen 1998: 43; Hashimoto 2006: 58). However, the support for this placement was soon questioned (Kristensen & Nielsen 1982) and is now definitely known to be spurious (Skalski 1995; Davis et al., unpubl.), as is also strongly indicated by preliminary analyses of 16SrRNA molecular data (Kobayashi et al. 2002; Gibbs et al. 2004; Hashimoto 2006). The phylogenetic posi- tion of this genus within the family Micropterigidae is still quite unclear, but while it is subordinate within the hierarchy of extant taxa, it is at least clearly external to the group comprising the other Northern Hemisphere members. There are 11 currently recognized genera in the family (including those recently described by Hashimoto (2006), but dis- regarding undescribed genera of which a number will be needed to render Sabatinca Walker, 1863 monophyletic). Micropterix shares with most or all of these genera the presence of fully functional, asymmetrical mandibles (with apical incisor cusps only on the left mandible), and presence of antennal ascoid type sensilla (see Kristensen, 1984a: 154 for a list of another eight characters synapomorphic for Micropterigidae). Movable mandibles are not, however, unique to Micropterigidae, also occurring in the primitive genera Agathiphaga and Heterobathmia. The use of mandibles for crushing pollen or fern spores, combined with the presence of filtering structures in the mouth cavity, ap- pears thus to have have been lost independently in ancestors of Agathiphaga (where the mandibles are functional only in the pharate adult) and Glossata (Kristensen & Skalski 1998). There is little doubt also that all the taxa currently placed within Micropterix 240 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe constitute a natural group (excluding for now fossil taxa, notably Micropterix anglica Jarzembowski, 1980, whose fragmentary forewing venation but not probable palaeoen- vironment is consistent with extant members of this genus, or any Baltic amber mate- rial that require reassessment). The genus Micropterix has at least three derived traits: (1) in the wing venation all Rs branches run to the costa; (2) the male tergum I has a special muscle set inserting on an anteromedial longitudinal costa; and (3) the male abdominal segment IX has special- ized dorsomedian and dorsolateral processes (Kristensen & Nielsen 1979). Also, more weakly (4) forewing R is unforked, but this trait occurs in other Northern hemisphere genera; (5) the dorsal part of the female abdominal sternite IX ring is unsclerotized (Kristensen & Nielsen 1982: 514); (6) Micropterix has lost the basal trait attributed to the Amphiesmenoptera of sternal V glands, although this loss has also happened elsewhere among the Micropterigidae (Kristensen 1984: 128) and (7) apyrene type of sperm is absent (so far checked in just a single species, Kristensen 1999). Diagnosis. A number of features are plesiomorphies widespread in other homoneurous moths: head rough scaled, densely covered; ocelli prominent (as in all extant microp- terigid genera); maxillary palpi long, with 5 segments, folded (as e.g. in eriocraniids). Other characters are more or less characteristic of Micropterigidae: full complement of sulci present on head; lacinia present, cardo irregular, longish; labial palpi shortened (2-segmented in Micropterix as in most micropterigids), with a sensory concavity (Von Rath’s organ) on the basal segment; antennal sensillae (including branched ‘ascoid’ basiconic type as also in Opostegidae) on all segments except the two basalmost ones and the distalmost one; mandibles present and fully functional in both pupa and adult; fore tibia with epiphysis; spur formula 0-0-2 as in other micropterigids (absence of mesotibial spurs shared uniquely with Heterobathmiidae); forewing with jugum, hind- wing with a row of non-coupling subcostal frenulum bristles. The Micropterix species of northern and central Europe, considered in this paper, have a wingspan of 5—12 mm; antennae are filiform or sub-moniliform, from 1/2 to almost forewing length, wing colouration is metallic shining, with two principal colour patterns only: golden with or without silvery spots and fasciae, or more or less purplish with golden spots and fasciae (Heath 1983; Scoble 1995; Kristensen 1999). The radial vein is unbranched in forewing as in “Micropterix” anglica, “Sabatinca” porphyrodes, and most E. Asian micropterig- ids (Hashimoto 2006) and the abdominal sternal V glands are absent, inferred lost, as occasional in a few tropical taxa such as Hypomartyria (Kristensen 1984b). Morphology of adults. The exoskeletal and internal adult morphology of the spe- cies Micropterix calthella (Linnaeus, 1761) has been more or less intensively studied and readers are referred here to a few principal works. Very little is known of the detailed morphology of other species in the genus. For characteristics in wing vena- tion, see for example Hashimoto (2006). Internal thoracic musculature of Micropterix aureatella shikotanica Kozlov, 1988 was treated by Kozlov (1986), whilst Kristensen (1984b) treated in detail the abdominal musculature and chordotonal organs of M. calthella, whose head musculature is described by Hannemann (1956). Kristensen (1984c) also treated the respiratory system of M. calthella. Sensillae of antennae and palps were described by Le Cerf (1926). Mouthpart sensillal ultrastructure of Nota lepid. 30 (2): 235-298 24] accessory membraneous lobes with additional spikes or setae Phallus Uncus Tegumen x] hair-like structure Terminal papillae Anal tube Sternum IX Vestibulum Oviduct : Ductus Accessory SU spermatecae gland Bursa copulatrix accessory claspers with modified spikes Receptaculum seminis Figs 2-3. Characters of genitalia referred to in the descriptions. 2. ©. 3. © (according to Kristensen in litt.). M. calthella was described in more detail by Chauvin & Faucheux (1981). In this paper, we focus on wing pattern and comparative features of abdominal sclerites and genitalia of Palaearctic species. Adult Micropterix have a forewing length of about 1.5-7 mm. On the head they bear a vestiture of hair-like scales, which, in most cases, are more or less yellow coloured, but sometimes may be dark brown or even black. The antennae of males are significantly longer in males, reaching 2/3—7/8 of the forewing length, whereas in females their length is only 1/2-2/3 of the forewing length. The forewings show a pattern of golden and usu- ally, also purple to violet markings, which are often very species-specific. The hindwings are devoid of any markings, more or less bronzy golden with the apex tinged purple. Pregenital abdomen. The pregenital abdomen consists of eight well developed segments. In both genders, the sclerites of segments I and II are heavily modified in shape, but the intraspecific variation of these characters is too great to be used for spe- cies-identifications. Also not useable for that purpose are all other segments including segment VIII, although sternites IIT-VI, as well as tergites VI- VIII often bear heav- ily sclerotized cross-ridges, or, seldom, a reticulate structure. In males, sternite VIII is always reduced to two tiny sclerotized platelets. Male genitalia. Segments IX and X form the core of the genitalia, which are not retractable into the abdomen. In a few species like Micropterix mansuetella Zeller, 1844, the two complexes consisting of vinculum and valvae on the one hand and uncus, tegu- men and two pairs of accessory claspers on the other hand, are clearly separated from each other. In most species however, the two parts are fused. Besides uncus and valvae, which are common in most other Lepidoptera, two additional clasping structures can be developed on the posterior margin of the tegumen. In Micropterix aureatella (Scopoli, 242 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe 1763) and its relatives, both clasping structures (Fig. 2) are developed and bear normal, hair-like (hereafter “unmodified”) or bristle-like/stout (hereafter “spinoid”) setae (Fig. 123). In Micropterix aruncella (Scopoli, 1763) and its relatives the upper pair is devel- oped prominently, whereas the lower pair (called accessory claspers) is more or less atrophied. In most other species of Micropterix, the upper pair is reduced or completely obsolete, whereas the accessory claspers are well developed. The accessory claspers normally bear up to 3 groups of thickened setae (Fig. 124) which may be on the inner surface. Often these setae are highly modified into T- or Y- shapes (Fig. 125). The valvae also bear spinoid setae, but these are not modified. The phallus, describe in detail by Hannemann 1957, is somewhat enlarged at its base and shows a delicate inner structure at the posterior end. Nevertheless, these structures are not informative at species level. Female genitalia. Segments IX and X are normally retracted into the abdo- men in order to form a short ovipositor. Tergite IX is obsolete with the exception of M. mansuetella, where a minute fragment is left. In all other cases so far investigated only a group of setae is left instead of the tergite. Sternite IX is also reduced and modified in shape in females and, despite a reasonable intraspecific variability, often of shape in- formative at species-level (for example this more or less strongly sclerotized plate may show concavities at its proximal margin, or form a complete ring fused at the dorsum). Two lateral setose sclerites constituting the discrete segment X of Micropterigidae, form the terminal papillae, and form a more or less an evenly wide ring. In some cases, they also can contribute to species identification. The diagnostic features of segments IX and X are discussed in the species descriptions. The internal parts of the female genitalia are not sclerotized at all. Therefore most structures are of no value for recog- nisizing species. Only the receptaculum seminis is distinctly cross-striated and in many cases its shape and length can contribute to species identification (Fig. 3). Morphology of preimaginal stages (Heath 1983; Hashimoto 2006). Egg. The eggs are oval, about 0.38 x 0.24 mm to about 0.53 x 0.43 mm (Heath 1962), translucent white with rod like structures of variable length. The eggs are deposited between the vegetation on the soil. Larva. The body is elongate or barrel-shaped, whitish grey to dark grey, body length 2.5-4.5 mm; the head is almost twice as long as broad; the antennae are well developed, 3-segmented and bear a seta on their ends; they are almost as long as the width of the head; head and prothorax can be retracted into meso- and metathorax; the thoracic legs have only three free segments; the abdominal legs consist of a pair of conical append- ages, bearing claws similar to the thoracic legs; the anal sucker is trilobed; each seg- ment bears four longitudinal ridges with eight rows of paired, scale-like setae. The larvae have been assumed to feed on small pieces of rotten leaves or perhaps fungal hyphae although observations such as collections of larvae from tussocks of Dactylis glomerata (Poaceae) have been tenuous (Scoble 1995: 194), but M. calthella and M. aruncella have been recorded to eat Stellaria spp. (Caryophyllaceae) or other angiosperm tissue in captivity although they refused mosses or liverworts (Carter & Dugdale 1982). They have been found in the soil at a depth of 10 cm in stony woodland Nota lepid. 30 (2): 235-298 243 soil (Heath 1983) and those of M. aureatella have been found in a range of leaf litter where they could feed on fungal mycorrhizae (Carter & Dugdale 1982). There would appear to be only three instars (Heath 1983). Pupa. Body length is about 2.5mm; the mandibles usually conspicuous and func- tional (Mosher 1916); the head and thorax with long bifurcate setae; the first seven abdominal segments are movable and the appendages are not fused to the body (Scoble 1995); the pupa rests within a tough silken cocoon. There is a nice drawing of the pupa of M. calthella in Lorenz (1961). Life history. The adult moths feed on the pollen of many different plants (already detailed), and they cluster gregariously on flowers, especially on easily accessible blossoms, rather than searching for pollen grains on leaves as for example some syr- phid flies. It has been demonstrated that in some cases the pollination is successful (Schwartz-Tzachor et al., 2004): e.g. in Israel, Micropterix berytella de Joannis, 1886 and Micropterix elegans Stainton, 1867 are, besides species of thrips, bees and flies, the most important pollinators of Cyclamen persicum Miller. For Micropterix, blos- soms are also mating sites (Kozlov & Zvereva 2006). We have observed that copulation often lasts only a few seconds. No Micropterix are known to feed on fern spores as do some other micropterigid lineages in North America (Tuskes & Smith 1984) and in the Southern Hemisphere (e.g. ‘Sabatinca’ porphyrodes; Gibbs, pers. comm.), but pollen is also known as a food source for New Caledonian Sabatinca sensu stricto (Thien et al. 1985) and for some Australian micropterigids (Common 1990: 130). The biotope preferences of most species have been insufficiently studied so far, but a certain constancy of soil humidity seems to be necessary for the development of the larvae, which are apparently usually detritus-living or subterranean. Therefore, the ge- nus inhabits biotopes with higher humidity like woods and scrubland, but also moors and moist meadows, especially places where the soil is moist, whereas only few spe- cies are able to live in dryer regions. Furthermore, some correlation between the col- ouration of the forewing and the biotope seems to exist. Species, inhabiting wood- and bushland, i.e. places with less solar insolation, are primarily purplish to violet with golden markings, whereas species living in open habitats are more reddish golden to golden coloured without darker wing markings. Due to the higher reflection rate, such golden animals are less susceptible to overheating in direct sunlight. It is also suggested (Kozlov 1985) that visual cues are important in the absence of sternal V glands and lack of apparent pheromonal communication (despite suggestion to this effect by Pringruber (1944)) and so this could explain the bright shiny scale colours that could be the cues for conspecific aggregation irrespective of sex. The adults of Micropterix are active in sunshine, although sometimes they may come to light. Depending on latitude and elevation, they swarm during a period of two or three weeks in a single generation any time from late January (southern Mediterranean area, e.g. Israel) to August (highest sites in the Alps). The actual flight period only lasts a few weeks in any locality, but the individual lifespan is poorly known. According to an experiment to prove the pollination of Cyclamen persicum Miller in Israel, M. elegans and M. berytella were kept in captivity for four or five days before they died (Schwartz- Tzachor, pers. comm. ). 244 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Micropterix is one of two genera of Micropterigidae (Gibbs, unpublished) with a larva that typically lives in the leaf litter zone where it may even feed on fungal hyphae as well as a probably wide range of plant material that includes living angiosperm leaf tissue and are the only micropterigids known to do so (Lorenz 1961; Carter & Dugdale 1982). Despite the diversity of species in Europe, it is remarkable that almost nothing still is known about the early stage life history, and apart from descriptions of eggs laid by females, there is knowledge only about the larva and pupa of M. aruncella and M. calthella (and larva of M. aureatella), for which remarks below apply. Phylogenetic relationships. Up to now no consensus hypothesis has emerged as to the phylogenetic relationship of species within the genus Micropterix. The authors have identified several morphological characters, which are assumed to be autapomorphies of recognized species groups (unpublished results). This has led to an ordered arrange- ment of species that has been used in this paper, although for the time being this ar- rangement must be regarded as provisional and partly phenetic. Further studies, includ- ing DNA investigations performed by the authors will hopefully improve the presented picture. Key to the species The key is based on characters observable without dissection, as far as possible. la Forewings uniformly golden (Fig. 4), or with only minute reddish to Violet markings (Fig: 5, 8) desert EEE 2 lb: Forewingsditierent (Pigs 6, 7) sccc.chsccscsvensscssansanesa\oxqoaaeacs de sensssannsescnisvasessaaact REIN 6 2a Hair-like scales of head black, forwing markings see Figs 6, 8 nee M. mansuetella 2b Hair-like’scales of head yellow to dark brown .........44mas RSR 3 3a Base of forewing purple from costa to inner margin (Fig. 5)... M. calthella 3b _ Only base of costa slightly purple; three taxa distinguishable only by genitalia examination (Fig) 4) u.a asus none hen nenne she anna anne EEE 4 Figs 4-8. Illustrations for the “Key to the species”: Schemes of forewing drawing, © genitalia and 9 abdomen of Micropterix. 4. M. isobasella. 5. M. calthella. 6. M. mansuetella. 7. M. osthelderi. 8. M. man- suetella. Nota lepid. 30 (2): 235-298 245 4a 4b 5a 5b 6a 6b ma 7b male genitalia: membranous lobe between accessory claspers and uncus prominent (Fig. 9); female genitalia: sternite IX much reduced, almost interrupted in the middle (Fig. 10); males without silvery Markınos; KO ME LE OMS ardinia sac care ttes dues M. aruncella HAE femal ese rit ES RS ARE EE ine ieeeas 5 male genitalia: segments IX (tegumen) and X (uncus, accessory claspers) completely fused, acces- sory claspers with sickle-shaped spines (Fig. 11); female genitalia: not distinguishable from M. isobasella with certainty (Fig. 12); normally the species is deep purple violet, a golden form without markings is known only from high altitudes in the south western Alps (Marguareis), flying together with more or less marked specimens ..............nnnennnnen M. paykullella male genitalia: segments IX and X distinctly separated from each other (Fig. 13); female genitalia: not distinguishable from M. paykullella (Fig. 14); distribution confined to southern Switzerland (Simplon) and adjacent regions of northern Italy at high altitudes ...................... M. isobasella Forewings golden with extended purple markings, often more or less diffuse ÉTÉ SOS a eisernen 7 Forewings reddish, coppery (Fig. 16), purple or deep violet with silvery or golden DATE SORT) PA D cutie esate 9 Hair-like scales of head black, forewing markings see Figs 6, 8 ....... nee M. mansuetella Hair-like scales of head yellow to brownish insert erccrtrs serres 8 Figs 9-16. 9-10. M. aruncella. 11-12. M. paykullella. 13-14. M. isobasella. 15. M. tunbergella. 16. M. aruncella. 246 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe 8a 8b 9a 9b 10a 10b Base of forewing purple from costa to inner margin, other markings more or less dimmise. (Ki. 17) nn RS ae ee eee M. calthella Base of forewing not extensive purple; purple fascia in the centre of the wing distinctly forked at costa, embedding a small golden costal spot (Fig. 15)... M. tunbergella Forewings bronzy golden or coppery with more or less distinct silvery markings (Figs 16, 21); two taxa distinguishable in the male sex only by genitalia examination ...........euesennnnnennnnnn 10 Forewings reddish, brownish golden, purple to purple violet with more or less distinct light golden to bronzy golden wing markings (Figs. 7, 18) 24 RP 11 Males: lobes between uncus and accessory claspers prominent, accessory claspers spatulate (Fig. 9); females: without silvery wing markings (Fig: IN mr... M. aruncella Males: lobes between uncus and accessory claspers missing; accessory claspers very prominent (Fig. 20, after Heath & Kaltenbach 1984); Females: silvery wing markings distinct; distribution confined to Alpi Cozic (Figs 21) a as ee eee M. fenestrellensis Small species: wing expanse less than 6.3 mm... ts. Sen 12 Larser species una u ei ee ee a ee oe 13 Forewings with three broad transverse fasciae reaching from costa to inner margin (PISS: 22223): nase ee ee ee ee EA EIER M. rablensis Forewings with two broad transverse fasciae reaching from costa to inner margin and an outer prominent costal spot, reaching only to the middle of the wing (Fig. 18) ........... M. myrtetella Forewings with three transverse fasciae reaching from costa to inner margin; a small costal spot may be present (Figs-24, 25) ca... een a a ee 14 Forewings with additional or other wing markings (Fig. 7) 540 17 Outer transverse fascia triangular and. extremely broad (Fig. 24) 5... ee 15 Figs 17-24. 17. M. calthella. 18. M. myrtetella. 19. M. aruncella 9. 20-21. M. fenestrellensis (S genitalia after Heat & Kaltenbach 1984). 22-23. M. rablensis. 24. M. trifasciella. Nota lepid. 30 (2): 235-298 247 14b 15a 15b 16a 16b 17a Outer transverse fascia narrower, sometimes not completely reaching costa or IDE MAIRIE rar aneerelnen c ide esemsne amer mnmiscs 16 Small costal spot normally absent (Fig. 24); male genitalia: accessory claspers slightly shorter and broader; spines beyond uncus normally present (Fig. 26); female genitalia: not distinguishable from M. allionella; distribution confined to a small region in the French-Italian border area SR eh RN REEL aida Uh a cad Ce rach Serena ART NERRERS EEE TEERSUEELEERNNN M. trifasciella Small costal spot normally present, often remnants of a bronzy golden outer margin also present (Fig. 27); male genitalia: accessory claspers slightly longer and smaller (Fig. 28); female geni- talia: not distinguishable from M. trifasciella ....ccccccceccccccssccceessectseseeeeesseeeessseees M. allionella Outer transverse fascia always reaching costa and inner margin (Fig. 25); male genitalia: lobes between uncus and accessory claspers completely atrophied (Fig. 29); female genitalia: un- known; distribution confined to high altitudes in the south western Alps (only known from type OC A Mlarpuage kc) I eat tee ee neta ee M. huemeri Outer transverse fascia normally not reaching costa and inner margin (Fig. 30); male genitalia: lobes between uncus and accessory claspers distinctly developed (Fig. 31); female genitalia: receptaculum seminisimoderately long)... M. aureatella Posterior margin not bronzy golden (Fig. 32), or if bronzy golden, this colouration not extending to SC ee covers ne ei te ee 18 Figs 25-33. 25. M. huemeri. 26. M. trifasciella. 27-28. M. allionella. 29. M. huemeri. 30-32. M. aure- atella. 33. M. aglaella. 248 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe 17b 18a 18b 19a 19b 20a 20b 2la 21b 22a 22b 23a 23b Posterior margin more or less distinctly bronzy golden from base to fascia at 1/2 (Fig. 7) ......... 22 Forewings markings: Two transverse fasciae at 1/4 and 1/2 and an outer spot, often almost reaching costa and posterior margin.(see also 13).(Eig, 32)... een etecnee nace eee M. aureatella Additional markings developed (Figs 33, 27) 44.44... ak en eee 19 Posterior margin without any bronzy golden area; outer transverse fascia very broad and triangular, extending across the whole wing width (Pig: 27) 4,22... ER 20 Posterior margin with bronzy golden area (Fig. 33), sometimes almost reaching fascia at 1/2 (Fig. 34); outer transverse fascia narrower, often not extending across the whole wing width and normally not triangular tm te 21 Transverse fasciae extremely broad, sometimes fused with costal spot; outer bronzy golden margin well developed (Fig. 35); male genitalia: accessory claspers broad triangular, proximal spines not on a separate stylus (Fig. 36); female genitalia: not distinguishable from M. allionella MINEN EEE REBLEBELS RUE EEE ANEEINERERCRUR nants IDEEN RER SREEE RIESTER causa LEE 0005501000 M. rothenbachii Transverse fasciae at 1/4 and 1/2 smaller; outer bronzy golden margin often reduced (especially males) (Fig. 27); male genitalia: accessory claspers not broad triangular, two proximal spines on a separate stylus (Fig. 28); female genitalia: not distinguishable from M. rothenbachii, D: c-ooooococ M. allionella Bronzy golden wing base almost extending to fascia at 1/2 at posterior margin; outer margin of the bronzy golden area and fascia at 1/2 therefore not parallel (Fig. 34) ................. M. paykullella Bronzy golden wing base not extending towards fascia at 1/2; outer margin of the bronzy golden area and fascia at 1/2 therefore parallel (Fig. 33) a... udn mn M. aglaella Ground colour often more or less reddish, light golden markings often indistinct; small costal spot HUSSNe (PIE, 37) re see M. aureoviridella Ground colour distinctly purple to bluish violet; markings distinctly golden; small costal spot always. present) OR sen een ee SR Oe eee 23 Inner transverse fascia light golden in its centre above bronzy golden inner margin (FGFS) ren uen Rd die RONDE M. schaefferi Inner transverse fascia and inner margin completely bronzy golden (Fig. 7) ............ M. osthelderi Figs 34-38. 34. M. paykullella. 35-36. M. rothenbachii. 37. M. aureoviridella. 38. M. schaefferi. Nota lepid. 30 (2): 235-298 249 Checklist of northern and central European species Micropterix mansuetella Zeller, 1844 = Lampronia ammanella (Hübner, 1813) sensu Wood, 1839 Micropterix calthella (Linnaeus, 1761) = Tinea urticella Costa, 1834 = Eriocephala sulcatella Bentley, 1845 = Micropteryx silesiaca Toll, 1942 Micropterix isobasella Staudinger, 1871 Micropterix aglaella (Duponchel, 1838) Micropterix aureatella (Scopoli, 1763) = Tinea paykullella Thunberg, 1794 = Tinea ammanella Hübner, 1813 Micropterix aruncella (Scopoli, 1763) = Tinea seppella Fabricius, 1777 = Tinea podevinella Hübner, 1813 = Lampronia concinnella Stephens, 1834 = Micropteryx eximiella Zeller, 1850 = Eriocephala atricapilla Wocke, 1877 = Micropteryx nuraghella Amsel, 1936 Micropterix tunbergella (Fabricius, 1787) = Tinea helwigella Hübner, 1805 = Tinea rubrifasciella Haworth, 1828 = Micropteryx depictella Herrich-Schäffer, 1851 Micropterix aureoviridella (Höfner, 1898) = Micropterix liogierella Real, 1987 Micropterix paykullella (Fabricius, 1794) = Tinea anderschella Hübner, 1813 = Micropterix paykulella [sıc!] f. rosarum Müller-Rutz, 1927 Micropterix allionella (Fabricius, 1794) = Tinea tricinctella Costa, 1836 Micropterix trifasciella Heath, 1965 Micropterix rothenbachi (Frey, 1856) = Micropterix australis Heath, 1981a = Micropterix vallebonnella Réal, 1988 Micropterix huemeri Kurz, Kurz & Zeller, 2004 Micropterix schaefferi Heath, 1975 = Micropteryx anderschella (Hübner, 1813) sensu Herrich-Schäffer, 1851 Micropterix osthelderi Heath, 1975 Micropterix fenestrellensis Heath & Kaltenbach, 1984 Micropterix rablensis Zeller, 1868 Micropterix myrtetella Zeller, 1850 250 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Micropterix mansuetella Zeller, 1844 Micropterix mansuetella Zeller, 1844: 16. Type locality: “Groß-Glogau” (now: Glogéw, Poland). Lectotype: in coll. BMNH. Lampronia ammanella (Hübner, 1813) sensu Wood, 1839: 231, pl. 50. Misidentification. Description of adults. Examined: 90°, 109. Forewing length: © 3.4-3.9 mm; 9 3.8- 4.2 mm. Head black; vestiture of hair-like scales on the head black-brown to black, golden fuscous; antennae 4/5 (©), respectively, 1/2 (9) of forewing length, dark brown at base, distal parts lighter brown, golden fuscous; thorax dark golden, prothorax poste- riorly with purple and bluish scales; tegulae purple, posteriorly edged bluish; forewing golden with bronzy golden to reddish golden pattern, especially in the southern part of the species’ geographic range the reddish golden markings often only faintly developed or, in males, almost oblique: a purple costal spot near the base; a broad, diffuse fascia at 1/3, narrowing towards inner margin, sometimes disrupted near the costa, sometimes purplish at the anterior margin and in the middle; a very broad, diffuse fascia at 3/4, purplish at costa, usually hardly distinguished from the bronzy golden apical part; cilia brightly golden; hindwings bronzy golden, especially at apex with purple tinge; cilia brightly bronzy golden with faint purple tinge; legs dark golden fuscous; abdomen golden brown. SO Genitalia. Uncus moderately long, broad, stout, distal with some short setae, forming a separate, clearly distinguishable unit with accessory claspers; accessory clas- pers moderately long, enlarged distally, anterior margin almost S-shaped, with short, stout thickened setae, which have spatulate bases and short, hook-like ends, pointing backwards; a row of shorter, unmodified setae near anterior margin; valvae rather short, stout, strongly narrowed near mid-length, with clearly distinguished ends; a small, ba- sal group of short setae at inner surface, postbasally a larger, elongated group of short setae; several rows of very short to moderately long, spinoid setae at inner surface of distal ends. 9 Genitalia. Sclerites of segment IX reduced, forming a complete sclerotized ring: ventral half of normal width, ventrolaterally the proximal part lobe-shaped and en- larged; dorsal third of normal width, laterally slightly constricted; distal margin of ring with straight, sharp border, proximal margin of ring dorsolaterally irregular, faintly sclerotized. Terminal papillae strongly sclerotized, distal margin straight, proximal margin irregular; ductus receptaculi short, sharply bent, very narrow at onset of re- ceptaculum seminis; onset of receptaculum seminis slightly enlarged, upper third con- stricted and slightly bent; afterwards forming a curved and slender sac; lower end with characteristic, semicircular appendix. Diagnosis. The species might be confused with worn specimens of M. tunbergella or with M. calthella. It differs, however, from all European species by the blackish scales on the head. Only M. aruncella from alpine locations has comparable dark scales. M. aruncella usually inhabits different biotopes (nutrient-poor meadows versus swamp- land) and is found at higher elevations (although sometimes syntopic). The wing col- our is greener in M. aruncella, having a distinct purple base at the costa. Adults are Nota lepid. 30 (2): 235-298 251 also usually smaller than M. mansuetella, with hair-like scales on the head less dark. Distribution. According to Heath (1996) and Karsholt (2004) this species occurs in northern, eastern, central and western Europe (including Great Britain and Ireland). Heath (1983) reports this species local in northern Europe as far south as Bavaria (Germany). Meyrick (1912) depicts Europe, without further specification. Meeß (1910) reports the species from central, western (including England) and northern Europe. The records from Italy (Heath 1996; Hartig 1964) respectively seem to be doubtful and those from Portugal (Meeß 1910; Heath 1996) belong to the recently described Micropterix herminiella Corley, 2007. The investigated specimens from the collection of Klimesch (now ZSM) are from the Austrian alpine regions: Edlbach-Moor (Upper Austrian) and Selzthal-Moor (Styria). Deutsch (pers. comm.) also collected the species ın Eastern Tyrol (Austria), which is as far as we know the most southern occurrence of this species. Life history. We collected specimens from flowering sedges (Carex ssp.) in open wood- land (Fraxinus-Salix-association) at the border of fens. Klimesch (pers. comm.) found the species favoured swamplands, feeding on sedges ın birch-groves. Deutsch (pers. comm.) reports its occurrence in a ditch with Alnus incana L. next to a small woody moorland slope. The species seems to be absent from higher mountain regions. Preimaginal stages. According to Heath (1962, 1983), the egg has an oval shape, a length of 360-490 um and a width of 260-330 um, with up to 140 um long, rod-like structures. Immediately before hatching, the translucent white colour turns to grey. The larva and pupa are unknown. Micropterix calthella (Linnaeus, 1761) Phalaena (Tinea) calthella Linnaeus, 1761: 367. Type locality: Sweden. Lectotype: in coll. LSUK. Tinea urticella Costa, 1834: 10-11, pl. 2 figs la—c. Type locality: Camaldoli (Capodimonte, Napoli, Italy). Junior subjective synonym. Eriocephala sulcatella Bentley, 1845: 1086-1087. Type locality: London (Great Britain). Junior subjective synonym. Micropteryx silesiaca Toll, 1942: 171. Type locality: Ustron (Poland). Junior subjective synonym. Description of adults. Examined: 130°, 439. Forewing length: © 3.1-3.7 mm; 9 3.6- 4.6 mm. Head black-brown; vestiture of hair-like scales on the head dirty white to rusty yellow; antennae brown, golden shining, with reddish tinge, 4/5 (©), respectively, nearly 3/5 (Q) of forewing length; thorax bronzy golden to coppery, posteriorly some- times purple mixed; tegulae coppery to bluish violet; forewing bronzy golden, brown- ish golden to slightly greenish golden, usually without any markings; the basal area from costa to inner margin, more or less extended, purple to purplish violet, sometimes with single bluish scales; ground colour occasionally intensely tinged purple or with rudiments of vague purple markings present; these consist of a costal spot near 1/4 and a similar spot at 2/3, the latter extending almost across the whole width of the wing; fringe light bronzy golden to whitish golden, sometimes with a purple tinge; hindwing light bronzy golden, sometimes more or less purplish at the apex; fringe light bronzy 252 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe golden, brighter at the distal margin, sometimes with a purple tinge; legs and abdomen brown, golden shining, sometimes slightly purplish. A great deal is known about the internal anatomy and exoskeletal ultrastructure of the adult of this species (see section: Morphology of adults). Below we describe the broad morphology of the abdomen only. OS Genitalia. Uncus moderately long and slender; ventrally beyond the uncus a tuft of long hair-like setae; accessory claspers well developed, nearly trapezoid; their rounded anterior margin with a row of seven or eight very short, thickened setae of spoon-like to spatulate shape; at the lower apex of the anterior margin a row of three similar but longer, straight thickened setae, which are partly hook-shaped at their ends; valvae moderately long, distinctly constricted medially, the distal fourth spatulately enlarged, bent upwards; at their inner surface postbasally two or three shorter, straight spinoid setae; the distal fourth at the inner surface with two to four irregular rows of moderately long to very short, straight spinoid setae. Q Genitalia. Tergite IX missing; sternite IX much reduced, moderately sclero- tized, sometimes with indistinct margins, but of relatively characteristic shape. Segment X (terminal papillae) with sclerotized band; ductus receptaculi very thin; receptaculum seminis long and slender, at the beginning slightly thickened, the first half straight, very slender, without transverse striation; the second half enlarged into a curved, elongate sac with somewhat irregular transverse striation; a thin appendix at the end. Diagnosis. M. calthella may be confused with other unicolorous golden species with- out markings like M. isobasella., unmarked forms of Micropterix sicanella Zeller, 1847 and M. paykullella, Micropterix garganoensis Heath, 1960 and the female of M. arun- cella. From these, M. calthella is separated mostly by its darker (more greenish or brownish) bronzy golden ground colouration, as well as by its distinct purple coloura- tion of the forewing base, reaching from costa to the inner margin. M. mansuetella with indistinct markings also may be very similar, but is readily distinguished by its black scalés on the head. In the male genitalia, M. calthella is well characterized. Distribution. According to Heath (1983) and Karsholt (2004), the species is distrib- uted across Europe to central Siberia. M. calthella has not been recorded from Iceland, the Iberian Peninsula, southern Balkans and from the Mediterranean islands (Heath 1996). Life history. M. calthella inhabits moist but not saturated locations such as marshy areas, forest tracks and outskirts of the forest. The adults feed on easily accessible pollen of a wide variety of different herbaceous plants (such as Ajuga, Cardamine, Mercurialis) and even occasionally trees (Crataegus, Acer), but they strongly favour sedges (Carex spp.), kingcup (Caltha palustris) and buttercups (Ranunculus Spp.). Preimaginal stages. According to Heath (1983), the egg has an oval shape, a length of 450-480 um and a width of 350-370 um, with up to 60 um long, rod-like structures. Immediately before hatching, the translucent white colour is turns to grey. Chauvin & Chauvin (1980) also describe the egg. The larva (see general description above, under Life history) is described by Martinova (1950) and Hamon & Chauvin (1995). Nota lepid. 30 (2): 235-298 253 The pupa rests in a compact cocoon. More about the pupa can be found in Lorenz (1961) and Hamon & Chauvin (1995). Remarks. We found this species near Sıena (Tuscany, Italy), which is as far as we know the most southern occurrence of this species. The records from Napoli (Campania, Italy) (for Tinea urticella Costa, 1834) should be verified. Micropterix isobasella Staudinger, 1871 Micropteryx isobasella Staudinger, 1871: 289. Type locality: Italy, western Alps, Macugnaga. 6 syntypes in coll. ZMHB. M. isobasella f. weberi Müller-Rutz, 1927; infrasubspecific. Description. Examined: 20°, 59. Forewing length: © 3.5-3.8 mm; 9 3.0-4.4 mm. Head black-brown; vestiture of hair-like scales on the head rusty yellow; antennae dark brown, coppery shining, slightly more than 3/4 (0°), respectively, slightly more than 1/2 (Q) of forewing length; thorax bronzy golden; tegulae bronzy golden, posteriorly with single purple scales; forewing golden with a greenish tinge; costal margin purplish up to 3/4; almost invisible bronzy golden transverse fasciae or spots at 1/3 and 2/3, seldom of diffuse reddish colour (f. weberi Müller-Rutz); base slightly darker, the darker col- ouration extending from costa across 1/3 of forewing width; fringe light golden, with a purple tinge, especially at its base; hindwing bronzy golden, fringe golden, both with a purple tinge; legs and abdomen brown, golden shining. CO Genitalia. Uncus moderately long, somewhat stout, with a broad tip; a paired association of hair-like setae ventrally beyond the uncus; accessory claspers moder- ately long, nearly keel-shaped, with about 11-12 elongate, straight, unmodified setae at the rounded distal margin; another irregular row of five or six shorter, finer setae, which are more inwardly located (shape not characterised as mostly folded in the per- manent preparation); an area of long, straight spinoid setae dorsal of the accessory claspers; valvae moderately long, stout, distal third enlarged and strongly bent upwards (constricted at the point of inflection); on the inner surface of the valvae three shorter setae postbasally; on the inner surface of the lower margin two to three irregular rows of shorter spinoid setae and some longer setae on the distal fourth. 9 Genitalia. Tergite IX missing; sternite IX reduced, constricted medially, with strongly fringed lateral margins, without diagnostic features. Terminal papillae with two sclerotized plates forming an undiagnostic band; ductus receptaculi thin and strongly bent, at the beginning of the receptaculum seminis short and straight; receptaculum seminis long and slender, constricted in the first third, with typical transverse striation. Diagnosis. M. isobasella can be separated from M. calthella and from females of M. aruncella by its almost complete lack of purple colouration at the forewing base. M. calthella has an extended purple basal area, whereas females of M. aruncella have only the base of the costa purplish coloured (males of M. aruncella are characterized by their silvery fasciae). M. isobasella has only a slight purple colouration along the costa. In the highly mountainous southwestern Alps, a form of M. paykullella has been found, more or less without any markings. This form can be distinguished from M. isobasella 254 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe only by examination of the male genitalia. Also very similar and therefore externally not distinguishable is a form without markings of Micropterix sicanella Zeller, 1847 (f. obsoleta Heath, 1963). All above-mentioned species can be easily identified by ex- amination of the male genitalia. In the female genitalia, only M. aruncella can be rec- ognized with some certainty by its reduced sternite IX and by the somewhat protruding terminal papillae. Females of the other three species cannot be identified with certainty due to their intraspecific variability. Further species without wing markings do not occur in the distribution range of M. isobasella. These are restricted to central Italy and northern Africa. Distribution. According to Heath (1996) this species is restricted to Italy and Switzerland, where it seems to be distributed only in southern Switzerland and the adjacent northern Italy (MeeB 1910; Meyrick 1912). Records from Sicily probably belong to Micropterix sicanella Zeller, 1847 (f. obsoleta Heath, 1963), records from North Africa to Micropterix constantinella Heath, 1986 or to Micropterix eatoniella Heath, 1986. The examined specimens were found in Switzerland (area of Simplon). Life history. Two of the examined specimens were found swarming around Lonicera sp.. Further records are from tall herbaceous vegetation in a mixed larch-pine forest at 1650-1800 m elevation. Preimaginal stages. The early stages are unknown. Remarks. M. isobasella f. weberi Müller-Rutz, 1927 from Switzerland (Laquintal) has two slight reddish cross-fasciae on the forewing. Micropterix aglaella (Duponchel, 1840) Adela aglaella Duponchel, 1840: 627, pl. 312 fig. 14. Type locality: Central and Southern France (includ- ing ‘Fonscolombe’, north of Aix-en-Provence). Syntype in coll. MNHN (Minet in litt.). Description. Examined: 5, 129. Forewing length: © 3.3-3.6 mm; 9 3.5—4.3 mm. Head black brown, vestiture of hair-like scales on the head white to rusty yellow; an- tennae dark brown, golden shining with a purple tinge, nearly 4/5 (S'), respectively, nearly 3/5 (9) of forewing length; thorax bronzy golden, posteriorly reddish to purple, tegulae coppery to purple violet; ground colour of forewing reddish golden to purple violet, distal half sometimes purplish brown, outer margin sometimes reddish golden again, apex rarely also of this colour; a bronzy golden colouration from the base to 1/4, leaving a purple violet basal spot at costa; markings light golden to golden, delicately bordered in bronzy gold: a broad fascia at 1/2, slightly bent outwards, extending across the whole width of the forewing; sometimes a small costal spot at 3/5 (found in 7 of 17 specimens); a larger, almost round to slightly oval spot at 3/4, extending from costa across more than half, sometime even across whole forewing width (in the latter case the posterior part of this fascia bronzy golden); fringe golden, basally purple coloured, outwards whitish; hindwing bronzy golden, with an intense purple tinge; fringe bronzy golden, outwards whitish; legs and abdomen brown, golden shining. CO Genitalia. Uncus moderately long, stout, with a broad, rounded tip; beyond the uncus a weak structure of hair-like setae; between uncus and accessory claspers are Nota lepid. 30 (2): 235-298 235 situated weakly sclerotized, elongated, spatulate-like lobes, somewhat variable in the length, at the anterior margin of the tegumen; these lobes with very long hair-like setae at their ends, as well as on a small appendix at their lower margin; accessory claspers spoon-like, with a row of nearly 13 moderately long to long, mostly sickle-shaped thickened setae; near the dorsal margin anteriorly two shorter, straight spinoid setae anterior and basally a row of about 6 strongly modified, very broad T-shaped thick- ened setae; valvae moderately long, stout, strongly constricted medially; at their inner margin a very long and a shorter seta basally, on the distal part a group of very short to rather long spinoid setae, clustered proximally towards the constriction; a row of short spinoid setae along the rounded anterior margin. Q Genitalia. Tergite IX missing, only indicated by a group of setae; sternite IX strongly reduced, weakly sclerotized, constricted medially. Terminal papillae consist- ing of two somewhat weakly sclerotized plates forming a band; receptaculum seminis more or less short and stout, the second half like a sac, with typical striation; vestibu- lum a large sac, without any special characters. Diagnosis. This species can be confused with M. paykullella, but in most cases it can be recognized by its less intense purple colouration. In contrast to M. paykullella and M. aureoviridella, the golden inner margin of M. aglaella does not reach the fascia in the middle (the border of the bronzy golden inner part of the wing and the fascia in the mid- dle are often nearly parallel). The outer spot at 3/4 extends across the entire width of the forewing in many cases. M. aglaella can also be separated from M. aureoviridella by the normally more acute shape of its markings, the darker purple colouration and by its consistently bronzy golden forewing base. The male genitalia resemble somewhat those of M. aureatella, but can be easily dis- tinguished. Also the female genitalia can be recognized quite well. In particular, the degree of sclerotization of sternite IX and of the terminal papillae of M. aglaella is distinctly weaker than that of M. paykullella. The receptaculum seminis seems to be shorter and stouter, but these differences are too minor to be useful Distribution. According to Heath (1996) and Karsholt (2004) this species occurs in Spain, France, Italy, Germany and Switzerland. The records from Portugal (Heath 1996; Karsholt 2004) seem to be doubtful (Corley 2007). To our present knowledge, M. aglaella is distributed in southern France, western Switzerland, the southern Alps (eastwards as far as Mt. Baldo, Italy), as well as in the Pyrenees. The record for the Schwäbische Alb (Germany) (Pröse 1987) has also been confirmed. Life history. We have no modern information so are unable to confirm Duponchel’s (1840: 628) records of his species from flowers of ‘troéne’ (=Ligustrum vulgare), ‘sureau’ (=Sambucus) and ‘cornouiller sanguin’ (=Cornus sanguinea). Preimaginal stages. The early stages are unknown. Remarks. According to Minet (in litt.) the year of description has to be corrected to 1840 (all types are labelled as Micropterix aglaella Duponchel, 1840 instead of 1838). 256 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Micropterix aureatella (Scopoli, 1763) Phalaena aureatella Scopoli, 1763: 254. Type locality: Slovenia, Carniola. Type: Lost or destroyed in coll. IEUP (Evenhuis 1997). Tinea paykullella Thunberg, 1794: 89. Type locality: “Vestrogothia” (Sweden). Homonym of Alucita paykullella Fabricius, 1794. Tinea ammanella Hübner, 1813: pl. 57 fig. 388. Type locality: None given. Junior subjective synonym. Description. Examined: 190°, 329. Forewing length: © 3.9-4.6 mm; Q 4.2-4.8 mm. Head black-brown; vestiture of hair-like scales on the head brownish yellow to light ochre; antennae dark brownish golden with a light purple tinge, 4/5 (©), respectively, 4/7 (Q) of forewing length; thorax dark bronzy golden, posteriorly partly purplish, tegulae purplish violet, posteriorly sometimes with single bluish scales; forewing pur- plish violet, to some extent with single bluish scales, sometimes bronzy golden with a purplish violet tinge; markings golden, delicately bordered in bronzy gold and of vari- able width; an indistinct and minute bronzy golden spot at the base; a fascia near 1/4, slightly bent inwards and slightly narrowed at the costa or in the middle, sometimes not reaching the costa; a fascia at 1/2, equal in width, somewhat bent outwards and some- times narrowed in the middle; at 3/4 a broad, more or less oval spot of irregular shape, mostly reaching costa but not inner margin; fringe whitish golden; hindwing bronzy golden, more or less with a purplish tinge; fringe bronzy golden, distally whitish; legs light brownish golden; abdomen golden brown. SO Genitalia. Uncus short, somewhat stout; ventrally beyond the uncus two un- sclerotized small lobes bearing humps; between these and the accessory claspers two lobe-shaped appendices at the anterior margin with long hair-like setae, above them a row with shorter setae; accessory claspers long, narrow, spatulate, at the apex with a small dorsally oriented lobe with short spinoid setae; accessory clasper on the ventral margin with numerous, long, bent spinoid or sickle-shaped setae, which are ventrally oriented; valvae long, strong, constricted medially, with many setae postbasally; the last third bent upwards and distinctly enlarged with many spinoid setae (two clusters of spinoid setae, one at the point of inflection and one at the apex ). © Genitalia. Tergite IX missing, sternite IX reduced, distinctly constricted medi- ally, laterally enlarged into a lobe, strongly sclerotized, usually making the margins distinct. Terminal papillae with sclerotization in a band; receptaculum seminis long, narrow, at the beginning of the ductus receptaculi somewhat enlarged, forming a sac in the second half with typical striation; receptaculum seminis with a short appendix at the end. Diagnosis. M. aureatella can be separated from most other purple-violet and golden species of Micropterix by its lack of a golden inner margin of the forewing, the more regular golden fasciae as well as the lack of the small golden spot at the costa at about 3/5. M. trifasciella, also without this spot, usually has broader golden fasciae. In some cases M. aureatella and M. trifasciella can be separated only by examination of the gen- italia. The type locality of M. trifasciella is in the area of Torino (Fenestrelle, Val Susa, the border region of Italy and France). Also M. huemeri, found in the Alpes Maritimes (France), has a similar wing pattern. M. aureatella can also be separated with certainty by the male genitalia from the somewhat smaller species M. rablensis and M. croatica. Nota lepid. 30 (2): 235-298 257 Sometimes the costal spot of M. allionella can be missing and then it can be confused also with M. aureatella. Micropterix wockei Staudinger, 1870, occurring in Greece, also looks superficially very similar to M. aureatella. The structures of the female abdomen (segment IX and X) of M. aureatella sometimes look similar to those of M. aureoviridella, but sternite IX of M. aureatella is normally more strongly sclerotized with thus more distinct lateral margins. Distribution. According to Heath (1996) and Karsholt (2004), M. aureatella is dis- tributed throughout Europe except Spain, Iceland, Luxembourg, Albania, Bulgaria, the European part of Turkey and the Mediterranean islands. Also Heath (1983) records this species throughout the Palaearctic region, except northern Africa (Heath 1983). The records from Portugal (Heath 1996) seem to be doubtful (Corley 2007). The typical subspecies is replaced by ssp. shikotanica Kozlov in the eastern part of the Palaearctic region. This subspecies seems to differ clearly from the typical ssp. aure- atella and therefore may present a separate species (Kozlov 1988, 1989; Moriuti 1982). The presence of the species in Japan is remarkable considering its long geological separation; the two populations in Hokkaido and Honshu have slightly different wing pattern but have not been placed to any subspecies (Hashimoto, 2006). Life history. This species occurs especially in high moorland, where the moths have been found swarming around flowering Pinus mugo mugo Turra, Vaccinium myrtillus L. and also Carex spp. in full sunshine. In Hokkaido, Japan, the species is recorded on flowers identified as Heracleum lanatum Michx. var. lanatum (Hashimoto 2006: Fig. 11 O; U. Jinbo, pers. comm.). In mountain areas, M. aureatella inhabits mainly in elfin woodland in similar biotopes. This species can sometimes be found at the edges of forest openings and forest tracks, on more or less acid soils. Preimaginal stages. According to Heath (1962, 1983), the egg has an oval shape, a length of 470-530 um and a width of 370-430 um with rod-like structures. Immediately before hatching, the translucent white colour turns grey. The larvae have been found in the strongly mycorrhizal leaf litter of bilberry plants and of oak and beech woodland (Carter and Dugdale, 1982). The pupa is unknown. Remarks. The male genitalia figured by Viette (1948) as M. aureatella probably be- longs to M. trifasciella. Micropterix aruncella (Scopoli, 1763) Phalaena aruncella Scopoli, 1763: 254. Type locality: Slovenia, Carniola. Type: Lost or destroyed in coll. IEUP (Evenhuis 1997). Tinea seppella Fabricius, 1777: 296. Type locality: England (Karsholt in litt.). Junior subjective syno- nym. Tinea podevinella Hübner, 1813: pl.50, fig.342. Type locality: None given. Junior subjective synonym. Lampronia concinnella Stephens: 361. Type locality: Darenth wood (Great Britain). Junior subjective synonym. Micropteryx eximiella Zeller, 1850: 62. Type locality: Montenero (near Livorno, Tuscany, Italy). Junior subjective synonym. Eriocephala atricapilla Wocke, 1877: 52. Type locality: Stelvio (Italy). Junior subjective synonym. Micropteryx nuraghella Amsel, 1936: 364. Type locality: Tempio Pausanias (Sardinia, Italy). Junior sub- jective synonym. 258 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Description of adults. Examined: 90 ©, 71 9. Forewing length: © 2.6-3.6 mm; Q 2.6-4.0 mm. Head black-brown, vestiture of hair-like scales on the head dirty white to rusty yellow, sometimes brownish golden to blackish (f. atricapilla Wocke); anten- nae dark brown, reddish golden shining, nearly 4/5 (©), or somewhat over 1/2 (9) of forewing length; thorax bronzy golden to coppery, posteriorly often purple; tegulae purple to bluish; forewing golden to bronzy golden (seldom light golden), with more or less reddish tinge, which is sometimes missing; in specimens from the northern and central Apennines ground colouration often reddish bronzy golden to coppery; basal area mostly distinct purple at the costa, to some extent with single bluish scales; this purple colouration sometimes extending to 1/4 along the costa, sometimes nearly com- pletely missing; apex in most specimens slightly more reddish than ground colour; Q mostly without any other markings, but specimens from the northern and central Apennines similar to males; © with silvery white, often diffusely, but in specimens from the northern and central Apennines markings very distinct (markings missing in f. nuraghella Amsel): a narrow fascia near 1/4, not reaching costa but extending across more than half of the forewing width (strongly bent inwards and sometimes reduced to a small spot); a narrow, quite straight fascia, mostly extending across the whole width of the forewing at 1/2; sometimes a small, round spot in the anterior half of the wing at 3/4 (f. seppella Fabricius), distinctly pronounced in specimens from the northern and central Apennines; fringe light golden, mostly with a slight purplish tinge; hindwing golden to bronzy golden, more or less purplish, especially at the apex; fringe light golden with a reddish tinge; legs and abdomen brown, golden shining. SO Genitalia. Uncus long and narrow, somewhat enlarged at the tip; tegumen nar- row, ventrally with a long, broadly hatchet-shaped terminal appendix, weakly sclero- tized especially at the anterior margin; this hatchet-shaped appendix with some short setae; accessory claspers small, slipper-shaped, inside the above mentioned appendices and ventrally beyond their beginnings; at the dorsal margin of the inner surface of the distal end of the accessory claspers four or five short, slightly bent spinoid setae, which are multiply split at their ends; mostly at the lower margin 10 long, straight spinoid setae, bent at their ends; in the middle of the accessory claspers some short setae; valvae stout, beyond the middle strongly constricted, the distal ends spoon-like and slightly bent dorsad; on their inner surface an elongated patch of setae postbasally, a row of longer setae at their distal end and two or three irregular rows of shorter, straight spinoid setae. © Genitalia. Tergite IX missing; sternite IX only moderately sclerotized, reduced to two nearly halfmoon-shaped sclerotized plates with a narrow, weaker sclerotized junction. This sternite IX, the receptaculum seminis and also the terminal papillae are more or less characteristic for this species, although sternite IX is not so distinctive in specimens from central Italy. Terminal papillae sclerotized in a medially broader band; receptaculum seminis very long and narrow, narrowest above the middle; the last part somewhat enlarged; receptaculum seminis with a distinct striation along the whole length and a short appendix. Diagnosis. Males with an intense, reddish colouration may be confused with M. au- reoviridella in central Europe, but they can be separated by their lack of the golden Nota lepid. 30 (2): 235-298 259 colouration of the inner margin and by their narrower and more silvery fasciae. Similar Mediterranean species, like Micropterix corcyrella Walsingham, 1919, Micropterix erctella Walsingham, 1919, Micropterix italica Heath, 1981 and Micropterix renatae Kurz, Kurz & Zeller, 1997, but also the alpine M. fenestrellensis can be recognized with certainty only by examination of the genitalia, although they are normally more red- dish and show more distinct silvery whitish markings (except M. fenestrellensis). We also found Micropterix myrtetella idae Rebel, 1902 from Peloponnes with dark scales on head, which looks also very similar to M. aruncella f. atricapilla. Females can be separated from other species without wing markings by their purple colouration at the base of the forewing costa. In M. calthella, this colouration reaches the inner margin, whereas in M. isobasella and Micropterix sicanella Zeller, 1847 f. obsoleta Heath, 1963, it is almost missing. In the Alpes Maritimes we have found a form of M. paykul- lella that also lacks markings and with a less extended purple colouration of the base of the forewing. M. mansuetella has black scales on the head, similar to M. aruncella f. at- ricapilla, although the latter never has such an intense black colouration. Furthermore, these two species prefer different biotopes. M. mansuetella inhabits swampland, where- as M. aruncella is normally found on sunny, dry, nutrient-poor meadows. Nevertheless, both species can also be found more or less syntopically, where these biotopes intersect. There are further similar species in the Sierra Nevada in Spain and in northern Africa, but so far M. aruncella has not been found there. In all cases, males can be easily rec- ognized by their characteristic genitalia, and also the female genitalia seem to be fairly diagnostic. Distribution. Heath (1996) and Karsholt (2004) have reported this species throughout Europe except Portugal, Sicily, Malta, Iceland, Romania, Albania, Bulgaria, Crete and the European part of Turkey. According to Heath (1983) and Meeß (1910), M. aruncel- la is distributed throughout Europe except the Iberian Peninsula, northwards to Sweden and Finland, eastwards to Russia. Life history. M. aruncella is found in meadows, in bushland, at grassy forest margins, but not inside taller forests, except within larger open clearings with grasses. This spe- cies prefers extensively used, more or less dry and bushy meadows, where the moths can be found feeding on flowering grass and also on blossoms of other plants (Heath 1960a; Meyer et. al. 2002). In the Alps M. aruncella occurs also in elfin woodland where we found it on flowering Rosa pendulina L. and Pinus mugo mugo Turra. In other habitats we found this species feeding on the flowers of Crataegus sp., Sambucus sp., Urtica sp., Cytisus spp., Lychnis flos-cuculi, Veronica chamaedrys and Plantago media. The moths are on the wing from May to August, depending on elevation. They occur from near sea level up to more than 2000 m. Preimaginal stages. According to Heath (1962, 1983), the egg has an oval shape, a length of 400-430 um and a width of 310-350 um, with up to 60 um long, rod-like structures. Immediately before hatching, the translucent white colour turns to grey. According to Klausnitzer (2002), the body length of the final larval stage is 4.0-4.5 mm. The larva was found on Dactylis sp., where it probably feeds on detritus (Luff 1964; Heath 1983). The pupa rests in a robust cocoon. 260 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Remarks. This species shows a distinctive sexual dichroism and is one of the most variable species of the genus. Besides f. atricapilla mentioned above, which is com- mon in the mountainous regions of the Alps, and f. seppella, specimens are known from the northern and central Apennines with distinct silvery white markings on a coppery ground colour. Furthermore, from Sardinia f. nuraghella has been described, charac- terized by males without any wing markings. Nevertheless, considering the genitalia, all these forms have been proven to belong to M. aruncella. The forms atricapilla and nuraghella, as well as the specimens from the northern and central Apennines may therefore present distinct subspecies. Micropterix tunbergella (Fabricius, 1787) Tinea tunbergella (Fabricius, 1787): 253. Type locality: Denmark, Funen, Faborg. Neotype @: Alléskov, Faaborg, 7.5.1926; coll. Larsen, genit. no. 842 Kristensen; in coll. ZMUC. Tinea thunbergella auct. nec Fabricius, 1794. Tinea helwigella Hiibner, 1805: pl. 38 fig. 263. Type locality: None given. Junior subjective synonym. Tinea rubrifasciella Haworth, 1828: 572. Type locality: Kent (Great Britain). Junior subjective syno- nym. Micropteryx depictella Herrich-Schäffer, 1851: fig. 7. Type locality: None given. Junior subjective syno- nym. Description of adults. Examined: 90°, 209. Forewing length: © 3.7-4.1 mm; Q 3.9- 4.9 mm. Head black-brown; vestiture of hair-like scales on the head yellow, rusty yel- low at the base of the antennae; antennae light brownish, 3/4 (9), respectively 1/2 (9) of forewing length; thorax golden; tegulae bronzy golden to purple, posteriorly purple violet; forewing with golden ground colouration and coppery to purple markings: a basal spot, extending from costa across nearly half of the forewing width; at 1/3 an elongated spot, extending from costa across 2/3 of the forewing width; distal of 1/2, a more or less broad fascia extending across the whole forewing width broadly bifur- cated and purple violet at the costa, often interrupted in the middle and sometimes also bifurcated at the inner margin; at the posterior third of this fascia, mostly connected with another fascia directed to, but not always reaching the apex, narrow at the junction and spoon-like, apically broadened, sometimes bifurcated; apex and outer margin with bronzy golden to purple scales; fringe bronzy golden, golden distally; hindwing bronzy golden, often with a purple tinge apically; fringe golden, whitish outwards; legs brown- ish, golden shining; abdomen brownish golden. SO Genitalia. Uncus stout, in the form of a broad beak; accessory claspers well developed, with pocket-like lobes at the ventral margin, which bear many, relatively long, slightly bent spinoid setae at the anterior part of the ventral margin; a group of shorter thickened setae at the anterior tip; a bunch of long hair-like setae at the poste- rior part of the accessory claspers; at the anterior margin of the accessory claspers a long, lobe-like appendix on both sides extending beyond the uncus ventrally with long, hair-like setae; valvae long, very slender, upwardly bent, slightly enlarged at the tip; on the inner surface two irregular rows of shorter spinoid setae anteriorly and some setae postbasally, as well as long hair-like setae medially. Nota lepid. 30 (2): 235-298 261 9 Genitalia. Tergite IX completely atrophied, leaving only a small remnant with a patch of setae, sternite IX typically reduced, 1/3 of the width of the other sclerites in the middle, laterally enlarged into a lobe and with indistinct border. Terminal papillae typically sclerotized; ductus receptaculi enlarged at the beginning of the receptaculum seminis and strongly bent; receptaculum seminis long and narrow, somewhat enlarged in the second half, with a longer, thin appendix at the end. Diagnosis. M. tunbergella can be sometimes confused with M. mansuetella. Also simi- lar is Micropterix kardamylensis Rebel, 1903 (from Peloponnes, Greece), but it has no golden spot at the costa at 2/3 (the outer purple fascia is not bifurcated). There is an undescribed species in Greece, which is very similar to M. tunbergella, but can be separated with certainty only by examination of the male genitalia. Distribution. According to Heath (1996, 1983) and Karsholt (2004), the species is distributed across Europe. M. tunbergella has not been recorded from Portugal, Italy, Bulgaria and Finland. In the Balkans, we recorded this species with certainty from Macedonia, Bosnia and Greece. Life history (Heath 1983). M. tunbergella mostly inhabits deciduous woodland. The adults feed on pollen of Quercus, Acer and Crataegus as well as on other plants. Sometimes M. tunbergella has been found swarming around tree-tops. Preimaginal stages. According to Heath (1983), the egg has an oval shape, a length of 380-390 um and a width of 240-250 um, with up to 100 um long, rod-like structures. Immediately before hatching, the translucent white colour turns grey. The larva and pupa are unknown. Remark. The identity of the species named tunbergella by Fabricius in 1787 was settled by Heath et al. (1979) who designated a neotype. The name thunbergella (Fabricius, 1794) is a synonym of Caloptilia alchimiella (Scopoli, 1763) (Gracillariidae). Micropterix aureoviridella (Höfner, 1898) Eriocephala aureoviridella Höfner, 1898: 73. Type locality: Austrian Alps, Carinthia, Petzen near Bleiburg. 4 syntypes (2 © [1 destroyed, only 1 forewing and parts of body left], 2 9) in coll. NHMK [exam- ined]. Micropterix liogierella Real, 1987: 377-378. Type locality: Cret de la Neige (France, Jura Mountains). Junior subjective synonym. Description of adults. Examined: 10 ©, 27 9. Forewing length: © 3.4-3.9 mm; Q 3.6-4.7 mm. Head black-brown; vestiture of hair-like scales on the head rusty yellow to dirty white; antennae golden brown, purple at the base, 4/5 (S'), respectively, 4/7 (9) of the forewing length; thorax golden, posteriorly purple violet; tegulae bronzy golden to purple violet, bluish in the posterior part; forewing bronzy golden to reddish golden, a purple violet spot near the base at the costa, basally and between the fasciae mixed with purple scales, sometimes also completely purple to purple violet in the anterior half of the forewing; inner margin broad golden to bronzy golden from the base to the middle of the wing; markings more or less silvery golden to golden, indistinctly 262 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe bordered: a fascia extending from costa across more than half of the forewing width at 1/5, sometimes poorly developed; a narrower, bent fascia across the entire width of the wing at 1/2, often narrowed in the anterior third, sometimes nearly completely inter- rupted; an elongated, more or less large spot at 3/4, usually not reaching the costa and the inner margin; a small bronzy golden costal spot at 3/4, sometimes very ambiguous; fringe bronzy golden to whitish bronzy golden; hindwing bronzy golden, more or less with a purple tinge; fringe lightly bronzy golden; legs and abdomen brownish, golden shining. SO Genitalia. Uncus moderately long, distinctly separated and stout; at the lower margin convex, with an area of setae; accessory claspers well developed, nearly tri- angular, with a row of many moderately long, partly modified spinoid setae, as well as inwardly oriented, nearly Y-shaped setae at the ventral margin of the anterior half; valvae moderately long, broad, distinctly constricted medially, strongly bent dorsad at the distal end; some setae postbasally on the inner surface of the valvae, two irregular rows of many shorter spinoid setae at the lower margin of the inner surface distally and a row of longer setae medially. © Genitalia. Tergite IX missing; sternite IX atrophied, only 1/3 of the normal width in the middle, laterally enlarged into a weak lobe, but variably, so poorly diag- nostic; distal and proximal margin distinctly bordered; lateral margin inwardly bent, indistinctly bordered. Terminal papillae with well developed sclerotized band, weakly diagnostic: proximally straight or slightly sinuate; ductus receptaculi short; receptacu- lum seminis small, at the beginning of the ductus receptaculi somewhat enlarged, af- terwards constricted; receptaculum seminis forming a sac in proximal half, with an inconspicuous, typical striation; a thin appendix at the end. Diagnosis. M. aureatella is coloured more purple violet (not as reddish as M. aureo- viridella) without golden inner margin and with distinctly bordered, more golden fas- ciae. M. aureoviridella can be separated from Micropterix facetella Zeller, 1850 (from Balkans) by its lack of the small costal spot near 3/5. Sometimes M. aureoviridella can be mixed up with males of M. aruncella. Very similar are furthermore M. aglaella and especially M. paykullella, which therefore they can be identified with certainty only by examination of the genitalia. Distribution. According to Heath (1996) and Karsholt (2004) this species has been found in Italy, Switzerland, Germany, Austria, Poland, Slovenia and Slovakia. The records from Greece (Heath 1996; Karsholt 2004) and from Romania (Karsholt 2004) seem to be doubtful. To our present knowledge, this species occurs in the northern and southern calcare- ous Alps and other upland areas (Germany: Harz; Switzerland and France: Jura [M. liogierella Réal, 1987]). Furthermore a series of females from the Dinarian Mountains (Croatia) probably belong to M. aureoviridella, suggesting the occurrence of this spe- cies across the Balkans (Zeller-Lukashort, Kurz & Kurz 2002). Life history. So far this species has been found on alkaline ground (carbonate sub- strate) at 900-2000 m elevation. M. aureoviridella prefers open and dry habitats on southwardly directed slopes, margins of light spruce forest with interspersed rocks (with ground vegetation: besides grasses, especially Mercurialis perennis L.), margins Nota lepid. 30 (2): 235-298 263 of forest and scrubs in mountainous areas, as well as in elfin woodland (vegetation: Pinus mugo Turra, Rhododendron hirsutum L., Vaccinium myrtillus L., Erica carnea L., Juniperus communis alpina (Suter), etc.). M. aureoviridella has also been found in sub-alpine dwarf scrub. The adults feed on flowering shrubs including P. mugo. According to the elevation, the adults occur from end of May to July. Preimaginal stages. The early stages are unknown. Remarks. A series of females from the Dinarian Mountains (Croatia) remains of un- certain taxonomic status. Superficially they resemble M. aureoviridella, but their wing markings are overall dark bronzy golden and very diffuse. Their ground colouration is more reddish to purple. M. aureoviridella has often been described as “golden green”, which is the translatıon of its Latin name. But the anımals are mostly reddish golden and often show a more or less intense purple colouration, which is already stated in the original description by Höfner (1898). Micropterix paykullella (Fabricius, 1794) Alucita paykullella Fabricius, 1794: 340. Type locality: Southern Europe. Holotype @: in coll. ZMUC (Karsholt in litt.). Tinea anderschella Hiibner, 1813: pl. 51, fig. 352. Type locality: none given. [Synonymy suggested by Heath (1987), but not clear from Hiibner’s illustration]. Micropterix paykulella [sie] f. rosarum Müller-Rutz, 1927. 532-533. Type locality: Above Törbel, 1600 m (Valais, Switzerland). Junior subjective synonym. Description of adults. Examined: 16 ©, 27 9. Forewing length: © 3.0-3.7 mm; 9 3.44.3 mm. Head black-brown; vestiture of hair-like scales on the head dirty white to rusty yellow; antennae dark brown, golden shining, 3/4 (©), respectively, 4/7 (Q) of forewing length; thorax bronzy golden, tegulae reddish bronzy golden, partly mixed purple; inner margin of forewing broad with bronzy gold extending from the base near- ly to the median fascia; a broad bronzy golden fascia nearly at 1/4, connected with the bronzy golden inner margin; the rest of the forewing reddish bronzy golden, more pur- ple at the costa and at the apex, in some specimens completely purple to purple-violet; wing markings whitish golden to golden, delicately bordered in bronzy gold: a narrow, slightly outwardly bent fascia nearly at 1/2 across the whole width of the wing, some- times narrowed anterior of the middle and more golden coloured; rarely a small costal spot at 3/5; a slightly ovate spot at 3/4, seldom reaching the outer margin but mostly reaching the costa, sometimes a broader, golden fascia across the entire width of the wing, whitish golden coloured in the anterior half of the wing, somewhat narrowed in the posterior third; fringe bronzy golden, somewhat purple shining, lighter distally; hindwing bronzy golden, apically with a purple tinge; fringe bronzy golden, lighter distally; legs and abdomen brown, golden shining. JS Genitalia. Uncus short, moderately broad, with a broadly rounded tip; ven- trally beyond the uncus and inside the tegumen, a tuft of longer hair-like setae; ac- cessory claspers relatively broad, rounded distally; a row of about 10 longer, straight to slightly bent spinoid setae on the inner surface of the anterior end of the accessory 264 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe claspers, which are partly hook-shaped at their ends; basally a second row of two longer, nearly straight spinoid setae and three to four sickle-shaped thickened setae; valvae moderately long, constricted beyond the middle, the last fourth enlarged, triangle- shaped and bent dorsad; a group of about five shorter spinoid setae and some setae on the inner surface postbasally; one or two rows of about 10 shorter, straight spinoid setae and a row of longer setae on the inner surface of the enlarged end of the valvae. 9 Genitalia. Tergite IX missing; sternite IX reduced, weakly sclerotized, with indistinct lateral margins. The pregenital abdomen noticeably but weakly differs from other species. Terminal papillae with a sclerotized band, without any specific charac- teristics; ductus receptaculi very narrow at the beginning of the receptaculum seminis; receptaculum seminis relatively long and narrow, striated, somewhat enlarged in the second half. Diagnosis. M. paykullella can be confused with M. aureoviridella, the typical form of Micropterix sicanella Zeller, 1847 (M. sicanella does not occur in the area of distribu- tion of M. paykullella so far) and especially with M. aglaella. M. paykullella can be separated from M. aureoviridella by its broader, bronzy golden fascia at 1/4, by its mostly narrower fascia at 1/2 and by its mostly stronger purple colouration especially at the apex. It can be separated from the typical form of M. sicanella by its bronzy golden colouration at the costa, which does not reach the base, by its narrower fascia at 1/2 and the mostly more intense purple colouration of the forewing, but also by lacking a cos- tal spot at 2/3. Nevertheless, both species are very variable, so they can be recognized with certainty only by examination of the genitalia. M. aglaella is also very similar; it shows a mostly more purple-violet colouration and sometimes a small costal spot at 2/3. The bronzy golden colouration at the base of the forewing nearly reaches the median fascia in M. paykullella, whereas in M. aglaella this colouration ends far before the median fascia. In M. aglaella, the outer border of the bronzy golden colouration at the base and the median fascia are nearly parallel. Since M. paykullella and M. aglaella may occur together in the western Alps, for proper identification genitalic examina- tion is needed. All above-mentioned species can easily be distinguished by their male genitalia. Females of M. paykullella can be separated genitalically from M. aureoviridella and M. sicanella particularly by their distinctly slender and longer receptaculum seminis, and from M. aglaella by their distinctly stronger sclerotization of sternite IX and ter- minal papillae. Furthermore, M. aglaella has a slightly shorter and stouter receptacu- lum seminis. Females of M. aruncella can be easily identified by their characteristic sternite IX. Distribution. M. paykullella seems to be distributed locally across the whole Alps. According to Heath (1996) and Karsholt (2004), this species occurs in France, Italy, Austria and Switzerland. Following Meeß (1910), M. paykullella occurs in southern France (also Viette 1948; Leraut 1980), in Switzerland (also Whitebread 1992), in Tyrol (Austria), in the Alps of Lower Austria and in Italy. Furthermore this species occurs in Vorarlberg (Austria) (Burmann & Huemer 1984) and Bavaria (Germany) (Osthelder 1951; Pröse 1987). Nota lepid. 30 (2): 235-298 265 The records from Dalmatia (Croatia/Montenegro) seem to be misidentifications prob- ably of Micropterix facetella Zeller, 1851. According to Karsholt (2004) the records from Germany (Heath 1996) seem to be doubtful. Records from southern Italy and Sicily belong to Micropterix sicanella Zeller, 1847. In Italy, M. paykullella is restricted only to the Alps according to present knowledge. Life history. Specimens without any markings were found above the timberline (2200 m) feeding on pollen of Helianthemum sp. Individuals with typical wing pattern were swarming around dwarf shrubs in clearings and on the outskirts of forest at lower montane level. Preimaginal stages. The early stages are unknown. Remarks. Individuals from the Alpes Maritimes (border between France and Italy, Marguareis, 2200 m) with golden colouration and fasciae at 1/2 and 3/4 absent or only weakly observable are confirmed genitalically to belong to this species. Micropterix allionella (Fabricius, 1794) Tinea allionella Fabricius, 1794: 321. Type locality: Southern Europe. Type: Not designated, in coll. ZMUC (Karsholt in litt.). Tinea tricinctella Costa, 1836: 223, pl. 2 fig. 2. Type locality: Napoli (Italy). Junior subjective synonym. Micropteryx rothenbachii auct. nec Frey, 1856. Description of adults. Examined: 50%, 39. Forewing length: © 3.6-4.5 mm; 9 4.7-4.8 mm. Head black-brown; vestiture of hair-like scales on the head dirty white to rusty yellow; antennae somewhat longer than 3/4 (©), or nearly 2/3 (Q) of the forewing length, sexually dichroic, with the © brown, golden shining with more or less purple tinge, with the © distinctly bi-coloured: light golden at the base to nearly 1/3, apically similar to the ©’; thorax golden, tegulae coppery bronzy golden to purple; forewing brown golden, especially at the costa mixed with purplish to purplish violet with a golden basal area and light golden to golden markings, bordered delicately in bronzy gold, broader in males, more slender in females: a broad fascia nearly at 1/4, narrowed at the costa; a broad fascia at nearly 1/2, somewhat narrower medially and more or less bent distad; mostly a small costal spot at 2/3; a fascia at 3/4, more or less triangle- shaped; wing often bronzy golden at the apex and sometimes partially so at the inner margin; tip of the apex purple; fringe whitish golden; hindwing greenish bronzy golden to bronzy golden, more or less with a purple tinge; fringe light golden, with a purple tinge at the basal half; legs and abdomen brown, golden shining. CO Genitalia. Uncus moderately long, somewhat stout, a long tuft of hair-like se- tae ventrally beyond the uncus; accessory claspers moderately long, about four nearly Y- or T-shaped and moderately long and thickened setae at the tip; the posterior margin of the accessory claspers slightly bent outwards (bearing some longer spinoid setae); at the lower end of the posterior margin a small separated lobe, also with one or two longer spinoid setae; another two longer spinoid setae in the middle of the accessory claspers, distinctly behind the inner margin and in line with the Y- or T-shaped thick- 266 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe ened setae; valvae moderately long, distally constricted medially, weakly bent; basally a longer spinoid seta on the inner surface as well as some setae postbasally; tip of valvae slightly spatulate with two or three irregular rows of shorter spinoid setae and a row of longer setae on the inner surface; another two shorter spinoid setae, displaced towards the constriction. © Genitalia. Tergite IX missing; sternite IX strongly reduced, constricted medi- ally, with a distinct border, although weakly sclerotized. Terminal papillae with a strong sclerotized band; ductus receptaculi starting broadly, then very narrow at the beginning of the receptaculum seminis; receptaculum seminis moderately long, the first third a sac that abruptly narrows; the last third an elongated sac; receptaculum seminis with distinct striation; a sac-like vestibulum. Diagnosis. M. allionella can be separated from most similar species by its broad golden fasciae, its small costal spot at 2/3 and almost complete absence of a golden inner mar- gin. Sometimes M. trifasciella has also a small costal spot at 2/3 and also sometimes this spot is missing in M. allionella, and therefore M. allionella, especially males with reduced golden outer margin, could be misidentified as M. trifasciella. M. allionella is also very similar to M. rothenbachii and cannot always be separated externally for sure. M. rothenbachii has a more distinct purple to purplish violet colouration, broader golden fasciae (fasciae of males of M. allionella are nearly as broad as the fasciae of females of M. rothenbachii) and a more intense bronzy golden colouration at the apex and at the outer margin. M. allionella without costal spot at 2/3 can also be confused with M. aureatella and M. rablensis. Micropterix hartigi Heath, 1981 (from southern Italy) is also usually very similar to reddish or brown-golden coloured forms of M. al- lionella (especially male), but can be distinguished by its genitalia. M. allionella has a somewhat longer uncus than M. rothenbachii and also differently shaped accessory claspers. M. allionella has four Y- or T-shaped thickened setae at the tip of the accessory claspers (M. rothenbachii only three, but these are also more dis- tinct). M. allionella has valvae which are only weakly bent dorsad (M. rothenbachii has more slender valvae and the distal third is distinctly bent dorsad). In the male genitalia, M. trifasciella is very similar to M. allionella, but can be separated as follows: The uncus is more distinctly separated and not as broad as in M. allionella, the accessory claspers are somewhat shorter and broader at the base. The distal third of the valvae of M. trifasciella is distinctly broader than in M. allionella, and somewhat triangle-shaped (in M. allionella narrow, spatulate). The differences in the genitalia to M. trifasciella are very small, since the kind of spinoid setae on the accessory claspers seems not to be a constant character. The abdomen of the female of M. allionella is indistinguishable from M. rothenbachii and M. trifasciella. Distribution. According to Heath (1996) and Karsholt (2004) this species occurs in France, Italy, Germany, Switzerland, Czech Republic, Slovakia, Bulgaria, Croatia, Slovenia, and Yugoslavia. Following Meeß (1910) and Meyrick (1912) M. allionel- la (noted as M. rothenbachi(ii)) occurs in Switzerland (see also Whitebread 1992), in northern and central Italy and in Austria. Furthermore, this species occurs in the Nota lepid. 30 (2): 235-298 267 Bavarian Alps (Osthelder 1951; Pröse 1987), together with M. rothenbachii according to Pröse (1987). Leraut (1980) has recorded this species from France, Belgium and Corsica. But these records seem partially to be misidentifications of M. rothenbachii. The records from Turkey (Heath 1996) seem to be doubtful. This species has also been recorded from southern Italy (Zeller-Lukashort, Kurz & Kurz 2002; Whitebread 1995). Life history. We found this species in clearings and on the outskirts of forest flying in tall herbaceous vegetation. The examined specimens were found at 250-1700 m eleva- tion. Preimaginal stages. The early stages are unknown. Micropterix trifasciella Heath, 1965 Micropteryx trifasciella Heath, 1965: 243-245. Type locality: Italy, Piemonte, Fenestrelle. Holotype: © Alpi Cozie, Val Chisone, Fenestrelle, 1300 m, leg. Della Beffa, agosto 1923 (examination of geni- talia Heath No. 368); in coll. MSNM. Micropteryx aureatella (Scopoli, 1763) sensu Viette 1948: 37, fig. 26. Misidentification. Description of adults. Examined: 160, 139. Forewing length: © 3.4-4.7 mm; 9 4.2-4.8 mm. Head black-brown, vestiture of hair-like scales on the head light to rusty yellow; antenna nearly 4/5 (©), respectively 1/2 (9) of forewing length, brown, golden shining with a slight purple tinge; thorax golden to bronzy golden, tegulae coppery bronzy golden to purple violet; forewing brownish golden to blue violet, at the base with little coppery to bronzy golden spots, forewing with three whitish golden to deep golden fasciae of variable width, delicately bordered with bronzy gold: a moderately broad and straight fascia at nearly 1/4, narrowing against the costa and slightly oriented distad; a moderately broad and more or less straight fascia at 1/2, sometimes bent dis- tad; a fascia at % of variable shape, often more or less triangle-shaped, sometimes only slightly so, sometimes broadly reaching costa and inner margin; sometimes a small cos- tal spot at 3/5; fringe purple bronzy golden, distally golden; hindwing bronzy golden, with a strong purple tinge and with bronzy golden fringe, which is purple at the base; legs and abdomen brown, golden shining. CO Genitalia. Uncus short, somewhat stout, with a moderately broad tip; ventrally beyond the uncus and inside the tegumen short tufts of hair-like setae; between the uncus and the accessory claspers an indistinctly bordered area of very long, flat setae, which are easily lost during preparation; accessory claspers short, enlarged at the tip; anterior margin slightly bent outwards, at the ventral margin a small, distinctly separat- ed appendix; accessory claspers with 19-24 thickened setae on the inner surface: At the tip about 7-10 strongly modified, hatchet-shaped and spatulate-shaped thickened setae in an upper row, a row of very long, slightly bent spinoid setae (about 9-12, two or rarely only one of them on the aforementioned posterior appendix) on the anterior mar- gin; three to five long, L-shaped thickened setae which extend the upper row; valvae moderately long, strongly constricted medially, the distal third spatulate or somewhat 268 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe triangle-shaped, with two or three irregular rows of shorter spinoid setae and some long setae on the inner surface; on the inner surface two or three short spinoid setae distally of the middle of the valve and one longer, straight spinoid seta basally. © Genitalia. Tergite IX missing; sternite IX strongly reduced, constricted medi- ally, not diagnostic. Terminal papillae with a sclerotized band; receptaculum seminis moderately long, very slender, somewhat enlarged in the second half, with typical stria- tion. Diagnosis. Due to the lack of the small costal spot at 2/3 this species can easily be con- fused with M. huemeri (separable with certainty only by examination of the genitalia), and also with M. aureatella (see Viette 1948). M. aureatella can be distinguished by its mostly distinctly smaller spot or fascia at 3/4 and by its mostly more slender fasciae at 1/4 and 1/2. M. trifasciella differs externally from M. allionella mostly by lack of the costal spot at 3/5. But this character is sometimes missing too in M. allionella, whereas it may seldom be present in M. trifasciella. Therefore both species can only be sepa- rated with certainty by genitalic examination. So far, they have not been found together syntopically. There are no differences in the female genitalia between M. trifasciella and M. al- lionella. Distribution. To present knowledge, M. trifasciella has been recorded with certainty only from the Italian and French Alps (Zeller-Lukashort, Kurz & Kurz 2002). Life history. This species inhabits tall herbaceous vegetation around boulders between montane and sub-alpine elevations (1400-1900 m). At the type location (Fenestrelle, Piemonte, Italy) this species was found again in 2003 at the roadside, also in tall her- baceous vegetation. Preimaginal stages. The early stages are unknown. Remarks. Compared with the original description, the examined specimens show dis- tinctly more slender fasciae and the shape of the fascia at 3/4 is variable. The male genitalia figured by Viette (1948) as M. aureatella probably belongs to M. trifasciella. Micropterix rothenbachii Frey, 1856 Micropteryx rothenbachii Frey, 1856: 52. Type locality: Switzerland. 2 syntypes in coll. BMNH. Micropterix australis Heath, 1981a: 99. Type locality: Maroggia (Switzerland, Ticino). Junior subjective synonym. Micropteryx rothenbachi auctt. (incorrect subsequent spelling). Micropterix germanica Heath. Nomen nudum (see remarks). Micropterix vallebonnella Real, 1988: 3-9. Type locality: Bonnevaux (Doubs, France). Junior subjective synonym. Description of adults. Examined: 270, 129. Forewing length: © 3.6-4.6 mm; 9 4.5- 5.0 mm. Head dark greyish brown, vestiture of hair-like scales on the head dirty white to rusty yellow; antennae 3/4 (©), respectively, slightly more than 1/2 (Q) of the forew- ing length, distinctly bi-coloured: light golden at the base (in 9 reaching nearly to 1/3), brownish, more or less with a distinctly purple tinge apically; thorax bronzy gold- Nota lepid. 30 (2): 235-298 269 en; tegulae coppery to purple violet, sometimes with single bluish scales posteriorly; forewing brownish golden to purple violet with whitish golden to golden markings of variable width (distinctly broader in © than in Q), delicately bordered in bronzy gold: forewing bronzy golden at the base; a fascia at 1/4, sometimes slightly broadened medi- ally or narrowed in the upper third; a broad fascia close to 1/2, strongly bent distad and often broadened at the proximad margin; a small costal spot at 3/5; opposite, sometimes a very indistinct, very small spot at the proximad margin; a broad, triangle-shaped fas- cia at 3/4; this fascia sometimes enlarged along the costa towards the base and reach- ing the costal spot at 3/5, merging into a very large rectangular spot; this confluence often only subcostally developed, leaving at the costal margin a very small purple area; bronzy golden at the apex and at the outer margin, the bronzy golden colouration form- ing a slender fascia; tip of the apex coppery to purple; fringe whitish golden, bronzy to purple basally; hindwing bronzy golden with a variable purple tinge, especially at the apex; fringe bronzy golden, tips whitish; legs light bronzy golden; abdomen brownish, golden shining. SO Genitalia. Uncus short, stout, with a broadly rounded tip; ventrally beyond the uncus a brush of hair-like setae; accessory claspers broad proximally, acuminate dis- tally, with three approximately Y-shaped, shorter thickened setae at the tip (mostly fold- ed in permanent preparation and therefore hardly visible) and with 9-11 rigid, longer spinoid setae, sometimes slightly bent (six to eight on the apical part, two more further inside at the ventral margin, one somewhat dorsad) (see Remark); valvae distinctly bent, constricted beyond the middle, spatulate-shaped at the tip, with two irregular rows of shorter spinoid setae distal of the constriction at the ventrad margin of the mesad surface; medially and postbasally a similar seta each. 9 Genitalia. Tergite IX missing; sternite IX reduced, more constricted medially than laterally, strongly sclerotized with distinct margins; lateral margins irregular, may- be diagnostic for this species. Terminal papillae with distinctly bordered sclerotized band, not protruding; ductus receptaculi at the beginning of the receptaculum seminis very narrow, before enlarging into a sac; receptaculum seminis moderately long, the first third small, then elongated into an enlarged sac; receptaculum seminis with typical striation. Diagnosis. M. rothenbachii can be recognized by lack of the broad golden colouration of the inner margin. M. allionella is however very similar. Distribution. Following Heath (1996) and Karsholt (2004), this species has been record- ed from Italy, Sicily, Austria, Switzerland and Germany. According to Heath (1981a), this species occurs in Germany (Schwarzwald), Switzerland, Austria (Klagenfurt) and Italy (incl. Sicily). Karsholt (2004) reports this species also from Croatia and Slovenia (see also Zeller-Lukashort, Kurz & Kurz 2002). Life history. This species inhabits light, dry and somewhat rocky, mixed beech forests, where it occurs in open places, mainly with natural cover such as brambles, grasses, etc. In central Italy, M. rothenbachii occurs together with Micropterix vulturensis Heath, 1981, and in the northern Apennines is syntopic with M. schaefferi and Micropterix zangheriella Heath, 1963. DATA) ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Preimaginal stages. The early stages are unknown. Remarks. This species was recognized as M. australis until 1987, whereas M. rothen- bachii Frey, 1856 was said to be a synonym to M. allionella. In the original description Of M. australis, Heath described 13 setae on the accessory clasper: “...with a marginal series of eight fairly long, stout, more or less curved setae and an inner row of five sımilar, straight setae.” In 1987, M. rothenbachii was re-established as a good species by Heath based on the examination of the type by Whitebread (1992) and M. australis became synony- mous to it. Originally, Heath wanted to describe a Micropterix germanica, but he changed the name before printing to M. australis, since the holotype did not come from Germany (Sattler, pers. comm.). Therefore, there are specimens in collections labelled as “Micropterix germanica Heath”. Micropterix huemeri Kurz, Kurz & Zeller, 2004 Micropterix huemeri Kurz, Kurz & Zeller-Lukashort, 2004: 111-114. Type locality: France, Alpes Maritimes, Marguareis. Holotype ©: France, Dep. Alpes Maritimes, Marguareis, west slope, Navela, 2100-2200 m; 18.7.1991, GU MIC2 © P. Huemer, ID-Nummer HdN-2289, in coll. TMLEF. Description of adults. Examined: 30, 19. Forewing length: © 3.9 mm; 9 4.4 mm. Head blackish, vestiture of hair-like scales on the head yellow; antennae approximately 3/4 (©) or almost 1/2 (Q) of forewing length, golden fuscous; thorax coppery to bronzy golden, tegulae purple violet with bronzy golden edges; forewings purple violet to blu- ish violet; base of costa bronzy golden; wing markings golden, delicately bronzy gold- en bordered; a fascia at 1/4, slightly bent, moderately broad on inner margin, distinctly narrowed from centre of wing to costa; a fascia at 1/2, slightly bent outwards, moder- ately broad, sometimes narrowed medially, sometimes distinctly broadened at costa; a fascia at 3/4, slightly broader than the other ones with distinctly curved inner margin; sometimes a residual costal spot at 3/5; cilia bronzy golden, apically whitish; hindwing coppery to bronzy golden, apically distinctly tinged purple; cilia bronzy golden; legs and abdomen golden fuscous. SO Genitalia. Uncus short, slightly stout with moderately broad tip; ventrally be- yond uncus a tuft of hair-like setae; a small area with several faint, straight setae at the posterior margin of tegumen between uncus and accessory claspers; accessory claspers proximally broad, distally tapered, on inner surface with an upper row of six sickle- shaped thickened setae and a lower row of nine more or less straight, moderately long spinoid setae, with the two proximal spinoid setae being slightly apart; valvae slightly bent, constricted beyond middle with a triangular distal end and two or three irregular rows of shorter spinoid setae on inner surface beyond the constricted part of the valve; post-basally a distinctly robust seta on inner surface. © Genitalia and pregenital abdominal exoskeleton. Noattempt has been made to prepare the genitalia of the single available female of this species. Nota lepid. 30 (2): 235-298 27] Diagnosis. M. huemeri belongs to a group of closely related species which is character- ized by the following characters: The accessory claspers, seen laterally, bear two rows of thickened setae. In the ventral row, the distal thickened setae are strongly modified (Y- or T-shaped), and the one or two proximal most ones are distinctly separated from the rest of the row. These characters are shared by M. rothenbachii, M. allionella and M. trifasciella. Externally, M. huemeri is quite well separated from the other species of this group by its three complete golden fasciae on the forewing and the absence of any further mark- ings. One exception is M. trifasciella, which has very similar wing pattern elements with only slightly broader fasciae on the forewing. However, the male genitalia of M. huemeri differ in the shape of the accessory clasper which is conspicuously narrower proximally and has a different arrangement of the thickened setae. Furthermore, on the inner surface of the valve, the row of the short and thickened setae extends further towards the base into the constricted part of the valve. In M. huemeri, the structures of the male genitalia are most similar to those of M. rothenbachii. M. huemeri can be distinguished superficially from M. rothenbachii by both the lack of a small costal golden spot and the golden tinge on the outer margin of the forewing. Concerning the male genitalia, M. rothenbachii has a distally club-shaped uncus and longer, distally more spatulate valvae with only one stout spinoid seta in the middle of the constriction. Distribution. M. huemeri seems to be an endemic of the geologically isolated region of the Marguareis (France). Life history. This species was found in high alpine grassland (at elevations higher than 2000 m) in tall herbaceous vegetation around boulders in July (P. Huemer, pers. comm. ). Preimaginal stages. The early stages are unknown. Micropterix schaefferi Heath, 1975 Micropterix schaefferi Heath, 1975: 253-254, figs 1-2. Type locality: Austria, Upper Austria, Linz. Holotype: © Oberösterreich, Linz, 25.1v.1934, Klimesch (genitalia preparation Heath no. 213); coll. Heath, in coll. BMNH. Micropteryx anderschella (Hübner, 1813) sensu Herrich-Schäffer, 1851: 392, pl. 1 fig. 4. Misidenti- fication. Micropteryx ammanella auctt. nec Hübner, 1813. Description of adults. Examined: 250°, 279. Forewing length: © 3.8-5.1 mm; Q 4.7- 5.8 mm. Head black, vestiture of hair-like scales on the head brownish yellow, yellow, whitish yellow or pale greyish yellow; antennae 3/4 (©), respectively slightly more than 1/2 (9); thorax dark golden to bronzy golden, tegulae bronzy golden, purplish to purplish violet tinged; forewings purplish bronzy golden to deep bluish violet with whitish golden to golden, finely bronzy golden bordered markings: inner margin broad bronzy golden reaching to 1/2; its basal part with an oblong spot of ground colour; apex, with exception of the outermost tip of the wing, as well as outer margin light bronzy golden; at 3/4 a small, whitish golden, often indistinct spot at the inner margin, Ze ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe which is embedded in the apical golden colouration; at 1/4 a transverse fascia, nar- row at costa, then broadening and reaching the bronzy golden inner margin; a broad, posteriorly somewhat narrower transverse fascia at 1/2, slightly bent outwards; a very broad transverse fascia at 3/4 reaching from costa and somewhat oblique to the centre of the wing, but not reaching the bronzy golden colouration at the outer margin; a small costal spot at 2/3, often connected with the outer transverse fascia, seldom also with the fascia at 1/2; fringe whitish golden, bronzy golden to purplish at its base; hindwing dark bronzy golden, more or less suffused with purple; fringe light bronzy golden; legs light brownish golden; abdomen brown, golden shining. SO Genitalia. Uncus more or less long, slender and somewhat flattened; acces- sory claspers broad, with a row of quite long, partly bent spines at the lower margin as well as with a shorter row of strongly modified, more or less Y-shaped, short thickened setae; valvae long, somewhat spatulate, narrower medially, their ends distinctly bent upwards; at the inner surface with a greater group of basal setae as well as with an ir- regular row of shorter spines at the lower margin and a row of long, flexible setae in the centre of the distal third of the valvae. Q Genitalia. Tergite IX missing; sternite IX much reduced, constricted in its mid- dle, with indistinct lateral margins, similar to M. osthelderi; all sclerites with slightly fringe-like lateral margins. Terminal papillae with centrally somewhat enlarged band of sclerotization, whose form is not diagnostic; ductus receptaculi at the onset of the re- ceptaculum seminis straight and very narrow for a short distance; receptaculum seminis short and a little stout, the last part an enlarged sac, with a typical transverse striation and a very narrow appendix at the end. Diagnosis. M. schaefferi is easily recognized by its golden inner margin and by the costal spot at 2/3: M. osthelderi has a much more extensive bronzy golden colouration at the base of the forewing (including the fascia at 1/4 which is also bronzy golden and not whitish golden); Micropterix facetella Zeller, 1851 (from Balkans) often lacks the golden colouration at the apex and at the outer margin. Micropterix vulturensis Heath, 1981 and Micropterix zangheriella Heath, 1963, being very similar superficially, can be distinguished from M. schaefferi with certainty only by investigating the male geni- talia. The former of these two species seems to be allopatric from M. schaefferi, being distributed in central and southern Italy, whereas the latter has been found sympatri- cally in the northern Apennines. Concerning the male genitalia, M. facetella is somewhat similar, but is distinguished by its longer uncus, the more distinctly pronounced and narrower accessory claspers and by having only three basal setae at the inner surface of the valvae (M. schaefferi having a group of more numerous setae). Concerning the structures of the female genitalia, M. osthelderi and M. schaefferi can be separated quite easily by the characteristics described for M. osthelderi. Distribution. According to Karsholt (2004) the species is recorded from France, Corsica, Italy, Belgium, the Netherlands, Germany, Switzerland, Austria, Hungary, Czech Republic, Bulgaria and Denmark. Heath (1996) reports this species also from Poland and former Yugoslavia. Following Heath (1975) this species occurs with cer- tainty also in Hungary and Czech Republic. Nota lepid. 30 (2): 235-298 215 Older records concerning Asia Minor (Meeß 1910; Meyrick 1912) are very doubtfully identified. In Italy this species seems to reach the border between the regions of Emilia Romagna and Tuscany, whereas to the South the very similar species Micropterix vulturensis Heath, 1981 can be found. Life history. The species inhabits open beech and coniferous mixed woodland, but also can be found in very wet situations in Fraxinus-dominated ravine forests as well as in dry situations in pine forests with Erica. Almost always the herbaceous layer in such woods is well developed, consisting in central Europe of different grasses, Mercurialis perennis L. and Dentaria enneaphyllos L., and in dry locations also Erica carnea L.. In elfin woodland in the mountains, the herbaceous layer is dominated by Vaccinium species. Furthermore, M. schaefferi can be found in cuttings rich in shrubs, in black- berry thickets and on sunny wood margins. The species is absent from dense woodland, especially spruce forests without herbaceous layer. Preimaginal stages. The early stages are unknown. Micropterix fenestrellensis Heath & Kaltenbach, 1984 Micropteryx fenestrellensis Heath & Kaltenbach, 1984: 22-23, figs 3-4. Type locality: Italy, Piemonte, Val Chisone, Fenestrelle. Holotype: Alpi Cozie, Val Chisone, Fenestrelle, 1300 m, leg. Della Beffa, agosto 1923 (examination of genitalia Kaltenbach GU 204); in coll. MSNV. Description of adults. Examined: 10,59. Forewing length: © 3.3 mm; 9 3.0-3.2 mm. Head black-brown, vestiture of hair-like scales on the head black; antennae dark-brown with a weak coppery tinge, about 3/4 (©), respectively, 2/3 (Q) of forewing length; tho- rax anteriorly bronzy golden, posteriorly purple to purplish violet; tegulae purple-violet; forewing bronzy golden to reddish bronzy golden, purple at the base of the costa, some- times reddish along the costa, with silvery white markings: an oval to longish, oblique spot near 1/4, not reaching costa and inner margin; a narrow fascia at 1/2, slightly bent outwards across the whole width of the wing; a round spot at 3/4 across the half width of the wing, near, but not quite reaching costa; fringe coppery bronzy golden, lighter outside; hindwing bronzy golden, with a purple tinge, especially at the apex; fringe cop- pery with lighter tip; legs and abdomen dark brown, golden shining. SO Genitalia. Uncus moderately long, club-shaped, rounded; tegumen and acces- sory claspers fused together; accessory claspers broad, trapezoid; at the anterior margin on the tip of the accessory claspers a group of short, acute, bent spinoid setae, along the anterior margin a row of five longer, acute spinoid setae, the one to two distalmost distinctly separated; at the end of this row another group of a few short, acute, bent spinoid setae; valvae moderately long, somewhat stout, constricted medially, enlarged and spatulate-like at the end; on the inner surface two longer, acute, straight spinoid setae postbasally, a very short spinoid seta medially and two or three rows of short, straight spinoid setae with some long setae at the end. 9 Genitalia. Tergite IX missing; sternite IX reduced to two very characteristic, elongated sclerotized platelets, which fuse together ventrally. Terminal papillae sclero- 274 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe tized in a band, somewhat bent outwards proximally; receptaculum seminis very long and slender, the second half enlarged, with typical striation. Diagnosis. M. fenestrellensis can only be confused with M. aruncella in its geographic range. The ground colouration of M. aruncella is generally less reddish, the silvery drawings are smaller and less distinct, and females do not show any markings. The male genitalia of M. fenestrellensis cannot be confused with any other species, and the sternite IX of the female is very characteristic for this species. Distribution. So far, this species has been recorded only from the type locality (Italy, Piemonte, Fenestrelle, 1300-1600 m) and from Monte Tanarello (Briga (fr.)) at 2000 m (French and Italian border area, south of Cuneo, Piemonte). Life history. At the type locality, the species occurs together with M. trifasciella in tall herbaceous vegetation around bushes. Preimaginal stages. The early stages are unknown. Micropterix osthelderi Heath, 1975 Micropterix osthelderi Heath, 1975: 256-258., figs 5-6. Type locality: Bavaria, Fürstenrud [Fürstenried] near Munich. Holotype @: Bav.mer., Fiirstenrud [Fürstenried] b. München, Daniel, 24.iv.26 (examina- tion of genitalia Heath No. 261); in coll. ZSM [examined]. Description of adults. Examined: 10°, 79. Forewing length: © 4.7 mm; 9 5.0-5.6 mm. Head dark-brown; vestiture of hair-like scales on the head light to dirty yellow; antennae brownish, bronzy golden shining, 3/4 (©), respectively, 1/2 (Q) of forewing length; thorax bronzy golden; tegulae bronzy golden, mixed with purple scales; forew- ing purplish violet to bluish violet: the basal fourth dark golden to bronzy golden, ex- cept small purplish violet streaks at the base of the costa and towards the dorsal margin lying within the basal bronzy golden colouration which reaches the median fascia, and a bronzy golden streak from the costa along the outer margin, just leaving the tip of the apex purplish violet; the rest of the markings whitish golden to golden, finely bordered in bronzy gold: a moderately broad fascia of equal width at 1/2, slightly bent outwards; a small costal spot at 2/3; a big costal spot at 3/4, bent inwards and sometimes enlarged towards the centre of the wing; fringe light bronzy golden; hindwing bronzy golden with a strong purple colouration, especially at the apex; fringe bronzy golden; legs and abdomen brown, bronzy golden shining. SO Genitalia. Uncus short, stout, with a small tip; accessory claspers with a lower row of shorter spinoid setae, partly bent at the end, and an upper row of strongly modi- fied Y-shaped thickened setae; valvae long, constricted medially, with probably two longer and some shorter, basal setae at the inner surface; two irregular rows of shorter spinoid setae and a row of longer setae at the distal third of the inner surface and many small spinoid setae at the tip of the valvae. 9 Genitalia. Tergite IX missing, sternite IX reduced and very similar to M. schaefferi, just somewhat enlarged medially. Terminal papillae with a very charac- teristic sclerotization: margin towards segment IX medially enlarged outwardly, with smooth margins; ductus receptaculi at the beginning of the receptaculum seminis very slender; receptaculum seminis long, constricted medially and strongly bent, only the Nota lepid. 30 (2): 235-298 275 last part enlarged like a sac, with a typical striation; receptaculum seminis with three very small spinoid setae at the end, lying side by side (this character was found in no other species); the shape of the receptaculum seminis and the number of spinoid setae are diagnostic. Diagnosis. M. osthelderi can easily be separated from all other similar species, includ- ing M. schaefferi, by its broad, dark golden colouration at the forewing base, and by its fascia at 1/4, which is not brightened in the middle but shows the same (bronzy) golden colouration as the inner margin. Poorly preserved specimens can easily be recognized by the very characteristic male and female genitalia. Distribution. According to Heath (1996) and Karsholt (2004) this species occurs in Italy, Germany, Switzerland, Austria, Poland, Czech Republic and Denmark. Heath (1975) records this species from the Alps (Austria, Switzerland, Bavaria) and from the uplands (Germany: Rheingau; Poland: Sudeten). Life history. This species seems to inhabit mixed coniferous forest especially at mon- tane elevations. Preimaginal stages. The early stages are unknown. Remarks. The paratypes of M. osthelderi (deposited in ZSM) include also specimens of M. schaefferi. Micropterix rablensis Zeller, 1868 Micropteryx rablensis Zeller, 1868: 133. Type locality: Italy, Alps, south of Tarviso, Raibl. Holotype ©: in coll. BMNH. Description of adults. Examined: 100°, 49. Forewing length: © 3.0-3.4 mm; 9 3.25- 3.8 mm. Head black-brown, vestiture of hair-like scales on the head dirty white, light yellowish grey to yellow; antennae brown, light reddish golden shining, 3/4 (©), re- spectively, 2/3 (Q) of forewing length; thorax bronzy golden, tegulae bronzy golden, posteriorly coppery to purple; forewing reddish bronzy golden to purple, with three whitish golden to light golden, sometimes diffusely bordered fasciae across the whole width of the wing: a fascia at 1/4, slightly bent inwards, narrow at the costa and con- stantly broadening towards the inner margin; a broad, more or less straight fascia at 1/2, sometimes slightly enlarged inwards at the anterior third; sometimes a small costal spot at 2/3, which can be joined with the median fascia or with the outer fascia; a broad fascia at 3/4, often enlarged inwards in the middle; fringe coppery proximally, distally whitish golden; hindwing bronzy golden, sometimes with a slightly reddish to purple tinge; fringe slightly reddish to purple proximally, distally light golden; legs and abdo- men light brown, golden shining. CO Genitalia. Uncus very short; beyond the uncus and inside the tegumen tufts of long, very acute hair-like setae; tegumen and accessory claspers fused, strongly devel- oped; accessory claspers with a row of short, isolated, straight spinoid setae at the ante- rior margin, starting near the uncus; a row of short, acute, proximally straight, distally increasingly curved spinoid setae inside the lower posterior margin; anteriorly a short row of small, strongly modified, broad T-shaped (or hatched-shaped) thickened setae (starting at the anterior margin of the accessory claspers); proximally of these some 276 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe very small, acute, straight spinoid setae; valvae moderately long, slender, strongly constricted medially, the distal part spatulate, slightly bent upwards; a row of longer setae and two to three rows of short to very short, straight spinoid setae (proximally intergrading to a row of spinoid setae reaching the constriction) at the distal part on the inner surface. Q Genitalia. Tergite IX missing, sternite IX strongly reduced, weakly sclerotized in the middle distally, here partly without sclerotization (maybe characteristic for this species), partly with only indistinctly lateral margins. Terminal papillae with a scle- rotized band; receptaculum seminis very short, narrow, at the beginning of the ductus receptaculi enlarged like a knot, distinctly constricted before the middle, enlarged into a sac in the last part; receptaculum seminis with striation along the entire length, but not very regular; a longer appendix at the end. Diagnosis. Micropterix croatica Heath & Kaltenbach, 1984 is externally very similar to M. rablensis, but can be separated by its darker purple-violet ground colouration of the forewing, as well as by its more distinct, sharper markings. M. rablensis could be confused with small specimens of M. aureatella or M. aureoviridella. M. aureatella is darker purple-violet with smaller and distincter fasciae, M. aureoviridella shows mostly a less reddish colouration than M. rablensis, with indistinct markings and a bronzy golden inner margin. M. rablensis can be separated from both species by its broad outer fascia, which is mostly only an oval spot in M. aureatella and M. aureo- viridella. Sometimes, small M. allionella can be confused with M. rablensis, which is normally smaller. The male genitalia of M. croatica are very similar to those of M. rablensis, but can be separated besides other characters by their row of spinoid setae on the valvae, which reach the basal third. M. myrtetella is also related to M. rablensis and shows also a row of spinoid setae on the valvae postbasally, but the outer fascia of the forewing is reduced to a spot. Distribution. M. rablensis is most probably restricted to Carinthia (Austria) and to the adjacent areas of Styria (Austria), of Italy (type locality) and potentially of Slovenia. According to Heath (1996) and Karsholt (2004), this species occurs in Italy and Austria. Records from Romania (Heath 1996; Karsholt 2004) and Croatia (Karsholt 2004) are doubtful and probably belong to M. myrtetella. Also records from France (Viette 1948) turned out to be a misidentification (Heath in litt. according to Leraut 1980). All exam- ined animals from Trieste (Italy) belong to Micropterix croatica Heath & Kaltenbach, 1984. A record of M. rablensis from the Traunstein (Upper Austria) in the ZOOBODAT seems to be very doubtful too. Prése (1987) probably also refers to this record. An examination of this record has not been possible so far, but probably this is a misiden- tification of M. aureoviridella. Life history. We have found this species at outskirts of forest with tall herbaceous veg- etation and bushes, congregating on Aruncus dioicus (Walter) and Vicia sylvatica L.. In competition with M. rablensis on flowers of Aruncus dioicus (Walter), we also have found M. aruncella, M. aureatella and M. rothenbachii. Preimaginal stages. The early stages are unknown. Nota lepid. 30 (2): 235-298 27 Micropterix myrtetella Zeller, 1850 Micropteryx myrtetella Zeller, 1850: 62. Type locality: Italy, Tuscany, Montenero near Livorno. Holotype CO: in coll. BMNH. Description of adults. Examined: 25¢ (incl. slide of holotype), 119. Forewing length: CO 2.2-2.8 mm, 9 2.7-3.0 mm. Head black-brown, vestiture of hair-like scales on the head dark yellow; antennae brown, bronzy golden shining, more than 3/4 (©), respec- tively, 2/3 (Q) of forewing length; thorax bronzy golden; tegulae purple, mixed with bronzy golden scales; forewing purple to purplish violet, distally often lighter, with golden, delicately bronzy golden bordered markings: a broad fascia at 1/4 across the whole width of the wing, narrowing towards the costa and sometimes not quite reach- ing it; a broad fascia nearly at 1/2, bent outwards, sometimes narrowed or even inter- rupted in the middle; seldom a small costal spot at 2/3; a broad, irregularly formed spot at 3/4, extending from costa across somewhat more than the middle of the wing, this spot sometimes also indistinct or nearly atrophied; fringe golden, outside whitish; hindwing bronzy golden with a purple tinge; fringe bronzy golden, distally lighter; legs and abdomen brown, golden shining. CO Genitalia. Uncus short with a broad rounded tip; ventrally beyond and inside the tegumen a tuft of hair-like setae; accessory claspers very broad; along their an- terior margin on the inner surface a row of short, straight spinoid setae, slightly bent at their end; at the lower end, somewhat separated basally, a second row of five or six short, straight spinoid setae; particularly stout setae have not been found; valvae moderately long, slender, constricted medially, the distal fourth spatulate to some- what triangle-shaped and somewhat bent upwards; postbasally a row of three or four moderately long, nearly straight spinoid setae at the inner surface of the valvae; at the inner surface of the enlarged end a row of six or seven straight, short spinoid setae and some setae. 9 Genitalia. Tergite IX missing, sternite IX reduced, strongly constricted me- dially, not characteristic. Terminal papillae with sclerotization in an indistinct band; receptaculum seminis short and stout, the second half like a sac, with typical striation; vestibulum moderately large, simple like a sac. Diagnosis. M. myrtetella is very similar to M. rablensis and Micropterix croatica Heath & Kaltenbach, 1984 (all three species are of about the same size), but can be separated most easily by its golden spot at 3/4, which spreads across the whole width of the forewing. The male genitalia of all three species are also very similar. M. myrtetella can be recog- nized by the less stout shape of the whole genitalia and by its distinctly pronounced un- cus. Furthermore, the species can be separated from M. rablensis by its postbasal row of spines on the valvae. Very similar is Micropterix trinacriella Kurz, Zeller & Kurz, 1997 from Sicily, which can be distinguished easily by its genitalia. An undescribed species from central Italy (Micropterix wockei sensu auctorum) can be recognized for certainty only by examination of the genitalia. Individuals from southern and south-eastern Greece show a strong reduction of the wing markings (lacking the spot at 3/4 and the fascia in the middle), also often with 278 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe a brighter purple colouration. They are considered to be a distinct subspecies (Micro- pterix myrtetella idae Rebel, 1902). Distribution. According to Heath (1996) and Karsholt (2004) this species occurs in Italy, Austria, Hungarıa, Croatia, Macedonia, Yugoslavia, Romania, Bulgaria, Albania and Greece (see also Heath 1965b). Some of these records may refer to M. rablensis or Micropterix croatica Heath & Kaltenbach, 1984. The records from Slovakia and Czech Republic (Karsholt 2004) seem to be doubtful. The male specimen found in Austria (Gumpoldskirchen) has been verified by genitalic examination. Further confirmed records are from the type location in Italy, Croatia, Montenegro and Greece (NW-Greece, Pilion, Peloponnes). Life history. This species inhabits outskirts of medium canopy height to tall, dense mediterranean woody shrubland. Also at the type locality (Italy, near Livorno, Monte- nero) we found a female flyıng in Mediterranean maquis shrubland. Preimaginal stages. The early stages are unknown. Remarks. The holotype shows a small, but distinct costal spot at 2/3, a character miss- ing in most other specimens, especially those from the Balkans. A male of Micropterix myrtetella idae Rebel, 1902 from Peloponnes with dark scales on head (very similar to M. aruncella) also belongs to this species as recognized by its genitalia. Nota lepid. 30 (2): 235-298 279 Figs 39-42. Typical habitats of some Micropterix species. 39. Slope of moor with Alnus sp. association (M. mansuetella). 40. Nutrient-poor meadow with grass, Galium sp., Rumex sp., and shrubs along the side of a path at the fringe of a spruce forest in the montane zone (M. aruncella). 41. Edge of a spruce forest with areas of Vaccinium sp. and Frangula alnus in the lowlands (M. aureatella) 42. Moist meadow with Ranunculus sp. and shrubs at the fringe of a lowland coppice forest (M. calthella). ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Figs 43-46. Micropterix adults in their natural habitat. 43. M. mansuetella. Specimen resting on Alnus sp.: Austria, Eastern Tyrol, Lienzer Dolomiten, Lavant, Kienbichl, June 10, 2004. 44. M. aruncella. Specimen feeding on Plantago media. Austria, Vorarlberg, Bregenzer Wald, near Bezau, Sienspitze, upper Hinteregg- Alpe. 45. M. aureoviridella. Specimen feeding on Pinus mugo . Austria, Vorarlberg, Rätikon, Lünersee, near Douglashütte, 2000 m, July 30, 2004. 46. M. calthella (9, ©). In copula. Austria, Salzburg, Flachgau, Köstendorf, Tannberg, from the summit to the Lassbergweg, May 15, 2005. Nota lepid. 30 (2): 235-298 Figs 47-54. Forewings golden without any or with small silvery markings (first 0, second 9). 47-48 Micropterix calthella. 49-50. M. isobasella. 51-52. M. paykullella, unicolourous form with strongly re- duced markings, see also Figs 65-66. 53-54. M. aruncella, typical form, see also Figs 55-56. ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Figs 55-62. Forewings golden with silvery or coppery to purple markings (first d', second 9). 55-56. Micropterix aruncella f. atricapilla, see also Figs 53-54. 57-58. M. fenestrellensis. 59-60. M. mansuetel- la. 61-62. M. tunbergella. Nota lepid. 30 (2): 235-298 Figs 63-70. Forewings reddish golden to purple violet, with 2 golden fasciae and a spot at 3/4 (first ©, second ©). 63-64. Micropterix aureoviridella. 65-66. M. paykullella, typical form, see also Figs 51-52. 67-68. M. aglaella. 69-70. M. aureatella. ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Figs 71-78. Forewings brownish golden to blue-violet with 3 fasciae (first d', second 9). 71-72. Micropterix rablensis, © with aberration on forewing. 73-74. M. myrtetella. 75-76. M. trifasciella, @ With aberration on forewing. 77-78. M. huemeri, not mounted. Nota lepid. 30 (2): 235-298 Figs 79-86. Forewings with addional golden drawings at the outer and/or at the inner margin (first ©, second Q). 79-80. Micropterix allionella. 81-82. M. rothenbachii. 83-84. M. schaefferi. 85-86. M. ost- helderi. 286 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Figs 87-92. S genitalia. 87. Micropterix mansuetella. 88. M. calthella. 89. M. isobasella. 90. M. aglaella. 91. M. aureatella. 92. M. aruncella. Nota lepid. 30 (2): 235-298 287 Figs 93-98. © genitalia. 93. M. tunbergella. 94. M. aureoviridella. 95. M. paykullella. 96. M. allionella. 97. M. trifasciella. 98. M. rothenbachii. 288 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Figs 99-104. © genitalia. 99. M. huemeri. 100. M. schaefferi. 101. M. osthelderi. 102. M. rablensis. 103. M. myrtetella. 104. M. fene. trellensis (uncus destroyed, see Fig. 20). 289 235-298 Nota lepid. 30 (2) tella. 106. M. calthella. 107. M. isobasella. 108. M. cropterix mansue i 105-110. © abdomen. 105. M aglaella. 109. M. aureatella. 110. M. aruncella. Figs 290 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Figs 111-116. 9 abdomen. 111. M. tunbergella. 112. M. aureoviridella. 113. M. paykullella. 114. M. al- lionella. 115. M. trifasciella. 116. M. rothenbachii. Nota lepid. 30 (2): 235-298 291 Figs 117-121. 9 abdomen. 117. M. schaefferi. 118. M. osthelderi. 119. M. rablensis. 120. M. myrtetella. 121. M. fenestrellensis. 292 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Fig. 122. Geographical area considered for this work. 128 124 Figs 123-125. Different forms of thickened setae on the accessory claspers. 123. M. aureatella. 124. M. sicanella (distinctly three groups of setae) 125. M. schaefferi, T- or Y-shaped. Nota lepid. 30 (2): 235-298 293 Acknowledgements We are grateful to the following persons for providing data, images or literature, but also for many valu- able hints and for improving our English (in alphabetical order): Dr. Don Davis, Washington; Helmut Deutsch, Lienz; Prof. Gernot Embacher, Salzburg; Markward Fischer, Dresden; Dr. Mike Fitton, London; Prof. Dr. George Gibbs, New Zealand; Johannes Gillmann, Enzelsdorf; Dr. Theodor Grünewald, Landshut; Mag. Fritz Gusenleitner, Linz; Dr. Satoshi Hashimoto, Japan; Dr. Peter Huemer, Innsbruck: Ole Karsholt, Copenhagen; Dr. Josef Klimesch, Linz; Prof. Dr. Niels Peder Kristensen, Copenhagen; Jan Liska, Praha; Dr. Martin Lödl, Vienna; Fritz Mairhuber, Salzburg; Dr. Joël Minet, Paris; Dr. Matthias Nuss, Dresden; Norbert Pöll, Bad Ischl; Dr. Pierre Réal, Aix en Provence; Dr. Gaden Robinson, London; Dr. Klaus Sattler, London; Racheli Schwartz-Tzachor, Israel; Dr. Andreas Segerer, Munich; Dr. Wolfgang Speidel, Munich; Andreas Stübner, Jänschwalde-Ost; Francesca Vegliante, Dresden; Dr. Pierre Viette, Paris; Dr. Christian Wieser, Klagenfurt; Dr. Steven Whitebread, Quincy, USA; Josef Wimmer, Steyr. We particularly ac- knowledge many helpful comments during the review process from Ole Karsholt, Niels P. Kristensen and Matthias Nuss. References Adler, W., K. Oswald & R. Fischer 1994. Exkursionsflora von Österreich. — Ulmer, Stuttgart, Wien, I“ Ed. Amsel, 1936. Zur Kenntnis der Kleinschmetterlingsfaua Sardiniens. — Veröffentlichungen aus dem Deut- schem Kolonial- und Übersee Museum Bremen 1 (3): 344-365, pl. 15. Bentley, W. 1845. Description of Eriocephla sulcatella, a small new British moth of the family Tinei- dae. — The Zoologist: A Popular Miscellany of Natural History 3: 1086-1087. Burmann, K. & P. Huemer 1984. Die Kleinschmetterlingssammlung von Prof. Franz Gradl in der Vor- arlberger Naturschau, Dornbirn. — Berichte des naturwissenschaftlich-medizinischen Vereins in Inns- bruck, Suppl. 1: 1-64. Carter, D. J. & J. S. Dugdale 1982. Notes on collecting and rearing Micropterix (Lepidoptera: Micro- pterigidae) larvae in England. — Entomologist’s Gazette 33: 43-47. Chauvin, J. T. & G. Chauvin 1980. Formation des reliefs externes de l’oeuf de Micropteryx calthella L. (Lepidoptera: Micropterigidae). — Canadian Journal of Zoology 58: 161-766. Chauvin, G. & M. Faucheux 1981. Les pieces buccales et leurs recepteurs sensoriels chez l’imago de Micropterix calthella L. (Lepidoptera: Micropterigidae). — International Journal of Insect Morpho- logy and Embryology 10 (5/6): 425-439. Common, I. F. B. 1990. Moths of Australia. — Brill Academic Publishers, 535 pp. Corley, M. F. V. 2007. A brief review of the Micropterigidae of Portugal, with description of a new species of Micropterix Hiibner. — Nota lepidopterologica 30 (1): 71-78. Costa O.-G. 1834 [“1832-1836"]. Lepidotteri. — In: O. G. Costa & A. Costa [eds], Fauna del regno di Napoli ossia enumerazione di tutti gli animali che abitano le diverse regioni di questo regno e le acque che le bagnano contenénto la descrizione de nuovi 0 poco esattamente conosciuti. — Napoli (Dai Torchi del Trameter): pp. [1 ]-[314], pls 1-14. Curtis, J. 1839. British entomology: being illustrations and descriptions of the genera of insects found in Great Britain and Ireland: containing coloured figures from nature of the most rare and beautiful spe- cies, and in many instances of the plants upon which they are found, vol. 6. — London. Duponchel, P.-A. J. [1838-1840]. Nocturnes, 8. — /n: J.-B. Godart, Histoire Naturelle des Lépidoptères ou Papillons de France 11. — Paris, Méquignon-Marvis. 720 pp., pls 287-314. Embacher, G., M. Kurz & H. C. Zeller-Lukashort 2004. Beitrag zur Microlepidopterenfauna Salzburgs (Lepidoptera). — Beiträge zur Entomofaunistik 5: 57-66. Evenhuis N. L. 1997. Litteratura taxonomica dipterorum. (1758-1930). — 2 vols. 871 pp. Leiden Backhuys. Fabricius, J.C. 1777 [imprint “1776”]. Genera insectorum eorumque charcteres naturales secundum nu- merum, figuram, situm et proportionem omnium partium oris adiecta mantissa specierum nuper detec- tarum. — Chilonii, pp. [1]-[xii], 1-310. Fabricius, J. C. 1787. Mantissa Insectorum sistens species nuper detectas adiectis synonymis, observa- tionibus, descriptionibus, emendationibus 2. — Hafniae, pp. 1-382. Fabricius, J. C. 1794, Entomologica systematica emendata et aucta. 3 (2). 349pp. Hafniae. 294 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Frey, H. 1856: Die Tineen und Pterophoren der Schweiz. — Zürich. Gibbs, G. W. 1983. Evolution of Micropterigidae (Lepidoptera) in the SW Pacific. — GeoJournal 7.6: 505-510. Gibbs, G., Y. Kobayashi, H. Suzuki, S. Hashimoto, D. C. Lees, M. Sugimoto, & T. Saigusa 2004. Mole- cular phylogeny of Micropterigidae (Lepidoptera). — Proceedings of the XII International Congress of Entomology, Brisbane 15-21 August 2004. [Abstract 15_3304, on published CD]. Hamon, C. & G. Chauvin 1995. Larval integument amd its differentiations in Micropterix calthella L. (Lepidoptera: Micropterigidae): Anatomy and ultrastructure. — International Journal of Insect Morphology and Embryology 24: 213-222. Hannemann, H. J. 1956. Die Kopfmuskulatur von Micropteryx calthella (L) (Lep) Morphologie und Funcktion. — Zoologisches Jahrbuch für Anatomie 75: 177-206. Hannemann, H. J. 1957. Die männlichen Terminalia von Micropterix calthella L. (Lep. Micropterygi- dae). — Deutsche Entomologische Zeitschrift (Neue Folge) 4: 209-222. Hartig, F. 1964. Microlepidotteri della Venezia Tridentina e delle regioni adiacenti, parte III (Fam. Gelechii- dae - Micropterigidae). — Studi Trentini di Scienze Naturali 41 (3, 4): 276-278. Hartig, F. 1973. Beitrag zur Kenntnis der Verbreitung der Micropterygiden. — Nachrichtenblatt der Bayeri- schen Entomologen 22 (4): 75-78. Hashimoto, S. 2006. A taxonomic study of the family Micropterigidae (Lepidoptera, Micropterigoidea) of Japan, with the phylogenetic relationships among the Northern Hemisphere genera. — Bulletin of the Kitakyushu Museum of Natural History and Human History, ser. A (Natural History) 4: 39-109. Haworth, A.H. 1828. Lepidoptera Britannica, sistens digestionem novam insectorum lepidopterorum quae in Magna Britannia reperiuntur, lavarum pabulo, temporeque pascendi; expansione alarum; mensibus- que volandi; synonymis atque locis observationibusque variis. Part 4. Londini, pp. 513-609. Heath, J. 1957. The British Eriocraniidae and Micropterygidae. — Proceedings of the South London Ento- mological and Natural History Society: 115-125, pl. 8. Heath, J. 1960a. The food plants of adult Micropterygids (Lep.). — The Entörmologıet s Monthly Magazine 95: 188. Heath, J. 1962. The eggs of Micropteryx (Lep., Micropterygidae). — The Entomologist‘s Monthly Magazine 97: 179-181. Heath, J. 1965a. A new species of Micropterix (Lepidoptera Zeugloptera: Micropterigidae). — Estratto da- gli Atti della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano 104 (ID: 243-245. Heath, J. 1965b. Ergebnisse der Albanien-Expedition 1961 des Deutschen Entomologischen Institutes, 33. Beitrag, Lepidoptera: Micropterigidae. — Beiträge zur Entomologie 15: 641-647. Heath, J. 1975. The ammanella complex of the genus Micropterix Hübner [1825] (Lepidoptera: Zeugloptera, Micropterigidae). — Entomologist’s Gazette 26: 253-258. Heath, J., N. P. Kristensen & E. S. Nielsen 1979. On the identity of Tinea tunbergella Fabricius, 1787 and Tinea thunbergella Fabricius, 1794 (Lepidoptera: Micropterigidae, Gracillariidae). — Entomologica Scandanavica 10: 9-12. Heath, J. 1981a. Two new species of Micropterix Huebner (Lepidoptera, Zeugloptera: Micropterigi- dae). — Entomologist’s Gazette 32: 99-102. Heath, J. 1983. The moths and butterflies of Great Britain and Ireland, Vol.I: Micropterigidae — Heliozeli- dae. — Harley Books, Martins, Great Horkesley, Colchester, Essex. Heath, J. & Th. Kaltenbach 1984. New species of Micropterix Hübner (Lepidoptera; Zeugloptera: Micro- pterigidae) from Italy and Yugoslavia. — Entomologist’s Gazette 35: 21-23. Heath, J. 1986. The Micropterigid Fauna of North Africa. — Entomologist’s Gazette 37: 17-32, 126. Heath, J. 1987. A check list of the genus Micropterix Hübner [1825] (Lepidoptera: Zeugloptera, Micro- pterigidae). — Entomologist’s Gazette 38: 205-207. Heath, J. 1996. Family Micropterigidae — In: O. Karsholt & J. Razowski (eds), The Lepidoptera of Europa. A distributional checklist. — Apollo-Books, Stenstrup. Herrich-Schäffer, G. A. W. 1847-1855 [“1853-1855”]. Systematische Bearbeitung der Schmetterlinge von Europa, zugleich als Text, Revision und Supplement zu Jakob Hübner’s Sammlung eu- ropäischer Schmetterlinge. 5. Die Schaben und Federmotten: [1]-2-394 + [1]-2-52, 124 + 7 + 1 pl. Regensburg. Nota lepid. 30 (2): 235-298 295 Höfner, G. (1898): 3 neue Schmetterlingsarten. — Societas Entomologica 13 (9/10): 1-6. Hübner, J. 1796-1836 [,,1796°“] a. Sammlung europäischer Schmetterlinge. Horde 8. Tineae-Schaben: [1]-[12]+13-70+[71]-[78], pls 2-71. Augsburg. Continued by C. Geyer. Hübner, J. 1816-1826 [“1816”] b. Verzeichniß bekannter Schmettlinge [sic]: 1-431 + [1]-2-72. Augs- burg. Huemer, P. & G. Tarmann 1993. Die Schmetterlinge Österreichs (Lepidoptera). — Beilageband 5 zu den Veröffentlichungen des Museum Ferdinandeum, 224 pp., Tiroler Landesmuseum Ferdinandeum, Innsbruck. Karsholt, ©. 2004. Families Acanthopteroctetidae, Axiidae, Castniidae, Cossidae, Drepanidae, Eriocotti- dae, Eriocraniidae, Gelechiidae, Heterogynidae, Limacodidae, Lypusidae, Micropterigidae, Roesler- stammiidae, Somabrachyidae, Uraniidae. /n: Karsholt, O. & E. J. van Nieukerken (eds.). Lepidoptera, Moths. — Fauna Europaea version 1.1, http://www.faunaeur.org [online 16 December 2004]. Klausnitzer B., E. Meyer, W. Kössler & G. Eisenbeis 2002. On the larval morphology of Micropterix aruncella (Scopoli, 1963). — Beiträge zur Entomologie 52 (2002) 2: 353-366. Klimesch, J. 1990. Microlepidoptera (Kleinschmetterlinge) I. 17-20. - Jn: K. Kusdas & E. R. Reichl (ed.), Die Schmetterlinge Oberösterreichs, Teil 6. — Linz. Kobayashi, Y., H. Suzuki, S. Hashimoto, G. W. Gibbs, D. C. Lees, T. Saigusa & N. Sugimoto 2002. Phylogeny of the Micropterigidae of Europe and Circum-Pacific areas inferred from mitochondrial DNA sequences. — Annual Meeting of the Entomological Society of Japan [Abstract]. Moriuti, S. 1982. Micropterigidae. vol. 1: 41-43; vol. 2: 111, 113, 153, pls 235, 245, 246. — In: H. Inoue, S. Sugi, H. Kuroko, S. Moriuti & A. Kawabe. Moths of Japan 1+2.— Kodansha, Tokyo. Kozlov, M. V. 1985. Precopulatory behaviour of lower Lepidoptera. — Entomologicheskoye Obozre- niye 3: 493-505. Kozlov, M. V. 1986. Muscles of the pterothorax of primitive moths (Lepidoptera: Micropterigidae — Tischeriidae). — Vestnik Zoologii 1986 (1), 59-70 [in Russian, English summary]. Kozlov, M. V. 1988. Kratkij ozor ı opredelitelonaja tablitsa vidov roda Micropterix (Lepidoptera, Micro- pterigidae) palearktiki, part 1. — Vestnik zoologii 1988 (4): 8-14 [In Russian]. Kozlov, M. V. 1989. Kratkij ozor 1 opredelitelonaja tablitsa vidov roda Micropterix (Lepidoptera, Micro- pterigidae) palearktiki, part 2. — Vestnik zoologii 1989 (6): 26-31 [In Russian]. Kozlov, M. V. 1990a. Kratkij ozor 1 opredelitelonaja tablitsa vidov roda Micropterix (Lepidoptera, Micro- pterigidae) palearktiki, part 3. — Vestnik zoologii 1990 (2): 21-26 [In Russian]. Kozlov, M. V. 1990b. Kratkij ozor 1 opredelitelonaja tablitsa vidov roda Micropterix (Lepidoptera, Micro- pterigidae) palearktiki, part 4. — Vestnik zoologii 1990 (3): 28-33 [In Russian]. Kozlov, M. V. 1995. Wing pattern variation and population structure of Micropterix maschukella Alphé- raky (Lepidoptera: Micropterigidae). — Entomologist’s Gazette 46: 243-252. Kozlov, M. & E. Zvereva 2006. Aggregation of Micropterix maschukella moths on inflorescences of common elder: mating at foraging sites (Lepidoptera Micropterigidae). — Ethology Ecology & Evolution 18: 147-158. Kristensen, N. P. 1984a. Studies on the morphology and systematics of primitive Lepidoptera (Insec- ta). — Steenstrupia 10: 141-191. Kristensen, N. P. 1984b. The pregenital abdomen of the Zeugloptera (Lepidoptera). — Steenstrupia 10: 113-136. Kristensen, N. P. 1984c. Respiratory system of the primitive moth Micropterix calthella (Linnaeus) (Lepidoptera: Micropterigidae). — International Journal of Insect Morphology and Embryology 13: 137-156. Kristensen, N. P. 1998. The non-glossatan moths: 41-49. — In: N. P. Kristensen (ed.), Lepidoptera, Moths and Butterflies. Vol. 1: Evolution, systematics, and biogeography. — In: M. Fischer (ed.), Handbook of Zoology. Vol. IV Arthropoda: Insecta, Part 35. — Walter De Gruyter, Berlin and New York. Kristensen, N. P. & E. S. Nielsen 1979. A new subfamily of Micropterigid moths from South America. A contribution to the morphology and phylogeny of the Micropterigidae, with a generic catalogue of the family (Lepidoptera: Zeugloptera). — Steenstrupia 5: 69-147. Kristensen, N. P. & E. S. Nielsen 1982. South American micropterigid moths: two new genera of the Sabatinca-group (Lepidoptera: micropterigidae). — Entomologica Scandanavica 13: 513-529. 296 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Kristensen, N. P. & A. W. Skalski 1998. Phylogeny and paleontology. pp. 7-25. — In: N. P. Kristensen (ed.), Lepidoptera, Moths and Butterflies. Vol. 1: Evolution, systematics, and biogeography. — In: M. Fischer (ed.), Handbook of Zoology. Vol. IV Arthropoda: Insecta, Part 35. — Walter De Gruyter, Berlin and New York. Kurz, M. A., M. E. Kurz & H. C. Zeller-Lukashort 1994a. Neue und interessante Schmetterlingsfunde aus Salzburg und Oberösterreich (Lepidoptera). — Zeitschrift der Arbeitsgemeinschaft Österreichi- scher Entomologen 45 (1993): 113-116. Kurz, M. A., M. E. Kurz & H. C. Zeller-Lukashort 1994b. Neue und interessante Schmetterlingsfunde aus Salzburg: 4. Beitrag zur Landesfauna (Lepidoptera). — Entomologisches Nachrichtenblatt 1 (3/4): 18-23. Kurz, M. A., M. E. Kurz & H. C. Zeller-Lukashort 1997a. A new Micropterix species from northern Italy (Micropterigidae). — Nota lepidopterologica 20 (3/4): 293-298. Kurz, M. A. 2003. NKIS - Naturkundliches Informationssystem. — Beiträge zur Entomofaunistik 3 (2002): 190-191. Kurz, M. A., M. E. Kurz & H. C. Zeller-Lukashort 2004a. A new Micopterix species from southern France (Micropterigidae). — Nota lepidopterologica 26 (3/4): 111-114. Kurz, M. A., M. E. Kurz & H. C. Zeller-Lukashort 2000-2007. Naturkundliches Informationssystem. — URL: http://www.nkis.info [visited on February 1, 2007]. Le Cerf, F. 1926. Contribution à l’étude des organes sensoriels des Lépidoptères. — Encyclopédie Entomologique. B HI (Lepidoptera) 1 (3): 133-158. Linnaeus, C. 1761. Fauna Suecica (2nd Edition). — Stockholm. 578 pp. Leraut, P. 1980. Liste systématique et synonimique des Lépidoptères de France, Belgique et Corse. — Supplément 4 Alexanor et Bulletin de la Société Entomologique de France. 334 pp. Lorenz, R. E. 1959. Vorläufiger Bericht über meine Zuchtversuche mit Micropteryx calthella L. (Lep. Micr.). — Mitteilungen der Deutschen Entomologischen Gesellschaft 18 (3): 42-45. Lorenz, R. E. 1961. Biologie und Morphologie von Micropterix calthella (L.) (Lep. Micropterygidae). — Deutsche Entomologische Zeitschrift 8 (VID: 1-23. Luff, M. L. 1964. Larvae of Micropteryx (Lepidoptera, Micropterigidae). — Proceedings of the Royal Entomological Society of London (C) 29: 6. Martinova, E. F. 1950. On the structure of the larvae of Micropteryx (Lepidoptera, Micropterygi- dae). — Entomologicheskoye Obozreniye 31: 142-150 [In Russian]. Meeß, A. 1910. Micropterygideae. pp. 483-484. — In: A. Spuler (1903-1910), Die Schmetterlinge Euro- pas. — E. Schweitzerbart#sche Verlagsbuchhandlung, Stuttgart. 523 pp., 91 pls. Meyer E., W. Kössler & G. Klausnitzer 2002. Zur Kenntnis der Biologie und Ökologie von Micropterix aruncella (Scopoli) an der zentralalpinen Waldgrenze (Lep. Micropterigidae). — Entomologische Nachrichten und Beiträge 46 (1): 17-22. Meyrick, E. 1912. Adelidae, Micropterygidae, Gracilariadae. — /n: H. Wagner (ed.), Lepidopterorum Catalogus, pars 6, Berlin. Mitterberger, K. 1909. Verzeichnis der im Kronlande Salzburg bisher beobachteten Mikrolepidopteren (Kleinschmetterlinge). — Mitteilungen der Gesellschaft für Salzburger Landeskunde 49: 195-552. Mosher, E. 1916. A classification of the Lepidoptera based on characters of the pupa. — Bulletin of Illinois Laboratory 12: 17-159. Müller-Rutz, J. 1927. Die Schmetterlinge der Schweiz. — Mitteilungen der Schweizerischen Entomolo- gischen Gesellschaft 13: 532-533. Osthelder, L. 1951. Die Schmetterlinge Südbayerns und der angrenzenden nördlichen Kalkalpen. II. Die Kleinschmetterlinge. 2. Heft. — Beilage zu Mitteilungen der Münchner Entomologischen Gesell- schaft 41. Pankhurst, R. (ed.) 1999. Database of the Flora Europaea. — Royal Botanical Garden of Edinborough. — URL http://rbg-web2.rbge.org.uk/FE/fe.html [visited on September 26, 2007]. Primary types of Lepidoptera in the ZMUC collection — URL: http://www.zmuc.dk [visited on February 22007: Pringruber, ©. 1944. Blütenökologische Untersuchungen über einige Falter der Gattung Micropteryx Hbn. — Verhandlungen der kaiserlich - königlichen zoologisch-botanischen Gesellschaft in Wien 90-91: 129-220. Nota lepid. 30 (2): 235-298 297 Pröse, H. 1987. Kleinschmetterlinge: Wissensstand, Erhebungen und Artenschutzproblematik, Anhang: Artenliste der in Bayern und den angrenzenden Gebieten nachgewiesenen Microlepidoptera (Klein- schmetterlinge). — Schriftenreihe Bayerisches Landesamt für Umweltschutz 77: 37-102. Réal, P. 1988. Lépidoptères nouveaux principalement jurassiens. - Mémoires du Comité de Liaison pour les Recherches Ecofaunistiques dans le Jura (CL.E.R.J.), No. 4: 3-9. Robinson, G. S. & E. S. Nielsen 1983. The Microlepidoptera described by Linnaeus and Clerck. — Syste- matic Entomology 8: 191-242. Skalski, A. W. 1995. Study on the Lepidoptera from fossil resins. Part XI. Baltimartyria, a new genus for Micropteryx proavittella Rebel, 1936, with redescription of this species (Lepidoptera, Zeugloptera, Micropterigidae). — Amber and Fossils 1 (1): 26-37. Schwartz-Tzachor R., D. Eisikowitch & A. Dafni 2004. Non syndromic pollination of Cyclamen persicum in Israel. - Medecos, 10" International Conference on Mediterranean climate ecosystems, Rhodes island, Greece, 25 April — 1 May, 2004. Scoble, M. J. 1995. The Lepidoptera: Form and function and Diversity. — Oxford University Press, 424 pp. Scopoli, J. A. 1763. Entomologica Carniolica exhibens Insecta Carnioliae Indigena et Distributa in Ordines, Genera, Varietates. Methodo Linnaeana. — Vindobona, 421 pp, 43 pls. [Reprinted in Graz 1972]. Sherborn’s Index Animalium — URL. http://uio.mbl.edu/Sherborne/index.html [visited at April 10, 2007]. Stainton, H. T. 1850. A monograph of the British species of the genus Micropteryx of Zeller. — Transactions of the Entomological Society of London (N.S.) 1: 26-40. Staudinger, ©. 1870-1871. Beschreibung neuer Lepidopteren des europäischen Faunengebiets. — Berliner Entomologische Zeitschrift 14: 97-132, 193-208 (1870); 273-330 (1871). Stephens, J. F. 1834. Illustrations of Britsh Entomology or, a Synopsis of Indigenous Insects: contain- ing their generic and specific dinctions with an account of their metamorphoses, times of appear- ance, localities, food, and economy, as far as prcticable. — Insecta Haustellata 4: 436 pp, pls 23-40. London. The Global Lepidoptera Names Index — URL: http://internt.nhm.ac.uk/jdsml/research-curation/projects/ lepindex/index.dsml [visited on April 10, 2007]. Thien, L. B., P. Bernhardt, G. W. Gibbs, O. Pellmyr, G. Bergstrom, I. Groth & G. McPherson 1985. The pollination of Zygogynum (Winteraceae) by a moth, Sabatinca (Micropterigidae) - an ancient associa- tion. — Science 227 (4686): 540-543. Thunberg, C. P. 1794. Dissertatio Entomologica sistens Insecta suecica 7: 83—98, | pl. Upsaliae. Toll, S. 1942. Neue Microlepidopteren. — Zeitschrift der Wiener Entomologischen Gesellschaft 27: 166-173, pl. Tuskes, P. M. & N.J. Smith 1984. The life history and behavior of ,‘ Epimartyria pardella” (Micropterigidae). — Journal of the Lepidopterists’ Society 38 (1): 40-46. Viette, P. 1948. Lépidopteres homoneures. — Faune de France, Paris 49. Whitebread, S. 1992. The Micropterigidae of Switzerland, with a key to their identification (Lepido- ptera). — Nota lepidopterologica, suppl. 4: 129-143. Whitebread, S. 1995. Micropterigidae. — /n: O. Karsholt, E. J. van Nieukerken, S. Whitebread & S. Zang- heri 1995. Checklist delle specie delle fauna Italiana. — Edizione Calderini Bologna Fascicolo 80: 4-5. Wocke, M. F. 1877. Über einige wenig bekannte oder neue Falter der deutschen Fauna — Zeitschrift für Entomologie, Breslau 52. Wood, W. 1839. Index Entomologicus; or a complete Illustrated Catalogue consisting of 1944 figures of the Lepidopterous Insects of Great Britain. London, 1-266, pls 54. Zagulajev, A. K. 1978. Micropterygidae — In: G. S. Medvedev (ed.), Keys to the insects of the europaean part of the USSR, Vol. IV: Lepidoptera, part 1 [English translation, Oxonian Press Pvt. Ltd., New Dehli, 1987]. Zagulajev, A. K. 1983. New and little known species of the moths (Lepidoptera: Tineidae, Micropterigi- dae, Pterophoridae) from the USSR and adjacent territories. — Entomologicheskoye Obozreniye 62: 106-122 [In Russian]. Zeller, P. C. 1839. Versuch einer naturgemäßen Eintheilung der Schaben. - Isis, Leipzig [32]: 167-219. 298 ZELLER-LUKASHORT et al.: Micropterix of Northern and central Europe Zeller, P. C. 1844. Klassifikation und Benennung der Schmetterlinge nach Ochsenheimer — Treitschke bis zu den Wicklern einschließlich, von da nach Zeller. Fünfter Bericht des Schlesischen Tauschver- eins für Schmetterlinge, Brieg: 1-20. Zeller, P. C. 1850. Verzeichnis der von Herrn Jos. Mann beobachteten Toscanischen Microlepidoptera. — Stettiner Entomologische Zeitung 11: 59-64. Zeller, P. C. 1851. Drei Schabengattungen: /ncurvaria, Micropteryx und Nemophora. — Linnaea Entomo- logica 5: 301-362, tab. 1. Zeller, P. C. 1868. Lepidopterologische Ergebnisse einer Reise nach Oberkärnthen. — Stettiner Entomolo- gische Zeitung 29 (4-6): 121-149. Zeller, P. C. 1868 [“1869”]. Beitrag zur Kenntnis der Lepidoptern-Fauna der Umgegend von Raibl in Oberkärnthen und Preth im angrenzenden Küstengebiet. — Verhandlungen der kaiserlich-königlichen zoologisch-botanischen Gesellschaft in Wien 18: 563-628. Zeller-Lukashort, H. C., M. E. Kurz & M. A. Kurz 2002. Neue Funddaten von Arten der Gattung Micro- pterix Hübner, [1825] (Lepidoptera: Micropterigidae). — Zeitschrift der Arbeitsgemeinschaft Öster- reichischer Entomologen 54: 35-38. Zeller-Lukashort, H. C., M. E. Kurz & M. A. Kurz 2006. Die Micropterigidae der Alpen. — 73. Ento- mologentagung, Linz, Austria, November 3 — 5, 2006 — URL: http://www.nkis.info/nkis/vortrag/ DieMicropterigidaeAlpen.pps [visited on September 16, 2007]. ZOBODAT 1972-2007. Zoologisch-botanische Datenbank, Oberösterreichsiches Landesmuseum. — URL: http://www.zobodat.at [visited on January 1, 2007]. Nota lepid. 30 (2): 299-313 299 Mating behaviour and copulation mechanisms in the genus Scopula (Geometridae: Sterrhinae) PASI SIHVONEN Käärmekuusenpolku 4 C 11, 02880 Veikkola, Finland; e-mail: pasi.sihvonen@aka.fı Abstract. The mating behaviour and functional morphology of the copulatory organs in the sterrhine genus Scopula Schrank, 1802 (Lepidoptera: Geometridae) were studied in order to determine the role the male 8th sternite’s specialized structures (sclerotized cerata and membranous mappa) play during copula- tion. The study included field observations, i.e. specimens studied in situ, and laboratory experiments using the Palaearctic species S. immorata (Linnaeus, 1758) and S. frigidaria (Möschler, 1869). It was found for S. immorata that mate locating behaviour is vagrant, 1.e. males search for females, courtship behaviour is simple, copulation is carried out in the end-to-end position, taking place early in the morning (between 6 a.m. to 10 a.m.) and lasting between 60 and 90 minutes. Sclerotized cerata in S. immorata, especially the stout setae at their distal end, were found to brush against the sclerotized lamella antevaginalis of the female genitalia during copulation. The interaction of these structures suggests that the male organs are involved in the tactical sexual stimulation of the female. In S. frigidaria the structures associated with the 8" sternite of the male are rudimentary and do not touch the female at all. For neither species a mechanic function was observed for the mappa, and females of both species were found to be polygamous. During copulation the sclerotized parts of the internal genitalia of the male and female (cornutus and ductus bur- sae, respectively) were aligned to each other. Introduction Structures of the sclerotized parts of the genitalia are among the most important sources of character information in Lepidoptera systematics, their value being especially pro- nounced in alpha taxonomy. Despite the large amount of information that has been published on the morphology of these structures, we know little about their functional morphology. This mismatch has been acknowledged in the few papers that have been published on the subject (e.g. Miller 1988). Pioneering works on copulatory mechanisms in Lepidoptera are those by Chapman (1916) and Stekolnikov (e.g. 1967) and Stekolnikov and Kuznetsov (1982), but un- doubtedly the most remarkable contributions are those by Callahan (1958, 1960) and Callahan and Chapin (1960). These papers deal with a detailed analysis of copulation, spermatophore production, and egg formation in Noctuidae. A total of 11 species were examined, and a method of serial homology, or serial dissection, of moth pairs at vari- ous stages during copulation was presented. Scopula Schrank, 1802 is a widespread and most species-rich genus in the geometrid subfamily Sterrhinae (Sihvonen & Kaila 2004). Currently over 800 species are consid- ered valid, and the majority of species are presumably tropical (Sihvonen & Siljander 2005). Nearly all Scopula species possess sclerotized lateral appendices, called cerata (singular: ceras), on the 8" sternite of the males. Often these appendices are slightly curved inwards, of unequal length, and the apex of either one or both cerata bear a few stout setae (Fig. 4) (Hausmann 2004; Sihvonen 2005). Between the cerata lies a soft, flexible, membranous structure termed the mappa (Fig. 4). In most instances it is a bare structure, but a few species have long, flattened, scale-like setae in its caudal margin. The basal region of the 8" sternite, i.e. its cephalic part, often takes the form of Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 300 SIHVONEN: Copulation in the genus Scopula a sclerotized, convex, plate-like structure (Fig. 4). The cephalic margin of 8" sternite may be medially elongated, medially invaginated or trifid in shape (Sihvonen 2005). Sımilar structures are also found in closely related genera of Scopulini, for example in Lipomelia Warren, 1893, Problepsis Lederer, 1853, and Somatina Guenée, 1858 (Sihvonen 2005). These structures of the male 8" sternite are species-specific in shape, and their value as a diagnostic tool in alpha-taxonomy is well recognized. As a result, they are routinely illustrated in taxonomic works (e.g. Covell 1970; Hausmann 2004). Interestingly, in a number of species the cerata are polymorphic, i.e. their length varies within a species, and relative proportions of different length morphs vary geographically (Hausmann 1999; 2004). The function of these peculiar, specialised structures has remained un- known, but Hausmann (1999) assumed that the 8" sternite may play a mechanical role during copulation, perhaps similar to that noted in the geometrid genus Eupithecia Curtis, 1825 (Larentiinae) (Mikkola 1994). Indeed, in fresh specimens or specimens that have been macerated with potassium hydroxide, the cerata of a number of species can be moved laterally. This movement is allowed by a loose, membranous abdominal cuticle at a point where the cerata join the base of the 8" sternite. In other words, the abdominal skin flexes when the cerata are moved laterally (unpublished, personal ob- servation). The purpose of this study is to document the mating behaviour of S. immorata (Linnaeus, 1758), and in particular to investigate what function the above-mentioned specialized pregenital structures may play during copulation. The latter aspect was studied with the aid of two species, S. immorata (Linnaeus, 1758) and S. frigidaria (Möschler, 1869). Material and Methods The term ‘functional morphology’ has been used in the literature in various ways. I fol- low Naumann (1987) and use it to describe the interaction of male and female genitalia structures during copulation, 1.e. the mechanism of copulation. The terminology for genitalia follows Covell (1970), Klots (1970) and Kristensen (2003). Species studied and their genital structures The two species were chosen for the study on the grounds that morphological structures of their genitalia are quite different (Figs 1-9). It was hoped that inclusion of morpho- logically different species might help to better understand the copulation mechanisms and the possible role of male’s 8" sternite structure and so to allow wider interpretation of the results. S. immorata. A Euro-Siberian species, which is widely distributed in temperate areas. It is a thermophilous species preferring open habitats, including waste lands, mead- ows and road sides. The larva is polyphagous, feeding on withered leaves of herba- ceous plants, for example on Plantago (Plantaginaceae), Taraxacum (Asteraceae) and Hieracium (Asteraceae) (Ebert 2001; Hausmann 2004). The species is usually bivoltine in June and August, except in the northern and mountainous areas of its distribution, where the second generation is incomplete or absent. Nota lepid. 30 (2): 299-313 301 ad SOCIUS valvula _ sacculus sclerotized apex lamella antevaginalis 7 7 Figs 1-5. Male and female genitalia of Scopula immorata. 1. Male genitalia and phallus in ventral view, coremata removed (slide PS 1040). 2. phallus, vesica everted (PS1041). 3. 8th abdominal tergite (PS1041). 4. 8th abdominal sternite (PS1041). 5. Female genitalia in ventral view (PS1043). 302 SIHVONEN: Copulation in the genus Scopula Male genitalia (Fig. 1). Ovoid, symmetrical; socii membranous, covered with short setae; valvae consist of ventral sacculi, sclerotized and acute; and dorsal valvuli, membranous, blunt ending, setose. The sacculi have a small membrane at their proxi- mal part allowing lateral movement. Juxta with large lateral arms, widest apically; vin- culum large, U-shaped; transtilla weakly developed, membranous bridge. Phallus (Figs 1, 2) wide, straight; apex bearing one large and several small sclerotized teeth dors- oventrally; caecum slightly curved ventrally; vesica opens ventrally, with one large, blunt-ended diverticulum that opens to left (when viewed ventrally); cornutus large, acute; ductus ejaculatorius opens ventrally. 8" tergite weakly developed (Fig. 3); cerata of unequal length, apices covered with stout setae; mappa round; base partly concealed by membrane (Fig. 4). The cerata of this species have not been recorded to be polymor- phic (Hausmann 2004). Female genitalia (Fig. 5). Papillae anales soft, setose; apophyses posteriores and a. anteriores long; lamella postvaginalis sclerotized, narrow ring; lamella antevaginalis large, sclerotized, proximal membrane allows ventral movement; lamella antevaginalis conceals ostium bursae; ductus bursae wide, short, partly sclerotized; ductus seminalis ventrally from proximal part of ductus bursae; corpus bursae elongated sac; signum small, weakly spinulate. Female abdominal segments weakly sclerotized, undifferenti- ated. S. frigidaria. A Holarctic species, its distribution in the Palaearctic region being con- fined to its northernmost areas, whereas in the Nearctic region it is widespread from Alaska to Newfoundland. It occurs in open tundra habitats, but is also found in co- niferous and mixed forests. In the Palaearctic region the larva has been recorded to feed on the leaves of Vaccinium myrtillus (Ericaceae), in the Nearctic region immature stages are unrecorded (Covell 1970). The species is univoltine from early June to late August. Male genitalia (Fig. 6). Ovoid, symmetrical; valvae short, resulting in fused ap- pearance; socii long, narrow, covered with setae; sacculi melanized, short, acute; valvuli short, membraneous, setose; juxta round, lateral arms short; vinculum large, U-shaped; transtilla weakly develoed membranous bridge. Phallus (Figs 6, 7) long, narrow, cae- cum slightly curved ventrally; vesica simple sac, opening ventrally, without cornutus; base of ductus ejaculatorius weakly sclerotized. 8" tergite weakly developed (Fig. 8), cerata short, symmetrical; mappa round; base partly concealed by a membrane. The cerata of this species have not been recorded to be polymorphic (Hausmann 2004). Female genitalia (Fig. 9). Papillae anales soft, setose; apophyses posteriores and a. anteriores long; lamella postvaginalis reduced; lamella antevaginalis narrow, horse-shoe shaped; ostium bursae small, sclerotized, cap-shaped lobe; ductus bursae unsclerotized; ductus seminalis ventrally from proximal part of ductus bursae; corpus bursae elongated sac; signum large, consisting of separate spinules or absent altogether. Female abdominal segments weakly sclerotized, undifferentiated. Copulation experiments Field experiments using S. immorata, including observations and material acquisition, were carried out at various sites in southern Finland between 2000-2005. Laboratory- Nota lepid. 30 (2): 299-313 303 Ä pillae anales SOCIUS arm of juxta : ostium Ji valvula Te cag if bursae 1€ sacculus | \ , lamella _ | antevaginalis _cerata base — Figs 6-9. Male and female genitalia of Scopula frigidaria. 6. Male genitalia and phallus in ventral view, coremata removed (PS1044). 7. Phallus, vesica everted (PS1058). 8. 8th abdominal segment (PS1058), 9. female genitalia (PS1047). bred specimens were reared in plastic cages, about 25 x 25 x 15 cm in size. When moths had emerged, the breeding cages were taken outdoors. The cages had small holes on their sides and on top, allowing female pheromones to dispense. No field or laboratory experiments of live specimens were carried out for S. frigidaria. Entire mounts of copulating pairs were obtained from specimens caught in situ in the field, and from specimens that were bred from ova and which copulated in breeding cages. The specimens were anesthetized with chloroform and stored in a freezer. The abdomens were carefully separated from the thorax with the aid of fine scissors and macerated in a hot (7 mins at 92 °C) potassium hydroxide (KOH) solution. The ab- domens and genitalia were dissected in 5% aqueous ethanol. Scales were removed from abdomens with help of fine brushes, the organs were stained with a weak solu- tion of Chlorazol Black, and prepared following routine techniques (Hardwick 1950). 304 SIHVONEN: Copulation in the genus Scopula The male vesica was everted via the caecum that was cut open by placing the phallus inside a hypodermic syringe (Sıhvonen 2001). The structures were preserved tempo- rarily in glycerol, which allowed examination from various perspectives. They were subsequently transferred into ethanol tubes for 24 hours for glycerine removal and then mounted in Euparal. The cover glass was supported by small glass props. Structures were prepared using a Wild MZ6 stereomicroscope (maximum magnification 64x). Characters were examined using a Wild MIO (512x) and Leitz Diaplan phase contrast compound microscope (1560x). The number of matings was assessed by counting the number of spermatophores stored in the female bursa copulatrix. The method relies on the assumption that only one sper- matophore is transferred during each copulation (for criticisms, see Cordero 1999). Results Mating behaviour Males of S. immorata were found to express female-searching behaviour early in the morning, approximately from 6 to 10 a.m. This vagrant behaviour started soon, be- tween one to five minutes, after sun had started to warm up the observation site. The males flew rapidly, just above the low vegetation, in a zigzagging manner towards the wind. Every now and then males landed in the vegetation, beat their wings swiftly a few times, and continued flying after a while. Most females emerged from the pupa in the evening or during the night. They remained still and did not start to attract males until early in the morning, around 6 a.m. They stayed hidden in the vegetation, head facing upwards, and held the tip of the abdomen in an elevated position. After a male had located the female, they both remained with their heads upwards for a few seconds (Fig. 10), whereupon the male tried to locate the tip of the abdomen of the female, turning 180 degrees. To end up in this position, males always turned from the left-hand side (Figs 11, 12). It was when in this end-to-end posi- tion that the male attached its genitalia to the female and copulation started (Fig. 12). Copulation was observed to last 60 to 90 minutes (n = 9), and three phases were more or less identifiable each time: 1) Male moving its genitalia and 8" sternite back and forth (up and down), rubbing the cerata against the lamella antevaginalis of female. Cerata did not move laterally (Fre 12): 2) Male moving its abdomen from side to side in a snake-like manner including both lateral and dorso-ventral moves (Fig. 12). 3). When viewed ventrally (as in Fig. 13), male turned slightly to ‘left’, thus male and female were at an angle of about 160 degrees. Stages one and two lasted about 15 minutes each, stage three lasted from five to ten minutes, after which the male de- tached itself from the female. After copulation was over, the female remained still until the following night, when oviposition took place. In three instances copulating pairs Nota lepid. 30 (2): 299-313 305 \ N | \ | Figs 10-13. Sequence of events occurring during copulation in Scopula immorata in ventral view. Pictures were taken through transparent plastic. 10. Male approaches female by wing fanning, heads upwards (length of this phase is about 10 seconds). 11. Male contacts and flexes abdomen toward female genitalia and starts to turn to end-to-end position (approx. 10 seconds). Male always turned to end-to-end position from left-hand side (when viewed ventrally). 12. Copulation, end-to-end position (60 to 90 minutes). Male rubbed the cerata against the lamella antevaginalis of female, both back-and-forth and side-to-side movements were observed. 13. End of copulation, male has tilted to left, abdomens are at an angle of 160 degrees (5-10 minutes). remained attached to each other, in phase three (Fig. 12), until the following night. I found two copulating pairs in the field in the afternoon that were apparently in phase three of copulation. When disturbed, these separated from each other immediately and the male took to the wing. It was noted that males were more active in sunny weather, on cloudy mornings most males were not eager to fly at all. Vagrant behaviour of males ended by 10 a.m. The same behaviour pattern was observed in the laboratory-bred specimens, when breeding cages were taken outdoors. Both males and females were active at dusk and during the night, and both sexes are attracted to light. Despite an intensive search in the field, and observations made in the laboratory, no mate searching behaviour was observed at dusk 306 SIHVONEN: Copulation in the genus Scopula and during the night. Neither sex was observed to feed on nectar plants. Females were observed to lay eggs from dusk until around midnight. Field observations and laboratory experiments showed that in Scopula both sexes can copulate several times. Up to two spermatophores were found inside the corpus bur- sae of both studied species. One male S. immorata was observed to copulate with two different females under laboratory conditions. A further three males tried to copulate for the second time with a different female, but the proper copulation posture was not assumed. Interaction of male and female genitalia structures during copulation S. immorata. The membrane between the 8" abdominal segment and the genitalia is loose and partly folded. Prior to copulation, the male everts its genitalia from this mem- brane pocket, so that the end of its abdomen projects in a telescopic manner. Before the male inserted its phallus into the ostium bursae of the female, the valvae changed their position remarkably. The sclerotized, ventral sacculi were curved about 90 degrees me- dially and came into contact with the region of the female genitalia between the ventral margin of the ostium bursae and its ventral sclerotized ring. The membranous dorsal valvuli were curved even more, about 180 degrees ventrally and opened the lamella antevaginalis of the female genitalia, thus exposing the ostium bursae (Fig. 14). The lateral arms of the juxta came into contact with the areas lateral of the ostium bursae, apparently performing a support function. The socii were inserted into the membranous region between papillae anales and ostium bursae (Fig. 14). During copulation phases one and two (Fig. 12), when the male moved its body from side to side and back and forth, it rubbed its cerata against the sclerotized lamella an- tevaginalis of the female (Fig. 14) (Note: The illustrated specimens were slightly dis- torted when they were placed under a cover slip for photography. As a result the male cerata do not touch the lamella antevaginalis in the picture). No lateral or pincer-like movements of the cerata were observed. At the same time the male inserted its phallus into the ostium bursae and started to evert its vesica. Also, the mappa was found to be adjacent to the lamella antevaginalis during copulation. The vesica was everted into the cephalic part of the ductus bursae, beyond the branch- ing point of the ductus seminalis (Fig. 15). The sclerotized teeth at the apex of the phal- lus and cornutus were found to rest in a position at the inner wall of the ductus bursae, corresponding to the sclerotized parts on the inner wall of the ductus bursae of the female genitalia (Fig 15, Table 1). This position was noted in all pairs examined. When fully everted, the vesica blocks the entire ductus bursae but it does not reach the corpus bursae. The spermatophore is formed in about 30 minutes and it is stored inside the corpus bursae (Fig. 5). The spermatophore is a simple, unsclerotized sac, which loosely fills the entire corpus bursae. The number of spermatophores found in the corpus bursae varied from zero to two, with most specimens having one. The material examined did not allow for conclusions to be made about the function of the signum. S. frigidaria. As in S. immorata, the valvae of S. frigidaria changed their position quite remarkably (Figs 17, 18). The ventral sacculi were curved by about 90 degrees Nota lepid. 30 (2): 299-313 307 papillae anales 8th tergite scerotized margin of ostium bursae base of valva : valvula da # base of 8th sternite # lamella antevaginalis ee Fig. 14. Lateral view of Scopula immorata in copula (unstained) (PS 1059). Female is on the left, male is on the right. Valvula of male valvae have turned ventrally by about 180 degrees and mechanically push open the lamella antevaginalis of female genitalia allowing phallus to penetrate into the ostium bursae. Afterwards, sclerotized cerata rub against sclerotized lamella antevaginalis. medially and came into contact with the cap-shaped sclerotizations of the ostium bursae. The dorsal valvuli were curved even further, by about 180 degrees, and fit into the horse-shoe shaped pocket of the lamella antevaginalis. The socii were inserted on the sides of the papillae anales, the latter being pushed inwards towards the female abdomen. The proximal part of the ductus bursae has a dorsal, half-ringed weak sclerotization, ventrally the ductus wall is membranous. The ductus bursae is concave in shape (when viewed through a cross-section of the ductus bursae). When the male inserts its phallus into the ductus bursae, the weakly sclerotized dorsal wall opens and the ductus bursae assumes a round shape. The number of spermatophores found in the corpus bursae varied from zero to two, with most specimens having one. Unlike in S. immorata, the rudimentary cerata and large mappa of male 8" sternite were not observed to have any mechanic function during copulation (Fig. 16). Discussion Mating behaviour. For a good number of Lepidoptera species, especially for those that are nocturnal and small, we do not know the basic attributes of their biology. Among those aspects are behavioural traits, and to give a concrete example, the timing of mat- ing of the Small Tortoiseshell (Nymphalis urticae) was discovered only a few years 308 SIHVONEN: Copulation in the genus Scopula ago, despite this being a common and familar species that occurs widely throughout the Palaearctic region. Males of this species search for females in bright daylight, and when a female is found, the pair fly into a sheltered place and the actual copulation takes place around midnight (Mikkola er al. 2005). A similar lack of basic biological information for S. immorata made it difficult to study the mechanisms of its copula- tion because it was unknown at what time of the day copulation takes place. Rearing experiments revealed that mate location and copulation take place in the early morning sunshine. This was against the a priori assumption that a nocturnal species is likely to copulate during the dark hours of the day. It is unknown what kind of mate locating behaviour is predominant in the large, cos- mopolitan genus Scopula. The strategy where males search for passive females in the morning sunshine, is here reported for S. immorata. In addition, I have made simi- lar field observations for the Palaearctic S. ternata (Schrank, 1802) and S. floslactata (Haworth, 1809) (unpublished). The vagrant strategy, combined with early morning activity, may therefore be a common strategy at least for the North Palaearctic species of the genus. The courtship behaviour of S. immorata was found to be simple; after the male had located the female, copulation occurred in a tail-to-tail position (the male remained fac- ing in the opposite direction as the female). The tail-to-tail position is typical for most Lepidoptera (for an overview, see Phelan & Baker 1990; for exceptions see Charlton & Cardé 1990). It is likely, however, that more complex and interactive courtship behav- iour prior to copulation is widespread in Scopula and related genera of Scopulini. The reason for this is that unlike in S. immorata, which has vestigial hair pencils on its hind legs, males of a number of other Scopulini species possess specialized, eversible scent emitting structures in their hind tibiae, termed hair pencils (Hashimoto 1992, Sihvonen 2005). Some of these secondary sexual structures are very pronounced, e.g. in Scopula urnaria (Guenée, 1858, illustrated in Sihvonen 2005). Further, a few species of closely related genera possess eversible scent structures on the male 8" abdominal segment (e.g. Pseudasellodes fenestraria (Guenée) or on the hind wings (e.g. Crypsityla quin- quelineata (Dognin), illustrated in Sihvonen & Kaila 2004). Possibly, species whose males either lack or have vestigial scent-emitting secondary sexual characters, such as S. immorata, have a simple courtship behaviour whereas species with such structures have a more complex and interactive courtship behaviour. In an extensive comparative study that was carried out on phycitine moths (Pyralidae), it was found that all spe- cies displaying the interactive courtship pattern possessed male scent-disseminating structures on the forewing and/or the eight abdominal segment, whereas those species engaging in a simple courtship had neither of those structures or only vestiges of them (Phelan & Baker 1990). Functional morphology of the male and female genitalia. The specific function of the male 8" sternite in Scopula and closely related genera during copulation is still unclear. Mikkola (1994) suggested for certain species of North American Eupithecia, which possess species-specific structures similar to cerata, that the recognition func- tion between sexes during the early phase of copulation has shifted from the valvae to Nota lepid. 30 (2): 299-313 309 N Fig. 15. Lateral view of female genitalia (left, PS1060) and male phallus (right, PS1041) of Scopula immorata. The structures are turned to show the anatomical correspondence during copulation. During copulation sclerotized structures of male phallus and female ductus bursae were adjacent to each other, see Table 1 for details. Tab. 1. Corresponding anatomical structures of male and female internal genitalia of Scopula immorata. Male phallus and vesica Female bursa copulatrix le do cine a sclerotized projection in the ductus Apex of phallus with dorsal teeth Ductus bursae slightly sclerotized dorsally Diverticulum in phallus Bi. Ventral margin of phallus angled Ventral margin of ductus bursae angled Ductus ejaculatorius opens ventrally Ductus seminalis opens ventrally Cornutus at apex of vesica (direction in Ductus bursae sclerotized ventrally and slides variable) laterally these structures. This hypothesis is supported by the observation that valvae of those Eupithecia Species are simple and uniform, compared to Lepidoptera valvae in general, which are often species-specific in structure. The material I examined does not allow to draw conclusions for or against this proposed recognition function. The hypothesis of Hausmann (1999) that the male 8" sternite may play a mechanical role during copulation is confirmed but the exact function remains unexplained. It was observed that sclerotized cerata of the male, especially the stout setae at their distal end, come into contact with the sclerotized lamella antevaginalis of the female during copulation. The male moves the distal end of the abdomen laterally and back-and-forth in a snake-like manner, with the result that the cerata rub against the sclerotized lamella antevaginalis. This mechanical movement suggests that the male organs are likely to be involved in tactical stimulation of the female. Besides sclerotizations of the female, I have failed to find any other morphologically detectable surface or cuticular structures 310 SIHVONEN: Copulation in the genus Scopula that could be sensory in nature. To achieve this conclusion, I have studied unsclerotized areas in 15 female S. immorata, in addition to routine dissections of female genitalia of more than 400 Scopula species. It is emphasized that the cerata of S. immorata are non-articulated at their cephalad part, therefore I doubt that the male can move its cerata independently without mov- ing its abdomen. There are a number of Scopula species, e.g. S. floslactata (Haworth, 1809), S. transsecta (Warren, 1898) and S. submutata (Treitschke, 1828), to mention a few, whose cerata are mediocephalically unsclerotized and which are articulated to a loose abdominal skin fold at the base of the 8" sternite. It is likely that those species are capable of moving their cerata passively in a pincer-like manner when they come into contact with female abdomen. This kind of ability to move sclerotized structures of the male 8" sternite, similar to cerata, has been suggested for certain North American spe- cies of Eupithecia (Mikkola 1994). The study was not based on in situ experiments, and it remained unverified whether the male could move those structures actively or not. As was mentioned in the Introduction, in a number of Scopula species the cerata are polymorphic, 1.e. their length varies within a species, and relative proportions of dif- ferent length morphs vary geographically (Hausmann 1999, 2004). This polymorphism of genitalia observed by Hausmann (1999, 2004) is unique among insects; I have man- aged to find data only on a species of Thysanoptera (thrips), in which winged and apterous male specimens differ remarkably with respect to the size of their genitalia (Mound er al. 1998), on water striders (Arnqvist & Danielsson 1999) and on Selenia te- tralunaria (Hufnagel, 1767) (Geometridae: Ennominae), in which the genitalia showed negative allometry in relation to body size (Mutanen & Kaitala 2006). The latter argue, however, that although genital dimorphism is rarely documented, it is quite expected. The scarcity of observations may be partly as a result of circular reasoning as species are often delimited by differences in genital characteristics (Mutanen & Kaitala 2006). The observed scrubbing function of the cerata in S. immorata does not offer clues that could explain the possible role of cerata in polymorphic species. If the main function of cerata is mechanic in nature, then in specimens with short cerata these most likely can- not perform the same mechanic function as in specimens with long cerata. If the main function of cerata is sexual stimulation or titillation, then polymorphism of this organ may be quite common within the species, or even expected, if one accepts that the main evolutionary force affecting genitalic diversification is sexual selection (Arnqvist 1997; Jocqué 2002; for a review on the subject see Hosken & Stockley 2004). Specifically, according to the cryptic female choice hypothesis of Eberhard (1985, 1996), polyan- drous females are able to control the sperm that fertilize their eggs, and males with the best ability to stimulate females during copulation are favoured. A study on the subject by Arnqvist and Danielsson (1999) dealt with fertilization success and the shape of male genital sclerites (plates) in a water strider when females mated with two males. After controlling for body size, it was found that sclerite shape influenced fertiliza- tion success. Further, the sexual selection hypothesis predicts that genitalia of species with polyandrous mating systems should be more divergent than those of monandrous species (see Arnqvist 1998). These findings and hypotheses are relevant in the context of this paper, because against this theoretical setting species of Scopula, and also of Nota lepid. 30 (2): 299-313 311 8th tergite papillae anales socil tegumen base of phallus rudimentary ceras ase of 8th sternite valvula _ valvula sacculus #4 sacculus & “ | base of phallus 16 | 17 Y oe 148 Fig. 16. Lateral view of Scopula frigidaria in copula (unstained). Female is on the left, male is on the right. Rudimentary cerata do not touch female genitalia during copulation. Figs 17-18. Male genitalia in ventral view. 17. Position of sacculus, valvula and phallus before copulation (PS1040). 18. Position of sacculus, valvula and phallus during copulation (PS414). Scopulini in general, seem a very attractive target group for experiments studying the mechanisms of genitalia diversification. It would be interesting to know how monandry (if it exists in Scopula and in Scopulini) vs. polyandry of Scopula and Scopulini species correlates with polymorphic vs. non-polymorphic genitalia (see Arnqvist 1997). For this purpose detailed experiments on their basic biology need to carried out. In non- polymorphic S. frigidaria the cerata are rudimentary and apparently do not come into contact at all with the female genitalia during copulation. The male genitalia have a rather fused appearance in Scopula, in a number of species this is due to the reduction in length of the valvae and enlargement of the immobile vin- culum. Thus it was quite surprising to learn that the sacculi and valvuli of the valva are flexible structures (Figs 17, 18). Especially the latter undergoes a quite drastic change in posture when the valvuli turn by 180 degrees ventromedially and they mechanically ‘open’ the lamella antevaginalis of the female genitalia before copulation (Fig. 15). Prior to copulation, the socii provide a dorsal support, and the lateral arms of the juxta provide lateral support for the male and female genitalia. The support function of the juxta in general is pronounced in many North American species of Scopula, e.g. in S. inductata (Guenée, 1858), S. luteolata (Hulst, 1880) and S. sideraria (Guenée, 1858) (illustrated in Covell 1970), where the juxta is transformed into an elongated, round tube that corresponds to the sclerotized ductus bursae of the female genitalia. This is not unique in Lepidoptera, however, but more typically precopulation support is pro- 312 SIHVONEN: Copulation in the genus Scopula vided by a combination of uncus and valvae. When copulation proper occurs, i.e. the male has everted its vesica into the bursa copulatrix of the female, the support function is further facilitated by internal genitalia. This is also true for S. immorata. Acknowledgements This study was financially supported by the Lepidopterological Society of Finland and Societas pro Fauna et Flora Fennica. Karoliina Reunanen is thanked for editing the figures and two anonymous referees are thanked for valuable comments, including linguistic corrections. References Arngvist, G. 1997. The evolution of animal genitalia: distinguishing between hypotheses by single species studies. — Biological Journal of the Linnean Society 60: 365-379. Arngvist, G. 1998. Comparative evidence for the evolution of genitalia by sexual selection. — Nature 393: 784-786. Arngvist, G. & Danielsson, I. 1999. Copulatory behavior, genital morphology and male fertilization suc- cess ın water striders. — Evolution 53: 147-156. Callahan, P. S. 1958. Serial morphology as a technique for determination of reproductive patterns in the corn earworm, Heliothis zea (Boddie). — Annals of the Entomological Society of America 51: 413-428. Callahan, P. S. 1960. A morphological study of spermatophore placement and mating in the subfamily Plusiinae (Noctuidae, Lepidoptera). — Proceedings of the XI International Congress of Entomology, Vienna, 1: 339-345. | Callahan, P. S. & Chapin, J. B. 1960. Morphology of the reproductive systems and mating in two repre- sentative members of the family Noctuidae, Pseudaletia unipuncta and Peridroma margaritosa, with comparisons to Heliothis zea. — Annals of the Entomological Society of America 53: 763-782. Charlton, R. E. & Cardé, R. T. 1990. Behavioral interactions in the courtship of Lymantria dispar (Lepi- doptera: Lymantriidae). — Annals of the entomological Society of America 83: 89-96. Chapman, T. A. 1916. On the pairing of the Plebeiid Blue Butterfly (Lycaeninae, tribe Plebeiidi). — Trans- actions of the entomological Society of London 1916: 156-180. Cordero, C. 1999. Is spermatophore number a good measure of mating frequency in female Callophrys xami (Lycaenidae). — Journal of the Lepidopterists’ Society 53: 169-170. Covell, C. V. 1970. A revision of the North American species of the genus Scopula (Lepidoptera: Geome- tridae). — Transactions of the American Entomological Society 96: 101-221. Eberhard, W. G. 1985. Sexual selection and animal genitalia. — Harvard University Press, Cambridge. Eberhard, W. G. 1996. Female control: sexual selection by cryptic female choice. — Princeton University Press, Princeton. Ebert, G. (ed.) 2001. Die Schmetterlinge Baden-Wiirttembergs, Band 8: Nachtfalter VI, Geometridae. — Verlag E. Ulmer, Stuttgart, 541 pp. Hardwick, D. F. 1950. Preparation of slide mounts of lepidopterous genitalia. - Canadian Entomologist 82: 231-235. Hashimoto S. 1992. Tibial scent organ and its related structures in the genera /daea and Scopula of the subfamily Sterrhinae (Lepidoptera: Geometridae). — Akitu 130: 1-7. Hausmann, A. 1999. Falsification of an entomological rule: polymorphic genitalia in Geometrid moths. — Spixiana 22: 83-90. Hausmann, A. 2004. Sterrhinae. Pp. 1-600. — In: A. Hausmann (ed.), The Geometrid moths of Europe 2. — Apollo Books, Stenstrup. Hosken, D. J. & Stockley, P. 2004. Sexual selection and genital evolution. — Trends in Ecology and Evolution 19: 87-93. Jocqué, R. 2002. Genitalic polymorphism: a challenge for taxonomy. — Journal of Arachnology 30: 298-306. Nota lepid. 30 (2): 299-313 313 Klots, A. B. 1970. Lepidoptera. Pp. 115-130. - In: S. L. Tuxen (ed.), Taxonomists’ glossary of genitalia in insects. — Copenhagen, Munksgaard. Kristensen, N. P. 2003. Skeleton and muscles: adults. Pp. 39-122. In: N. P. Kristensen (ed.), Lepido- ptera, Moths and Butterflies, vol. 2: Morphology, Physiology, and Development. Handbook of Zoo- logy, volume IV, Arthropoda: Insecta, part 36. — Walter de Gruyter, Berlin. Mikkola, K. 1994. Inferences about the function of genitalia in the genus Eupithecia, with description of a new organ (Lepidoptera, Geometridae). — Nota lepidopterologica, supplement No. 5: 73-78. Mikkola, K., Murtosaari, J. & Nissinen, K. 2005. Perhosten lumo. Suomalainen perhostieto [in Finnish]. — Tammi, Helsinki, 343 pp. Miller, J. S. 1988. External genitalic morphology and copulatory mechanism of Cyanotricha necyria (Felder) (Dioptidae). — Journal of the Lepidopterists’ Society 42: 103-115. Mound, L. A., Crespi, B. J. & Tucker, A. 1998. Polymorphism and kleptoparasitism in thrips (Thysano- ptera: Phlaeothripidae) from woody galls on Casuarina trees. — Australian Journal of Entomology 37: 8-16. Mutanen, M. & Kaitala, A. 2006. Genital variation in a dimorphic moth Selenia tetralunaria (Lepidoptera: Ennominae). — Biological Journal of the Linnean Society 87: 297-307. Naumann, C. M. 1987. Functional morphology of the external male and female genitalia in Zygaena Fabricius, 1775 (Lepidoptera: Zygaenidae). — Entomologica Scandinavica 18: 213-219. Phelan, P. L. & Baker, T. C. 1990. Comparative study of courtship in twelve Phycitine moths (Lepidoptera: Pyralidae). — Journal of Insect Behavior 3: 303-326. Sihvonen, P. 2001. Everted vesicae of the Timandra griseata group: methodology and differential features (Geometridae, Sterrhinae). — Nota lepidopterologica 24: 57-63. Sihvonen, P. 2005. Phylogeny and classification of the Scopulini moths (Lepidoptera: Geometridae, Ster- rhinae). — Zoological Journal of the Linnean Society 143: 473-530. Sihvonen, P. & Kaila L. 2004. Phylogeny and tribal classification of Sterrhinae with emphasis on deli- miting Scopulini (Lepidoptera: Geometridae). — Systematic Entomology 29: 324-358. Sihvonen, P. & Siljander, M. 2005. Species diversity and geographical distribution of the Scopulini moths on a worldwide scale: (Lepidoptera: Geometridae, Sterrhinae). — Biodiversity and Conservation 14: 703-721. Stekolnikov, A. A. 1967. Phylogenetic relationships within the Rhopalocera on the basis of the functional morphology of the genital apparatus. — Entomological Review 46: 1-11. Stekolnikov, A. A. & Kuznetsov, V. I. 1982. Functional morphology of the male genitalia and a new tribal division of the Ennominae (Lepidoptera: Geometridae). — Entomological Review 61: 92-122. Nota lepid. 30 (2): 315-327 35 The nomenclature of the family-group names of Eupterotidae (Bombycoidea) WOLFGANG A. Nässıc !:* & ROLF G. OBERPRIELER ? ' Entomologie II, Forschungsinstitut und Museum Senckenberg, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany; e-mail: wolfgang.naessig@senckenberg.de ? Zimmerman Fellow, CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia; e-mail: rolf.oberprieler@csiro.au 3 Studies in Eupterotidae no. 6 Abstract. The origin and historical use of all family-group names pertaining to the bombycoid family Eupterotidae are determined. The family-group name Eupterotidae takes Swinhoe, 1892 as authorship, although the name was evidently originally coined by Hampson, 1893. However, three older family-group names are available for the taxon: Striphnopterygidae Wallengren, 1858, Phialidae Wallengren, 1865 and Janidae Aurivillius, 1892. Striphnopterygidae has been used as the valid name for the family by at least ten authors until at least 1965 and again in 2006, and as a valid subfamily name throughout, but the younger Eupterotidae gained almost universal acceptance as the name of the family in the literature since about 1928. Application of Article 35.5, introduced in the current, 4th edition of the Code (ICZN 1999), allows the retention of the younger name Eupterotidae as the valid name of the family. The authorship of another bombycoid family name, Lemoniidae, is corrected from Hampson, 1918 to Neumoegen & Dyar, 1894. Introduction The bombycoid family Eupterotidae contains about 53 genera, the majority of them occurring in the Ethiopian and Oriental regions but a few also in the eastern Palaearctic, the Australian region and Central America. The classification and phylogeny of the group remains inadequately studied, the major recent works being Forbes (1955), Minet (1994) and Oberprieler et al. (2003). The family was deemed to be monophyletic by Minet (1994) and Lemaire & Minet (1998) based on three synapomorphies, but Oberprieler et al. (2003) demonstrated that two of them cannot be regarded as such, leaving the family defined on a single (weak) character and its monophyly under some doubt. However, a recent study of the phylogenetic relationships of the Anthelidae (Zwick 2006) provides some molecular support for the monophyly of the Eupterotidae. The division of the family into natural subfamilies and tribes is also not yet settled. Following Aurivillius (1901a), early classifications (e.g., Distant 1903; Gaede 1927) generally recognised only two subfamilies, Striphnopteryginae and Janinae, although American authors often also included Apatelodes Packard, 1864 and related genera in Eupterotidae as a subfamily Apatelodinae (e.g., Grote 1896; Forbes 1923; Remington 1954). In his revision of the entire family, Forbes (1955) added to it the Australian Panacelinae and Asian Prismostictinae and divided the Eupterotinae into five tribes, Janini, Tissangini, Eupterotini (including Striphnopteryx), Phialini and Cotanini. Minet (1994) again restricted the concept of the family by excluding Apatelodinae and Prismostictinae and recognised five subfamilies: Hibrildinae, Tissanginae, Janinae, Eupterotinae (including Striphnopteryginae) and Panacelinae (including Cotanini). Oberprieler et al. (2003) included Tissanginae and Hibrildinae in Janinae and Cotanini in Eupterotinae, treated Striphnopteryginae as a separate subfamily and identified an informal “Ganisa group” that does not fit into any of the formal subfamilies. Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 316 NAssic & OBERPRIELER: On eupterotid family group names Apart from these taxonomic issues, the nomenclature of the family is also subject to some confusion, regarding both its valid name and its precise authorship. This issue resurfaced recently when the older name Striphnopterygidae was again used for the family (Mielke & Casagrande 2006), despite changes introduced to the latest edition of the International Code of Zoological Nomenclature (ICZN 1999) designed to preserve the stability of widely used family-group names when threatened by older but less often used ones. In this paper we clarıfy the nomenclature and synonymy of all the family- group names applying to the taxon Eupterotidae. The authorship of another bombycoid family name, Lemoniidae, is corrected at the same time. The family-group names The reference point to the use of the family name Eupterotidae in all recent litera- ture appears to be Fletcher & Nye (1982: viii), who gave its authorship as “Swinhoe, 1892”. Earlier works, however, either explicitly (Aurivillius 1894b, 1901a; Dyar 1895; Distant 1903; Bryk 1944; Griveaud 1962) or seemingly implicitly credited the name to “Hampson, 1892”. Neither Swinhoe nor Hampson stated specifically that they established a new family, but they were evidently in contact with each other at the time and seemingly coordinated their classification systems. In his preface, Hampson (1893b: 111) wrote that he hoped to have “evolved” a natural classification scheme of the Heterocera, to replace the outdated one of Guenée, and he provided a description of the family Eupterotidae, whereas Swinhoe (1892) only gave the family name and, in the preface to his work, thanked Hampson for “working out the genera’. Shortly before he had still treated Eupterote in Lasiocampidae (Swinhoe 1891), the usual classifica- tion by British authors at the time (e.g., Butler 1881; Moore 1883; Cotes & Swinhoe 1887; Kirby 1892). Also Distant (1892), describing a new species in Eupterotidae, thanked Hampson for deciding its “true generic position”. No contemporary author ever credited Swinhoe with the family name, and therefore it appears that it was, in fact, Hampson who originally coined the name Eupterotidae. However, the Moths Volume I of the Fauna of British India series, in which Hampson’s description of the family Eupterotidae was published, only became available in 1893 (see Fletcher & Nye 1982, Nassig & Nye 1991, and the bibliographical notice of Butler 1893), and hence both Swinhoe (1892) and Distant (1892) published the name Eupterotidae before Hampson. In both these works the name meets the conditions of Art. 11.7.1 of the Code (ICZN 1999) and is thus available. Swinhoe’s Catalogue only bears a publication date of 1892, but the preface is dated | October and the copy in the Macrolepidoptera Sectional Library of the Natural History Museum, London, is annotated ”Nov 4th* in Hampson’s handwriting (Kitching, pers. comm. 2007), taken here as the earliest day on which the work is demonstrated to have been in existence as a published work (Art. 21.3). Hampson’s (1893a) bibliographical notice of Swinhoe’s Catalogue, issued in January 1893, confirms that the latter was indeed published in 1892. The publication date of Distant’s (1892) paper is given in the journal as “November 1892”, for nomenclatu- ral purposes deemed to be the 30th November (Art. 21.3.2). Consequently, the name Eupterotidae must take Swinhoe, 1892 for its authorship. Nota lepid. 30 (2): 315-327 37 Hampson may have simply based the name of the family on the oldest included genus, Eupterote Hübner, 1820, but, although no such origin is cited, he may also have taken the concept from Hübner’s original name “Eupterotae”. In fact, Grote (1896: 45) ex- plicitly credited the name Eupterotidae to Hübner, not to Hampson. Hübner (1820: 187) used the plural noun “Eupterotae” in his category of Verein, or Coitus, and included in it two species, Eupterote fabia Cramer and E. petosiris Cramer. Hemming (1937: 16) es- tablished that Hübner’s Coitus is equivalent to the genus category in modern nomencla- ture, and “Eupterotae” must therefore be regarded as the original citation of the genus name Eupterote Hübner, 1820 (“1816”). Furthermore, plural nouns merely referring to members of a genus are not considered valid family-group names (Art. 11.7.1.2). Therefore, the family name Eupterotidae cannot take Hübner, 1820 for ıts authorship. Describing numerous new genera of moths particularly from the expeditions of Delegorgue and Wahlberg in southern Africa, Wallengren (1858: 210) proposed a new family, “Fam. Striphnopteryges”, for two new genera, Striphnopteryx from “Caffraria” (South Africa) and Festra from “N. Hollandia” (Australia), the latter a synonym of Chelepteryx Gray, 1835 (“1836”), now placed in Anthelidae. A few years later he sup- plied a proper description of the family (Wallengren 1865: 28) and added the genus Homochroa Wallengren, 1858 (a junior synonym of Phyllalia Walker, 1855, and a jun- ior homonym; see Fletcher & Nye 1982). In the same paper he also described another new family, “Fam. Phialide Wallengr.”, for a “new” genus, Phiala (already validly de- scribed by Wallengren 1860, see Fletcher & Nye 1982, and a junior subjective synonym of Euchera Hübner, 1825 (“1816”), see Vari et al. 2002: 150, 219). Wallengren (1875) later described another species of Phiala in the Phialidae and Aurivillius (1879) a new genus, Trichophiala. Aurivillius (1892: 195) proposed another new family, Janidae, for a new species of Jana Herrich-Schäffer, 1854 (“1850-1858”) from Cameroon. The Häfte 2-3 of vol. 13 of the Entomologisk Tidskrift, in which this family name was pub- lished, was issued on | June 1892, Janidae thus also taking priority over Eupterotidae Swinhoe, 1892. Aurivillius (1893) placed some other African genera in Janidae and also the Asian genus Melanothrix C. & R. Felder, 1874 (Aurivillius 1894a) but then synonymised both Phialidae and Janidae with Striphnopterygidae (Aurivillius 1894b). The family name Janidae was afterwards apparently only used three times, by Strand (1911a, b) and, somewhat surprisingly, Aurivillius (1921). Aurivillius was a firm proponent of the principle of priority in taxonomic nomenclature, which was, at the time, not yet officially entrenched in a universal code of zoological no- menclature (ICZN 1999: XXI). Being familiar with the African as well as the Asian fau- nas of bombycoids, Aurivillius further concluded that Wallengren’s Striphnopterygidae and Phialidae, his Janidae and Hampson’s Eupterotidae together constituted a “natural unit”, which had to take the oldest Striphnopterygidae as its valid name, and he clearly tabulated this synonymy of family names (Aurivillius 1894b: 185). He consistently used Striphnopterygidae in all his later publications dealing with this group of moths (Aurivillius 190 1a, b, 1904, 1905a, b, 1906, 1909, 1910, 1911, 1914, 1925a, b), and this concept and name was also adopted by many contemporary German-speaking authors (e.g., Karsch 1895, 1898a, b; Weymer 1909; Grünberg 1910; Strand 1909, 1911c, 1927; Schultze 1915; Krausse & Wolff 1919), but not by, e.g., Semper (1896), Rebel (1900, 318 NAssic & OBERPRIELER: On eupterotid family group names 1914) and the authors of the chapters on Eupterotidae in Seitz’ Gross-Schmetterlinge der Erde (Grünberg 1911; Strand 1922; Gaede 1927; Draudt 1928). English-speak- ing authors, however, used the name Eupterotidae throughout, often dealing only with the Asian or Australian fauna (the “true” eupterotines) (e.g., Hampson 1896, 1918; Bethune-Baker 1904, 1908; Swinhoe 1905; Rothschild 1917a; Turner 1922, 1947) but sometimes also including African (striphnopterygine and janine) genera (Butler 1898; Hampson 1910; Rothschild 1917b; Platt 1921). Distant (1903) was an exception in using Striphnopterygidae for the South African fauna. The American authors, too, fol- lowing Schaus (1894), placed their Apatelodes (or Apatelodinae) in Eupterotidae and apparently always used this name (e.g., Dyar 1895, 1901; Grote 1896; Fracker 1915; Forbes 1923), except Holland (1920), who placed African taxa in Striphnopterygidae. In some cases the use of the name Eupterotidae may have arisen out of ignorance of Aurivillius’ work, but in others (e.g., Rebel 1900; Hampson 1901; Gaede 1927) it evi- dently flowed out of a rejection or disregard of the priority principle. After Aurivillius’ death in 1928, Striphnopterygidae almost completely disappeared from the literature as a family name in favour of Eupterotidae (e.g., Candèze 1927; de Joannis 1929; van Eecke 1930; Mell 1930, 1937; Golding 1940; Bryk 1944; Sevastopulo 1944; Remington 1954; Forbes 1955; Berger 1958, 1980; Griveaud 1962; Kuznetsov 1967; Fletcher 1968; Brock 1971; D’Abrera 1974; Fontaine 1975; Pinhey 1975; Holloway 1976, 1987; Dall’Asta 1979; Fletcher & Nye 1982; Inoue et al. 1982; Barlow 1983; Vari & Kroon 1986; Holloway et al. 1987, 2001; Nässig 1989, 1995, 2000; Viette 1990; Common 1990; Nielsen & Common 1991: Scoble 1992; Kishida 1992; Minet 1994; Oberprieler & Duke 1994; Becker 1996; Edwards 1996; Lemaire & Minet 1998; Heppner 1998; Kroon 1999; Deml & Nässig 2001; Robinson et al. 2001; Vari et al. 2002; Oberprieler et al. 2003; Fu & Tuzuoo 2004), although Striphnopteryginae per- sisted as a valid subfamily name throughout (e.g., Gaede 1927; Fontaine 1975; Pinhey 1975; Vari & Kroon 1986; Oberprieler et al. 2003). A lone exception was Taylor (1949, 1950, 1951, 1957, 1961, 1965) in South Africa, who continued using Stripnopterygidae as the name of the family. No further uses of this family name have come to light until the recent bibliographic catalogue of ordinal names in Lepidoptera (Mielke & Casagrande 2006), which again listed Stripnopterygidae as the valid family name over Eupterotidae. Additional family-group names were proposed in the Eupterotidae by Forbes (1955) — Tissangini, Cotanini and Panacelinae (seemingly also Phialini, not cited as of Wallengren, 1865) —, and Minet (1994) added Hibrildinae, proposed as a family by Berger (1958). None of the family-group names pertaining to this taxon of moths is contained in the official lists of valid names in zoology (Hemming & Noakes 1958a; Melville & Smith 1987; Smith 2001; ICZN 2006) or in the index of rejected and invalid family-group names (Hemming & Noakes 1958b). Their priority is as follows: Striphnopterygidae Wallengren, 1858: 210 (type genus: Striphnopteryx Wallengren, 1858) Phialidae Wallengren, 1865: 33 (type genus: Phiala Wallengren, 1860 [= Euchera Hübner, 1825 (“1816”)]) Nota lepid. 30 (2): 315-327 319 Janidae Aurivillius, 1892 [1 June]: 195 (type genus: Jana Herrich-Schäffer, 1854 (“1850-1858”)) Eupterotidae Swinhoe, 1892 [4 November]: 408 (type genus: Eupterote Hübner, 1820 (“1816”)) Tissangini Forbes, 1955: 98 (type genus: Tissanga Aurivillius, 1903) Panacelinae Forbes, 1955: 120 (type genus: Panacela Walker, 1865) Cotanini Forbes, 1955: 131 (type genus: Cotana Walker, 1865) Hibrildidae Berger, 1958: 73 (type genus: Hibrildes Druce, 1883). The valid family name From the above overview of the history of the family-group names pertaining to the taxon in question we conclude that: the name Eupterotidae, although evidently orıginally coined by Hampson, takes Swinhoe, 1892 for its authorship; when the genera Striphnopteryx, Euchera (= Phiala), Jana and Eupterote are placed in the same family-group taxon, Striphnopterygidae Wallengren, 1858 ıs the oldest available family-group name for it; Striphnopterygidae Wallengren, 1858 was used as the valid name for the family by at least ten authors until at least 1965, and again in 2006, and as a valid subfamily or tribe name continuously into the present; the junior synonym Eupterotidae Swinhoe, 1892 gained almost universal accept- ance as the name of the family after about 1928, the priority of Striphnopterygidae, Phialidae and Janidae being overlooked or ignored. In strict compliance with the Principle of Priority ICZN 1999, Art. 23), the name of the family taxon in question should therefore be Striphnopterygidae. The moderation of this principle as regulated by Art. 23.9 (reversal of precedence) does not apply, as the conditions of Art. 23.9.1.1 are not met since Striphnopterygidae has been used as a valid family-group name well after 1899. A further moderation of the priority principle specifically in the family-group category was added in the current, 4th edition of the Code (ICZN 1999), preserving, after 1999, prevailing usage of a younger name when threatened by an older name in use for a category of lower rank (Art. 35.5). However, since this moderation is not specifically included in Art. 23, and Art. 35.5 does not ex- plicitly state that it moderates Art. 23, only the general statement in Art. 23.1 “unless ... another name is given precedence by any provision of the Code” provides grounds to regard Art. 35.5 as a valid moderation of Art. 23. Further ambiguity exists in Art. 35.5 in that it stipulates that the older name (of the lower-ranked taxon) has to be “in use” and, in the example given, that its precedence over the younger name (of the higher-ranked taxon) is maintained (only) as long as both names “are used for different subfamilies [= lower-ranked taxa] within the [family, = higher-ranked taxon]”. This implies that, if the older name is not in use for a separate taxon (i.e. is a plain older synonym of the same taxon), Art. 35.5 does not apply and the matter has to be settled by application of Art. 23. 320 NAssic & OBERPRIELER: On eupterotid family group names In Eupterotidae, Art. 35.5 is readily applicable as, in the current classification system (Oberprieler et al. 2003), the older name is in use for a different lower-ranked taxon (subfamily Striphnopteryginae) than that denoted by the younger name (subfamily Eupterotinae), and the name in prevailing usage for the higher-ranked taxon (family Eupterotidae) is therefore to be maintained. Before 2000, i.e. before Art. 35.5 came into existence, the family would have had to be called Striphnopterygidae, in compliance with Art. 23. In this group of moths, the current family-group names and their synonymies (follow- ing Oberprieler et al. 2003) therefore are: Family Eupterotidae Swinhoe, 1892 Subfamily Janinae Aurivillius, 1892 (= Tissanginae Forbes, 1955; = Hibrildinae Berger, 1958) Subfamily Striphnopteryginae Wallengren, 1858 (= Phialinae Wallengren, 1865) Subfamily Eupterotinae Swinhoe, 1892 (= Cotaninae Forbes, 1955) Subfamily Panacelinae Forbes, 1955 The confusion surrounding the names of this family, and the time and effort spent on resolving it, highlight the urgent need for stabilisation of the Lepidoptera family-group names and their authorships. The only currently available attempt to do so (Fletcher & Nye 1982, and other volumes of the Generic Names of Moths of the World series) is clearly inadequate; only for the Noctuoidea a more complete catalogue has recently been compiled (Speidel & Naumann 2005). As a further example, the bombycoid family name Lemontidae does not take Hampson, 1918 for its authorship (Fletcher & Nye 1982: viii) but in fact Neumoegen & Dyar, 1894 (as Lemoniinae — Neumoegen & Dyar 1894: 121), thus avoiding the invocation of Art. 35.5 to preserve this family name over Sabaliidae Hampson, 1901 (in the current concept of the family Lemoniidae). Likewise, the correction of the family-group name Lemoniidae Kirby, 1871, based on the riodinid butterfly genus Lemonias Hubner, 1807, to Lemoniadidae by Hall & Heppner (1999, as Lemoniadini) had already been effected a century earlier by Rebel (1900), who then also proposed a new family Lemoniidae for Lemonia Hübner, 1820 (“1816”). Rebel (1900) therefore did not create a homonym with Kirby’s name, but he evidently did not know that Neumoegen & Dyar (1894) had already proposed this family name earlier, and a homonymy between the Kirby and the Neumoegen & Dyar family-group names thus existed only between 1894 and 1900 — even though Rebel (1914) himself later incorrectly used “Lemoniidae” for the but- terfly group again. An analogous situation involving family-group names in Coleoptera was addressed by the compilation of a comprehensive synopsis of family and subfamily names (Lawrence & Newton 1995). While necessitating a number of name changes at the time (under the previous edition of the Code), this list provides an invaluable reference point to beetle family-group names and their authorships and greatly stabilises their nomenclature. We believe that a similar effort is urgently needed in Lepidoptera. The recent bibliographic catalogue of ordinal names (Mielke & Casagrande 2006) provides a first step in this direction but needs to be expanded to the family level. Nota lepid. 30 (2): 315-327 3 Acknowledgments We sincerely thank Bruce Halliday, Andreas Zwick and Ted Edwards (CSIRO Entomology, Canberra) for thoughtful discussion and constructive comments on this matter and for reading the manuscript; Miguel Alonso-Zarazaga (Museo Nacional de Ciencias Naturales, Madrid) for advice on zoological nomen- clature; Katherine Drayson (Oxford University Museum of Natural History, Oxford), Bert Gustafsson (Naturhistoriska Riksmuseet, Stockholm) and Ian Kitching (The Natural History Museum, London) for tracing critical publication dates; Melanie Grimm (Forschungsinstitut und Museum Senckenberg, Frankfurt am Main), the staff of the university library (Universitätsbibliothek) Johann Christian Senckenberg (Frankfurt am Main) and Trudi Prideaux (CSIRO Black Mountain Library, Canberra) for procuring vari- ous old publications; and two anonymous referees for their comments on the manuscript. References Aurivillius, C. 1879. Lepidoptera Damarensia. Förteckning pä fjärilar insamlade i Damaralandet af G. de Vylder ären 1873 och 1874 jemte beskrifning öfver förut okända arter. — Ofversigt af Kongliga Veten- skaps-Akademiens Förhandlingar 36 (7): 39-69. Aurivillius, C. 1892. Verzeichniss einer vom Herrn Fritz Theorin aus Gabun und dem Gebiete des Came- runflusses heimgebrachten Schmetterlingssammlung. — Entomologisk Tidskrift 13 (3): 169-177. Aurivillius, C. 1893. Diagnosen neuer Lepidopteren aus Afrika. — Entomologisk Tidskrift 14: 199-214. Aurivillius, C. 1894a. Neue Spinner aus Asien. — Entomologisk Tidskrift 15: 169-177. Aurivillius, C. 1894b. Die palaearktischen Gattungen der Lasiocampiden, Striphnopterygiden und Mega- lopygiden. — Deutsche Entomologische Zeitschrift “Iris” 7: 121-192. Aurivillius, C. 1901a. On the Ethiopian genera of the family Striphnopterygidae. — Bihang till Kongliga Svenska Vetenskaps-Akademiens Handlingar 27 Afd. 4, (7): 1-33. Aurivillius, C. 1901b. Diagnosen neuer Lepidopteren aus Afrika. — Entomologisk Tidskrift 22: 113-128. Aurivillius, C. 1904. New species of African Striphnopterygidae, Notodontidae, and Chrysopolomidae in the British Museum. — Transactions of the Entomological Society of London 1904 (4): 695-700, col. pl. XXX. Aurivillius, C. 1905a. Beiträge zur Kenntnis der Insektenfauna von Kamerun. No. 11. Lepidoptera Hetero- cera. — Arkiv för Zoologi 2 (4): 1-68, 1 col. pl. Aurivillius, C. 1905b. Leutnant A. Schultzes Sammlung von Lepidopteren aus West-Afrika. — Arkiv för Zoologi 2 (12): 1-47, 5 col. pls. Aurivillius, C. 1906. Verzeichnis von Lepidopteren, gesammelt bei Mukimbunga am unteren Kongo von Herrn E. Laman. - Arkiv för Zoologi, 3 (1): 1-16. Aurivillius, C. 1909. Diagnosen neuer Lepidopteren aus Afrika. — Arkiv för Zoologi 5 (1): 1-29. Aurivillius, C. 1910. Lepidoptera. Jn: Wissenschaftliche Ergebnisse der schwedischen zoologischen Ex- pedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Ostafrikas 1905-1906 unter Leitung von Prof. Dr. Yngve Sjöstedt. 2. Band, Abteilung 9: 1-56. — Stockholm (P. Palmquist). Aurivillius, C. 1911. New genera and species of Striphnopterygidae and Lasiocampidae in the British Mu- seum. — Transactions of the Entomological Society of London 1911 (1): 161-167. Aurivillius, C. 1914. New species of African Lasiocampidae and Striphnopterygidae from English collec- tions. — Arkiv för Zoologi 9 (11): 1-9. Aurivillius, C. 1921. 4. — Descriptions of some South African Heterocera (Lepidoptera). — Annals of the South African Museum 18: 235-244. Aurivillius, C. 1925a. Zoological results of the Swedish expedition to Central Africa 1921. Insecta. 12. Lepidoptera. 1. — Arkiv för Zoologi 17A (32): 1-20. Aurivillius, C. 1925b. Lepidoptera. IV. Teil. Lieferung 18. — Pp. 1243-1359 in: Schubotz, H. (ed.). Ergeb- nisse der Zweiten Deutschen Zentral-Afrika-Expedition 1910-1911 unter der Führung Adolf Fried- richs, Herzogs zu Mecklenburg. Band I: Zoologie. — Leipzig (Klinkhardt & Biermann). Barlow, H. S. 1983 (“1982”). An introduction to the moths of South East Asia. — Kuala Lumpur (Malayan Nature Society); x + 305 pp., 50 pls. [including: Taxonomic appendix by J. D. Holloway, pp. 174-253 plus genitalia drawings]. Becker, V. 1996. 106. Eupterotidae. pp. 3, 13. — /n: J. B. Heppner (ed.), Atlas of Neotropical Lepidoptera. Vol. 5B, Checklist: Part 4B. Drepanoidea — Bombycoidea — Sphingoidea. — Gainesville (ATL, Scientif. Publ.). 322 NAssic & OBERPRIELER: On eupterotid family group names Berger, L. A. 1958 (“1957”). Clé pour la détermination des Familles de Macrolépidoptères et des groupes supérieurs de Microlépidoptères (Faune Ethiopienne). — Lambillionea 57 (9/10): 72-84. Berger, L. A. 1980. Description d’un nouveau Jana Walker (Lep. Eupterotidae). — Lambillionea 80 (5/6): 36-37. Bethune-Baker, G. T. 1904. New Lepidoptera from British New Guinea. — Novitates zoologicae 11 (2): 367-429. Bethune-Baker, G. T. 1908. New Heterocera from British New Guinea. — Novitates zoologicae 15 (1): 175-243. Brock, J. P. 1971. A contribution towards an understanding of the morphology and phylogeny of the Di- trysian Lepidoptera. — Journal of Natural History 5: 29-102. Bryk, F. 1944. Entomological results from the Swedish expedition 1934 to Burma and British India. Lepi- doptera: Saturniidae, Bombycidae, Eupterotidae, Uraniidae, Epiblemidae und Sphingidae. Gesammelt von René Malaise. — Arkiv för Zoologi 35A (8): 1-55, 6 pls. Butler, A. G. 1881. Illustrations of typical specimens of Lepidoptera Heterocera in the collection of the British Museum, Part V. — London (Trustees of the BMNH), xii + 74 pp., pls LXXVIII-C. Butler, A. G. 1893 [April]. Bibliographical notice. The Fauna of British India, including Ceylon and Bur- ma. — Annals and Magazine of Natural History (6) 11 (64): 410-411. Butler, A. G. 1898. On a collection of Lepidoptera made in British East Africa by Mr. C. S. Betton. — Pro- ceedings of the Zoological Society of London 1898: 395-444. Candèze, L. 1927. Lépidoptères Hétérocères de l’Indochine Française. — Encyclopédie entomologique, Ser. B, II, Lepidoptera 2 (2, 3/4): 73-133. Common, I. F. B. 1990. Moths of Australia. — Leiden, New York, Kobenhavn, Köln (E. J. Brill), 535 pp. + 32 col. pls. Cotes, E. C., & C. Swinhoe 1887. A catalogue of the moths of India. Part II, Bombyces. — Calcutta (Trustees of the Indian Museum), 812 pp. D’Abrera, B. 1974. Moths of Australia. — Melbourne (Lansdowne Press), 79 [+ 5] pp. de Joannis, J. 1929. Eupterotidae, in: Lépidopteres hétérocères du Tonkin. — Annales de la Société ento- mologique de France 98: 447-449 [separate reprint of 1930, pp. 215-217]. Dall’ Asta, U. 1979. Le genre Janomima Auriv. (Lepidoptera, Eupterotidae). — Revue de zoologie africaine 93 (4): 912-918. Deml, R., & W. A. Nässig 2001. Secondary compounds in larvae of Eupterotidae (Lepidoptera). — Zoo- logischer Anzeiger 240: 209-210. Distant, W. L. 1892. Contributions to a knowledge of the entomology of the Transvaal. — Annals and Magazine of Natural History (6) 10: 407-408. Distant, W. L. 1903. Order Lepidoptera, suborder Heterocera. pp. 49-96, pls IH-VII. — Jn: W. L. Distant (1900-1911), Insecta Transvaaliensia. A contribution to a knowledge of the entomology of South Africa, Vol. 1.— London (W. L. Distant), iv + 299 pp. Draudt, M. 1928. 8. Familie, Eupterotidae. pp. 629, pl. 86. — In: A. Seitz [ed.] (1913-1940), Die Gross- Schmetterlinge der Erde. Eine systematische Bearbeitung der bis jetzt bekannten Gross-Schmetterlinge, 6. Band. Die Amerikanischen Spinner und Schwärmer. — Stuttgart (A. Kernen), VIII + 1452 pp., VIII pp. + pls 1-185. Dyar, H. G. 1895. Remarks on Apatelodes suggested by an article by Mr. Schaus. — The Canadian Ento- mologist 27 (6): 159-160. Dyar, H. G. 1901. Life history of Callidapteryx dryopterata Grt. — Proceedings of the Entomological So- ciety of Washington 4: 414-418. Edwards, E. D. 1996. 78. Eupterotidae. — In: E. S. Nielsen, E. D. Edwards & T. V. Rangsi (Eds.), Checklist of the Lepidoptera of Australia. - Monographs of Australian Lepidoptera, 4: 261 (CSIRO Publishing: Collingwood). Fletcher, D. S. 1968. Cossidae, Metarbelidae, Psychidae, Limacodidae, Drepanidae, Uraniidae, Lasio- campidae, Eupterotidae, Bombycidae, Saturniidae & Sphingidae. — Ruwenzori Expedition 1952 1 (8): 325-348. Fletcher, D. S., & I. W. B. Nye 1982. Bombycoidea, Castnioidea, Cossoidea, Mimallonoidea, Sesioidea, Sphingoidea, Zygaenoidea. — In: I. W. B. Nye (ed.), The generic names of moths of the world, vol. 4. - London (Trustees of the British Museum (Natural History)), xiv + 192 pp. Nota lepid. 30 (2): 315-327 525 Fontaine, M. 1975. Un élevage ab ovo de Sabalia tippelskirchi Karsh [sic!] (Lép. Eupterotidae — Stryphnopteryginae [sic!] selon M. Gaede in Seitz, F. Eth. pp. 301-302). — Lambillionea 98 bis (volume jubilaire): 36-39. Forbes, W. T. M. 1923. The Lepidoptera of New York and neighboring states. Primitive forms, Micro- lepidoptera, Pyraloids, Bombyces. — Cornell University Agricultural Experiment Station Memoir 68: 1-729. Forbes, W. T. M. 1955. The subdivision of the Eupterotidae (Lepidoptera). — Tijdschrift voor Entomologie 98 (2): 85-132. Fracker, S. B. 1915. The classification of lepidopterous larvae. — Illinois Biological Monographs 2 (1): 1-169. Fu,C.M. & H.R. Tuzuoo 2004. Moths of Anmashan, Part 2. — Taichung (Taiwan, Taichung Nature Research Society), 263 pp. Gaede, M. 1927. 11. Familie: Eupterotidae. pp. 293-311, pls 41-47. — In: A. Seitz (ed.) 1926-1930, Die Gross-Schmetterlinge der Erde, Band 14, Die afrikanischen Spinner und Schwärmer. — Stuttgart (A. Kernen), 599 pp., 80 pls. Golding, F. D. 1940. Further notes on the food-plants of Nigerian insects. V. — Bulletin of Entomological Research 31: 127-130. Griveaud, P. 1962 (“1961”). Insectes. Lépidoptères Eupterotidae et Attacidae. — Faune de Madagascar 14: 64 pp., 12 pls. Grote, A. R. 1896. List of North American Eupterotidae, Ptilodontae, Thyatyiridae, Apatelidae and Agro- tidae. - Abhandlungen, Naturwissenschaftlicher Verein zu Bremen 14: 44-128. Grünberg, K. 1910. H) Lepidoptera. — Jn: Schultze, L. (ed.) Zoologische und anthropologische Ergeb- nisse einer Forschungsreise im westlichen und zentralen Südafrika. — Denkschrift der medizinisch- naturwissenschaftlichen Gesellschaft zu Jena 16: 91-146. Grünberg, K. 1911. 9. Familie: Eupterotidae. pp. 185-199, pls 29-30. - In: A. Seitz (ed.), 1909-1913, Die Gross-Schmetterlinge der Erde, Band 2, Die palaearktischen Spinner & Schwärmer. — Stuttgart (A. Kernen), VII + 479 + [III] + 479 pp., 56 pls. Hall, J. P. W., & J. B. Heppner 1999. Lemoniadini, a corrected tribal name in the Riodinidae (Lepidoptera: Riodininae). — Tropical Lepidoptera 10 (1): 30. Hampson, G. F. 1893a [January]. Bibliographical notice. Catalogue of Eastern and Australian Lepidoptera Heterocera ...— Annals and Magazine of Natural History (6) 11 (61): 58-59. Hampson, G. F. 1893b (“1892”). The Fauna of British India, including Ceylon and Burma. Moths, vol. 1. — London (Taylor & Francis), xxiii + 527 pp. Hampson, G. F. 1896. The Fauna of British India, including Ceylon and Burma. Moths, vol. 4. — London (Taylor & Francis), xxvin + 594 pp. Hampson, G. F. 1901. A classification of a new family of the Lepidoptera. — Transactions of the Ento-mo- logical Society of London 1901: 187-192. Hampson, G. F. 1910. Zoological collections from Northern Rhodesia and adjacent territories: Lepidoptera Phalaenae. — Proceedings of the Zoological Society of London 1910: 388-510. Hampson, G. F. 1918. Some small families of the Lepidoptera which are not included in the key to 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. — Novitates zoologicae 25: 366-394. Hemming, F. 1937. Hübner. A bibliographical and systematic account of the entomological works of Jacob Hiibner and of the supplements thereto by Carl Geyer, Gottfried Franz von Frohlich and Gottlieb August Wilhelm Herrich-Schäffer. Vol. I. — London (Royal Entomological Society of London), xxxiv + 605 pp. Hemming, F., & D. Noakes 1958a. Official list of family-group names in zoology. First instalment: Names 1-236. — London (International Trust for Zoological Nomenclature), 38 pp. Hemming, F., & D. Noakes 1958b. Official index of rejected and invalid family-group names in zoo- logy. First instalment: Names 1-273. — London (International Trust for Zoological Nomenclature), 38 pp. Heppner, J.B. 1998. Classification of Lepidoptera. Part 1. Introduction. — Holarctic Lepidoptera 5 (Suppl. 1): v + 148 + 6 pp. 324 NÄssıG & OBERPRIELER: On eupterotid family group names Holland, W. J. 1920. Lepidoptera of the Congo, being a systematic list of the butterflies and moths col- lected by the American Museum of Natural History Congo Expedition, together with descriptions of some hitherto undescribed species. — Bulletin of the American Museum of Natural History 43 (6): 109-369. Holloway, J. D. 1976. Moths of Borneo with special reference to Mount Kinabalu. — Kuala Lumpur (Malay. Nature Soc./Sabah Found.); 264 pp. Holloway, J. D. 1987. The moths of Borneo, part 3, [internal title: Superfamily Bombycoidea], Lasiocam- pidae, Eupterotidae, Bombycidae, Brahmaeidae, Saturniidae, Sphingidae. — Kuala Lumpur (South- dene), 199 pp., 163 b & w pls, 20 col. pls. Holloway, J. D., J. D. Bradley & D. J. Carter 1987. CIE guides to insects of importance to man (ed. C. R. Betts), 1. Lepidoptera. — London (CAB Internat. Inst. Entomol./Brit. Mus. Nat. Hist.), 262 pp. Holloway, J. D., G. Kibby & D. Peggie 2001. The families of Malesian moths and butterflies. (Fauna Malesiana Handbook 3.) — Leiden, Boston, Köln (Brill), x1 + 455 pp. Hübner, J. 1816-[1826]. Verzeichniß bekannter Schmettlinge [sic]. — Augsburg (the author), pp. 1-16 (1816), pp. 17-176 ([1819]), pp. 177-208 ([1820]), pp. 209-256 ([1821]), pp. 257-304 ([1823]), pp. 305-431 ([1825]), Anzeiger, pp. 1-72 ([1826]). ICZN 1999. International code of Zoological Nomenclature, fourth edition, adopted by the International Union of Biological Sciences. — London (International Trust for Zoological Nomenclature, BMNH), XxIX + 306 pp. ICZN 2006. Official lists and indexes of names and works in zoology 2001-2005. — London (Internatio- nal Trust for Zoological Nomenclature), http://www.iczn.org/Official_Lists_Indexes_names2005.htm, update May 2006 (accessed 29 January 2007). Inoue, H., S. Sugi, H. Kuroko, S. Moriuti & A. Kawabe 1982. Moths of Japan, 2 vols. [most parts in Japanese]. — Tokyo (Kodansha), 966 + 552 pp., 392 pls. Karsch, F. 1895. Aethiopische Heteroceren. I. — Entomologische Nachrichten 21 (22): 337-360. Karsch, F. 1898a. Über die Arten der Lepidopteren-Gattung Sabalia WIk. nebst Beschreibung einer neuen Art: Sabalia tippelskirchi. — Entomologische Nachrichten 24 (19): 289-295. Karsch, F. 1898b. Gibt es ein System der recenten Lepidopteren auf phyletischer Basis? — Entomologische Nachrichten 24 (19): 296-303. Kirby, W.F. 1892. A synonymic catalogue of Lepidoptera Heterocera (moths). Vol. I. Sphinges and Bombyces. — London (Gurney & Jackson), xii + 951 pp. Kishida, Y. 1992. 106. Eupterotidae. p. 153. — /n: J. B. Heppner & H. Inoue (eds.), Lepidoptera of Taiwan, vol. 1, Part 2, Checklist. — Gainesville (ATL, Scientif. Publ.), xlix + 276 pp. Krausse, A. & M. Wolff 1919. Eine Übersicht über die bisher aufgestellten fossilen und rezenten Insek- tenordnungen. — Archiv für Naturgeschichte 85 (3): 151-171. Kroon, D. M. 1999. Lepidoptera of southern Africa. Host-plants & other associations. A catalogue. — Sasolburg (the author/Lepidopterists’ Society of Africa), xi+ 159 pp. Kuznetsov, N. Y. 1967. Fauna of Russia and adjacent countries. Lepidoptera. Vol. 1. Introduction. (Trans- lated by A. Mercado). — Washington (U. S. Department of Agriculture), 305 pp. Lawrence, J. F.,& A. F. Newton 1995. Families and subfamilies of Coleoptera (with selected genera, notes and dates on family-group names). pp. 779-1006. - In: J. Pakaluk & S. A. Slipinski [eds.], Biology, phylogeny and classification of Coleoptera. Papers Celebrating the 80th Birthday of Roy A. Crow- son. — Warszawa (Muzeum 1 Instytut Zoologii PAN). Lemaire, C., & J. Minet 1998 (“1999”). 18. The Bombycoidea and their relatives. pp. 321-353. — In: N. P. Kristensen (ed.), Lepidoptera, moths and butterflies. Volume 1: Evolution, systematics, and biogeog- raphy. (Handbook of zoology, vol. IV, part 35). — Berlin, New York (W. de Gruyter). Mell, R. 1930 (“1929”). Beiträge zur Fauna sinica. V. Die Brahmaeidae und Eupterotidae Chinas. — Deut- sche Entomologische Zeitschrift 1929 (5): 337-494, pls. 1-12. Mell, R. 1937. Beiträge zur Fauna sinica. XIV. Ergänzungen zur Sphingiden-, Brahmaeiden- und Eupte- rotidenfauna Chinas (Lep.). — Deutsche Entomologische Zeitschrift 1937 (1/2): 1-19. Melville, R. V., & J. D. D. Smith 1987. Official lists and indexes of names and works in zoology. — London (International Trust for Zoological Nomenclature), 366 pp. Mielke, O. H. H., & M. M. Casagrande 2006. Catalogo bibliografico dos taxa superiores da ordem Lepi- doptera. — Revista brasileira de Zoologia 23 (1): 1-41. Nota lepid. 30 (2): 315-327 325 Minet, J. 1994. The Bombycoidea: Phylogeny and higher classification (Lepidoptera: Glossata). — Entomo- logica scandinavica 25 (1): 63-88. Moore, F 1883. The Lepidoptera of Ceylon, vol. IL. — London (L. Reeve); 162 p. + pls. 72-143. Nässig, W. A. 1989. A new species of the genus Eupterote Hübner [1822] from Sumatra (Lepidoptera, Eupterotidae). — Heterocera sumatrana 2 (7): 169-174. Nässig, W. A. 1995. Revisional notes on Philippine Eupterotidae: Sarmalia Walker 1866 another new synonym of Eupterote Hübner [1822] (Lepidoptera: Eupterotidae). — Nachrichten des Entomologi- schen Vereins Apollo, Suppl. 14: 119-124. Nässig, W. A. 2000. A new and remarkable species of Eupterote from the mountains of West Sumatra (Lepidoptera: Eupterotidae). — Heterocera sumatrana 12 (2): 67-77. Nässig, W. A., & I. W. B. Nye 1991. Case 2737. Brahmaea Walker, 1855 (Insects, Lepidoptera): pro- posed confirmation of Bombyx certhia Fabricius, 1793 as the type species. — Bulletin of Zoological Nomenclature 48 (2): 137-139. Neumoegen, B., & H.G. Dyar 1894. A preliminary revision of the Bombyces of America north of Mexico. — Journal of the New York Entomological Society 2 (1): 1-30, (2): 57-76, (3): 109-132, (4): 147-174. Nielsen, E. S., & I. F. B. Common 1991. Lepidoptera (moths and butterflies). pp. 817-915. — Zn: I. F. B. Common (ed.). The Insects of Australia, 2 vols. — Melbourne (Melbourne Univ. Pr., CSIRO Ento- mology), 1137 pp. Oberprieler, R.G., & N. J. Duke 1994. The life history and immature stages of Spiramiopsis comma Hampson, 1901 (Lepidoptera: Bombycoidea), with comments on its taxonomic position and on preimaginal characters of the Bombycoidea. — Nachrichten des Entomologischen Vereins Apollo, N.F. 15 (3): 199-244. Oberprieler, R. G., W. A. Nässig & E. D. Edwards 2003. Ebbepterote, a new genus for the Australian ‘Eupterote’ expansa (T. P. Lucas), with a revised classification of the family Eupterotidae (Lepido- ptera). — Invertebrate Systematics 17: 99-110. Pinhey, E. C. G. 1975. Moths of southern Africa. — Cape Town (Tafelberg), 273 pp. Platt, E. E. 1921. List of foodplants of some South African lepidopterous larvae. — South African Journal of Natural History 3 (1): 65-138. Rebel, H. 1900. Zur Auffassung der Lemoniiden als selbstständige Lepidopterenfamilie. — Entomologi- sche Nachrichten 26 (4): 49-51. Rebel, H. 1914. Wissenschaftliche Ergebnisse der Expedition R. Grauer nach Zentralafrika, Dezember 1909 bis Februar 1911. Lepidopteren. — Annalen des K. K. naturhistorischen Hofmuseums 28 (2): 219-294, pls. xvii—xxiv. Remington, C. L. 1954. Order Lepidoptera (Glossata). pp. 226-305. — In: C. T. Brues, A. L. Melander & F. M. Carpenter [eds.], Classification of insects. Keys to the living and extinct families of insects, and to the living families of other terrestrial arthropods. — Bulletin of the Museum of Comparative Zoology at Harvard College, 108: v + 917 pp. Robinson, G. S., P. R. Ackery, I. J. Kitching, G. W. Beccaloni & L. M. Hernandez 2001. Hostplants of the moth and butterfly caterpillars of the Oriental Region. — London (BMNH), Kuala Lumpur (Southdene), 744 pp. Rothschild, Lord [W.] 1917a. On the genera Melanothrix, Drepanojana, Melanergon, Paracydas, Cotana, Hypercydas, Epicydas, and Nervicompressa of the family Eupterotidae with descriptions of new forms. — Novitates zoologicae 24: 463-474, pls xi-xil. Rothschild, Lord [W.] 1917b. Some new moths of the families Arctiidae and Eupterotidae. — Novitates zoologicae 24: 475-492. Schaus, W. 1894. On new species of Heterocera from tropical America. — Proceedings of the Zoological Society of London 1894: 225-243. Schultze, A. 1915. Zur Kenntnis der ersten Stände von einigen west- und zentralafrikanischen Hetero- ceren. — Archiv für Naturgeschichte 80A (1): 144-163, pls i-iv. Scoble, M. J. 1992. The Lepidoptera. Form, function and diversity. - London (Natural History Museum Publications), Oxford (Oxford Univ. Pr.), 404 pp. Semper, G. 1896. Heterocera, Familie Eupterotidae. pp. 387-390. — In: G. Semper (1896-1902), Die Schmetterlinge der philippinischen Inseln. Beitrag zur indo-malayischen Lepidopteren-Fauna. Zwei- 326 NÄssıG & OBERPRIELER: On eupterotid family group names ter Band [Zweite Abtheilung]: Die Nachtfalter. Heterocera. [Edited by C. Semper under the title: Rei- sen im Archipel der Philippinen, zweiter Theil, Wissenschaftliche Resultate, Sechster Band.] — Wies- baden (C. W. Kreidel), pp. 381-728, pls. C-V [preimaginals], I-Ixvi [imagines]. Sevastopulo, D. G. 1944. The early stages of Indian Lepidoptera. — Journal of the Bombay Natural History Society 44 (1/2): 78-87. Smith, J. D. D. 2001. Official lists and indexes of names and works in zoology. Supplement 1986-2000. — London (International Trust for Zoological Nomenclature), 136 pp. Speidel, W., & C. M. Naumann 2005. A survey of family-group names in noctuoid moths (Insecta: Lepi- doptera). — Systematics and Biodiversity 2 (2): 191-221. Strand, E. 1909. Schmetterlinge aus dem Sambesi-Gebiet, gesammelt und dem Berliner Museum ge- schenkt von Herrn Franz Seiner. — Archiv für Naturgeschichte 75 (1-3): 375-386. Strand, E. 1911a. Sechs neue exotische Grossschmetterlinge. — Archiv für Naturgeschichte 77 (1, Suppl. 3): 98-102. Strand, E. 1911b. Zoologische Ergebnisse der Expedition des Herrn Hauptmann a. D. Fromm 1908/09 nach Deutsch-Ostafrika. 1. Lepidoptera. — Mitteilungen aus dem Zoologischen Museum in Berlin 5 (2): 275-304. Strand, E. 1911c. Beschreibungen afrikanischer Lepidopteren, insbesondere Striphnopterygiden. — Anna- les de la Société entomologique de Belgique 55: 145-164. Strand, E. 1922 [some parts of the family treatment were written by A. Seitz]. 10. Familie, Eupterotidae. pp. 417-432, pls 31, 36, 37, 56 B, 57. — In: A. Seitz (ed.) 1911-1933, Die Groß-Schmetterlinge der Erde, 10, Die indoaustralischen Spinner und Schwärmer. — Stuttgart (A. Kernen), ix + ii + 909 pp, 104 pls. Strand, E. 1927. Enumération des Hétérocères exotiques (à l’exclusion des Noctuides, des Microlépido- pteres, des espèces de l’Île Formosa et des espèces que j’ai décrites dans Seitz “Macrolepidopteres”) qui jusqu’à l’année 1926 sont nommés et publiées. — Entomologisches Nachrichtenblatt 1 (2): 12-25, 1 (3): 42-55. Swinhoe, C. 1891. New species of moths from Southern India. — Transactions of the Entomological Society of London 1891 (1): 133-154. Swinhoe, C. 1892. Catalogue of the Eastern and Australian Lepidoptera Heterocera in the collection of the Oxford University Museum. Part 1, Sphinges and Bombyces. — Oxford (Clarendon), viii + 324 pp., 8 pls. Swinhoe, C. 1905. XV. — Notes on Eastern and Australian Heterocera, with descriptions of one new genus and thirteen new species. — Annals and Magazine of Natural History (7) 16: 142-155. Taylor, J. S. 1949. Notes on Lepidoptera in the eastern Cape Province (Part I). - Journal of the Entomological Society of southern Africa 12: 78-95. Taylor, J. S. 1950. Notes on Phiala patagiata Aur., the Karoo Tent Caterpillar. - Entomology Memoir, Department of Agriculture, Union of South Africa 2 (8): 219-229. Taylor, J. S. 1951. Notes on Lepidoptera in the Eastern Cape Province (Part ID). — Journal of the Entomological Society of southern Africa 14 (2): 94-126. Taylor, J. S. 1957. Notes on Lepidoptera in the Eastern Cape Province. IV. — Journal of the Entomological Society of southern Africa 20 (2): 315-332. Taylor, J. S. 1961. Some records of Tachinidae and their hosts in the Eastern Cape Province. — Technical Communication No. 3, Entomology Series No. 46: 1-15. Taylor, J. S. 1965. Notes on Lepidoptera in the eastern Cape Province, 5. — Journal of the Entomological Society of southern Africa 28: 137-154. Turner, A. J. 1922. Revision of Australian Lepidoptera. Saturniadae, Bombycidae, Eupterotidae, Noto- dontidae. — Proceedings of the Linnean Society of New South Wales 47: 348-390. Turner, A. J. 1947. A review of the phylogeny and classification of the Lepidoptera. — Proceedings of the Linnean Society of New South Wales 71 (5/6): 303-338. van Eecke, R. 1930. De Heterocera van Sumatra, eerste deel. — Leiden (E. J. Brill), 456 pp., 13 pls [Reprinted from a series of publications in Zoologische Mededeelingen, 8 (3/4), 1925, to 12 (3/4), 1929.] Väri, L., & D. Kroon 1986. Southern African Lepidoptera. A series of cross-referenced indices. — Pretoria (The Lepidopterists’ Society of Southern Africa, The Transvaal Museum), x + 198 pp. Nota lepid. 30 (2): 315-327 >21 Vari, L., D. M. Kroon & M. Krüger 2002. Classification and checklist of the species of Lepidoptera recorded in southern Africa. — Sasolburg (the authors), Chatswood (Simple Solutions Australia), xxi + 385 pp. Viette, P. 1990. Liste récapitulative des Lépidoptères Hétérocères de Madagascar. A provisional check-list of the Lepidoptera Heterocera of Madagascar. — Faune de Madagascar, Suppl. 1: 263 pp. Wallengren, H. D. J. 1858. Nya Fjäril-slägten. — Ofversigt af Kongliga Vetenskaps-Akademiens Förhand- lingar 15: 75-84, 135-142, 209-215. Wallengren, H. D. J. 1860. Lepidopterologische Mittheilungen. I. — Wiener Entomologische Monatschrift 4 (6): 161-176. Wallengren, H. D. J. 1865 (“1863”). Heterocer-Fjärilar, samlade i Kafferlandet af J. A. Wahlberg. — Kong- liga Svenska Vetenskaps-Akademiens Handlingar 5 (4): 1-83. Wallengren, H. D. J. 1875. Insecta Transvaaliensia. — Bidrag till Transvaalska Republikens ı Södra Afrika Insektfauna. — Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar 32 (1): 83-137. Weymer, G. 1909. Exotische Lepidopteren. — Deutsche Entomologische Zeitschrift „Iris” 22: 1-35. Zwick, A. 2006. Phylogenetic analyses and taxonomy of Anthelidae (Lepidoptera). A dissertation sub- mitted for the degree of Doctor of Philosophy at the School of Botany and Zoology, Australian Na- tional University, Canberra. — Canberra (ANU, unpublished thesis), xiv + 679 pp. Nota lepid. 30 (2): 329-353 329 Geographical variation and Pleistocene history of the Erebia pandrose - sthennyo complex (Nymphalidae; Satyrinae) FRANS CUPEDO Processieweg 2, NL-6243 BB Geulle, Netherlands; e-mail: frans@cupedo.eu Abstract. The Erebia pandrose-sthennyo complex consists of two taxa: pandrose, with a Eurosiberian, arctic-alpine disjunct distribution, and sthennyo, hitherto considered a Pyrenean endemic. Their taxonom- ic status is uncertain because of their strictly allopatric distributions. The discovery of a sthennyo-like population in the Alps was cause for the present re-examination of the geographical distribution of typi- cal pandrose and sthennyo characters, in 43 populations covering the whole geographic range of the E. pandrose-sthennyo complex. The investigated populations indeed split up into two taxonomical units, pandrose and sthennyo. The latter ıs not confined to the Pyrenees, but also inhabits the southeastern Alps, the Durmitor (Montenegro) and the southern Altai (Kazakhstan). No sympatric occurrence of pandrose and sthennyo was found. In the Alps, pandrose and sthennyo are separated by a belt of intermediate popu- lations, resulting from intermingling in a zone of contact. Apparently they are not reproductively isolated. Pleistocene history. Both pandrose and sthennyo have a Eurosiberian distribution, resulting from spread in the glacial trans-palaearctic tundra belt, and subsequent shrink and disruption of the occupied area during interglacial times. As, however, pandrose and sthennyo are not reproductively isolated, they cannot have occupied the tundra simuitaneously. Consequently their expansion has to be dated back to different glacial periods. Two arguments point to sthennyo as the first to have established its actual distribution: (1) The sthennyo populations of the Pyrenees, the Alps and the Altai show morphological differentiations, in wing design, wing shape, and in one genital character, whereas pandrose is uniform throughout its territory. The pandrose populations differ merely in the extent of sthennyo traits, resulting from former gene exchange. (11) Today, pandrose is still inhabiting the northern region of the Alps, bordering the previous tundra zone, whereas sthennyo occupies refugial areas in the southern Alps, completely cut off from its former dis- persion route. It is argued that the spread of sthennyo must date back to (at least) the Mindel glaciation (MIS8), and the spread of pandrose to (at least) the Riss period (MIS6). The main argument being that colonization of the Apennines (from the western Alps) and the Balkans (from the southern Carpathians) must date (at least) from the Würm glacial (MIS2-4), which implies that pandrose already lived in the Alps and the Carpathians during the Riss-Wiirm interglacial (MIS5). Scandinavian populations are completely intermediate. Sthennyo, present since the end of the Mindel glaciation, survived the Riss glaciation on the southern edge of the polar ice cap, where it interbred with pandrose. At the end of Riss, Scandinavia was repopulated by these mixed populations. Taxonomy and nomenclature. As pandrose and sthennyo are not reproductively isolated, they have to be considered conspecific. The hierarchical structure of E. pandrose is best reflected in taxonomy by creating two groups of subspecies, according to art. 6.2 of the ICZN, which should be named Erebia pandrose (pandrose) and Erebia pandrose (sthennyo). Alpine and Scandinavian intermediate populations are arbitrarily placed in the pandrose group. Within the pandrose group, ingana Fruhstorfer, 1911 is shown to be a junior synonym (syn. n.) of pandrose (Borkhausen, 1788). Within the sthennyo group, infraclara Verity, 1953 is shown to be a junior synonym (syn. n.) of marmolata Dannehl, 1927. The South Altai population of the sthennyo group is described as narymica ssp. n. Introduction The geographical distribution pattern of oreo-tundral species is the result of repeated cycles of areal expansions and contractions, generated by Pleistocene climatic fluc- tuations (De Lattin 1967). During glacial times the arctic tundra shifted southwards, extending into a continuous trans-palaearctic belt between the polar ice sheet and the southern mountain systems (Pyrenees, Alps, Carpathians, Caucasus, and Altai). Not only tundral species, but also part of the populations inhabiting these mountain systems spread into the tundra. In many cases this led to oreo-tundral (arctic-alpine) disjunction, and/or to long-distance expansion in an east-west direction, resulting in a Eurasian dis- Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 330 Cupepo: Erebia pandrose — sthennyo complex Fig. 1. Typical upperside, underside and valve of pan- drose (left) and sthennyo (right). Fig. 2. Ocelli position. The ratio AB/AE was calcu- lated from the measured AD and BC. tribution (De Lattin 1967; Kostrowicki 1969). Arctic-alpine disjunction and Eurasian distribution are strongly cor- related (Varga 1996), as both result from tundral dispersion during glacial periods. In addition, populations inhabiting the southern mountain ranges could survive in spatially restricted peripheral refugia. These were characterised by the verti- cal distance between the ice-surface and the snowline being sufficient to enable permanent settlement of alpine organ- isms in between. Through the pioneer- ing work of Penck & Bruckner (1909, first published in separate issues 1901- 1908) detailed data on glacier altitude and snowline altitude on the outskirts of the Alpine ice sheet became avail- able for the first time. Soon, locations of potential refugia were reconstructed, and distributional areas of alpine plant species were related to them (Chodat & Pampanini 1902; Briquet 1906; Brock- mann-Jerosch & Brockmann-Jerosch 1926). Meanwhile, the location of those potential refugia has been re- fined (Stehlik 2000; Schönswetter et al. 2005), and postglacial dispersal routes in the Alps have been reconstructed for a considerable number of plants (Stehlik 2003, Tribsch & Schönswetter 2003) and a few alpine animal taxa. In the latter, such reconstructions have been based on molecular markers (e.g., Schmitt & Hewitt 2004, Vila et al. 2005, Schmitt et al. 2006) or morphological characters (Holdhaus 1954, Cupedo 2004). This paper deals with the Erebia pan- drose-sthennyo complex, which con- sists of two, closely related taxa, pan- drose (Borkhausen, 1788) and sthennyo (Graslin, 1850). Because of strong morphological similarities they are widely consid- ered conspecific (Warren 1936, Manley & Allcard 1970, Gömez Bustillo 1974, Higgins 1975, Leraut 1980). Because of certain constant differences, however, both in genital Nota lepid. 30 (2): 329-353 53: characters and in Wing design, some authors consider them specifically distinct (Willien 1990, Kudrna 1986, Lafranchis 2000). Biological evidence for either of these views 1s lacking. Erebia pandrose is a tundral and high mountain species with a Eurosiberian distribu- tion (Warren 1936, Kostrowicki 1969). The European part of the area is oreo-tundral disjunct. The tundral component includes Scandinavia and the adjacent Russia, as far as the Kanin peninsula (Warren 1936) and Kolguev Island (Tatarinov & Dolgin 1999). The oreal area is fragmented: it comprises the Alps, Pyrenees, Apennines, Carpathians, and the Balkan mountains. The Asıan subarea is exclusively oreal, and is confined to the Altai-Sayan mountain range. The species is absent from the East Palaearctic tun- dra. Records from the Caucasus (Kostrowicki 1969) have not been confirmed and are supposedly erroneous (Nekrutenko 1990, Tuzov et al. 1997). Erebia sthennyo has only been recorded from the central Pyrenees, where it lives in close proximity to Erebia pandrose (De Lesse 1952). In the 1990’s, however, the author found several populations in the southern Alps that, according to both male genital characters and wing design, belong to sthennyo. They probably remained unnoticed because since Warren (1936) genital characters of alpine pandrose populations never received serious attention, as these were a priori consid- ered not to belong to sthennyo (Verity 1953, Varga 1971, Roos & Arnscheid 1976). As a result our actual knowledge of the geographical distribution of typical pandrose and typical sthennyo characters is incomplete. This paper presents the results of a re-exami- nation of a large number of populations, covering the whole geographical range of the E. pandrose-sthennyo complex, and based on a predefined set of differentiating mor- phological characters. It aims to establish the actual geographical distribution of typical pandrose and typical sthennyo characters, in order (1) to determine the validity of the taxa pandrose and sthennyo, (1) to examine to what extent their Pleistocene history can be reconstructed, based on distributional data, and (111) to work out the taxonomic and nomenclatural implications of the results. Abbreviations ZMAN = Zoological Museum Amsterdam (NL); MNHL = Nationaal Natuurhistorisch Museum Leiden (NL); CFC = Collection Frans Cupedo, Geulle (NL); CSC = Collection Sergei Churkin, Moscow (RU); SP = Number of spines per valve; OP = Ocelli Position; FWD = Forewing Design; HWD = Hindwing Design; MIS = Marine Isotope Stage. Material and Methods Samples were studied from 43 localities. See table 1 and figure 3. (Tables in the appendix). Five differentiating sthennyo characters have been quantified and measured as follows (figure 1; see also note 1): Male valve: the sthennyo male has prominent spines on the dorsal edge of the valve, which are lacking in pandrose. The number of spines on the dorsal ridge of the valve (SP) was counted. Asymmetry being rule rather than exception, left and right valve were scored separately. (Stereo- microscope, 20 x) 332 CupEpo: Erebia pandrose — sthennyo complex Male forewing upperside: in sthennyo the apical ocelli are situated closer to the wing margin than in pandrose. Well set males were photographed and printed on a 2: | scale. The position of the apical ocelli relative to the wing margin (OP) was expressed in the ratio AB/AE in Figure 2. Male forewing upperside: in sthennyo, the postdiscal band is poorly devel- oped, its edges are suffused, and the fine submarginal zigzag line, common in pandrose, is missing. The development of the postdiscal field was quantified as follows: 0 = Band completely obscure, in ground colour. | = Lighter rings around ocelli. 2 = Lighter colour more extended than merely rings; less than 3. 3 = Lighter colour filling the postdiscal band completely, at least in cells 4 and 5, eventually in cell 3. 4 = Idem, including cell 2. Distal edge of the postdiscal band: 0 = fading into submarginal field; 1 = distinct. Denticulate transverse line separating submarginal and postdiscal band: 0 = absent; | = present. The sum of the three scores is used as a measure for forewing upperside design (FWD). Male hindwing underside: In sthennyo the design of the hindwing under- side 1s poorly developed, and marginal elements (chevrons and its dark filling) are lacking. Transverse lines. 0 = absent; | = faintly indicated; 2 = present Ocelli. 0 = absent; | = one point in cell 2; 2 = more than one point Chevrons. 0 = absent; | = indicated; 2 = present The dark filling of the chevrons. 0 = absent; | = indicated; 2 = present The sum of the four scores is used as a measure for hindwing underside design (HWD). Female genitalia. In sthennyo the antevaginal plate is wider than in pandrose. The width of the antevaginal plate was measured. (Measuring microscope, 30 x) Deviations from normality were calculated using the Kolmogorov-Smirnov test with Lilliefors correction. For cluster analysis three algorithms were applied: Ward-method, average linkage within groups and average linkage between groups. Z-scores were standardized and squared Euclidian distance was measured. Homogeneity of variances was determined with Levene’s test, and analysis of variance was performed with one- way ANOVA or through the non parametric Kruskal-Wallis test. All statistical tests Nota lepid. 30 (2): 329-353 3353 EN A CSS { { | =< | Î / \ Jh / )) Altai-Sayan. Light grey = mountains >1500 m. Dark grey = lakes and rivers. Black = political boundaries between Kazakhstan (W), Russia (N), Mongolia (E) and China (S). C. Europe. Light grey = main moun- tain systems. The numbers refer to Tab. 1. were performed with the software SPSS 12.0. Pleistocene timetable 1s based on oxygen isotope dating (MIS stages). Results and conclusions 1.1. Male characters (Fig. 4) The population average values for each of the four variables are listed in Table 2 and graphically presented on Figure 4. Only the distribution of the number of spines per valve deviates significantly from normality (Kolmogorov-Smirnov p <0.001). Itiseven discontinuous, with values either < 0.5 or > 1.0. Hierarchical cluster analysis (Ward method), based on the four male characters, resulted in the dendrogram in Figure 5. The two main clusters were designated A and B. For forewing design and hindwing design (equal variances, Levene’s p = 0.445 and 0.421 respectively) ANOVA gave a very good discrimination between the clusters A and B (F = 15.4 and 23.5 respectively; p <0.001 334 Cupepo: Erebia pandrose — sthennyo complex => © Frequency Frequency N 0 00 05 19 15 20 25 30 0,24 026 0,28 Number of spines per valve Ocelli position > [ex] w Frequency Frequency N 3,0 4,0 0 20 30 40 26002602240 Fore wing design Hind wing design Fig. 4. Frequency distribution of the population means of the four differentiating characters in males. in both cases). For the number of spines and ocelli posotion (variances significantly unequal, Levene’s p <0.001 and =0.019, respectively) Kruskal-Wallis test confirmed significant difference between both clusters (p <0.001 for both variables). Average link- age within groups resulted in identical clustering. Using between-groups algorithm, sample 33 was transferred to the B-cluster. The main conclusion is that, on the basis of the morphological characters that are tradi- tionally used to discriminate between pandrose and sthennyo, the populations of the E. pandrose-sthennyo complex indeed split up into two distinct groups. As the populations with typical pandrose characters belong to the A-cluster, and populations with sthen- Nota lepid. 30 (2): 329-353 535 nyo characters to the B-cluster, these will provisionally be referred to as the pandrose group and the sthennyo group, in short pandrose and sthennyo, without any taxonomi- cal bearing. 1.2. Female genitalia There is a discrepancy between the Pyrenean sthennyo population (sample 1) and all other populations investigated. Because of the extremely small sample size (n=1-5 per population), these results were omitted from the cluster analysis. If the available data were included, sample | would branch off from the B-cluster at a distance of 7 scale- units (not shown). 1.3. Geographical distribution The geographical distribution of the taxon sthennyo, hitherto considered a Pyrenean endemic, is far more extended than ever thought. It inhabits three major mountain sys- tems and at least one small one: the Pyrenees, the southeastern Alps, the Altai, and the Durmitor in Montenegro. Within the pandrose group, each of the studied characters showed a continuous vari- ation, the extremes of which can be characterised as “typical pandrose” and “more or less sthennyo-like” respectively. To what degree each of the populations of the pandrose group tends towards a sthennyo habitus is best visualised by a simple ranking (Table 3), based on the rescaled (O—100) sum of the rescaled scores for the four characters. The most typical pandrose (sum = 0) ranks at the top. Geographical distribution in relation to ranking was not random. In Figure 6, populations with a sum < 50 and those with a sum > 50 are plotted in different symbols. The most sthennyo-like populations turned out to be concentrated in the eastern Alps, in a zone bordering the area of the sthennyo group. Beyond the Alps such populations were found in Scandinavia only. The other extreme, the most typical pandrose populations, were found in the southern Carpathians and the Balkans (Bucegi and Rila), followed by the Siberian populations (Altai and Sayan). Discussion 1. Monophyly of the pandrose group and the sthennyo group The pandrose group and the sthennyo group share three mountain systems: Pyrenees, Alps, and Altai. It is quite unlikely that the morphological differences between both groups arose independently in each of these regions. Obviously, pandrose and sthen- nyo form two monophyletic groups, 1.e. the differences between them must date back to their respective common ancestors. The primary dichotomy in the dendrogram thus reflects the oldest detectable geographical disjunction within the Erebia pandrose- sthennyo complex, which has resulted in two isolated populations that subsequently differentiated into the pandrose ancestor and the sthennyo ancestor. The fact that the most sthennyo-like populations in the Alps are concentrated in a zone bordering the area of the sthennyo group, strongly suggests that they originate from 336 CupEDo: Erebia pandrose — sthennyo complex postglacial intermingling in a contact zone. Apparently, pandrose and sthennyo did interbreed where they met. That was emphatically the case in the eastern Alps. To a minor extent, however, intermingling with sthennyo genes has taken place anywhere in the Alps: populations with a relatively high score for one of the studied characters are found throughout the Alps, and in the Apennines (Table 3). 2. Pleistocene history 2.1. Würm-glacial refuges in the Alps, the Apennines, the Carpathians, and the Balkans Each population of the sthennyo group, and some populations of the pandrose group, can easily be related to one of the well known glacial refugia. For the remaining pan- drose populations, there is a logical relation with a refugium, but they cannot be mor- phologically linked to it, as this study focuses merely on differentiating characters be- tween pandrose and sthennyo. Those refugia are printed in small font. The Bergamasque Alps The Bergamasque Alps remained largely unglaciated during the Wiirm glaciation (MIS2-4, 75-10 ky BP). They harboured an isolated refugial flora and fauna, including sthennyo. During Wiirm, the valleys of Adda and Oglio attained their actual depth by glacial erosion, which largely prevented post-glacial expansion. Sthennyo could just infiltrate the southern Ortler group across the Aprica pass, which accounts for the in- termediate population found there (sample 15, see table 3). The Bergamasque sthennyo populations show endemic differentiations (see the taxonomy and nomenclature sec- tion). The Brescian refugium The area between Lake Iseo and Lake Garda is considered here a separate refugium, called Brescian refugium. From this refuge the Adamello-Presanella group has been repopulated in postglacial times. It has been noticed for a long time that the populations of the Adamello-Presanella group and those of the adjacent Brenta group are strikingly different (Hartig 1937). That is confirmed by the results of this study: the former has a pandrose habitus, the latter a prominent sthennyo habitus (sum= 49 and 91 respective- ly). This applies also to the southernmost known population in the Adamello group, on the Passo Croce Domini, which contributed substantially to sample 17. Thus, it cannot be excluded that a population with a pandrose habitus has survived the Würm-glacial in the Brescian refugium, whereas the adjacent refugia were inhabited by sthennyo. In addition, postglacial gene flow across the Passo Carlomagno must have been mini- mal or absent, a phenomenon that has also been reported for Erebia pluto (Cupedo 2004). The refugia east of Lake Garda The southern-alpine sthennyo populations East of Lake Garda have their origin in a series of well known small refuges on the southern border of the eastern Alps (Tribsch & Schönswetter 2003): the Lessinic Alps, the Venetian and Carnic Pre-Alps, and the Nota lepid. 30 (2): 329-353 Se. Julian Alps. Postglacially they just moved to higher altitudes, and spread more or less northwards, into the Dolomites and the mountains south of the High Tauern watershed, where they apparently intermingled with populations of the pandrose group. The popu- lations of the Lessinic Alps and the Venetian Pre-Alps (samples 22 and 27) have been living, up to present time, in unglaciated areas. The East-alpine refugium The vast refugial area that existed in the non-glaciated parts of the Styrian Alps (Penck & Bruckner 1909, Tribsch & Schönswetter 2003) was another dispersion centre of the pandrose group. Populations on the Zirbitzkogel (Reichl 1992) and in the eastern Karawank mountains (JakSi¢ 1998) have even persisted here within the refugial area to date. At least the populations of Gurktal Alps and Niedere Tauern (samples 29, 30, 31) must have their origins in this refugium. The Apennines During glacial stages the Apennines were ecologically contiguous with the Ligurian and Maritime Alps, and must have acted as an important refugium for many species. This is reflected in the morphological resemblance of their Erebia species with western Alpine forms. For E. cassioides this relation has been confirmed by molecular data (Lattes et al. 1994). The small witness population in the Laga mountains proves that also pandrose withdrew into the Italian peninsula. As there is only one pathway, there is no doubt that the Apennines were colonised from the western Alps. (Note that this does not necessarily imply a postglacial colonisation of the western Alps from the Apennines!) Northern refugia The northern limestone Alps must logically be colonised from refuges at its northern outskirts, or even from the periglacial tundra. Western alpine refugia Vast refugia existed in the French Pre-Alps (Vercors, Diois, Provence), many narrow refugial areas were found at the eastern slopes of the Cottian and Graian Alps (Briquet 1906). Undoubtedly these refugia contributed to the present-day pandrose populations in the western Alps, but this cannot be inferred from the present study. Carpathians and Balkans Varga (1975b) demonstrated that populations of many species in the Bulgarian moun- tains (Stara Planina, Rila, Pirin), including Erebia pandrose, were morphologically closely related to Carpathian populations. He made clear that the Balkan mountains were colonised from the southern Carpathians, and that the oldest differentiations among Balkan Erebia dated to the Würm-glacial. The present data (samples 37 and 39) agree with that hypothesis. In the western Balkans on the other hand, the Dinarics and the mountains of Bosnia-Herzegovina, Montenegro, and Macedonia were connected with the Julian Alps, which harbour closely related butterfly faunas (Varga 1975b). The fact that the Julian Alps and the Durmitor currently are both inhabited by sthennyo sup- ports such a connection. 336 Cupepo: Erebia pandrose — sthennyo complex 2.2. Pre-Würm history Dispersion routes. Both the pandrose group and the sthennyo group show distribution patterns that range from the European mountains to Central Asia. This is due to the fact that dispersion of both groups took place along the same pathway: the glacial trans- palaearctic tundra belt (De Lattin 1967, Kostrowicki 1969). As, however, pandrose and sthennyo would have intermingled wherever they met, they can not have occupied the tundra belt simultaneously. Consequently, pandrose and sthennyo must have spread in two waves, at different moments in history, 1.e. during different glacial cycles. Sequence. The populations of the pandrose group are rather uniform throughout their territory. They differ merely in the extent of the sthennyo influence. The sthennyo popu- lations are morphologically far more differentiated: the Kazakhstan population shows a different arrangement of forewing ocelli and a different forewing shape; the Pyrenean population developed an apomorphic genital character in the females; the populations of the Bergamasque Alps differ in their wing design from the remaining populations in the south-alpine subarea (see the taxonomy and nomenclature section). That suggests that spread and subsequent disjunction of the pandrose group is of more recent date than that of the sthennyo group. This is confirmed by their actual distribution in the Alps: pandrose is widespread in the northern regions, bordering the glacial tundra belt along which both groups have reached the Alps, whereas sthennyo has been completely cut off from the previous tundra zone: it has been forced into refugial areas, “dead ends”, on the southern outskirts of the Alps. Dating. The witness population in the Apennines unequivocally proves that the pan- drose group was present in the Alps before the beginning of the Würm glacial, i.e. dur- ing the Riss-Würm interglacial (MIS5, 128-75 ky BP). The same goes for the southern Carpathians, in view of the presence of pandrose in the Rila mountains. Thus, the spread of pandrose in the tundra belt has to be dated at its latest to the Riss glaciation (MIS6, 185-128 ky BP) [Note 2+3]. Consequently, the spread of sthennyo along the tundra belt took place at its latest during the Mindel glaciation (MIS8, 300-242 ky BP). That means that more or less unmixed sthennyo populations persisted in the southeastern Alps for at least two glacial cycles. This is probably due to the predominant east-west orientation of mountain chains in this part of the Alps. In the western Alps, however, no geographic barrier prevented intensive gene flow between pandrose and sthennyo. As a result, no sthennyo populations exist there today, but a number of pandrose populations show sthennyo traits to some degree (cf. the mean SP values of samples 4 and 7, table 3). Scandinavia. The Scandinavian populations must result from the same two colonisa- tion waves. During Riss, however, Scandinavian sthennyo populations were shoved south by the growing land ice cap and survived at its edge. Here they intermingled with pandrose already during glacial time, prior to the repopulation of Scandinavia. As a result Scandinavia nowadays is inhabited entirely by intermediate populations, in contrast with the Alps, where postglacial encounter led only to borderline mixing. Nonetheless, the Scandinavian populations are far from uniform, as was already stated by Henriksen & Kreutzer (1982). Therefore, it is not surprising that one of these popu- lations (sample 33) is placed in the sthennyo cluster when using a different algorithm for hierarchical cluster analysis. Nota lepid. 30 (2): 329-353 339 Rescaled Distance Cluster Combine 0 5 10 15 20 25 Sample Nr + =----4-------— + + = nn + Penninic.N 6 Ortler.W 14 Adamello 17 Dauphine 3 Ötztal 19 Gerlitzen 30 GriParad: 5 Sayan 42 Pt.S.Bernh 4 Penninic.S 7 Karwendel 20 Rofan 21 Bernina 1:0 Pyrenees.E 2 Albula 9 Bernese 8 Rila 39 Altai 41 Tatra 36 Bucegi 37 A Amberg 29 Sevo 40 N.Tauern 31 Norway.S 35 Sweden.N 33 Ortler.S 15 Brenta 18 H.Tauern 26 Dolom.Lat 23 Dolom.Mar 24 Ortler.N 16 Norway.N 34 Dolom.Pala 2 Julian 32 Narym 43 Venetian 27 B Carnic 28 Pyrenees.C ak Bergam.W ala Bergam.C 12 Lessinic 22 Durmitor 38 Bergam.E 18 Fig. 5. Dendrogram resulting from hierarchical cluster analysis (Ward algorithm). Centres of differentiation. The results of thıs study provide no clue to the geographic differentiation centres of the ancestors of the pandrose group and the sthennyo group, nor to the direction of their glacial expansion. From the above can only be inferred that, prior to the dispersion of pandrose, sthennyo already occupied the European mountains (Alps and Pyrenees), the European tundra (Scandinavia), and the Siberian mountains (Altai). That leaves the Siberian tundra as a hypothetical centre of origin for pandrose. The differentiation of pandrose and sthennyo might, theoretically, have resulted from an early Pleistocene oreo-tundral disjunction in the eastern Palaearctic. 340 Cupebo: Erebia pandrose — sthennyo complex 3. Taxonomy and nomenclature 3.1. The type locality of Erebia pandrose Borkhausen, 1788 Erebia pandrose was originally described as Papilio castor Esper, 1781. Borkhausen (1788) replaced this name (a primary homonym of Papilio castor Cramer, 1775) by pandrose. Description and type locality, however, were fixed by Esper. His material came from “Steiermark” (Stiria), without further specification. The only Stirian mas- sıfs harbouring E. pandrose are the Niedere Tauern and the northern Saualpe (Reichl 1992). In the material investigated for this study, sample 31, from the western end of the Niedere Tauern, is regarded to be closest to nomotypical Erebia pandrose. 3.2. The taxonomic status of the sthennyo group In the Pyrenees, despite intensive and targeted research by de Lesse (1952), sthen- nyo and pandrose were nowhere found living sympatrically. He just found one pos- sible hybrid, flying together with some pandrose, in the region separating the areas of sthennyo and pandrose. His observation has been interpreted as a sympatric occurrence (Lafranchis 2000), but is certainly not a proof of reproductional isolation. In the Alps, both groups have intermingled where they met. That implies that, at least in the Alps, no reproductive barriers exist between members of the pandrose group and members of the sthennyo group. Consequently, pandrose and sthennyo are to be considered con- specific. Nonetheless, there are constant anatomical differences between unmixed populations of the two groups, which result, as has been shown, from at least 240,000 year of iso- lation. The subspecies of Erebia pandrose thus can be grouped in two monophyletic aggregates of subspecies which, according to Art. 6.2 of the Code, should be designated as Erebia pandrose (group pandrose) and Erebia pandrose (group sthennyo) (ICZN 2000). 3.3. Erebia pandrose pandrose Borkhausen, 1788 and E. pandrose ingana Fruhstorfer, 1911 In the Alps, two subspecies of the pandrose group have been described. The validity of ssp. ingana, characterised by the intensively coloured postdiscal band and the promi- nent black spots on the forewing upperside, has long been questioned. Warren (1936) states that “such specimens occur wherever pandrose flies, and always in company with poorly marked specimens”. Von der Goltz (1938), after studying Fruhstorfer’s cotypes, concluded that the typical characters of ingana are individual characteristics, not typical of the population. Both Warren’s and Von der Goltz’s opinion are confirmed by the large sample 5, which was collected at exactly the same place where Fruhstorfer collected his type series. Fruhstorfer’s description applies to a minority of brilliantly coloured specimens, which occur equally in many other samples. This explains why distributional data for ingana of leading authors are so different, sometimes even con- tradictory, and lack any zoogeographical logic (compare Fruhstorfer 1911, Hartig 1937, Verity 1953, Varga 1975b, Roos & Arnscheid 1976 and Arnscheid 1981). In addition, Nota lepid. 30 (2): 329-353 34] none of the characters investigated in this study justifies a subdivision of the alpine populations of the pandrose group into two morphologically defined and geographi- cally coherent units. Consequently, Erebia pandrose ingana Fruhstorfer, 1911 is to be considered a subjective junior synonym of Erebia pandrose pandrose (Borkhausen, 1788). 3.4. Erebia pandrose marmolata Dannehl, 1927 and E. pandrose infraclara Verity, 1953 In the Alps, two subspecies of the sthennyo group have been described. The type local- ity of marmolata, in spite of its name, ıs the Pala group (Dannehl, 1927 p. 3), the type locality of ssp. infraclara ıs Val Milbach above Sappada in the Carnic Pre-Alps (Verity 1953, p. 132). Samples 25 and 28 have been collected at both type localities. Valve morphology and wing design are strikingly similar. Indeed, Dannehl’s description of the marmolata upperside ıs an accurate description of the sthennyo form inhabiting the southern Alps from the Adige valley to the Tagliamento valley, including Verity’s infra- clara. According to Warren (1936) the ground colour of the hindwing underside is the main differentiating character. In both sample 25 and 28, however, fresh males have the silvery grey hindwing underside which Verity considered typical of infraclara, whereas in worn males the ground colour has turned into a rusty grey-brown, fitting Dannehls description of marmolata. The only real difference is found in the sprinkling of dark scales on the hindwing underside, which is denser in Pala specimens than in Sappada specimens. Before describing infraclara (1953), Verity explicitly denied the subspe- cific nature of marmolata, stating that each of the marmolata characters mentioned by Dannehl, is found in other regions as well. As was shown earlier in this paper, this is correct: sthennyo characters do occur in many alpine populations. In fact, Verity misled himself by taking the description of marmolata as a reference, instead of material from the type locality. Had Verity’s collection contained specimens from the Pala group, ssp. infraclara would not have been described. Consequently, Erebia pandrose infraclara Verity, 1953 is to be considered a subjective junior synonym of Erebia pandrose mar- molata Dannehl, 1927. The currently known range of ssp. marmolata covers the Bergamasque Alps, the Lessinic Alps, the Venetian Pre-Alps, the adjacent Pala group, the Carnic Pre-Alps, the Julian Alps, and the Durmitor. As to the Monte Baldo: photographs of the (poor) existing material (Sala 1996, Sala pers. com.) suggest that the population belongs to the sthennyo group. However, this could not be confirmed by genital examination, as no material was available for dissection. The populations inhabiting the northern and western Dolomites, the Brenta group, and Eastern Tyrol are morphologically interme- diate between marmolata and pandrose. 3.5. Geographical variation within ssp. marmolata (Fig. 7) In the Bergamasque Alps 27% of the individuals have only the two apical spots on the forewing upperside. In all other investigated samples this feature was encountered only incidentally. In the Julian Alps the forewing design is extremely variable, which is typi- 342 Cupepo: Erebia pandrose — sthennyo complex group, the latter in relation to their ranking in table 3. Open circle: sum < 50; Circle with black centre: sum > 50. cal of populations of a mixed origin. The observed variation might be due to intermin- gling with pandrose from the Karawank mountains, which, however, is not supported by alower SP value. 3.6. The Asian populations of the sthennyo group Asıan populations of Erebia pandrose have been described as ssp. orientalis Goltz, 1930. This name, a junior homonym of Erebia epiphron orientalis Elwes, 1900, was replaced by yernikensis Korshunov, 1995 (Korshunov & Gorbunov 1995). Both the type locality (Sayan mountains) and the excellent description unmistakably apply to representatives of the pandrose group, represented in this study by samples 41 and 42. The population of the sthennyo group inhabiting the Kazakhstan part of southern Altai (sample 43), has not been described yet. It differs from the other known populations of the sthennyo group by its elongate forewing shape, and by the forewing ocelli being positioned as in the pandrose group (Figure 8). It is described here as a new subspecies within the sthennyo group. Nota lepid. 30 (2): 329-353 343 Erebia pandrose (sthennyo) narymica ssp. n. (Figs 8, Al-B$8) Material. Holotype: ©, label in Cyrillic script (figure 9), transliteration: “V[ostochno]-Kazlakstanskaya] obl[ast] Bolschenar.[ymskyi] r[ayo]n | s[elo] Novoberezovka 2800 [m]! A.G.Aniskovich | 24. VI 1986’, CFC. — Paratypes: 10, label in Cyrillic (figure 9), transliteration: ‘S.[elo] Novoberezovka 3000 m. | Bolshenarymskyi r-[ayo]n | Vostochno-Kazakst.[anskaya] obl.[ast’]| E. pandrose 2.7.1986.’, CFC; 10° ‘E. Kazakhstan | South Altai | Narymsky range | Novoberezovka v. 12800 m. 125-26.06.1986 | A. Aniskovich leg.’, CFC; 10°, 19 ‘3-7-1986 USSR | Vostoëno-Kazachst. | S. Novoberjezovka | 3000 m leg. Aniskovic’, ZMAN; 1, label in Cyrillic script, transliteration: ‘V-Kaz obl Bolschenar. rn | s. Novoberezovka 2800 | A.G.Aniskovich | 30.VI 1986’, CSC.; 20, label in Cyrillic script, transliteration: “V-Kaz obl Bolschenar. rn | s. Novoberezovka 2800 | A.G.Aniskovich | 2.VII 1986’, CSC.; 40°, 19, ‘E. Kazakhstan | South Altai | Narymsky range | Novoberezovka v. 12800 m. 125-26.06.1986 | A. Aniskovich leg.’, CSC. Holotype and paratypes form a series collected by A. Aniskovich & G. Makhat on a trip in the Narymsky Range from 24 June until 3 July 1986. The specimens were papered and dispersed among several collectors, who wrote their own labels, accord- ing to Aniskovich’s data. The holotype and three paratypes bear handwritten labels by Aniskovich. Description. Male forewing length: 21-23 mm. Forewing with pointed apex; upper- side pattern indistinct, submarginal zigzag line absent; position of forewing ocelli as in Erebia p. pandrose, not shifted towards the wing margin as in Erebia p. sthennyo. Hindwing underside design vague, ocelli and marginal elements absent. Male genitalia: Dorsal ridge of valve provided with a prominent spines, with mean of 2.8 spines per valve in studied sample. Diagnosis. Differs from all subspecies of the Erebia pandrose (pandrose) group by the spines on the valve and the indistinct pattern of the forewing upperside and the hind- wing underside. Differs from all other subspecies of the Erebia pandrose (sthennyo) group by the pointed forewing, and by the forewing ocelli being situated less close to the wing margin. Etymology. The name refers to the type locality, the Narym mountains (Narymsky Range). Geographical distribution. Apart from the type locality (South Altai, Narym Moun- tains in East Kazakhstan, Bolshenarymskoye district, south-east of the city of Novo- berezovka), the new ssp. is known from the adjacent Sarym-Sakty mountains (CSC). Remark. Males from the Lystviaga and Kholsun mountains (Russian Altai) have simi- lar valves, but the wing design tends towards yernikensis (V. Lukhtanov, pers. comm. ). Probably intermediate populations exist. 4. Checklist 4.1. Erebia pandrose (group pandrose) Diagnosis: mean number of spines per valve < 0.5. Design of forewing upperside and hindwing underside complete and distinct. Erebia pandrose (pandrose) pandrose (Borkhausen, 1788). (=Erebia pandrose (pandrose) ingana Fruhstorfer, 1911, syn. n.) Distribution: Alps. Absent from the area of the sthennyo group in the south-east (see below), and from the Chartreuse and the Vercors in the west (Willien 1990). 344 Cupebo: Erebia pandrose — sthennyo complex Fig. 7. European representatives of the sthennyo group of E. pandrose. A. ssp. sthennyo, Central Pyrenees (sample 1). B. ssp. marmolata, Bergamasque Alps (sample 11-13). C. ssp. marmolata, Pala group (sample 25). D. ssp. marmolata, Julian Alps (sample 32). e. ssp. marmolata, Durmitor (sample 38). Nota lepid. 30 (2): 329-353 345 Fig. 8. Erebia pandrose (sthennyo) narymica ssp. n., (A=upperside, B=underside) and Erebia pandrose (pandrose) yernikensis Korshunov, 1995, (C=upperside, D=underside). Row 1-7: males; row 8: females. Al and Bl: holotype, A2 — B8 paratypes. 346 Cupepo: Erebia pandrose — sthennyo complex Note: Most populations are in variable degree intermediate between the pandrose group and the sthennyo group. This is most prominent in the eastern Alps. Erebia pandrose (pandrose) sevoensis Willien, 1975. Distribution: Apennines. One single population on the Pizzo di Sevo in the Monti della Laga, Lazio (Chiavetta 2000). Erebia pandrose (pandrose) yernikensis Korshunov, 1995. Distribution: Altai and Sayan Mountains (Siberia), recently also found in the Barguzin Mountains in Transbaikalia (S. Churkin, pers. comm.). Erebia pandrose (pandrose) gracilis v.d. Goltz, 1930. Distribution: Pyrenees. Occupies a restricted region in the French provinces of Arrıege and Pyrénées Orientales, in the Spanish provinces of Lleida and Girona, and in Andorra (Gomez Bustillo & Fernandez-Rubio 1974, Willien 1990). Erebia pandrose (pandrose) roberti Peschke, 1920. Distribution: northwestern Carpathians. Found in the High Tatra, Lower Tatra, Belér Alps and Liptau Mountains (Moucha 1959, Krzywicki 1966, 1982). Erebia pandrose (pandrose) cibiniaca Dannehl, 1927 Distribution: eastern and southern Carpathians. Found in the Bihar, Retezat, Cibin, Bucegi, and Rodna Mountains (Hormuzaki 1901, Varga 1971). Erebia pandrose (pandrose) ambicolorata Varga, 1971. Distribution: Rila mountains (Varga 1971). Note: Subspecies roberti, cibiniaca, and ambicolorata are closely related. They share the pronounced design, especially of the marginal elements, on the hindwing underside. The differences have been worked out by Varga (1971). Erebia pandrose (pandrose) lappona Thunberg, 1791. Distribution: Scandinavia, Kanin peninsula and Kolguev Island (Warren 1936, Tatarınov & Dolgin 1999). Note: A rather heterogeneous group of populations that are morphologically inter- mediate between the pandrose group and the sthennyo group. They are provision- ally placed within the pandrose group. 4.2. Erebia pandrose (group sthennyo) Diagnosis: the mean number of spines per valve >1. Design of forewing upperside and hindwing underside incomplete and more or less suffused. Erebia pandrose (sthennyo) sthennyo (Graslin, 1850). Distribution: Pyrenees. Confined to the French departments of Pyrénées Atlan- tiques, Hautes Pyrénées, Haute Garonne and Arriège, and the Spanish provinces of Lleida and Huesca (Gömez Bustillo & Fernändez-Rubio 1974, Willien 1990). Erebia pandrose (sthennyo) marmolata Dannehl, 1927. (= Erebia pandrose (sthennyo) infraclara Verity, 1953, syn. n.) Nota lepid. 30 (2): 329-353 347 Kay et BOAguenap. pH ‘Qg €. O6oSapejsolKa 2800 > ATAHUCKOÖDUN Prwue Zuesoperiiekus Pr Boerousr- Kazarer, acte Fig. 9. Ereb drose (sth ; ; ye : ig. 9. Erebia pandrose (sthen- £ PrauTrE LE # og VEC is ma n., label of holotype (top) and of paratype no. | (bottom). Distribution: the ssp. occupies three separated subareas. a. The Bergamasque Alps. b. The southeastern Alps, from the Lessinic Alps (possibly from Monte Baldo) to the Julian Alps, and penetrating into the Pala group of the Dolomites. c. The Durmitor (Montenegro). Probably all populations in Bosnia-Herzegovina, Montenegro, Macedonia (JakSıc 1988), and in Albania (Thurner 1964) belong to this subspecies. Erebia pandrose (sthennyo) narymica ssp. n. Distribution: Hitherto known from the Narym and Sarym-Sakty mountains in the Kazakhstan part of the Altaı [Note 4]. Notes. l. Differentiating characters between pandrose and sthennyo have been taken from De Lesse (1952) and Warren (1936). De Graslin (1850), describing Erebia sthen- nyo, was not even aware of the existence of E. pandrose. He described the species as being different from what he called E. manto but what, according to his descrip- tion, most probably was E. pronoe. 2. From the present data it cannot be inferred whether pandrose colonised the Pyre- nees already during the Riss glaciation, or reached them from the Alps during Würm. 3. As far as distributional events (expansion, or withdrawal into refuges) are being related to a glacial or interglacial period, this should be regarded a minimum dating. Glacial cycles may have passed by without recognisably affecting the overall distri- bution of a taxon. 348 Cupepo: Erebia pandrose — sthennyo complex 4. It may well be that specimens of this subspecies have already been known to lepi- dopterists for a long time, possibly even from other localities. Warren (1936) writes that “in the Sayan the f. aglauros is the prevalent male form”, and Varga (1971) states that the Scandinavian and the Central Asiatic form are very similar, character- ised by its hazy, monotonous colouring and frequent occurrence of the f. aglau- ros. Both statements are in sharp contrast with v. Goltz’ description of ssp. orientalis (=yernikensis ) and with the habitus of the yernikensis populations investigated for this study, but do apply to narymica. Acknowledgements I am indebted to those who helped to bring together the material for this study: Mr. Sergei Churkin (Moscow, RU), Prof. Vladimir Lukhtanov (St. Petersburg, RU), Dr. Roman Yakovlev (Barnaul, RU), Dr. Oleg Kosterin (Novosibirsk, RU), Prof. Nils Ryrholm (Uppsala, S), Dr. Jostein Engdal (Saetre, N), Mr. Giinther Stangelmaier (Villach, A), Mr. Adriano Teobaldelli (Sforzacosta, I), Mr. Giovanni Sala (Salo, I), Mr. Jaap Poot (Beek, NL); to the curators who gave me access to museum collections or loaned me material: Dr. Rienk de Jong (RMNH Leiden, NL), Dr. Willem Hogenes (ZMAN Amsterdam, NL), Dr. Peter Huemer (TLM “Ferdinandeum” Innsbruck, A), Dr. Mihai Stänescu (NMNH “Grigora Antipa” Bukarest, RO) and Dr. Norbert Lenz (Löbbecke Museum Düsseldorf, D); to the Administration of the Gran Paradiso National Park (Torino, I) for the authorisation to collect a sample at the type locality of ssp. ingana within protected area. I am particularly grateful to Dr. Thomas Schmitt (Trier, D) and Dr. Marta Vila (A Coruña, E), whose comments led to substantial improvements of the manuscript. References Arnscheid, W. 1981. Die Macrolepidopteren-Fauna des Sonnental-Nonsberggebietes (Val di Sole und Val di Non in Oberitalien) (Insecta: Lepidoptera). — Studi Trentini di Scienze Naturali 57(1980): 95-245. Borkhausen, M. B. 1788. Naturgeschichte der europäischen Schmetterlinge nach systematischer Ord- nung. — Varrentrapp und Wenner, Frankfurt. Briquet, J. 1906. Le développement des flores dans les Alpes occidentales, avec aperçu sur les Alpes en général. Pp. 130-173 — In: Wettstein, R., Wiesner, J. & A. Zahlbruckner (eds): Wissenschaftliche Er- gebnisse des Internationalen Botanischen Kongresses Wien 1905. — Gustav Fischer, Jena. Brockmann-Jerosch, H. & M. Brockmann-Jerosch 1926. Die Geschichte der Schweizerischen Alpen- flora. Pp. 1110-1215 - Jn: Schröter, C. (eds): Das Pflanzenleben der Alpen. — Raustein, Zürich. Chiavetta, M. 2000. Le farfalle d’Italia. Atlante biogeografico. — Nuovo editoriale Grasso, Bologna, 111 pp. Chodat, R. & R. Pampanini 1902. Sur la distribution des plantes dans les alpes austro-orientales et plus particulièrement d’un choix de plantes des alpes cadoriques et venitiennes. — Le Globe 41: 1-40. Cupedo, F. 1997. Die geographische Variabilität und der taxonomische Status der Erebia manto bubastis- Gruppe, nebst Beschreibung einer neuen Unterart (Nymphalidae: Satyrinae). — Nota Lepidopterolo- gica 20: 3-22. Cupedo, F. 2004. Geographische Variabilität und spätglaziale Einwanderwege von Erebia pluto (de Prun- ner, 1798) in der Ortlergruppe und den Otztaler Alpen (Nymphalidae). — Nota Lepidopterologica 26: 137-152. Dannehl, F. 1927. Neue Formen und geographische Rassen aus meinen Rhopaloceren-Ausbeuten der letzten Jahren. — Mitteilungen der Münchner entomologischen Gesellschaft 17: 1-8. Esper, E. J. C. 1781. Die Schmetterlinge. Erster Teil, zweiter Band. — Walthers, Erlangen. Fruhstorfer, H. 1911. Neue paläarktische Rhopaloceren. — Societas Entomologica 25: 95-96. Goltz, [D]. H. v.d. 1930. Neue Erebienformen. — Deutsche Entomologische Zeitschrift Iris, Dresden 44: 78-82. Goltz, D. H. v.d. 1938-1939. Bemerkungen über Erebien. — Deutsche Entomologische Zeitschrift Iris, Dresden 52(1938): 160-171; 5301939): 36-48; 74-88. Nota lepid. 30 (2): 329-353 349 Gömez Bustillo, M. R. & F. Fernändez-Rubio 1974. Mariposas de la Peninsula Iberica. II Ropalöce- ros. — Ministerio de Agricultura, Madrid. 258 pp. Graslin, M. A. de 1850. Notice sur quelques lépidoptères nouveaux, trouvés dans les Pyrénées Orientales en 1847. — Annales de la société entomologique de France 19: 391-397. Hartig, F. 1937. I macrolepidotteri di Madonna di Campiglio. - Memorie della Societa Entomologica Italiana 16: 232-271. Henriksen, H.J. & I. Kreutzer 1982. The butterflies of Scandinavia in nature. — Skandinavisk Bogforlag, Odense, Denmark. 215 pp. Hewitt, G. M. 2000. The genetic legacy of the Quaternary ice ages. — Nature 405: 907-913. Higgins, L. G. 1975. The classification of European butterflies. — Collins, London. 320 pp. Holdhaus, K. 1954. Die Spuren der Eiszeit in der Tierwelt Europas. — Abhandlungen der zoologisch-bota- nischen Gesellschaft in Wien 63: 1-493. Hormuzaki, C. von 1901. Ueber die in den Karpaten einheimischen Arten der Gattung Erebia Dalm. Und deren Beziehungen zur pleistocänen Fauna Mitteleuropas. — Deutsche entomologische Zeitschrift Iris 15: 353-380. ICZN 2000. International code of zoological nomenclature, fourth edition. — Natural History Museum, London. 126 pp. JakSic, P. 1988. Privremene karte rasprostranjenosti dnevnih leptira Jugoslavije (Lepidoptera, Rhopalo- cera). — Societas Entomologica Jugoslavica, editiones separatae 1: 1-216. Korshunov, Y. & Gorbunov, P. 1995. Butterflies of the Asıan part of Russia. A guide. — Ekaterinburg. 202 pp. Kostrowicki, A. S. 1969. Geography of the Palaearctic Papilionoidea (Lepidoptera). — Drukarnia Uniwer- sytetu Jagiellonskiego, Kraköw. 371 pp. Krzywicki, M. 1966. Klucze do oznaczania owadöw Polski, XX VII (Lepidoptera), 63 (Satyrıdae). — Panst- wowe Wydawniktwo Naukowe, Warszawa. 41 pp. Krzywicki, M. 1982. Der gegenwärtige Stand der Tagfalterfauna Polens unter besonderer Berücksichtigung ihrere Bedrohung. — Nota Lepidopterologica 5: 3-16. Kudrna, O. 1986. Butterflies of Europe. Vol. 8: Aspects of the conservation of Butterflies in Europe. — Aula- Verlag, Wiesbaden. 323 pp. Lafranchis, T. 2000. Les papillons de jour de France, Belgique et Luxembourg et leurs chenilles. — Collec- tion Parthénope, Editions Biotope, Mèze (France). 448 pp. Lattes, A., P. Mensi, L. Cassulo & E. Balletto 1994. Genotypic variability in western European members of the Erebia tyndarus species group (Lepidoptera, Satyrıdae). — Nota Lepidopterologica Supplement No. 5: 93-104. Lattin, G. de 1967. Grundriss der Zoogeographie. — Fischer Verlag, Stuttgart. 499 pp. Leraut, P. 1980. Liste systématique et synonymique des Lépidoptères de France, Belgique et Corse. — Société entomologique de France, Paris. 334 pp. Lesse, H. de 1951. Contribution à l’étude du genre Erebia (4° note). Répartition de E. pandrose (Borkh.) et de sa sous-espèce sthennyo Grasl. dans les Pyrénées. — Vie et milieu 2: 267-277. Lesse, H. de 1952. Contribution à l’étude du genre Erebia (7° note). Erebia sthennyo Grasl. est-il une bonne espèce ? — Revue Française d’Entomologie 13: 217-219. Manley, W. B. L. & Allcard, H. G. 1970. A field guide to the butterflies and burnets of Spain. — E. W. Classey Ltd, Hampton. 192 pp. & 40 pls. Moucha, J. 1959. Neueste Forschungsergebnisse über unsere Lepidopteren-Fauna und deren Bedeutung für die Lösung zoogeographischer Fragen in der Tschechoslowakei (Lepidoptera). — Acta faunistica entomologica musei nationalis Pragae 4: 3-81. Nekrutenko, Y. P. 1990. Dnevnye babochki Kavkaza. Opredelitel’. Semeistva Papilionidae, Pieridae, Sa- tyridae, Danaidae. [Butterflies of the Caucasus. A Guide. Families Papilionidae, Pieridae, Satyridae, Danaidae] — Naukova Dumka, Kiev. 215 pp. 32 pl. [In Russian] Penck, A. & E. Brückner 1909. Die Alpen im Eiszeitalter. 3 Teile. - C. H. Tauchnitz, Leipzig. 1199 pp. Peschke, R. 1920. Erebia lappona v. roberti Peschke. — Internationale Entomologische Zeitschrift 145: 118-119. Reichl, E. R. 1992. Verbreitungsatlas der Tierwelt Osterreichs. Band 1: Lepidoptera — Diurna, Tagfalter. — Forschungsinstitut fiir Umweltinformatik, Linz. 6 pp. 216 pl. 350 Cupepo: Erebia pandrose — sthennyo complex Roos, P. & Arnscheid, W. 1976. Zur Verbreitung und subspezifischen Gliederung von Erebia pandrose Bkh. im Alpenraum (Lep., Satyridae). Beitrag zur Kenntnis der Erebien, III. — Nachrichtenblatt der Bayerischen Entomologen 25: 119-123. Sala, G. 1996. I lepidotteri del compensorio gardesano.— Societa Editoriale Multimediale, Salo (BS). 160 pp. Schmitt, T. & G. M. Hewitt 2004. Molecular biogeography of the arctic-alpine disjunct burnet moth species Zygaena exulans (Zygaenidae. Lepidoptera) in the Pyrenees and Alps. — Journal of Biogeo- graphy 31: 885-893. Schmitt, T., G. M. Hewitt & P. Müller 2006. Disjunct distribution during glacial and interglacial peri- ods in mountain butterflies: Erebia epiphron as an example. — Journal of Evolutionary Biology 19: 108-113. Schönswetter, P., I. Stehlik, R. Holderegger & A. Tribsch 2005. Molecular evidence for glacial refugia of mountain plants in the European Alps. — Molecular Ecology 14: 3547-3555. Stehlik, I. 2000. Nunataks and peripheral refugia for alpine plants during Quaternary glaciation in the mid- dle parts of the Alps. — Botanica Helvetica 110: 25-30. Stehlik, I. 2003. Resistance or emigration? Response of alpine plants to the ice ages. — Taxon 52: 499— oy LOR Tatarinov, A. G. & M. M. Dolgin 1999. Bulavousye CeSuekrylye. In: A. Estafyev (ed): Fauna Evropejskogo severo-vostoka Rossi, To. VII, ch. 1. [Club-antenned Lepidoptera. In: A. Estafyev (ed): Fauna of the European north-east of Russia, Vol. VII, part 1.] — Nauka, Sanct-Petersburg, 183 pp. Thurner, J. 1964. Die Lepidopterenfauna Jugoslavisch Mazedoniens. I. Rhopalocera, Grypocera und Noc- tuidae. — Prirodonaucen Muzej Skopje, 159 pp. Tribsch, A. & P. Schönswetter 2003. Patterns of endemism and comparative phylogeography confirm pal- aeoenvironmental evidence for Pleistocene refugia in the Eastern Alps. — Taxon 52: 477-497. Tuzov, V., P. Bogdanov, A. Devyatkin, L. Kaabak, V. Korolev, V. Murzin, G. Samodurov & E. Tarasov 1997. Guide to the butterflies of Russia and adjacent territories (Lepidoptera, Rhopalocera). Vol. 1. — Pensoft, Sofia-Moscow, 480 pp. incl. 79 pls. Varga, Z. 1971. Die Erebien (Lep.: Satyridae) der Balkanhalbinsel und der Karpaten III. Die Verbreitung und subspezifische Gliederung von Erebia pandrose (Bkh., 1788) nebst Beschreibung einer neuen Unterart: E. pandrose ambicolorata ssp. nova. — Acta biologica Debrecina 11: 227-235. Varga, Z. 1975a. Zoogeographische Gliederung der Paläarktischen Orealfauna. — Verhandlungen des Sech- sten Internationalen Symposiums über Entomofaunistik in Mitteleuropa: 263-385. Varga, Z. 1975b. Geographische Isolation und Subspeziation bei den Hochgebirgs-Lepidopteren der Bal- kanhalbinsel. — Acta entomologica Jugoslavica 11(1—2): 5-41. Varga, Z. 1996. Biogeography and evolution of oreal lepidoptera in the Palaearctic. — Acta Zoologica Aca- demiae Scientiarum Hungaricae 42(4): 289-330. Verity, R. 1953. Le farfalle diurne d’Italia, vol. V. - Marzocco S.A., Firenze. pp. 124-226. Vila, M. J. R. Vidal-Romani & M. Björklund, 2005. The importance of time scale and multiple refu- gia: Incipient speciation and admixture of lineages in the butterfly Erebia triaria (Nymphalidae). — Molecular Phylogenetics and Evolution 36: 249-260. Warren, B. C. S. 1936. Monograph of the genus Erebia. — British Museum, London. Willien, P. 1990. Contribution lépidoptérique française à la cartographie des invertébrés Européens (C.LE.) XVI. Le genre Erebia (Lépidoptères Nymphalidae Satyrinae). — Alexanor 16(5): 259-290. Willien, P. & T. Racheli 1975. Description d’une nouvelle sous-espèce d’Italie de Erebia pandrose Bork- hausen (Lep. Satyridae). — Entomops No 35: 102-104. Nota lepid. 30 (2): 329-353 351 Appendix Tab. 1. Sampling sites and sample sizes. Listed from west to east approximately. --- = sampled at more than one locality. WD = sample size for male wing design. MG = sample size for male genital characters. FG = sample size for female genital characters. sample | massif or region | locality | Pyrenees.E [Eastern Pyrenees f= Dauphiné _| Dauphiné Alps f= = SD DID IA le ID ID IR IS DIR IR ID Re D TIR IHR ID IR [WWW IR INO RK IRIS | Lote ID US DID IR | IS IR [Un DIRIRIOIRI00101R || IS ID IC | [00 BIO IS IO ID | OO ISA {OPO IN |) PAIN IS ID ID IR ID ICO IR IQUIS IS ID IQ Is ISO = Zee EE EE Se seele ee ed sey li ee OO | PN IS ICO IRIS IR IN IIS fol w {oo DIN IA IND IS | O ISA IN ISIN [BIW IC lO [W/E IR ICO] IS IS ID IS || | — ep) op) countr — S.Bernh 2 Penninic.N Wallis Penninic.S Val St. Barthélém [Bernese Alps [= AlbulaAlps f= | Bernina oam.W Pizzo Tre Signori gam.C Passo Vivione gam.E Passo Aprica Umbrailpass Gaviapass 7 Martelltal Adamello Pso Croce Domini Passo Grostè i Karwendel Karwendel mountains Rofan mountains =) we) DIX IT us) ao) ac) ale) A QC CIOICITIDIE es | BIS + N I® Eee © | un - Sais) = S |S @ |o |@ PS pe fe e |» a © OU =) Ele = Ron |I- — 10 © | > £ © a5 ® IN BIB] [Oo [Oo [Oo [Oo [Oo JOJO | 9 | Go [D [D IN ID IR ID III IN [Re lu lu le le lo be be bo cl Un | }oolpo =) —|/OlOlOINI NI NIB IS ID IS |OLOLOIN IAN! INI BIlWINO IS IS lola PR IS ID | ~ + "7 Pizzo Tre Signori Passo Vivione | Passo Aprica Umbrailpass | Gaviapass | Martelltal /Pso Croce Domini [Passo Groste SIEB | Karwendel mountains | [Rofan mountains Lessinic Dolom.Lat Dolom.Mar Dolom.Pala H. Tauern | Carnic Alps g Gerlitzen Julian [JulianAlps fT KZ 14 | 552 Cupepo: Erebia pandrose — sthennyo complex Tab. 2. Population means of the measured parameters. SP = number of spines per valve. OP = ocelli position. FWD = forewing design. HWD = hindwing design. AV = width antevaginal plate (mm). sd = standard deviation (for interval/ratio variables with adequate sample size). Sample Q ic Pyrenees.C 1.0623 al 6 Pyrenees.E 0.5150 |0. 5 A 0.72 Dauphine 0.2887 Pt.S.Bernh 0.6279 Gr. Parad. 0.2505 Penninic.N 10.154 [0.5547 Penninic.S 10.400 [0.6222 Bernese [0.107 10.2841 Albula 0.100 _ [0.2754 Bernina 10.042 10.1412 Bergam.W 0.9244 . a DÉS © i) DD | Bergam.C 0.7739 0.73 Bergam.E 0.5869 0.83 Ortler. W 0.2520 |0.266 |0.0184 [436 |500 [0.69 Ortler.S 0.6778 Ortler.N 0.5409 110.252 |00261 |3.147 MSN ON Adamello 0.4295 Brenta 0.292 [0.5526 Karwendel 0.5207 [0.288 |00260 1392 |396 |- | Rofan 0.1826 0.1398 | 3.63 0.84 Lessinic 0.8650 0.82 Dolom.Lat 0.7012 0.77 Dolom.Mar 0.4820 0.81 Dolom.Pala 1.3664 0.78 H. Tauern 0.4667 0.80 Venetian 0.9063 0.83 Carnic 1.3961 0.79 Amberg 0.2996 0.61 Gerlitzen 0.6463 0.72 N. Tauern 0.3737 0.73 Julian 1.1991 0.89 © Norway.S 10.261 [0.4510 0.74 Ötztal 0.4592 Sweden.N 0.3466 0.67 Norway.N 0.5075 0.71 I Un Tatra 0.119 10.3502 0.74 37 | Bucegi 0.0000 [0.271 10018 15.00 1696 [0. Durmitor 1.0535 0.84 39 0.1508 0.73 Sevo 0.1021 0.87 Altai 0.3349 Sayan 0.2306 43 | Narym 1.3076 [0.275 10.0098 [3.07 _ |ı00 [0.69 | Nota lepid. 30 (2): 329-353 35 | Go Tab. 3. Population means for SP, OP, FWD and HWD, rescaled to 0-100, and their rescaled sum. The populations (pandrose group only) are sorted according to increasing sum. 2 a: FWD SUM Bucegi OO OU > SI EN © Un Itai — \O — ofan atra Ibula Un Un ND ON Un I DD Un ioe) D ee) N I Ne) D |W|\O BIN N |W} | WIN IN Teele SLO TN INI DIN | D | NHN] oO u D ni = | OR oo I U9 Oo Un DD > NS) U) w Un Penninic.N Pyrenees.E auphiné N Un SS) Oo I Oo ie) ernina ee I Oo Ne) OU) N ernese Ötztal Penninic.S Pt.S.Bernh damello mberg Un Un Un es > O0 I N Un | Go > wlus0|Z|5/Zl0l0/z)> 2 \w | |O olrlel3lzlole 2! = © ba O oO |= 0 & & = = ate À 5 = = NE = w \O Un OO 3 >) SQ [Où OO | Un [Un © |Un (SY) Un N I erlitzen rtler.N orway.S olom.Mar orway.N olom.Lat Grim Un No) ey) ON Blolalian iN > —_ IS) un = > IR AIRJBEIR[OIKRIKR [Dp lolatie OININ/OlEIMIn!lwl]D alo oO co | aS U ON On DIRIRIDN i) os = I Ne DIN © à LUCE LO, [© = D I D = > =) Od Un Oo I OU) I DD Un Oo OO Un OO SI Un AN 5 Go No) — © © : Oo Sweden.N N — © >) Un > Wa |) | | | In | oo D [© | | ID IN |W |W IR In Wir [D | © ID |© = Se OO IQ ee) ON Un OO (oe) — © © I — — RP [OS ID ID WIN [W]e Jw OS ID | mm | m = |N WIN IRIS | 5 Ortler.S 100 58 19 100 Nota lepid. 30 (2): 355-359 355 Une nouvelle espece d’Hyponephele du Pakistan (Nymphalidae: Satyrinae) JERÖME PAGES Agrocampus, 65 rue de Saint-Brieuc, F-35042 Rennes cedex; e-mail: jerome.pages@agrocampus-rennes.fr Resume. Une nouvelle espèce, Hyponephele filistigma sp. n., est décrite de la région de Chitral, Pakistan. Elle appartient au groupe H. davendra (Moore, 1865) et se distingue immédiatement des autres espèces du groupe par une longue tache androconiale filiforme pour le mâle et, pour les deux sexes, au dessous des ailes postérieures, par une ligne postdiscale brune présentant un angle aigu vers l’extérieur très prononcé au niveau de l’espace 3. Abstract. A new species, Hyponephele filistigma sp. n., is described from the vicinity of Chitral (Pakistan). It can be distinguished readily from the other species of the H. davendra (Moore, 1865) group, to which it belongs, by its long and filiform androconial band for the male, and, for both genders, on the underside of the hind wing, by the postdiscal line with the outer acute angle prominent in area 3. Introduction On trouve dans la région de Chitral, quatre espèces d’Hyponephele (Muschamp, 1915) du groupe davendra (Moore, 1865). Trois sont connues et nous rappelons les principa- les caractéristiques de leur habitus. Une quatrième forme présente des caractéristiques bien marquées qui suggèrent qu’il s’agit la d’une espèce distincte des précédentes mais aussi des autres du groupe (des synthèses récentes se trouvent dans Carbonell (1997, 2000) et Samadurow et al. (1997)). Nous décrivons ci-après cette nouvelle espèce, en référence aux trois précédentes que l’on trouve dans des zones assez voisines mais aussi par rapport à H. brevistigma (Moore, 1892), espèce de haute altitude connue, pour le Pakistan, de la partie est (Hunza) et dont on trouve une description dans Carbonell (1997) et des représentations dans Tshikolovets (2005b). Hyponephele tenuistigma (Moore, 1892), avec la sous-espèce laspura (Evans, 1932) Localité type: Pakistan, Chitral, au-dessus de 9,000 pieds. Cette espèce de taille moyenne (©: 34-38 mm; 9: 39-41 mm) présente, pour le mâle, une tache androconiale assez longue (1 cm) et étroite (environ 2 mm). Au dessous des ailes antérieures, une ligne discale est généralement nette, critère cité par d’Abrera (1992) pour séparer tenuistigma de davendra mais qui est encore plus caractéristique chez chitralica (Evans, 1932), traitée ci-après. Une caractéristique de la sous-espèce laspura ne semble pas avoir été citée: les nervures de la face supérieure des ailes anté- rieures sont soulignées de noir. Les exemplaires que j’ai capturés dans les environs de Chitral sont bien conformes au type déposé au BMNH (Londres, Angleterre). Evans (1932) indique: « Chitral over 9,000 ft, rare ». Nous avons observé cette espèce entre 2700 m et 3200 m dans une seule zone des environs de Chitral. En revanche, je l’ai observée en plusieurs autres endroits des montagnes du Nord du Pakistan (Hunza, Astor). Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 356 Paces: A new Hyponephele from Pakistan Bu i 4 q fi ern Hi PI 1. De haut en bas : H. davendra (Chitral), H. chitralica (Chitral), H. laspura (Chitral), H. brevistigma (CS Passu, Pakistan; @ Sabu, Ladakh). Pour chaque ligne, de gauche à droite: mâle (dessus, dessous); femelle (dessus, dessous). Nota lepid. 30 (2): 355-359 Ss) | Un PI 2. H. filistigma n. sp. En haut, holotype male (Chitral) dessus dessous. En bas, allotype femelle (Chitral) dessus dessous. Hyponephele chitralica (Evans, 1923) Localité type: Khyber (Pakistan). Cette espèce de taille moyenne (©: 34-38 mm ; 9: 39-41 mm) est caractérisée, pour le mâle par une tache androconiale entrecoupée par les nervures qui ne longe que par- tiellement la cellule et, sur le revers des ailes antérieures des deux sexes, par une ligne discale bien marquée. Une autre caractéristique de cette espèce, assez nette lorsque l’on compare des séries, ne semble pas avoir été citée : la forme arrondie des ailes antérieu- res. Un syntype de cette espèce peu connue est représenté dans Tshikolovets (2005a). Evans (1932) indique: « Chitral, not rare ». Ceci est en accord avec le fait que j’ai ob- servé cette espèce en des endroits variés de 1600 m (pelouses sèches de la vallée de la Kunar river) à 2800 m (allées forestières). Je ne l’ai pas observée ailleurs qu’à Chitral. Hyponephele davendra (Moore, 1865) Localité type: Spiti (Tibet). Espèce de taille plutôt grande (S': 41-43mm ; 9: 45mm). Les exemplaires que j’ai cap- turés aux environs de Chitral ressemblent bien à ceux de la région de Kaboul figurés par Wyatt & Omoto (1966) ou par de Freina & Aussem (1986) ainsi qu’à ceux d’Afghanis- tan figurés par Sakai (1981). En première analyse, ils ne diffèrent pas particulièrement d'exemplaires trouvés en plusieurs endroits des montagnes du Nord du Pakistan. La 358 Packs: A new Hyponephele from Pakistan tache androconiale est grande mais pas au point de rattacher spontanément la forme de Chitral au taxon latistigma (Moore, 1893) (voir par exemple l’exemplaire figuré par Nazari (2003), ou ceux, méme si cela apparait un peu moins nettement, figurés dans Eckweiler (2005)). Toutefois, il est prudent de dire que ce rattachement n’est pas une certitude absolue. Hyponephele filistigma sp. n. PL 2 Matériel. Holotype ©, ‘Pakistan | Chitral 10 km W | 3200 m | 26 vii 1996’, leg. J. Pages, coll. Muséum National d’Histoire Naturelle de Paris. — Allotype 9, ‘Pakistan | Chitral 10 km W | 3200 m | 4 vii 2006’, Leg. et Coll. J. Pages. — Autres paratypes (40°, 39: Leg. et Coll. J. Pages): 10°, ‘Pakistan | Chitral 10 km W | 3500 m | 26 vii 1996’; 10°, ‘Pakistan | Bumburet | 3200 m | 26 vii 1997’; 10°, ‘Pakistan | Chitral 10 km W | 3500 m I 14 vii 2006’; 10°, ‘Pakistan | Kalam 7 km N | 2400 m | 1 vii 2006’; 29, ‘Pakistan | Chitral 10 km W | 3500 m | 10 vii 2006’; 19, ‘Pakistan | Kalam 7 km N | 2200 m | 1 vii 2006’. Description. Mâle. Envergure entre 39 et 44 mm. Dessus aile antérieure : fond jaune orangé bordé de brun-gris ; forme peu élancée ; la tache androconiale, longue (plus d’un cm de long) et filiforme (moins d’un mm de large) borde la cellule en allant de la base de l’aile au milieu de l’espace 3 ; franges grises à la base et blanches sur leur moitié extérieure. Dessus aile postérieure : couleur de fond brun-gris uniforme avec un fin liséré près de l’angle anal. Dessous aile antérieure : fond jaune orangé plus clair que le recto ; ligne postdiscale bien visible ; elle sépare une aire intérieure sensible- ment plus foncée que l’aire extérieure ; irrégulière, elle montre en particulier un angle aigu vers l’extérieur au niveau de l’espace 3. Ocelle apical généralement assez grand. Dessous aile postérieure : allure générale comme brevistigma mais avec des dessins plus nets ; fond gris clair ; ligne postdiscale brune, bordée extérieurement de blanc ; elle présente un angle aigu vers l’extérieur très prononcé au niveau de l’espace 3 ; deux lignes foncées, submarginale et marginales, délimitent un bande claire ; deux ocelles proéminents dans les espaces 2 et 5. Femelle. Envergure entre 41 et 45 mm. Dessus aile antérieure. Fond jaune orangé bordé de brun-gris ; deux ocelles bien marqués dans les espaces 2 et 5. Bande postdis- cale à peine visible. Dessus aile antérieure. Comme chez le mâle. Dessous. Identique à celui du mâle, excepté un second ocelle souvent présent dans l’espace 2. Variation individuelle. Faible. Au-dessous, la ligne postdiscale des anté- rieures est plus ou moins nette. Diagnose. Par rapport aux autres espèces du groupe, la tache androconiale, longue et filiforme, permet d’identifier le mâle au premier coup d’ceil. Pour les deux sexes, le dessous est proche de celui de brevistigma mais avec des dessins plus nets et une ligne postdiscale brune, bordée extérieurement de blanc présentant un angle aigu vers l’exté- rieur très prononcé au niveau de l’espace 3 (chez brevistigma cet angle existe mais est moins marqué). Deux lignes foncées, submarginale et marginales, délimitent un bande claire (chez brevistigma il y a un simple liséré clair submarginal). Biotope. Hyponephele filistigma n. sp. a été trouvé de la fin juin à la fin juillet sur des pentes rocailleuses à des altitudes plutôt élevées (3000-3500 m) dans la région de Chitral et à une altitude moindre (2200-2500 m) dans les environs de Kalam (Swat valley). Nota lepid. 30 (2): 355-359 359 Remerciements Il m’est agréable de remercier V. Tshikolovets qui m’a fourni une photographie d’un syntype de A. tenuis- tigma laspura. References D’Abrera, B. 1992. Butterflies of the holarctic region. Part Il : Satyridae (concl.) & Nymphalidae (partim) pls 85-334. Hill House, Victoria (Australia). 149 pp., 71 pls. Carbonell, F. 1997. Hyponephele shirazica n. sp. en Iran méridional (Lepidoptera: Nymphalidae, Saty- rinae). — Linneana Belgica 16: 99-104. Carbonell, F. 2000. Découverte de la femelle d’Hyponephele shirazica Carbonell, 1997. (Lepidoptera : Nymphalidae, Satyrinae). — Linneana Belgica 17 : 271-272. Eckweiler, W. 2005. Neue Taxa der Gattung Hyponephele Muschamp, 1915 aus Iran (Lepidoptera: Nym- phalidae, Satyrınae). — Nachrichten des Entomologischen Vereins Apollo, N. F. 26 (4) : 217-220. Evans, W. H. 1932. The identification of the Indian butterflies, 2nd ed., Bombay Natural History Society, Madras. 484 pp., 32 pls. Freina, de J. J. & B. Aussem 1986. Hyponephele urartua sp. n. eine neue Satyriden-Art aus Ostanato- lien. — Nota lepidopterologica 9: 200-212. Nazari, V. 2003. Butterflies of Iran. Teheran (National Museum of Natural History of the Islamic Republic of Iran, Dayereh-Sabz). 568 pp., 74 pls. Sakai S. 1981. Butterflies of Afghanistan. Tokyo, Kodansha. 272 pp., 48 pls. Samadurow, G. D., W. A. Korolew & W. W. Tschikolovez 1997. Eine Übersicht über die Satyriden der Gattung Hyponephele Muschamp, 1915. V. Die Arten Hyponephele latistigma, (Moore 1893), H. bre- vistigma (Moore, 1893), A. issykkuli Samodurow, 1996, H. korshunovi Lukhtanov, 1994, H. comara (Lederer 1870), H. tenuistigma (Moore, 1893) und H. argyrostigma Tuzov & Samodurow, 1997. — Atalanta 2 : 69-105. Tshikolovets, V. 2005a. The butterflies of Kyrgyzstan. — Nekrutenko édit. Brno-Kiev. 511 pp., 108 pls. Tshikolovets, V. 2005b. The butterflies of Ladak (N.-W. India). — Nekrutenko édit. Brno-Kiev. 176 pp., 30 pls. Wyatt C. & K. Omoto 1966. New Lepidoptera from Afghanistan. — Entomops 6: 169-200. 360 Book reviev Tuzov, V. K. & G. C. Bozano 2006. Guide to the butterflies of the Palearctic Region. Nymphalidae part II. Tribe Argynnini: Boloria, Proclossiana, Clossiana. — Omnes Artes, Milano. 72 pp. ISBN 8887989079, 40.00 €. This book is the latest addition to the series “Guide to the butterflies of the Palearctic Region” edited by G. C. Bozano, a series that is renowned for splendid figures and detailed taxonomic treatment of the various groups of Palearctic butterflies. The current volume deals with the Argynnini genera Boloria, Proclassiana, and Clossiana, commonly known as “small fritillaries”. The group reaches its highest diversity in the Eastern Palearctic region with several little known or doubtful species or subspecies. The book is for that reason alone a very welcome contribution to the study of Palearctic butterflies. The authors take a rather conservative but also refreshingly bold approach to the classification of the group. They are conservative with respect to the higher classification where they retain the three classic genera instead of merging all three into one unified Boloria. As the precise relationships between these genera at the present still are somewhat uncertain, this approach is fairly sound, though using a large unified Boloria for all three genera probably would have been a more straightforward approach. On the other hand, the authors make some quite bold statements when it comes to the taxonomic status of some of the species/subspecies, giving full species status to what they term “most of the ‘strong’ subspecies”. This approach is commendable as it forces students of the group and conservationists alike to focus on problematic taxa that could otherwise be dismissed as “mere” subspecies. Furthermore, though some of these species may later be “sunk” to subspecies again, a detailed analysis of the true status of these species falls well outside the scope of the book. And by giving full species status to what may or may not be a species, the authors point to future subjects for taxonomic research within the group. Along these lines the authors also recognize a wide range of subspecies. Though the use of subspecies is debated, this approach too seems sound, as it allows students and collectors to name different geographical forms. Furthermore, some of these subspecies may later prove to be full species. By recognizing them here, the authors enhance the chance that future studies will focus on these groups and perhaps recognize them as full species. Each genus is described with its diagnostic characters, and a full checklist of species and subspecies found in the Palearctic region is provided, including all species and several subspecies found outside the region. Each species is provided with a full list of synonyms, a description of diagnostic characters, a list of recognized subspecies (including synonyms), a discussion of taxonomic problems (if any), a brief summary of the distribution range of each subspecies, and an easily understood (albeit small) distribution map. Each species is richly illustrated with high quality photos (or in rare cases, drawings from older literature) of the upper and under sides of both sexes for almost all subspecies, and important recognition characters are clearly marked on the figures and explained in the text. Drawings of the male genitalia or parts of the male genitalia are provided for all species. These drawings are not quite of the same standard as the other illustrations. The drawing styles, level of details, and, in some cases, angle of view are not consistent between drawings. This can make comparing different drawings problematic at times. The bibliography at the end of the book is a very comprehensive (if not exhaustive) account of the literature on the group and as such a very important tool for any student of these genera. To sum up: this volume of the “Guide to the butterflies of the Palearctic Region” is a very high quality, and virtually indispensable tool for any amateur, student, or researcher who wishes to work on taxonomic or systematic aspects of Boloria, Proclossiana, and Clossiana. The book is by far the most comprehensive taxonomic treatment of the group to date. THOMAS SIMONSEN Nota lepid. 30 (2): 361-365 361 Une nouvelle espece de Pseudochazara du Pakistan (Nymphalidae, Satyrinae) JERÖME PAGES Agrocampus, 65 rue de Saint-Brieuc, F-35042 Rennes cedex; e-mail: jerome.pages@agrocampus-rennes.fr Resume. Pseudochazara annieae sp. n. a été découvert dans la vallée de Swat (Nord Pakistan). Le mâle est immédiatement reconnaissable par ses ailes antérieures de couleur brun gris foncé ne présentent qu’une faible trace de bande postmédiane et ses ailes postérieures présentant, sur le même fond brun gris foncé, une bande postmédiane fauve orangé. La femelle présente ces mêmes caractéristiques à un degré moindre avec un fond brun clair. Dans les deux sexes, le revers des postérieures est gris avec des lignes postmédiane et antémarginale bien marquées. L’habitus des mâle et femelle est illustré en couleur en vues dorsale et ventrale. L’habitus de Pseudochazara droshica (Tytler), l’espece la plus semblable a P. annieae, est éga- lement 1llustré. Abstract. Pseudochazara annieae sp. n., was discovered in the Swat Valley (Pakistan). The male can be distinguished immediately from the other Pseudochazara by its dark grey-brown forewings with only a trace of a postdiscal band and by a orange postdiscal band on the hindwings. The female presents the same features but less prominent and with a clear brown background. In the two genders, the verso of the hindwings is grey with marked postdiscal and sub-marginal lines The habitus is illustrated in colour for both sexes in dorsal and ventral view. Pseudochazara droshica (Tytler), the species most similar to P. annieae is also illustrated. Introduction On trouve dans la vallée de Swat (Nord Pakistan) une espèce se référant au genre Pseudochazara présentant un habitus bien caractéristique: de grande taille, les ailes antérieures du mâle sont de couleur brun gris foncé et ne présentent qu’une faible trace de bande postmédiane alors que les ailes postérieures montrent une bande postmédiane frappante fauve orangé. Bien que le Nord Pakistan ait donné lieu a des prospections entomologiques depuis plus d’un siècle, cette espèce ne semble pas avoir été signalée. Nous donnons ci-après quelques points de repère bibliographique. L'ouvrage récent de T. J. Roberts (2001) cite, pour le Pakistan, deux espèces du gen- re Pseudochazara (en les affectant au genre Eumenis): mniszechii gilgitica (Tytler) qu’il cite des provinces suivantes: Balouchistan, frontière du Nord Ouest (North West Frontier Province: NWFP) et régions du Nord. (Northern Areas: NA); il considère cette espèce synonyme de baldiva (Moore); telephassa (Hübner), qu’il cite du Balouchistan et de la région frontalière ouest de la province de la frontière du nord ouest. Il considère cette espèce synonyme de lehana (Moore). L'ouvrage ancien mais fort complet de Talbot (1947) mentionne quant à lui les Pseu- dochazara suivants pour l’ensemble Inde + Pakistan (en les affectant au genre Hip- parchia): mniszechii balucha (Evans), Balouchistan; mniszechii droshica (Tytler), Chitral, Gilgit, Baltistan; mniszechii gilgitica, Chitral, Gilgit, Baltistan; mniszechii lehana, Ladak; mniszechii baldiva, Shipki, Kunawur to western Tibet; et telephassa, Balouchistan Khyber pass. Ces taxa sont repris (dans le genre Hipparchia) par Mani (1986) qui n’en ajoute pas d’autres pour |’Himalaya. L’ouvrage de Wynter-Blyth (1957), dédié au sous-continent Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 362 Paces: A new Pseudochazara from Pakistan Fig. 1. Pseudochazara annieae n. sp. En haut, holotype ©, recto verso ; en bas, allotype 9, recto verso. indien, mentionne, quant à lui (dans ce groupe taxonomique mais en les rattachant au genre Eumenis), les seuls taxa mniszechii (sans mention de sous-espèce) et rele- phassa. Concernant la faune indo-pakistanaise, il convient enfin de mentionner le taxon Hip- parchia diffusa Butler, 1880; ce taxon n’a été repris par aucun des « grands » auteurs, Evans (1932) et Talbot (1947). Kudrna (1977) le mentionne parmi les types du BMNH qu'il a examinés et le considère comme synonyme de Hipparchia semele semele (Lin- naeus), en suggérant, à propos de la localité type (India: Punjab: Ravi River): « locality certainly erroneus ». Les spécimens de l’espèce décrite ici présentent un habitus très éloigné de celui de semele. Il en est de même dans les régions limitrophes. Tshikolovets (1997) mentionne du Pamir: droshica badachshana Wyatt & Omoto, droshica rajevskyi Tshikolovets, sa- gina (Staudinger) et baldiva. Koçak & Kemal (2006) mentionnent d’Afghanistan: kani- shka Aussem, telephassa, droshica, panjshira Wyatt & Omoto, porphyritica Clench & Soumanov, sagina et turkestana (Groum-Grshimailo). Sakai (1981) mentionne, pour l’Afghanistan: baldiva, droshica, lehana et telephassa. Enfin, Nazari (2003) ne men- tionne que felephassa pour les Pseudochazara du sud de l’Iran. En dehors des faunes « régionales », on trouve aussi (par exemple dans Funet (2007)) le taxon pakistana Gross décrit en 1978 à partir d'exemplaires de Ziarat (Balouchistan pakistanais) et représenté dans Tuzov (1997) selon lequel la distribution est assez vaste, incluant en particulier 1’ Hindukush. Nota lepid. 30 (2): 361-365 363 Fig. 2. Pseudochazara droshica Tytler. En haut, ©, recto verso, 6.v11.1997, Birmoglasht (Chitral) ; en bas, Q, recto verso, 9.v111.1997, Bumburet (Chitral). Ce n’est pas le lieu ici de trancher les options systématiques de ces auteurs. Il suffit pour notre propos de retenir que, dans tous les taxa mentionnés, les deux sexes présen- tent une bande postmédiane jaune bien marquée sur les deux ailes. En s’éloignant du Pakistan et en considérant l’ensemble des espèces du genre Pseu- dochazara, seuls mamurra (Herrich-Schaffer) (Turquie, Iran, Irak), beroe (Herrich- Schaffer) (Turquie, Elbourz et Kopet Dagh), /ydia (Staudinger) (Turquie) et shakuhen- sis (Staudinger) (Turquie, Elbours et Kopet Dagh), présentent une bande postmédiane a peine marquée ; mais ceci vaut pour les deux ailes ce qui, ajouté a leur plus petite taille, leur fond beaucoup plus clair, leur forme moins élancée et le revers de leurs ailes postérieures aux dessins confus leur confèrent une allure bien distincte de l’espece de la vallée de Swat (cf. par exemple les iconographies de D’ Abrera (1992), Hesselbarth et al. (1995), Tuzov (1997), Tshikolovets (1997, 1998) et Sakai (1981)). Tout ceci conduit a décrire cette nouvelle espece. Pseudochazara annieae n. sp. (Fig. 1) Matériel. Holotype ©, ‘Pseudochazara annieae | holotype | Pakistan | Kalam 2100m | 29 vi 2006 | leg. J. Pages’, déposé au Muséum National d'Histoire Naturelle de Paris. — Allotype 9, ‘Pakistan | Kalam 2100m | 4 vii 2006’, leg. et coll. J. Pagès. — Autres paratypes (309, 29: leg. et coll. J. Pages sauf 10 coll. J. Mairiaux et 10° coll. Kiev Zoological Museum) ; mêmes localités avec 150° (29 vii 2006), 150 (4.v11.2006) et 29 (4.vii.2006). Description. Mâle. Envergure entre 52 et 57 mm. Dessus aile antérieure : fond brun gris ; bande postmédiane à peine visible, légèrement plus claire que le fond ; deux 364 Pacés: A new Pseudochazara from Pakistan ocelles bien nets légèrement cerclés de jaune dans les espaces 2 et 5; quelquefois, deux petits points blancs dans les espaces 3 et 4. Taches androconiales, bien visibles sous certaines incidences, formant une bande médiane entrecoupée par les nervures; fran- ges blanches entrecoupées de noir à l’extrémité des nervures. Dessus aile postérieure: couleur de fond identique à celle des ailes antérieures; bande postmédiane, limitée aux espaces 2, 3, 4 et 5; cette bande est large de couleur fauve vif, ayant un bord interne zigzaguant mais bien délimité; petit ocelle net dans l’espace 2; fine ligne antémarginale noire très légèrement sinueuse. Dessous aile antérieure: fond gris; plage médiane fauve foncé séparée d’une large bande postmédiane plus claire par une fine ligne grise à peine visible; les deux ocelles apparaissent nettement. Dessous aile postérieure: fond gris clair avec des dessins noirs assez nets; deux lignes brisées, postmédiane et submargi- nale, apparaissent nettement en délimitant une aire légèrement plus claire; petit ocelle dans l’espace 2. Femelle. Envergure entre 58 et 60 mm. Dessus aile antérieure: fond brunâtre, plus clair que celui du mâle; éclaircies jaunes dans la partie discale; bande postmédiane plus apparente que chez le mâle, mais discontinue, réduite essentiellement à deux plages jaunes autour des deux ocelles proéminents. Dessous aile postérieure: comme chez le mâle mais couleur de la bande jaune orangé moins vif. Dessous identique à celui du mâle. Variation individuelle: très faible sur les exemplaires mâles, capturés, il est vrai, la même année et dans deux stations assez voisines de même altitude. Les trois femelles étudiées sont légèrement variables quant à la taille et l’étendue des plages claires. Diagnose. Espèce de grande taille, peut-être la plus grande dans le genre Pseudochazara. Chez le dessus du mâle, la bande claire postmédiane habituelle des autres Pseudochazara est, aux ailes antérieures, à peine plus claire que le fond et, aux ailes postérieures, est presque aussi large que chez droshica mais d’une nuance sensiblement plus orangée. Ce contraste entre aile antérieure et postérieure rend cette espèce immédiatement re- connaissable. Enfin, la forme de la plage androconiale, qui s’apparente à celle des espè- ces du groupe Hipparchia semele, n’a pas d’équivalent dans le genre Pseudochazara. Chez la femelle, sur le dessus des ailes antérieures, la bande postmédiane est plus appa- rente que chez le mâle mais reste beaucoup plus réduite que chez les femelles des autres espèces. Celle des ailes postérieures est large comme chez le mâle, de la même nuance orangée mais plus foncée, ici encore sans équivalent dans le genre. Etymologie. Cette espèce est dédiée à mon épouse Annie, qui m’accompagne dans bien des aventures. Biotope. Pseudochazara annieae n. sp. a été trouvé sur des pentes rocailleuses à rela- tivement faible altitude (2 100 m). En 2006, les mâles volaient dès la fin de juin, attirés par des chardons. Quelques femelles sont apparues début juillet. Très peu d’autres espè- ces volaient dans ces biotopes secs; citons seulement Strymonidia sassanides (Kollar), assez commun dans la région. Nota lepid. 30 (2): 361-365 505 Remerciements Il m’est agréable de remercier Bernard Landry pour l’excellent accueil qu’il a réservé à ce travail. Références D’Abrera, B. 1992. Butterflies of the holarctic region. Part II : Satyridae (concl.) & Nymphalidae (partim) pls 85-334. — Hill House, Victoria (Australia). 149 pp., 71 pls. Evans, W. H. 1932. The identification of the Indian butterflies, 2nd ed. - Bombay Natural History Society, Madras. 484 pp., 32 pls. Funet, 2007. Pseudochazara de Lesse 1951. — Liste disponible sur http://www.funet.fi/pub/sci/bio/life/ insecta/lepidoptera/ Hesselbarth, G, H. Van Oorschot & S. Wagener 1995. Die tagfalter der Türkei. Vol 3. — S. Wagener édit., Bocholt (Allemagne). 847 pp., 128 pls. Koçak, A. O. & M. Kemal 2006. Checklist of the butterflies of Afghanistan. Disponible sur http://www. members.tripod.com/entlep/Af.htm Kudrna, O. 1977. A revision of the genus Hipparchia Fabricius. — E. W. Classey Ltd, Faringdon, Oxon. 300 pp., 40 pls. Mani, M. S. 1986. Butterflies of the Himalaya. — Oxford & IBH Publishing Co, New Delhi. 181 pp. 25 pls. Nazari, V. 2003. Butterflies of Iran. — National Museum of Natural History of the Islamic Republic of Iran, Dayereh-Sabz, Teheran. 568 pp., 74 pls. Roberts, T. J. 2001. The butterflies of Pakistan. — Oxford University Press, Karachi. 200 pp., 57 pls. Sakai S. 1981. Butterflies of Afghanistan. — Kodansha, Tokyo. 272 pp., 48 pls. Talbot, G. 1947. The fauna of British India. Vol IT : Danaidae, Satyridae, Amathusiidae, Acraeidae. Indian reprint edition (1986). — Today & Tomorow Printers & Publishers, New Delhi. 506 pp., 2pls. Tshikolovets, V. 1997. The butterflies of Pamir. — Tshikolovets édit., Kiev-Brno. 282 pp., 46 pls. Tshikolovets, V. 1998. The butterflies of Turkmenistan. — Nekrutenko édit., Kiev. 237 pp., 34 pls. Tuzov (V. K.), 1997. Guide to the butterflies of Russia and adjacent territories. Vol I : Hesperiidae, Papilio- nidae, Pieridae, Satyridae. — Pensoft publishers, Sofia. 480pp., 79 pls. Wynter-Blyth, M. A. 1957. Butterflies of the Indian region. Indian reprint edition (1986). — Today & To- morow Printers & Publishers, New Delhi. 523 pp., 72 pls. Er > rm D i L fh Nota lepid. 30 (2): 367-373 367 A new Amata species from Israel (Arctiidae, Syntominae) Tuomas J. Witt !, VasiL1Y D. KRAVCHENKO ?, WOLFGANG SPEIDEL °, JosEF MoOoSsER *, AMY JUNNILA ° & GUNTER C. MULLER ® Museum Witt, Tengstr. 33, D-80796 Munich, Germany; e-mail: thomas@ witt-thomas.com Department of Zoology, Tel Aviv University, Tel Aviv, Israel; e-mail: vasiliy@post.tau.ac.il Museum Witt, Tengstr. 33, D-80796 Munich, Germany; e-mail: speidel-wolfgang@ web.de Seilerbruecklstr. 23, D-85354 Freising, Germany; e-mail: jomooser@aol.com Department of Parasitology, McGill University, Macdonald Campus, Ste-Anne-de-Bellevue, Québec H9X 3V9, Canada; e-mail: amyj@sympatico.ca Department of Parasitology, Kuvin Centre for the Study of Infectious and Tropical Diseases, The Hebrew University — Hadassah-Medical School, Jerusalem, Israel; e-mail: guntercmuller@hotmail.com vu F&F © NO — fo) Abstract. A new Amata species, Amata gil sp. n., is described from Mount Hermon, Israel. It is related to Amata libanotica (Bang-Haas, 1906) and A. mestralii (Bugnion, 1837), but can be easily discriminated from these species by its considerably smaller size and its lack of the yellow ring on the first abdominal segment which is normally present in the two other species, though it can be reduced in A. mestralii. Here, the distribution, ecology, and phenology of the new species are reported and the habitats are pictured in colour. Zusammenfassung. Eine neue Amata-Art, Amata gil sp. n., wird vom Berg Hermon, Israel, beschrieben. Sie steht Amata libanotica (Bang-Haas, 1906) und A. mestralii (Bugnion, 1837) nahe, aber kann von die- sen Arten leicht unterschieden werden durch beträchtlich kleinere Größe und durch das Fehlen des gelben Ringes am ersten Abdominalsegment, der bei den beiden anderen Arten normalerweise vorhanden ist, obgleich er bei einigen Individuen von A. mestralii auch reduziert sein kann. Die Verbreitung, Okologie und Phänologie werden dargestellt und Habitate werden auf einer Farbtafel abgebildet. Introduction Until recently, the name Amata Fabricius, 1807 has been widely used as the gener- ic descriptor of this group and the name Syntomis Ochsenheimer, 1808 treated as a junior synonym (e. g. Obraztsov 1966; Holloway 1988; Edwards 1996; Nielsen 1996). However, a separate genus Syntomis has been accepted by Schneider et al. (1999). The monophyly of the genus Syntomis sensu Schneider et al. is very well supported by the presence of androconial hairbrushes on the foreleg coxa and by molecular genetic data (Schneider et al. 1999). However, the monophyly of Amata, which lacks hairbrushes on the forecoxa, is not supported by the three published phylogenetic trees based on partial sequences of the mitochondrial 16S rRNA gene. These trees show Amata to be polyphyletic, if Syntomis and Hydrusa are accepted as separate genera (Schneider et al. 1999). Therefore, at present, it seems best to retain Amata in the wide sense as defined by Holloway (1988), but to recognize a monophyletic Syntomis section within that genus to which the new species belongs. A generic separation of Syntomis from Amata would also neces- sitate the splitting of Amata s. str. in several genera and make the group rather unpractical, especially in the Oriental tropics. The new taxon can be further at- tributed to the phegea species group in the sense of Obraztsov (1966) because of the dark front of the adults. Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 368 Witt et al.: À new Amata from Israel Material and methods Besides the specimens mentioned in the description of the new species, the following material (all from Museum Witt) of related species was examined: Amata sintenisi (Standfuss, 1892): 360°, 99 from Asia Minor (Turkey), Amata sintenisi aurivala (Schawerda, 1923): 99 (including 3 paratypes) from Iraq and Asia Minor (Turkey), Amata tanina (de Freina, 1982): 700 (including holotype and 6 paratypes), 729 (in- cluding 5 paratypes) from Asıa Minor (Turkey), Amata libanotica (Bang-Haas, 1906): 30° from Lebanon, Amata taurica (Turati, 1917): 10° from Asia Minor (Turkey), 200° from Syria, Amata antiochena (Lederer, 1861): 90 from Asia Minor (Turkey), Amata mestralii mestralii (Bugnion, 1837): 180°, 439 from Lebanon and Syria, Amata mestralii antilibanotica Obraztsov, 1966: 20°, 19 (paratypes) from Antilibanon (mountain ridge separating Lebanon and Syria, exact locality (Zebdani) in Syria), ' Amata mestralii palaestinae (Bang-Haas, 1906): 540°, 59 from Israel. The present paper 1s based on the revision of Obraztsov (1966) and follows his termi- nology, including the wing pattern elements. In that monograph, the full synonymy of all mentioned species ıs indicated and the male genitalia are figured. Amata gil sp. n. | (Figs 1-5, 9) Material. Holotype: ©, ‘Israel | Hermon | 2200 m | 21. 6. 1969 | leg. Bytinski-Salz’, ‘coll. | Daniel’, ‘Amata sp. n. | nahe sintensis Stdf. | det. F Daniel 1972’, ‘Genitalpräparat | Nr 3379 | Museum Witt München’, ‘Holotypus | Amata gil | Witt & Kravchenko & Speidel & Mooser & Junilla & Müller’. Deposited in Museum Witt, Munich (MWM). — Paratypes: 10, ‘Israel | Hermon | 2200 m | 21.6.1969’ (MWM). 19, ‘Israel, 1900 m | Mt. Hermon | 1.v11.1986 | A. Freidberg’ (deposited in Tel Aviv University). 10°, ‘Israel | Nord Galiläa, Mt. Hermon | Lower Cable Station, 1500 m | 18.6.87 | coll. Mooser’ (deposited in coll. Mooser). 10, ‘Israel | Mt. Chermon [sic., for Hermon], 2000 m | 26.V.1967 | coll. Nisan’ (deposited in Tel Aviv University). 20, ‘Syria | Mt. Hermon, ca. 2600 m | 10.7.2003 | coll. R. Preiss’ (deposited in coll. R. Preiss). Description. Adult male (Figs 1-5). Head, antennae and body black, with yellowish white ring on abdominal segment 5. Underside of body and legs black, with few white scales on thorax and rarely on abdomen laterally. Wingspan: 18-20 mm (Holotype 19 mm). Forewing black, with white spots: basal spot (m1) round, spot m2 small, round, spot m3 a narrow oblique stripe, m4 elongate, spots m5 and m6 very close to each other and small. Hindwing white, with margin at costa and termen broad, black; black border broader near anal angle; anal margin only with very narrow black border; wing base also with black scales. Female. Unknown. ' There is a ‘subspecies’ Amata mestralii palaestinae ‘Hampson, 1898’ according to Obraztsov (1966). However, this name was introduced as Syntomis mestralii ab[erration]. palaestinae and so is unavail- able (ICZN art. 45.6.2). It was first cited in the sense of a geographical population as var. palaestinae by Bang-Haas (1906) and so it became nomenclatorially available (ICZN, 4" ed., art. 10.2). Amata mestralii palaestinae (Bang-Haas, 1906) doubtfully represents a valid biological entity. The large ma- terial of A. mestralii at hand seems to demonstrate no constant geographical variability. Nota lepid. 30 (2): 367-373 369 Figs 1-8. Adult moths (Photos by Igor Kostjuk). 1. Amata gil sp. n.’Israel, Hermon, 2200 m, 21. 6. 1969, leg. Bytinski-Salz’, ‘coll. Daniel’, ‘Amata sp. n. nahe sintensis Stdf., det. F. Daniel 1972’, ‘Genitalpräparat Nr 3379 Museum Witt München’. Holotype. 2. Amata gil sp. n. Holotype. Underside. 3. Amata gil sp. n. ‘Israel, Hermon, 2200 m, 21. 6. 1969’, ‘Museum Witt? (MWM). Paratype. 4. Amata gil sp. n. ‘Israel, Nord Galiläa, Mt. Hermon, Lower Cable Station, 1500 m, 18. 6. 87, coll. Mooser’. Coll. Mooser, Freising. Paratype. 5. Amata gil sp. n. ‘Israel, 1900 m, Mt. Hermon, 1. vii. 1986, A. Freidberg’. Tel Aviv University. Paratype. 6. Amata libanotica (Bang-Haas, 1906) ‘Libanon, Franz Daniel’, ‘Amata libanotica B.-H. & det. N. Obraztsov 1948’, ‘Fig. in N. Obraztsov Monogr. Amata’. 7. Amata mestralii (Bugnion, 1837) © ‘Israel, Nord-Gallilaa, 10 km E Quiryat-Shemona, 350 m, leg. de Freina’, ‘coll. de Freina, Museum Witt, München’. 8. Amata mestralii (Bugnion, 1837) @ ‘Meron, Israel, 5. v. 1975, leg. Kugler’, ‘Amata mestralii (Bugnion, 1837), det. T. Witt 2003’. Tel Aviv University. 370 Wirt et al.: À new Amata from Israel Male genitalia (Fig. 9, one specimen dissected). Uncus stout; vinculum with small saccus; valvae similarly shaped, pointed at apex; phallus ba- sally very broad, bulbus ejaculato- rius rather short, inserting at base of phallus. Diagnosis. The new species is close- ly related to Amata libanotica (Fig. 6) and A. mestralii (Figs 7, 8) because in these two species both valvae are more or less pointed, the right and left one of approximately the same shape. The genitalia structure of A. sintenisi is somewhat sımilar, but the valvae are more rounded at the apex. Amata libanotica always has a quite Fig. 9. Amata gil sp. n. Holotype, male genitalia. clear basal abdominal yellow ring dorsally on abdominal segment 1, which is weak or sometimes absent in A. mestralii. In this respect, the new species is very similar to A. mestralii, as the yellow ring is absent in all known specimens. Amata mestralii is therefore regarded as possibly the closest relative of the new species. However, in A. mestralii and A. liban- otica, the basal processus of the left valva is longer than the processus of the right one (Obraztsov 1966), whereas in the new species the right processus is longer than the left one. Nevertheless, the new species can be clearly distinguished from A. mestralii by the much smaller size and the smaller forewing spots in combination with the larger white area in the hindwing. Amata mestralii males have a wingspan of 33-40 mm. A. antiochena and A. tanina have a different shape of each valva: the right valva is angled at outer margin, with shorter processus basalis, whereas the left one is pointed, with longer basal processus (Obraztsov 1966; de Freina 1982). Comparative figures of the genitalia of all species can be found in Obraztsov (1966) except N. tanina, which is figured in the original description (de Freina 1982). Etymology. The species is dedicated to Gil D. Reichstadt-Ofarim, the well-known mu- sician who now lives in Munich. Distribution, ecology and phenology. The new species has so far only been collected from Mount Hermon where it inhabits elevations from 1500 to 2600 m. However, if its distribution pattern extends beyond this at all, it is probably restricted to the Levant. Mount Hermon is a cluster of mountains, mostly made of limestone, with three distinct summits comprising the southernmost part of the Anti-Lebanon mountain range. The highest elevation in Israel is only 2224 m while the highest peak (2800 m) is in Syria. Five of the six records of this new species are from the Tragacanth high altitude zone, which is restricted to the areas above 1900 m. This area receives snow cover and has Nota lepid. 30 (2): 367-373 Sa Range, with snow line at about 1500 m , mid winter. 11. View from Syria, towards the south, tragacanth vegetation on Mt Hermon on a wind- facing slope, about 2200 m, mid summer. 12. North-facing slope of Mt Hermon in Syria, about 2500 m, mid summer. 13. Large dolinas, a traditional collecting place on the Israeli part of the mountain ridge, near the upper cable station, about 2000 m, mid summer. 14. Rocky, south-facing summit with scattered bushes, near the lower cable station, about 1600 m, mid spring. very low temperatures in winter, whereas summers are hot and dry. This situation cre- ates specific plant communities dominated by spiny, round, dense, cushion-like shrubs such as Astragulus and Onobrychis (Danin1988). The main water source in this area is melting snow, consequently most of this karstic mountain area is rather arid (Danin 1995). Only one specimen was collected in flight on a xerotherm, karstic slope about 1500 m with few scattered Rosa canina L. and Crataegus sp. bushes (Rosaceae) (see Fig. 1). The two specimens from Syria were collected sitting on flowers growing among Astragalus sp. bushes (see Fig. 3) in the late morning. A. gil appears to be a summer 372. Wirt et al.: À new Amata from Israel species that flies from June to July. The area is well known for its wealth of indigenous anımals and plants, however, it is a sensitive border area between Israel, Lebanon and Syria and as such, it was, in the last few decades, difficult to collect material there, espe- cially on the peaks on which military installations are found (Furth 1975; Kravchenko et al. 2006). Nevertheless, the Tel Aviv University Entomology Department was able to conduct considerable day-time collecting activities in this area. Because this rather distinctive species is absent in local collections, it is probably rare. It should also be mentioned that although numerous light traps were operated for years by the Israeli- German Lepidoptera Project, on the Israeli part of Mt Hermon, and hundreds of other Amata were caught, all the specimens of the new species were collected during the day (Müller et al. 2006). The host plants of A. gil are unknown, but the larvae of the closely related A. mestralii in Israel are polyphagous on low herbaceous plants (unpublished data of the authors). Remark. Amata tanina, which is very similar in size to A. gil, is found in similar habi- tats at high elevations and has brachypterous females with extremely small wing rudi- ments. The related species A. mestralii has reduced wings and the female of the related Amata libanotica has fully developed wings. Accordingly it is difficult to predict the characteristics of the wings of the female of the new species. Acknowledgements We are thankful to all the colleagues and to the many generous Israeli citizens who helped with this survey. Moreover, we are grateful to the Israeli Nature and Parks Authority (NPA) who supplied the col- lecting permits. The authors would like to specially thank Dr. Rueben Ortal (Science and Conservation Division), Dr. Didi Kaplan (NPA Northern District Biologist), Mr. Yiftah Sinai (NRA Carmel District Biologist), Mr. Zeev Kuller (Central District Biologist), the staff of the NPA- regional rangers, Nature Reserves and National Parks directors of Israel, Prof. J. Kugler, Dr. A. Freidberg (Entomological collec- tion, University of Tel Aviv), Prof. J. Fittkau (former Head of the Bayrische Zoologische Staatssammlung, Munich, Germany), Dr. Hedva Pener, Dr. Laor Orshan and Dr. Heather Bromly-Schnur (Entomological Laboratory, Ministry of Health), the late Dr. Shoshana Yatom (Volcani Center, Bet Dagan), Mr. Yossi Lev Ari, and Mr. Giora Gissis (Bet Ushishkin Museum, Qibbutz Dan), the late Mr. Zeev Shoam (Qibbutz Neot Mordachai), and particularly Benni & Aliza Ben David of Kefar Sabba. This study would not have been possible without the generous help of Prof. Y. Schlein, Hebrew University, Hadassah Medical School. References Bang-Haas, A. 1906. Neue oder wenig bekannte palaearctische Macrolepidopteren. — Deutsche entomolo- gische Zeitschrift Iris 19: 127-144. Danin, A. 1988. Flora and vegetation of Israel and adjacent areas. Pp. 251-276. — In: Y. Yom-Tov & E. Tchernov (eds), The Zoogeography of Israel. — Junk Publishers, Dordrecht, The Netherlands. Danin, A. 1995. Vegetation maps. Pp. 32-33. — In: The New Atlas of Israel. The Survey of Israel and The Hebrew University of Jerusalem, Tel Aviv [in Hebrew]. de Freina, J. J. 1982. 3. Beitrag zur systematischen Erfassung der Bombyces- und Sphinges-Fauna Klein- asiens. Neue Arten der Gattung Syntomis Ochsenheimer, 1808, aus Türkisch-Kurdistan und Aserbeid- jan. — Nachrichtenblatt der Bayerischen Entomologen 31: 47-64. Edwards, E. D. 1996. Arctiidae. Pp. 278-286. — In: E: S. Nielsen, E. D. Edwards & T. V. Rangsi (eds), Checklist of the Lepidoptera of Australia. — CSIRO Publishing, Collingwood. Furth, D. G. 1975. Israel, a great biogeographic crossroad. — Discovery 11: 3-13. Hampson, G. F. 1898. Catalogue of the Lepidoptera Phalaenae in the British Museum 1. Catalogue of the Syntomidae in the collection of the British Museum. — Taylor and Francis, London. xxi + 559 pp. Nota lepid. 30 (2): 367-373 373 Holloway, J. D. 1988. The moths of Borneo. 6: Family Arctiidae, subfamilies Syntominae, Euchromiinae, Arctiinae; Noctuidae misplaced in Arctiidae (Camptoloma, Aganainae). — Southdene Sdn. Bhd., Kuala Lumpur. 101 pp., 168 figs, 6 color plates. Kravchenko, V. D., M. Fibiger, J. Mooser & G. C. Miiller 2006. The Noctuinae of Israel (Lepidoptera: Noctuidae). - SHILAP Revista Lepidopterologica 34: 353-370. Müller, G. C., V. D. Kravchenko, D. Chikatunov, R. Ortal, ©. Orlova, C. Li, T. Witt, W. Speidel, J. Mooser & A. Hausmann 2006. General aspects of the Israeli light-trap network concerning Coleoptera. — Esperiana 12: 269-281. Nielsen, P. S. 1996. Syntominae. P. 297. - In: O. Karsholt & J. Razowski (eds), The Lepidoptera of Europe. A distributional checklist. — Apollo Books, Stenstrup. 380 pp. Obraztsov, N. S. 1966. Die palaearktischen Amata-Arten (Lepidoptera, Ctenuchidae). — Veröffentlichungen der Zoologischen Staatssammlung München 10: 1-383. Schneider, D., L. Legal, W. Dierl & M. Wink 1999. Androconial hairbrushes of the Syntomis (Amata) phe- gea (L.) group (Lepidoptera, Ctenuchinae): A synapomorphic character supported by sequence data of the mitochondrial 16S rRNA gene. — Zeitschrift für Naturforschung 54e: 1119-1139. 374 Book reviev Gershenson, Z. S, T. Pavlitek, V. Kravchenko & E. Nevo 2006. Yponomeutoid Moths (Lepidoptera: Yponomeutidae, Plutellidae, Argyresthiidae) of Israel. — Pensoft Series Faunistica 58, Sofia-Moscow. 200 pp. Hardcover ISSN 1312-0174, 60.00 €. This book is nicely produced in a hard cover and at first sight it appears to be a faunal list with descriptions of species. On the back of the cover, however, is a further clue: “This monograph, part of the “Evolution Canyon” model research program, ...” After Preface and Introduction there is a chapter headed “General Points”. This contains a full and well researched section on the biology and ecology of the group, drawing largely on European literature; however it is remarkable that in the section on larvae there is no mention of the major work on chaetotaxy by Werner, nor in that dealing with pupae is there mention of the pioneering work of Patocka & Turéani. There is then a section on distribution where the data from within Israel read more like a research paper in a scientific journal than what one would expect to find in a book of this kind. Economic significance is the final section which is adequately presented. A small section on classification makes reference to molecular methods, but no attempt has been made to use such methods for a comparison with the phylogeny established using conventional morphological methods. Yponomeutoid in the title makes one imagine that all Yponomeutoidea may be treated, instead it is just the three families listed in the subtitle following Moriuti in his choice of Yponomeutidae s./. Chapter 5 entitled Systematics is, as one would expect, the largest part of the book, but not massively so. Each genus and each of the 25 species is briefly described with illustrations which are line drawings, of variable quality, usually depicting the adult, male and female genitalia, and a distribution map. There is then a comprehensive list of references and a single coloured plate illustrating just six of the larger well known species described in the text. The data used in producing this book come from just 12 localities, from the years 2000-2003, which means that the distribution maps cannot give a very clear picture of the range of each species. One always welcomes treatment of a group of small moths from a part of the world less well worked entomologically, but it is over ambitious to think that an exhaustive faunal list can result from so few data. It is likely that with further work more species may be discovered in Israel, but the species descriptions and illustrations make it possible that they might be overlooked. In these days it is disappointing that adult moths are not comprehensively illustrated, and the taxonomic relationships not explored more deeply. At the same time it is good to find the biology of a group so thoroughly described. Overall the book rather falls between stools: the data result from a three-year research project added to which is general material about the group and brief taxonomic treatment of the genera and species encountered; these components together are inadequate for a country fauna. Despite these reservations a faunal survey is always a stimulus for further study and exploration; if it can encourage more work on the microlepidoptera of this part of the world it will have been well worth while. Davip AGASSIZ Nota lepid. 30 (2): 375-386 575 Creatonotos omanirana sp. n. aus dem Oman und dem Iran (Artiidae: Arctiinae) JOSEF J. DE FREINA Eduard Schmid-Str.10, 81541 München, Germany; e-mail: defreina.j@online.de Abstract. A distinct species of Creatonotos Hübner, 1816, Creatonotos omanirana sp. n., is described from NE-Oman and the Iranian side of the Gulf of Oman (Hormozgan, Kerman). Details on life history, breeding trials, habitat preferences, and distribution are given. The holotype, male and female habitus and genitalia, male androconial organs, as well as larval instar and habitat are figured. The habitus of both sexes, genitalia, and corematal organs of the closely related Creatonotos gangis (Linnaeus, 1763) are compared and illustrated. Zusammenfassung. Eine neue Art der Gattung Creatonotos Hübner, 1816, Creatonotos omanirana Sp. n. wird aus dem Nordoman und der iranischen Seite des Golfs von Oman aus den Provinzen Hormozgan und Kerman beschrieben. Der Holotypus, Tiere beider Geschlechter, die Genitalmorphologie, die männlichen Duftorgane (Coremata), die Raupe sowie der omanische Lebenraum der neuen Art werden abgebildet und mit Merkmalen der nächstverwandten Creatonotos gangis (Linnaeus, 1764) verglichen. Zur Lebensweise, den Präimaginalstadien, dem Lebensraum und der Verbreitung von C. omanirana werden ausführliche Daten geliefert. Einleitung Die Fauna der Arabischen Halbinsel kennt drei Arten der Gattung Creatonotos Hübner, 1816: C. leucanioides Holland, 1893 (Hacker et al. 1999) und C. albidior Wiltshire, 1986, beides aethiopische Faunenelemente, sowie eine aus dem Nordoman bekannt gewordene und als C. gangis (Linnaeus, 1763) fehl interpretierte Art (Wiltshire 1986: 292, fig.), die in folgendem als Creatonotos omanirana sp. n. beschrieben wird. Das Taxon arabicum Hampson, 1896, ebenfalls lange Zeit in der Gattung Creatonotos kombiniert, ıst jetzt Typusart der monospezifischen Gattung Creataloum Dubatolov, 2006. Die Konspezifität jemenitischer Tiere mit C. leucanioides (loc. typ. Westafrika, Kamerun, valley of the Ogove River) bedarf der Bestätigung. Creatonotos omanirana Sp. n. ist der südwestlichste Vertreter einer vorder- bis osta- siatisch tropischen „gangis“-Artengruppe, die nordostwärts über Nordpakistan und die himalajanische Region bis Nordchina, südostwärts über die indonesische und phil- ippinische Inselwelt bis Queensland verbreitet ist. Habituell unterscheiden sich diese Arten nur unwesentlich. Genitaliter finden sich im weiblichen Geschlecht deutliche, bei den Männchen weniger auffällige arttypische Strukturen (vgl. hierzu Goodger & Watson 1995; Wiltshire 1980; Dubatolov 2006). Auffälliges Artmerkmal sind die unterschiedlichen androkonialen Duftorgane der Männchen. Form und Anzahl der Coremaschläuche sind arttypisch. Die variable Größe der Coremata ist dagegen ta- xonomisch ohne Belang. Der Grund hierfür sind quantitativ unterschiedliche aufge- nommene Mengen an Pyrrolozidin-Alkaloiden während der Larvalphase (Boppré & Schneider 1985; Schneider & Boppré 1981; Schneider et al. 1982). Es ist naheliegend, dass sich die Arten auch in der chemischen Zusammensetzung ihrer Pheromone und pyrrolizidinen Alkaloide unterscheiden (Bell & Meinwald 1986; Wunderer et al. 1986; Hartmann 1999). Die neue omanische Art wird in folgendem beschrieben und mit der Typusart der Gattung Creatonotos, C. gangis (Linnaeus, 1764) verglichen. Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 376 DE FREINA: Creatonotos omanirana sp. n. Abkürzungen CDF coll. de Freina, München Gen.Präp. de Fr. Genitalpräparat de Freina Hflgl. Hinterflügel MWM Museum Witt, München Vfigl. Vorderflügel ZSM Zoologische Staatssammlung, München Creatonotos omanirana sp. n. Material. Holotypus ©: N-Oman, Westl. Hajar-Gebirge, ca. 45 km SSW Barka, Oase (Wadi) Abyadah, ca. 150-300 m, 04.-07.4.2006, leg. de Freina, ex coll. CDF(MWM). — Paratypen: 550°, 349, gleiche Daten wie Holotypus, davon 20° genital untersucht (Gen.Präp. de Fr. 2006/50, 2006/51), CDF (MWM); 259, 229, gleiche Daten wie Holotypus, Fl-Generation 08.-25.6.2006 (aus Muttertier 04.-07.4.2006, leg. de Freina), cult. de Freina, CDF (MWM), davon 20 genital untersucht (Gen.Präp. de Fr. 2006/52, 2006/53). 59,69, N-Oman, Jabal Nakhl-Gebirge, Oase Nakhl, ca. 450 m, 30.-31.3.2006, davon 10° genital unter- sucht (Gen.Präp. de Fr. 2006/54), CDF (MWM); 19, gleiche Daten, jedoch 12.12.2006, leg. et coll. de Freina (MWM); 240, 189, gleiche Daten, jedoch ex larva (aus L3-L5 04.-12.12.2006), 25.12.-20.1.2007, cult. et coll. de Freina (MWM), davon 10, 19 (Genitalien in Kopula vereint, Fig. 23) und 39 genital un- tersucht (Gen.Präp. de Fr. 2006/68, 2006/55 2006/56, 2006/66), ein Präparat Fühler+ Beine + Thorax + Abdomen (Präparat 2006/67); 250°, 219, gleiche Daten, F1-Generation 10.-20.4.2006 (Elterntiere 1.2007, cult. de Freina), cult. et coll. de Freina (MWM); 320°, 269, gleiche Daten, F2-Generation 10.-20.6.2007 (Elterntiere IV.2007, cult. de Freina), cult. et coll. de Freina (MWM); 439, 369, gleiche Daten, F3- Generation 16.—30.8.2007 (Elterntiere VI.2007, cult. de Freina), cult. et coll. de Freina; 110°, 69 gleiche Daten, jedoch 27.7.-10.8.2007 cult. et coll. Speidel, MWM; 1169, 849, gleiche Daten, F4-Generation 08.-30.10.2007 (Elterntiere VIII.2007, cult. de Freina), cult. et coll. de Freina (MWM). Weitere F4-Tiere mit gleichen Daten: 50,59 ZSM, 160°, 109, cult. et coll. Buchsbaum, ZSM, 200°, 109 cult. et coll. Hager (Wuppertal); 150, 129 cult. et coll. Pensotti (Mailand); 150°, 119 cult. et coll. Piatkowski (Hanau). 79, 69 N-Oman, Jabal-Akhdar-Siidauslaufer, ca. 28 km E Al Nizwa, 10 km E Izki, ca. 600 m, 04.04.2005, leg. et coll. de Freina (MWM); 1709, 129 N-Oman, Jabal Nakhl-Gebirge, ca. 16 km N Nakhl, Unteres Wadi Mistal, ca. 600 m, 08.-09.4.2005, leg. et coll. de Freina (MWM); 70°, 69, N-Oman, 18 km W Barka, Küstenbereich bei Ras al Sawadi, 0-10m, 01.-08.4.2006, leg. et coll. de Freina (MWM), davon 19 genital untersucht (Gen.Präp. de Fr. 2006/57); 18, 169, N-Oman, Jabal al Akhdar SO-Ausläufer, Umg. Birkat al Mawz, Eingang Wadi al Muaydin, ca. 700 m, 04.04.2005, leg. et coll. de Freina (MWM); 220°, 119, NO- Oman, Jabalal Akhdar, SO-Ausläufer, 10 km WNW Birkat Al Mawz, ca. 650 m, 04.04.2005, leg. et coll. de Freina; 169, 129, NO-Oman, Jabal Akhdar-Südausläufer, ca. 20 km NW Nizwa, Umg. Tanuf, Eingang Wadi Tanuf, ca. 850 m, 05.4.2005, leg. et coll. de Freina (MWM); 9, 39, Sultanat Oman, Tanuf, 630 m, N 23°03°08°E 57°27 27, V.2005, e. o. cult. Berger (MWM); 19, Nizwa 15 km North, 600 m, 11.2.2007, leg. Major, coll. Pensotti (Mailand); 10°, S-Iran, Hormozgan Prov., Beshagerd [Bashäkerd] Mts., 26°34°N 57°54’E, 900 m, 25.3.-05.04.2000, leg. Siniaev & Plutenko, ex coll. Schintlmeister (MWM) (Gen.Präp. de Fr. 2006/58); 19, Kerman, Senderk, 31.05.1977, leg. Sojak, ex coll. KruSek (MWM); 10°, Kerman, 30 km SSW Jiroft [= Sabzvaran], Mohamadabad, Maskün, 18.05.1977, leg. Sojak, ex coll. KruSek (MWM). Beschreibung (Fig. 1-9). Holotypus © Vorderfliigellange 18 mm, Paratypen © 16-22 mm,durchschnittlich 18mm. Fühlerschwarzgrau bis schwarz, filiform, Fühlergliederzahl 45; Vorderbein ohne, mittleres Bein mit einem, Hinterbein mit zwei Paaren kurzer, gedrungener Tibialspornen, Coxa, Femur und Tibia aller Beine an der Außenseite schwarz, innen rosabraun, die Tarsen schwarz; Thorax rosa ocker, Zeichnung von Kopf und Dorsum samtschwarz; Abdomen dorsal bis mediodorsal zinnoberrot, lateroventral dunkel lilabraun bis schwarzbraun, ventral rußig schwarz, das erste Abdominalsegment auffällig zitronengelb behaart, das Abdominalende lateral mit zwei schwarzgrauen, die Coremataöffnungen tiberdeckenden Haarbiischeln; Abdominalsegmente mediodor- sal mit schwarzem, fein gelb umrandetem Fleck, der des letzten Segments langlich mit deutlicher gelboranger Umrandung; mediolateral und medioventral sitzen weitere schwarze Punktfleckenreihen. Vorderfliigelgrundfarbe hell zimtfarben, postbasal bis Nota lepid. 30 (2): 375-386 S77 zur Zelle unterhalb der Mediane ein schwarzer, zur Zell hin erweiterter Keilfleck, da- hinter in den oberen beiden Medianfeldern ein schwarzer, von der Zelle ausgehend bis vor den Außenrand ein zweiter, kürzerer schwarzer Streifen, an den äußeren Enden der Diskodalader zwei fein rosa eingefaßte schwarze Punktflecken. Hinterflügel glasig rußig grau, subapikal und meist auch am Tornus je ein feiner schwarzer Fleck, der Saum dunkler rußig schwarz. Unterseite beider Flügelpaare rußig schwarz, Vflgl. mit schwacher Transparenz der oberseitigen Strichzeichnung. Weibchen. Vorderflügellänge 19 mm, Paratypen 18-22,5 mm, durchschnittlich 20,5 mm. Geschlechtsdimorphismus nicht sehr auffällig, Fühler wie beim Männchen filiform, die Flügelunterseiten jedoch deutlich heller als beim Männchen. Vorderflügelgrundfarbe lichter, mehr ockerfarben, Hinterflügel glasig weiß, die weißliche Unterseite beider Flügelpaare weiß mit deutlicheren schwarzen Submarginalflecken; Abdomen ventral licht grau mit deutlichen schwarzen Fleckenreihen, das Abdominalende um die Papilla analis hell ocker. Variabilität. Die Art variiert habituell nur unwesentlich. Die beiden Längswische nähern sich gelegentlich an, die feinen Diskalflecken im Vflgl. können sowohl fehlen als auch in der Anzahl zwischen einem und drei Flecken (selten) variieren, der Tornus gelegentlich mit feinem, dunklen Fleck. Die Submarginalflecken im Hflgl. Können bei beiden Geschlechtern fehlen, aber auch schwach bis sehr prominent entwickelt sein. In beiden Geschlechern betragen die Größenunterschiede die bis zu einem Drittel ihrer Körpergröße. Bei Generationen der kühleren und feuchteren Jahreszeit (Dezember bis April) ist der Geschlechtsdimorphismus deutlicher ausgeprägt, die Farbunterschiede sind deutlicher. Tiere der Sommergenerationen sind größer und in der Grundfarbe ge- ringfügig blasser (vgl. Fig. 1, 2 mit 3, 4). Aus in Mitteleuropa durchgeführten Zuchten hervorgegangene Tiere neigen zum Luxurieren. Die Vorderflügelgrundfarbe der ira- nischen Tieren ist heller als bei der Nominatunterart, mehr gelbocker bis sandfarben, die Hinterflügel sind in beiden Geschlechtern mehr weiß, der Apikalfleck bei beiden Geschlechtern punktförmig oder fehlend, die mediodorsale Thorakalzeichnung insge- samt feiner. SO Genitalien (Fig. 19-23, 28, 29). [Gen.Präp. de Fr. 2006/50, 51, 52, 53, 54, 58, 68 (letzteres OQ in Kopula)] Uncus schnabelförmig, apikal leicht gekrümmt, etwa doppelt so lang wie das flache Tegumen; Valven lang, schlank, das distale Drittel schwach nach innen gekrümmt, praeapikal an der Innenseite mit kürzerem zahnartigem Processus; Juxta flaschenförmig, an der Basıs annähernd rund, als sklerotisiertes Band bis zur tuba analis reichend; Vinculum breit vau-förmig, Saccus gerundet, schwach sklerotisiert; Phallus etwa um 4 länger als die Valven, das Coecum nur unwesentlich breiter als das kufenförmig gebogene distale Ende; Vesica massiv, partim verknorpelt, etwas langer als die halbe Lange des Phallus, mit vier unterschiedlich massiven, deut- lich getrennten Spiculaegruppen; Spiculae mittellang, stiftähnlich, stark sklerotisiert, die der prominentesten Gruppe longitudinal angeordnet, die der kleinsten aus lediglich zwei Nadeln bestehend. 9 Genitalien (Fig. 23, 35, 36). [Gen.Präp. de Fr. 2006/55, 57, 66, 68 (letzteres OQ in Kopula)] Corpus bursae länglich rund mit zwei kleineren ovalen, paarweise angeordneten, fein strukturierten Signa; appendix bursae auffällig massiv, kugelig, mit 378 DE FREINA: Creatonotos omanirana Sp. n. RTE Bi Nota lepid. 30 (2): 375-386 39 sieben kräftigen, krallenförmigen Spiculae; ductus seminalis lang, distal sackförmig; Antrum sehr lang (von der Länge des Phallus), bandförmig, verwunden und stark skle- rotisiert; ductus bursae am distalen Ende mit zwei Gruppen Spiculae, deren auffäl- ligere aus longitudinal angeordneten feinen Nadeln besteht, während sich die andere aus wenigen, aber größeren Nadeln zusammensetzt; lamela antevaginalis sehr kräftig, trichterförmig, ostium bursae halbkreisförmig, die posterior apophyses von der Länge der papillae analis. Coremata des 8. Sternits (Fig. 24-27, 30). Die Coremata bestehen aus einem Paar vergleichsweise kleiner, paarweise angeordneter, in unausgestülptem (luftleerem) Zustand knopfartig eingestülpten Schläuchen, denen distal jeweils ein um ein vielfaches längerer Haarschleier anhaftet. Ausgestülpt erreichen die Coremata etwa die 10fache Länge. Sie sind schlauchartig bei distaler Verengung und über die Gesamtlänge mit hautigen, Zahnkränzen ähnlichen Strukturen besetzt. Das 8. Sternit ist halbkreisförmig. Diagnose (Fig. 31-34, 37). Für die Differentialdiagnose wurden von C. gangis die fol- senden Tiere genital untersucht (und die Genitalabbildung von C. gangis bei Goodger & Hampson (1995) und Dubatolov (2006) herangezogen): 19 Nepal, Trisuli valley, 820 m, 3 km N Betrawati, at the Trisuli bridge, 8SS°11’E 27°59°N, 25.9.1995, leg. B. Herczig & G. M. Läszi6 (MWM): Gen.Präp. de Fr. 2006/59; 10° valley of Tamea Kosi river, 5 km S of Piguti, 950 m, 8/9.X.1995, leg. L. Németh (MWM): Gen.Präp. deFr 2006/60); 10° India, Assam, Kaziranga Wildlife res., 27°06°N 93°56° E, 200 m, 2.-3.7.1997, leg. Sinjaev (MWM): Gen.Präp. de Fr. 2006/69; 19 Assam, Nameri National Park, 40 km N Tepzur, 150 m, 27°20°N 93°15°E, 24.7.-2.8.1997, leg. Sinjaev & Murzin (MWM): Gen.Präp. de Fr. 2006/70; 19 China, Yünnan, Xishuangbanna Dai, auto- nom. Pref., Puwen, 30 km SSW Simao, 900 m, 22°30°N 100°02’E, 16.3.-10.4.2000, leg. Brechlin‘s einh. Coll. (MWM): Gen.Präp. de Fr. 2006/61; 19 Yünnan, Lincang distr., 10 km W Yunxiang, Daxing 120 km S Dali, 1200 m, 24°30°N 100°01’E, 16.3.-10.4.2000, leg. Brechlin’s einh. Coll. (MWM): Gen.Präp. de Fr. 2006/62. Creatonotos omanirana sp. n. ist in erster Linie durch die Form der Coremata und charakteristischer Strukturen des weiblichen Genitals zu unterscheiden. Habituell ist sie nicht auffallig von C. gangis verschieden. Mehrheitlich ist die Farbung von Thorax und Vorderfliigel oberseitig nicht zimtrosa wie bei dieser, sondern mehr rosa ocker, die Unterseite der Fliigel ist heller, die zitronengelbe Behaarung des ersten Abdominasegment ist markanter, die gelbe Fleckumrandung des distalen Abdominalflecks ist betonter. Im Bau des männlichen Genitals unterscheiden sich beide Arten nicht auffällig, jedoch konstant. C. omanirana besitzt proximal schlankere Valven, der praeapikale Processus Fig. 1-12. Adulte Creatonotos Tiere. 1-9. C. omanirana omanirana sp. n. 10-12. C. gangis (Linnaeus, 1764). 1. Holotypus ©. N-Oman, Westl. Hajar-Gebirge, Oase (Wadi) Al Abyadah, 04.-07.4.2006. 2. Paratypus 9. N-Oman, Jabal Nakhl-Gebirge, Oase Nakhl 30.-31.3.2006. 3. Paratypus ©, Fundort wie Holotypus, ex ovo F1, 08.-25.6.2006. 4. Paratypus 9, gleiche Daten wie Fig. 3. 5. Paratypus ©, gleiche Daten wie Holotypus (Gen.Präp. deFR 2006/50, Fig. 19, 20 24). 6. Paratypus 9, Fundort wie Fig. 2, je- doch 12.12.2006 (alle Tiere leg. bzw. cult. de Freina). 7. Paratypus ©. Iran, Hormozgan Prov., Beshagerd Mts., 900 m, 25.03.-05.4.2000, leg. Siniaev & Plutenko (Gen.Präp. deFr 2006/58). 8. Paratypus 9. Iran, Kerman, Senderk, 31.05.1977, leg. Sojak. 9. Paratypus ©, Iran, Kerman, 30 km SSW Jiroft, Mohamadabad, Maskün, 18.05.1977, leg. Sojak. 10. ©. Nepal, Trisuli valley, 820 m, 3 km N Betrawati, at the Trisuli bridge, 85°11 °E 27°59'N, 25.9.1995, leg. Herczig & Läszié. 11. ©. Nepal, valley of Tamea Kosi river, 5 km S of Piguti, 950 m, 8/9.10.1995, leg. Németh (Gen.Präp. deFr 2006/60). 12. ©. Gleiche Daten wie Fig. 10 (Gen.Prap. deFr 2006/59) (alle Tiere und Präparate MWM). 380 DE FREINA: Creatonotos omanirana sp. n. Fig. 13-18. Creatonotos omanirana sp. n. 13. Lebensraum Nordoman, Oase Nakhl. 14. Lebensraum Nordoman, NW Nizwa, Umg. Tanuf, Eingang Wadi Tanuf. 15. L1-Raupe (Fl), Orginalgröße 0,6 mm. 16. L2-Raupe (F2), Orginalgröße 1,1 mm. 17. Erwachsene Raupe, typische Farbvariante, natürliche Größe. 18. Erwachsene Raupen, rot- und gelbgefleckte Variante (unten). steht in deutlicherem, fast rechtem Winkel von der Valve ab, der apikale Finger ab dem Processusansatz ist kürzer. Die Juxta ist an der Basis weniger kugelförmig und am Übergang zum posterioren Band deutlich abgeschrägt. Der Phallus von C. oma- nirana ist distal stärker gekrümmt, die Vesica besitzt vier klar voneinander getrennte Nota lepid. 30 (2): 375-386 381 Spiculaegruppen, während diese bei C. gangis nur unwesentlich voneinander abgesetzt sind. Holloway (1988) spricht von „three fields of numerous moderate, long spines“, es dürften jedoch vier Gruppen Spiculae sein, was allerdings aufgrund der undeutlichen Abstände schwer zu diagnostizieren ist (siehe Fig.31 und 33). Auch die Abbildungen bei Goodger & Watson (1995: 37, fig. 100) und Dubatolov (2006: 146, fig. 27) spre- chen für vier Gruppen an Spiculae. Unverkennbare Artmerkmale sind die auffälligen Unterschiede in der Struktur der Coremaschläuche. Bei C. omanirana bestehen diese aus einem Paar an zwei kleineren Schläuchen, C. gangis hat zwei Paar weitaus größe- re, voluminösere, längere und mit mittellangen aber dichteren Haarschleiern bestück- te, paarweise angeordnete Schläuche (Fig. 32, 34). Die weiblichen Genitalien weisen mehrere arttypische Merkmale auf. C. omanirana besitzt schlankere papillae analıs, das ostium bursae und die lamela antevaginalıs sind enger und runder geformt, bei C. gan- gis sind letztere breiter und flacher; ductus bursae bei C. omanirana länglich, schlank trichterförmig mit zwei Gruppen Spiculae, bei C. gangis breiter und kürzer trichter- förmig bei schwach entwickelten Spiculaegruppen; appendix bursae bei C. omanirana mit sieben krallenförmigen Spiculae, bei C. gangis mit nur zwei Spiculae; Signa bei C. omanirana nebeneinander angeordnet, bei C. gangis deutlich getrennt. Verbreitung und Habitat (Fig. 13-14). C. omanirana ist in zwei geographisch ge- trennten Populationen bekannt. Im Oman ist die Art von der nördlichen Küstenebene bis in die Nord- und Südhänge des östlichen Jebel Akhdar-Gebirges verbreitet. Der Fund von Ras al Sawadı bestätigt das Vorkommen bis in Strandnähe, aus der Djebel Akhdar-Region stammt ein Nachweis aus 850 m Höhe. Auf der asiatischen Seite des Persischen Golfs ist C. omanirana aus den iranischen Provinzen Kerman und Hormozgan bekannt. Ein Paratypus stammt aus 900 m Höhe. C. omanirana ist im halb- wüstenreichen Nordoman bisher ausschließlich in oder am Rande von Oasen, in länger wasserführenden Wadis oder in bewässerten Kulturlandschaften der Batinah (nördliche Küstenebene) nachgewiesen. Sie ist thermophil, besiedelt aber in ihren Lebensräumen halbschattige, mehr oder weniger feuchte, kühlere und krautige Nischen. Am Rande von Oasen entwickelt sie sich in der Randvegetation von Bewässerungsgräben. Man kann die Art als Kulturfolger bezeichnen. Lebensweise (Fig. 15-18). Bei Zuchten erwiesen sich die Raupen als weitgehend polyphag, bevorzugten jedoch Löwenzahn (Taraxacum). In ihren angestammten Lebensräumen wurde die Raupe an Gräsern und Fabaceae-Arten (Leguminosae) gefun- den. Sie ist extrem überlebenstüchtig und robust. Die bisherigen Zuchtergebnisse zei- gen, dass die Entwicklungsdauer einer Generation zwischen 2 und 24 Monaten liegt. Unter Berücksichtigung der hochsommerlichen Trockenperioden des Nordoman kann man davon ausgehen, dass die Art pro Jahr in vier Generationen auftritt. Hinsichtlich der Geschlechterverteilung fällt der überproportional hohe Anteil an Weibchen auf, der eine hohe Reproduktion der Art garantiert. Der Geschlechterproporz Männchen zu Weibchen beträgt etwa 3 : 2. Die aus im Oman eingesammelten und in Deutschland gezüchteten Raupen stammenden Imagines erwiesen sich als sexuell nahezu inaktiv, so dass in zwei Fällen trotz einer höheren Anzahl an Tieren jeweils nur eine Kopula erfolgte. Dagegen zeigten bereits die F1- Nachkommen trotz kühlerer Temperaturen rege sexuelle Aktivität, die sich bei den F2-Tieren noch steigerte, so dass es zu zahl- 382 DE FREINA: Creatonotos omanirana Sp. n. Nota lepid. 30 (2): 375-386 383 reichen Paarungen kam. Dieses Verhalten deutet auf ein rasches Anpassungsvermögen der Art an neue Lebensbedingen hın. Der Paarungsflug setzt gegen 18°°Uhr ein, also zu einer Zeit, zu der im Oman be- reits die Dämmerung eingesetzt hat, und endet gegen 22°° Uhr. Die Kopula dauert ın der Regel bis in die Morgenstunden des folgenden Tages. Zwei Tage nach Lösen der Kopula beginnt das Weibchen mit der Eiablage. Die Anzahl der Eier schwankt zwi- schen 250 und 300, die Eidauer beträgt 10 Tage. Die elfenbeinfarbenen, bereits nach 2 Tagen gelblich verfärbten, runden Eier werden einschichtig geordnet in mehreren Spiegeln abgelegt. Auffällig ist die, gemessen an der Größe der Imagines, unverhält- nismäßig kleine Eiform, die der Art eine hohe Reproduktion aufgrund der höheren Eimenge ermöglicht. Die Eischale bildet die erste Nahrung der geschlüpften Raupe. Die Eiraupe (Fig. 15) ist auberginefarben. Alle Füße einschließlich dem Analpaar, Kopf, Mandibeln, Prothorakal- schildchen und Warzen sind schwarzbraun. Die Behaarung ist einfach, borstenartig, ungefiedert, schwarz glänzend. Die singulären Borstenhaare auf der Kopfkapsel und dem Nackenschild sowie die Borsten der dahinter liegenden Warzen sind auffällig lang und entsprechen etwa einem Viertel der Köperlänge. Noch länger sind die Borstenhaare der beiden letzten Abdominalsegmente von etwa ein Drittel der Körperlänge. Die bei- den vorderen Thorakalsegmente besitzen laterodorsal je zwei Warzenpaare mit jeweils zwei Borsten, die dahinter liegenden Segmente nur eine. Lateral sitzt jeweils eine mas- sive, sternförmig mit kurzen Borsten besetzte Warze. Bereits die L2-Raupe (Fig. 16) zeigt die für die Art typische Tracht mit heller Mediodorsallinie und größerem rostbraunen Fleck in der Rückenmitte jedes Segments, der in die Linie eingebunden ist. Die Warzen sind schwarzbraun und mit kräftigen, sternförmig angeodneten Borstenbüscheln besetzt. In ihrem weiteren Wachstum ändert sich das Raupenkleid bis ins letzte Stadium nur noch unbedeutend. Lediglich die me- diodorsalen Flecken werden flächiger, das mediodorsale Band wird breiter, der weiße Lateralfleck auf dem zweiten Thorakalsegment vergrößert sich deutlich. Die Warzen bleiben schwarzbraun, die dichten, sternförmigen Borstenbüschel sind ım letzten Larvalstadium rötlıchbraun. Die Grundfarbe der erwachsene C. omanirana-Raupe variiert zwischen braun und schwarz, die Farbe der Dorsalflecken zwischen gelbocker und rostrot (Fig. 17, 18). Wird die Raupe in humidem Milieu gezogen, entwickelt sich fast ausschließlich die melanistische dunkle Form, bei Trockenheit überwiegt die hellere Variante. Fig. 19-30. Creatonotos omanirana sp. n., © Genital- und Coremata-Strukturen des 8. Sternits. 19, 20. S Genital mit Phallus (vergrößert), N-Oman, Westl. Hajar-Gebirge, Oase (Wadi) Al Abyadah (Gen.Präp. deFr 2006/50). 21. © Genital mit Phallus, N-Oman, Jabal Nakhl-Gebirge, Oase Nakhl (Gen. Präp. deFr 2006/54). 22, 23. © Genital mit Phallus, Fundort wie Fig. 21 (Gen.Präp. deFr 2006/68). Phallus (23) in Kopula mit weiblichem Genital (Maßstab wie Fig. 20). 24-27. Corematastrukturen. 24, 25. Coremata in Ruhestellung: Gen.Präp. deFr 2006/50 (wie Fig. 19) und Gen.Präp. deFr 2006/52. 26. Coremata vergrößert, zu % ausgestülpt: Gen.Präp. deFr 2006/51. 27. Coremata in Ruhestellung, ver- größert: Gen.Präp. deFr 2006/68) (wie Fig. 21). 28-30. © Genital mit Phallus und Coremata-Struktur des 8. Sternits in Ruhestellung, ©. Iran, Hormozgan Prov., Beshagerd Mts., 900 m (Gen.Präp. deFr 2006/58). 384 DE FREINA: Creatonotos omanirana sp. n. Fig. 31-34. Creatonotos gangis (Linnaeus, 1764), © Genital und Coremata-Strukturen des 8. Sternits (in Ruhestellung). 31, 32. Nepal, Trisuli valley, 820 m, 3 km N Betrawati, at the Trisuli bridge (Gen. Präp. deFr 2006/59) (Wie Fig. 12). 33, 34. Nepal, valley of Tamea Kosi river, 5 km S of Piguti, 950 m, 8/9.X.1995 (Gen.Präp. deFr 2006/60) (Wie Fig. 11). Fig. 35-37. Creatonotos spp., 9 Genital. Fig. 35-36. C. omanirana sp. n. 35. N-Oman, Jabal Nakhl Ge- birge, Oase Nakhl (Gen.Präp. deFr 2006/56). 36. Wie Fig. 35 (Gen.Präp. deFr 2006/55). 37. C. gangis (Linnaeus, 1764), China, Prov. W-Yünnan, Lincang distr., 10 km W Yunxiang, Daxing, 120 km S Dali, 1200 m, 16.3.-10.4.2000 (Gen.Präp. deFr 2006/62). Nota lepid. 30 (2): 375-386 385 Die lichtscheue, überwiegend nachtaktive Raupe neigt im Endstadium bei Futtermangel zu kannibalischem Verhalten, wobei sowohl in Häutung befindliche Raupen als auch Puppen gefressen werden. Landwirtschaftlich genutzte Lebensräume der Art werden regelmäßig im Zuge von Bewässerungsmaßnahmen geflutet (siehe Fig. 13). Das dichte Borstenkleid verhindert jedoch ein Ertrinken der Raupen. Zusammengerollt sind sıe ın der Lage, zeitlich unbegrenzt auf der Wasseroberfläche zu treiben und dank ihrer guten Schwimmfähigkeit wıeder festen Boden zu erreichen. Die Verpuppung erfolgt bodennah in einem grauen, relatıv dichten, aber transparenten Gespinst. Die Puppe ruht dabei in der Mitte des Gespinstes. Die Raupenhaut wird nicht gänzlich abgestreift, sondern verbleibt, die hinteren beiden Segmente bedeckend, am Abdominalende der Puppe verankert. Die vorderen Abdominalsegmente der frischen rotbraunen Puppe sind zunächst trüb weiß gefleckt. Später dunkelt die Puppe schwarz ein, die Fleckung der Abdominalsegmente bleibt jedoch rötlich transparent, erhalten. Die Lateralpartie der vorderen Abdominalsegmente ist mit feinen Borsten besetzt. Der Kremaster fehlt, wird aber in beiden Geschlechtern durch eine Gruppe Hufnägeln ähnlichen Fortsätzen, etwa 14-18 an der Zahl, ersetzt. Die Puppenruhe variierte bei Zuchten zwischen 18-20 Tagen. Die Falter schlüpfen am frühen Nachmittag. Sie verhalten sich bis auf die Paarungs- aktivität sehr ruhig. Tagsüber verkriechen sie sich in der Vegetation. Frisch geschlüpfte bzw. noch nicht flugfähige Falter scheiden bei Störung zu ihrem Schutz den braunrosa Puppenharn aus, dem ein bitterer Geruch anhaftet. Bei Störung zeigen die Tiere aki- netisches Verhalten mit Zusammenklappen der Flügelpaare und Anspreizen der Beine. Dabei legen sie sich seitlich und präsentieren ihr rotes Abdomen bei gleichzeitiger Absonderung eines vermutlich toxischen Tropfen mit stark nikotinähnlichem Geruch. Die Thanatose hält mehrere Minuten an. Derivatio nominis. Die neue Art ist nach ihrer geographischen Herkunft benannt. Danksagung Der Verfasser dankt Ulf Buchsbaum (Zoologische Staatssammlung, München) und Wolfgang Speidel (Museum Witt, München) für ihre hilfreiche Unterstützung bei fototechnischen Arbeiten. Thomas Witt (München danke ich für Materialleihgaben. Hubert Abele, Augsburg, gilt mein Dank für die Betreuung von Zuchtmaterial. Literatur Bell, T. W. & J. Meinwald 1986. Pheromones of two arctiid moths (Creatonotos transiens and C. gangis): chiral components from both sexes and achiral female components. — Journal of Chemical Ecology 12: 385-409. Boppré, M. & D. Schneider 1985. Pyrrolizidine alkaloids quantitatively regulate both scent organ mor- phogenesis and pheromone biosynthesis in male Creatonotos moths (Lep.: Arctiidae). — Journal of comparative Physiology 157: 569-577. Dubatolov, V. V. 2006. New genera and species of Arctiidae from the Afrotropical fauna (Lepidoptera: Arctiidae). — Nachrichten des Entomologischen Vereins Apollo, N. F. 27 (3): 139-152. Goodger, T. & A. Watson 1995. The Afrotropical Tiger-Moths. — Apollo Books Aps., Denmark, 65 pp. Hacker, H., Aulombard, F., Bischof, A., Bittermann, J., Fibiger, M. & H. P. Schreier 2001. Lepidoptera of Yemen Arab Republic, collected during three expeditions in 1996, 1998 and 2000. — Esperiana 8: 597-632. 386 DE FREINA: Creatonotos omanirana sp. n. Hampson, G. F. 1901. Catalogue of the Helicopter Phalaenae in the British Museum 3: xix+690 pp. — London. Hampson, G. F. 1911. Description of new genera and species of Syntomidae, Arctiadae, Agaristidae and Noctuidae. — Annals and Magazine of Natural History (ser. 8) 8: 394-445. Hartmann, T. 1999. The chemical ecology of pyrrolizidine alkaloids. — Planta 207: 483-495. Holloway, J. D. 1988. The moths of Borneo 6: Family Arctiidae, subfamilies Syntominae, Euchromiinae, Arctiinae; Noctuidae misplaced in Arctiidae (Camptoloma, Aganainae). — Southdene Sdn. Bhd., Kuala Lumpur, 101 pp. Linnaeus, C. 1763. Centuria Insectorum. — Amoenitates Academiae 6: 384-415. Schneider, D. & M. Boppré 1981. Pyrrolozidin-Alkaloide als Vorstufen fiir die Duftstoff-Biosynthese und als Regulatoren der Duftorgan-Morphogenese bei Creatonotos. — Verhandlungen der Deutschen Zoo- logischen Gesellschaft 1981: 269. Schneider, D., M. Boppré, J. Zweig, S. B. Horsley, T. W. Bell, J. Meinwald, K. Hansen & E. W. Diehl 1982. Scent organ development in Creatonotos moths: regulation by pyrrolizidine alkaloids. — Science 215: 1254-1255. Strand, E. 1919. Arctiidae. — In: H. Wagner (Hrsg.), Lepidopterorum Catalogus 22. — W. Junk, Berlin. 416 pp. Wiltshire, E. P. 1980. The Larger Moths of Dhofar and their Zoogeographic Composition. — The Journal of Oman Studies, Special Report 2: 187-216. Wiltshire, E. P. 1986. Lepidoptera of Saudi Arabia. Fam. Cossidae, Sesiidae, Metarbelidae, Lasiocampidae, Sphingidae, Geometridae, Lymantriidae, Arctiidae, Nolidae, Noctuidae (Heterocera); Fa. Satyridae (Rhopalocera) (Pt. 5). — Fauna of Saudi Arabia 8: 262-323. Wunderer, H., K. Hansen, T. W. Bell, D. Schneider & J. Meinwald 1986. Sex pheromones of two Asian moths (Creatonotos transiens, C. gangis; Lepidoptera - Arctiidae): behavior, morphology, chemistry and electrophysiology. — The Journal of Experimenal Biology 46: 11-27. Nota lepid. 30 (2): 387-396 387 A revision of the clearwing moth species described by Zukowsky from China with additional notes on Sesiidae species from the Mell collection (Sesiidae) AXEL KALLIES The Walter and Eliza Hall Institute of Medical Research, 1G Royal Parade, Melbourne/Parkville, Victoria 3050, Australia; e-mail: kallies@wehi.edu.au Abstract. In the present paper the Chinese Sesiidae species described by Zukowsky from the Mell col- lection are reviewed. The following new combinations are introduced: Glossosphecia melli (Zukowsky, 1929) comb. n., Nokona semidiaphana (Zukowsky, 1929) comb. n., and Adixoa leucocyanea (Zukowsky, 1929) comb. n. Aegeria sangaica Zukowsky, 1932 is formally established as a synonym of Toleria abiaeformis Walker, 1865. Additional Sesiidae records published by Zukowsky (1929) from China are critically analyzed and mostly rejected as the specimens on which these records were based were found to be misidentified. Melittia inouei (Arita & Yata, 1987) and Macroscelesia japona (Hampson, 1919) are new records for the fauna of China. Introduction Examining the Sesiidae collection of the Zoological Museum of the Humboldt University, Berlin, Germany, the author came across a series of clearwing moth speci- mens, which had not been incorporated into the main collection. The majority of these moths carried only numbers for labels, but some of them had additional hand-written determination labels, which identified them as taxa described by Zukowsky (1929) from southern China. Comparing these specimens and their labels with the published descriptions and the handwriting of Zukowsky, confirmed that these specimens were holotypes of the taxa in question. In his paper on the clearwing moth species of the Mell collection from southern China Zukowsky (1929) named three species, Aegeria melli, Paranthrene semidiaphana and Synanthedon leucocyanea, and published records of an additional 19 species. Although some specimens listed in the paper are missing from the collection, most of them could be recovered and assigned to records published by Zukowsky (1929). Examination of the specimens revealed, that Zukowsky’s taxa are valid species but have to be trans- ferred to other genera. Most of the additional specimens, however, were misidentified and consequently the majority of Zukowsky’s faunistical records, some of which were subsequently cited by other researchers (Gaede 1933; Xu & Liu 1993), are rejected here. In addition to the above mentioned type material, another badly damaged speci- men could be identified as a type specimen of a further taxon named by Zukowsky, Aegeria sangaica Zukowsky, 1932. This taxon is here confirmed to be a synonym of Toleria abiaeformis Walker, 1865. Material and Methods All material is located in the Zoological Museum of the Humboldt University, Berlin, Germany (ZMHB). Locality data are quoted as on the labels. When original labels Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 388 KALLIES: On clearwing moths from China were absent locality data are given in square brackets, [...], according to Zukowsky (1929); new transcriptions of locality data, if available, are provided in round brackets, (...). The following abbreviations were used: ATA — Anterior Transparent Area; ETA — Exterior Transparent Area; PTA — Posterior Transparent Area. Taxa described by Zukowsky from China CISSUVORINI Glossosphecia melli (Zukowsky) comb. n. (Figs 1, 5) Aegeria melli Zukowsky, 1929: 34. Type locality: South China, Sin hang. Gaede 1933: 787. Heppner & Duckworth 1981: 32 (Synanthedon). Pühringer & Kallies 2004: 29 (Synan- thedon). Material. Holotype © [Sin hang] ,,11. Aeg. melli“ / „11.“, gen. prep. AK156. This species is a typical representative of the genus Glossosphecia Hampson 1919. This genus was redefined by Arita & Gorbunov (1998) and besides the species dealt with here it comprises four species, Glossosphecia contaminata (Butler, 1878) (the type species), Glossosphecia romanovi (Leech, 1889), Glossosphecia sherpa (Bartsch, 2003) and Glossosphecia huoshanensis (Xu, 1993). G. melli seems to be closely related to G. contaminata. It differs from the latter by the dark fuscous patagia (bright yellow in contaminata) and the abdomen (tergite 4 with a broad and bright yellow band in con- taminata). G. huoshanensis was described from the southern Chinese Huoshan County, Anhui Prov. (Xu 1993) and is possibly a junior synonym of G. melli. Description. Alar expanse 38 mm. Forewing length 16.5 mm. Body length 19 mm. Head. Frons light grey, laterally white; vertex fuscous with fine white scales subdor- sally; antenna brown, ventrally orange, unipectinate, ciliate; labial palps with basal segment fuscous, second segment deep yellow with a brown lateral line, apical seg- ment orange-yellow; scapus brown. Legs. Fore coxa yellow with brown margins; fore femur and tibia dark brown, dorsally with long scales, ventrally deep orange; fore tar- sus orange; mid and hind coxae yellow; mid and hind femurs yellow in distal portion; mid and hind tibiae ventrally black with yellow patches close to spurs, dorsally dirty orange; basal hind tarsomers with long and rough scaling, dorsally brown-orange, re- mainder of hind tarsus brown, ventrally yellow-orange. Forewing. Transparent with a rusty shine; discal spot narrow, with a long and narrow projection into ATA; costal margin and discal spot with individual orange scales; ETA with 6 cells, partly covered with minute, semitransparent scales; apical area not developed. Hindwing. Discal spot very small; transparent cells in distal portion of the wing with minute, semitransparent scales. Abdomen. Dark fuscous, tergites 1-3 with narrow yellow posterior margins, tergite 4 with yellow scales at anterior margin, sternites 2-7 with pale yellow posterior margins, anal tuft fuscous, apically pale yellow. Distribution. The position of the type locality could not be verified. It is likely to be a place in the Guangdong Province were most of Mell’s material was collected. Nota lepid. 30 (2): 387-396 389 Figs 1-4. Sesiidae types from the Mell collection (all in MNHB). 1. Glossosphecia melli comb. n., holo- type, male. 2. Toleria abiaeformis, female (syntype of Aegeria sangaica). 3. Nokona semidiaphana comb. n., holotype, male. 4. Adixoa leucocyanea comb. n., holotype, female. Scale bars 10 mm. Fig. 5. Glossosphecia melli, holotype, male genitalia. a. uncus-tegumen, lateral. b. right valva, ventral. c. phallus, lateral. d. saccus, ventral. 390 KALLIES: On clearwing moths from China Fig. 6. Nokona semidiaphana, holotype, male genitalia. a. uncus-tegumen, lateral. b. left valva, ventral. c. phallus, lateral. d. saccus, ventral. Habitat and Bionomics. Unknown; the specimen was collected ‘on a leaf in a forest’ (Zukowsky 1929). Remarks. Generic combinations of some of the above mentioned species were es- tablished during the course of this work but have been formally published earlier (Pühringer & Kallies 2004). Toleria abiaeformis Walker, 1865 (Fig. 2) = Aegeria sangaica Zukowsky, 1932: 316. Type locality: Shanghai. Heppner & Duckworth 1981: 28 (Sesia). Spatenka et al. 1999: 115. Pühringer & Kallies 2004: 12. Material. 10 (syntype, Fig. 2). “Shanghai (China), Juni 1918., (H. Höne)” / “det. Zukowsky” / “Type, H. Hone, 11.1.19, Aegeria sangaica Hone ?” Aegeria sangaica Zukowsky, 1932 and Sphecia sinensis Walker, 1865 were both con- sidered synonyms of Toleria abiaeformis Walker, 1865 although the type material of Zukowsky’s taxon had not been traced (Spatenka et al. 1999). Arita & Gorbunov (1998) resurrected sinensis from synonymy and assigned it to the re-established generic name Chimaerosphecia Strand, 1916; however, without establishing the identity of Aegeria sangaica. Examination of a syntype of sangaica confirms that this taxon is a junior subjective synonym of Toleria abiaeformis. The genus Toleria in the present view contains only two valid species, T. abiaeformis and the very similar 7. ilana Arita & Gorbunov, 2001. The structures of the female genitalia are only insufficiently known. The genus Chimaerosphecia Strand, 1916 on the other hand is known from females only. It was redefined and separated from Toleria Nota lepid. 30 (2): 387-396 391 Walker, 1865 by Arita & Gorbunov (1998) on the grounds of small differences in the venation and labial palpus. Apart from the type species, Ch. aegerides Strand, 1916, from Taiwan, it contains another two species, Ch. sinensis Walker, 1865 from Hong Kong and Ch. colochelyna Bryk, 1947 from Kiangsu, Southern China. The validity of the genus Chimaerosphecia is considered doubtful by the present author and more material, in particular clearly conspecific males and females, is needed to establish the identity of the genus. Remarks. Zukowsky (1932) mentioned three type specimens from the Hone collec- tion (today in Zoologisches Forschungsinstitut und Museum Alexander König, Bonn, Germany), all from Shanghai, one of which was found in the ZMHB. This specimen consequently is a syntype. The designation of the lectotype is avoided here as the present specimen is badly damaged by frass with major parts of the genitalia destroyed. If additional syntypes can be found in other parts of the Höne collection one should be selected as lectotype. The biology of the species of Toleria is not known; however, the larvae of a species of the closely related genus Chimaerosphecia, Ch. sinensis, were collected in the trunks of Ormosia pachycarpa (Fabaceae) at Ma On Chan, Hong Kong, in May and adults emerged in early June (Kendrick 2001). PARANTHRENINI Nokona semidiaphana (Zukowsky) comb. n. (Figs 3, 6, 7) Paranthrene semidiaphana Zukowsky, 1929: 36. Type locality: Mahn tsi shan. Gaede 1933: 794. Heppner & Duckworth 1981: 24. Liu & Shen 1992: 728. Piihringer & Kallies 2004: 21. Material. Holotype So “24, P. semidiaphana” / “24” [Mahn tsi shan 25.vii.1915] (Guangdon Prov, E Nanling, between rivers Si Kiang and Yangste), gen. prep. AK157. Additional material: 10° „China Canton, 9.1910, Mell S.V.“; 19 [Mahn tsi shan, 900 m, 31.vii.1915]; 19 [Lin-ping, 700 m, NO Kwangtung, 14.v11.1920]. This species is a typical representative ofthe genus Nokona Matsumura, 1931. It differs from most congeners by the conspicuously broad and dark margins of the hindwings. Broadened hindwing margins are known from some other species of Nokona such as N. powondrae (Dalla Torre, 1925) and N. inexpectata Arita & Gorbunov, 2001, both from Taiwan. These species, however, have distinctly different markings on the abdo- men (comp. Arıta & Gorbunov 2001) and significantly different genitalia (valva long and tapered, saccus short in the species compared; valva shorter, saccus long and nar- row in N. semidiaphana). Apart from the holotype of this species, the Mell collection contains an additional three specimens, which are here considered as belonging to N. semidiaphana. These speci- mens were misidentified by Zukowsky (1929) as Paranthrene davidi Le Cerf, 1917 and Paranthrene cupreivitta (Hampson, 1893), respectively. Description. Alar expanse 29 mm. Forewing length 13 mm. Body length 17 mm. Head. Frons black, laterally white to grey; labial palpus black, yellow ventrally and medially; vertex black; pericephalic scales white. Thorax. Black; with a yellow patch below forewings; patagia with individual yellow scales laterally. Legs. Fore leg black; 392 KALLIES: On clearwing moths from China mid coxa black (remaining parts of mid leg broken off); hind coxa yellow; hind femur with posterior margin white; hind tibia black with white lateral patch, yellow at poste- rior end, medially mainly white; hind tarsus black, basal tarsomer white at distal end. Forewing. Almost entirely opaque brown. Hindwing. Transparent, with a broad brown margin. Abdomen. Black, tergite 1 with individual yellow scales at posterior margin; tergites 2 and 4 with narrow yellow posterior margins; in tergite 4 broadened towards lateral; sternite 4 broad yellow in posterior half; anal tuft triangular, black, ventro-me- dial with some yellow scales; outer surface of valvae with white scales. Distribution. Known from the Guangdong Province, China. Habitat and Bionomics. Most records are from July, one specimen was collected in September. Adixoa leucocyanea (Zukowsky) comb. n. (Figs 4, 6) Synanthedon leucocyanea Zukowsky, 1929: 36. Type locality: Lung tao shan. Gaede 1933: 781. Heppner & Duckworth 1981: 31. Pühringer & Kallies 2004: 29. Material. 19 (holotype) “29. S. leucocyanea” / “29.” [Lung tao shan, 19.vii.1917] (Guangdong Prov., Mt. Longtou, ca 230 km N Guangzhou), gen. prep. AK191. This species belongs to the tribe Paranthrenini and is here assigned to the genus Adixoa Hampson, 1893. This association, however, remains tentative until more material, in particular males, is known. The genus Adixoa is insufficiently known. So far only the male of Adixoa trizonata (Hampson, 1900) has been examined in detail (Gorbunov & Arita 1995). The female genitalia of A. leucocyanea are unique amongst the Sesiidae in their formation of the ostium bursae and the most distal parts of the ductus (strongly sclerotized, curved and protruding from the sclerite). This structure distinguishes A. leucocyanea readily from all other known Paranthrenini and may prove to be an au- tapomorphy of the genus Adixoa. Habitually A. leucocyanea differs from other known species of Adixoa by the coloration of the abdomen (black, with white markings; with yellow markings in all other species). Description. Alar expanse 24 mm. Forewing length 10 mm. Body length 10.5 mm. Head. Frons light grey, laterally white; labial palps grey, ventrally with some white scales, relatively long and upcurved, almost reaching the scapus; vertex black, antenna black, scapus white ventrally; pericephalic scales black, laterally white. Thorax. Black; patagia creme laterally. Legs. Fore coxa creme, basally with a white spot, distally black; fore leg black, dorsally partly white; basal tarsomer with a white distal margin; mid and hind tibiae with a white spot ventrally; spurs black, medial side white; basal and subbasal tarsomers distally white. Forewing. With well-developed transparent areas; discal spot broad, near the cubitus protruding into the ATA, with yellow scales in distal portion; ETA consisting of 5 large cells and an additional small cell between R4 and RS; PTA reaching the discal spot, with some yellow scales in distal portion; apical area almost as wide as ETA. Hindwing. Discal spot well-developed, relatively broad and straight; outer margin well-developed, about as broad as the fringe. Black; tergites 2, 4, 6 each with a narrow white distal band; sternite | white in distal half; sternite 3 with a Nota lepid. 30 (2): 387-396 393 narrow, sternites 4 and 5 with broad white distal bands; sternite 6 with only some white scales; anal tuft black, with white scales dorso-laterally. Distribution. Known only from the Guangdong Province, China. Habitat and Bionomics. Unknown. The type specimen was found ‘on a leaf in a forest, in July’ (Zukowsky 1929). Additional Sesiidae records from the Mell collection The following species were identified from the material of the Mell collection. Trichocerota melli Kallies & Arita, 2001 Material. 19, 19 „Canton (China), Westfluss, Ting-Wu-San, 28.vi.1910, Mell S.G.” (Guangdong Prov.). Two specimens in the Mell collection were misidentified as Trichocertota brachythyra Hampson, 1919 (Zukowsky 1929); later these specimens became part of the type series of T. melli (comp. Kallies & Arıta 2001). Trichocerota tricolor Kallies & Arita, 2001 Material. 10, 19 [Tsha yuen shan, 19.v.1915]. Two specimens in the Mell collection were misidentified as Trichocerota dizona Hampson, 1919 (Zukowsky 1929); later these specimens became paratypes of T. tri- color (comp. Kallies & Arıta 2001). Caudicornia tonkinensis Kallies & Arita, 2001 Three specimens of this species in the Mell collection, marked only with “22”, could not be assıgned to any published records. Presumably these specimens were collected by Mell in southern China. Melittia inouei (Arita & Yata, 1987) Material. 10 [South China, Gao fung, 9.vi.1917, leg. Mell], gen. prep. AK188. This species has been known only from Japan and Korea (Spatenka et al. 1999; Arita et al. 2004) and thus, the specimen listed here represents a new record for China. The specimen in the Mell collection was misidentified as “M. bombyliformis Cramer” (Zukowsky 1929). The exact position of the collecting locality could not be estab- lished. Macroscelesia japona (Hampson, 1919) Material. 19 [South China, Gao fung, 9.vi.1917, leg. Mell], gen. prep. AK184. This species was known only from Japan and Korea (Spatenka et al. 1999; Arita et al. 2004). The specimen in the Mell collection was identified by Zukowsky (1929) correct- ly as M. eurytion sensu Bartel, 1912 (= japona Hampson, 1919). A second specimen, supposedly a male, mentioned by Zukowsky (1929) was not found in the collection. The exact position of the collection locality could not be established. 394 KALLIES: On clearwing moths from China Cyanosesia tonkinensis Gorbunov & Arita, 1995 Material. 19 [China, Shui yün shan, 2.v1.1917, coll. Mell] (probably Guangdong Province, Shaoguang City, Ongyuen County, 18 km ENE of Fongwan, 900 m, mountain forest), gen. prep. AK187. This specimen was misidentified by Zu- kowsky (1929) as Paranthrene trizonata Hampson, 1900 but its identity was es- tablished as Cyanosesia tonkinensis later (Kallies 2003). Nokona iridina (Bryk, 1947) Material. 19 [Kanton]; 19 „Canton (China), Westfluss, Ting-Wu-San, Mell S.G.“; IM [Lung tao shan, 3.v.1918]. The specimens in the Mell collec- tion were misidentified by Zukowsky (1929) as Paranthrene bicincta and Paranthrene pernix, respectively. Here they are regarded as N. iridina although the status of this taxon in respect to Nokona bicincta (Walker, 1865) needs validation. Gorbunov & Arita (2001) regarded iridina a distinct species while Spatenka et al. (1999) considered it a synonym of bicincta. Fig. 7. Female genitalia. a. Adixoa leucocyanea, holo- type. b. Nokona semidiaphana (?). Nokona regale (Butler, 1878) Material. 19 [Lung tao shan]. The single specimen present in the Mell collection was correctly identified as Paranthrene regale (Zukowsky 1929). Appendix Zukowsky (1929) recorded a number of Sesiidae taxa as new for the Chinese fau- na. Many of these records, however, have to be considered doubtful with respect to the known distribution of the species concerned. The true identity of some specimens present in the collection, as listed above, could be established here, but others were found to represent unnamed species or their identity remains elusive due to the poor Nota lepid. 30 (2): 387-396 395 condition of the material. On the grounds of lacking material, records of the following species as given by Zukowsky (1929) are here rejected: Paranthrenopsis polishana (Strand, 1916); Trichocerota brachythyra Hampson, 1919; Trichocerota cupreipennis (Walker, 1865); Oligophlebia cristata Le Cerf, 1916; Toleria abiaeformis Walker, 1865; M. chalciformis (Fabricius, 1793) (as “M. bombyliformis Cramer”); Nokona bicinc- ta Walker, 1865 (as Paranthrene); Nokona pernix (Leech, 1889) (as Paranthrene); Nokona davidi (Le Cerf, 1917) (as Paranthrene), Paranthrene cupreivitta (Hampson, 1893); Adixoa trizonata (Hampson, 1900) (as Paranthrene), Pseudosesia limpida (Le Cerf, 1916) (as Paranthrene); Ichneumenoptera auripes Hamson, 1893; Synanthedon subauratus Le Cerf, 1916, Synanthedon unocingulata Bartel, 1912; Synanthedon con- cavifascia Le Cerf, 1916. Acknowledgments I wish to express my cordial thanks to Wolfram Mey (ZMHB) for his permission to examine material under his care and to Daniel Bartsch, Stuttgart, for providing some of the photographs reproduced here. References Arita, Y. & O. G. Gorbunov 1998. A revision of Embrik Strand’s clearwing moth types (Lepidoptera: Sesiidae). — Chinese Journal of Entomology 18: 141-165. Arita, Y. & ©. G. Gorbunov 2001. Sesiidae of Taiwan. I. The Tribes Tinthiini, Similipepsini, Paraglosse- ciini, Pennisetiini, Paranthrenini and Cissuvorini. — Japanese Journal of Systematic Entomology 7: 131-188. Arita, Y., Y. S. Bae, C. M. Lee & M. Ikeda 2004. Sesiidae (Lepidoptera) of Korea. — Transactions of the Lepidopterist’s Society of Japan 55: 1-12. Gaede, M. 1933. Aegeriidae. Pp. 775-802, pls 94-95 — In: A. Seitz (ed.), Die Gross-Schmetterlinge der Erde, II. Abteilung: Exotische Fauna 10 (Die indo-australischen Spinner und Schwärmer). — A. Ker- nen, Stuttgart. Gorbunov, O. G. & Y. Arita 1995. A revision of Frederic Moore’s clearwing moth types (Lepidoptera, Se- siidae), at Humboldt University, Berlin. — Tinea 14: 204—224. Gorbunov, O. G. & Y. Arita 2001. A revision of Felix Bryk’s clearwing moth types (Lepidoptera, Sesiidae), at the Naturhistoriska Riksmuseet in Stockholm, Sweden. — Melittia 1: 9-51. Heppner, J. B. & W. D. Duckworth 1981. Classification of the Superfamily Sesioidea (Lepidoptera: Ditry- sia). — Smithsonian Contributions to Zoology 314: 1-144. Liu, Y. & G. Shen 1992. Lepidoptera: Sesiidae, pp. 728-731. - In: J. Peng & Y. Liu (eds), Iconography of forest insects in Hunan China. — Academica Sinica & Hunan Forestry Institute, Hunan, China. pp. 666-755, Peking. Kallies, A. 2003. Two new species of Sesiini from the Oriental Region (Lepidoptera: Sesiidae, Sesii- nae). — Nachrichten des entomologischen Vereins Apollo, N.F. 23: 161-166. Kallies, A. & Y. Arita 2001. The Tinthiinae of North Vietnam (Lepidoptera, Sesiidae). — Transactions of the Lepidopterist’s Society of Japan 52: 187-235. Kendrick, R. C. 2001. New moth species for Hong Kong, part 5: update and further 1999 records. — Por- cupine! (Newsletter of the Department of Ecology and Biodiversity, The University of Hong Kong) 22: 3-4. Liu, Y. & G. Shen 1992. Lepidoptera: Sesiidae, pp. 728-731. — In: J. Peng & Y. Liu (eds), Iconography of forest insects in Hunan China. — Academica Sinica & Hunan Forestry Institute, Hunan, China. pp. 666-755, Peking. [in Chinese] Pühringer, F. & A. Kallies 2004. Provisional checklist of the Sesiidae of the world (Lepidoptera, Ditrysia). — Mitteilungen der Entomologischen Arbeitsgemeinschaft Salzkammergut 4: 1-85. 396 KALLIES: On clearwing moths from China Spatenka, K., ©. G. Gorbunov, Z. Lastuvka, I. Tosevski & Y. Arita 1999. Sesiidae — Clearwing Moths. — In: C. M. Naumann (ed.), Handbook of Palaearctic Macrolepidoptera. Vol. 1, 569 pp. - Gem Publish- ing Company, Wallingford, England. Xu, Z. & Y. Liu 1993. Four new Species and two new Records of the Subfamily Tinthiinae from China (Lepidoptera: Sesiidae). — Acta Agriculturae Boreali-occidentalis Sinica 2: 1-5. Xu,Z. 1993. Notes of a new species Cissuvora huoshanensis Xu (Lep., Sesiidae). — Acta Agriculturae Boreali-occidentalis Sinica 2: 7-9. Zukowsky, B. 1929. Beiträge zur Fauna sinica (VIII). Die südchinesischen Aegeriiden der Sammlung Mell. — Internationale Entomologische Zeitschrift, Guben 23: 33-37. Zukowsky, B. 1932. Neue paläarktische Aegeriidae. — Internationale Entomologische Zeitschrift, Guben 26: 316-317. Nota lepid. 30 (2): 397-406 397 Rediscovery of Sciopetris melitensis Rebel, 1919 and description of its morphology and life history (Psychidae) PETER HÄTTENSCHWILER !, PAUL SAMMUT? & MICHAEL ZERAFA * ' Seeblickstrasse 4, 8610 Uster, Switzerland; e-mail: peter.haettenschwiler@dative.ch 2 137, ‘Fawkner/2’, Dingli Road, Rabat, RBT 9023, Malta; e-mail: farfett@onvol.net 3 16 ‘Agape’, Triq L-Imaghazel, Naxxar, NXR 02, Malta; e-mail: family_zerafa@hotmail.com Abstract. TThe original description of Sciopetris melitensis Rebel, 1919 is based on a strongly damaged male collected in 1915 by Adolf Andres at the former Verdala Barracks, Bormla. The species was re-de- scribed by Amsel (1955) based on four badly damaged males collected at Gharghur in 1953. Since then only these specimens are known in the literature. After a lapse of over 50 years the species was rediscov- ered and breeding from eggs succeeded. Based on this material, males, females, eggs, larvae, pupae and bag as well as the life history are described and compared with the other species of the genus. Zusammenfassung. Die Originalbeschreibung von Sciopetris melitensis Rebel, 1919 basiert auf einem stark beschädigten Männchen, das 1915 von Adolf Andres in den damaligen Verdala Barracks Bormla gefangen wurde. Die Art wurde von Amsel (1955) aufgrund von vier, in Gharghur 1953 gesammelten, sehr schlecht erhaltenen Männchen nachbeschrieben. Seit dem sind nur diese Männchen in der Literatur bekannt. Nach einer Unterbrechung von über 50 Jahren wurde die Art wieder gefunden und es gelang die Zucht aus dem Ei. Basierend auf diesem Material werden Männchen, Weibchen, Ei, Larve, Puppe und Sack sowie die Entwicklung beschrieben und mit den anderen Arten der Gattung verglichen. Résumé. La description de Sciopetris melitensis Rebel, 1919 est basée sur un mâle fortement endommagé, capturé par Adolf Andres en 1915 à Verdala Barracks, Bormla. Amsel l’a re-décrit de quatre mâles en très mauvais état, capturés à Gharghur en 1953. Depuis, seulement ces cinq exemplaires ont été mentionnés dans la littérature. Après une interruption de plus de 50 ans l’espèce a été retrouvée et élevée à partir de l’œuf. Des informations additionnelles sont données pour le mâle, la femelle, la larve, le fourreau et le développement de l’espèce, qui est comparée avec les autres espèces du genre. Introduction Today the genus Sciopetris Meyrick, 1891 incorporates six species: the type species M. technica Meyrick, 1891 described from Algeria, M. amseli Sieder, 1959 from Afghanistan, M. melitensis Rebel, 1919 from Malta, M. pretiosa (Stainton, 1872) from Morocco, M. hartigi Sieder, 1976 from Sardinia and M. karsholti Hättenschwiler, 1996 from Tunisia. All are small species and mostly insufficiently known. For most of the species only the male is known while female, larva, bag, and life history remain unde- scribed. The bag and life history are known only for S. hartigi. Several species are only known from a very limited area as are the islands of Sardinia and Malta. The descrip- tion of the genus Sciopetris is based on three males that were collected by Meyrick “from sheltered rock-faces in Algeria”. The genus and its type species M. technica are described very briefly: "Head rough-haired; ocelli present; tongue absent. Antennae two-thirds, in males fili- form, clothed with rather long pubescence (2), basal joint moderate, stout, with well de- veloped pectin. Labial palpi rather short, porrected or drooping, with loose projecting hair-scales. Maxillary palpi obsolete. Posterior tibiae with appressed scales. Fore-wing with vein 1 furcate, 2 from two-thirds of cell, 7 absent, 10 absent, 11 from before mid- dle of cell. Hind-wings under one, elongate-ovate, cilia 1; vein 3 remote from 4, 4 and 5 from a point or stalked, 6 and 7 parallel. (female probably apterous).” Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 398 HÄTTENSCHWILER et al.: On Sciopetris melitensis from Malta From this we can understand that there are eight veins from the discoidal cell on the forewing and six on the hindwing, that m2+3 are stalked, that ocelli are present, and that the antennae reach 2/3 of the wing- length and are filiform. The wing venation in several of the species is not constant and in some the ocelli are absent. The species also show variations in other identification criteria. In the descriptions of the vari- ous species, due to the limited number of specimens available, only a few of the identification criteria could be compared. In this study we shall concentrate on S. melitensis, the species from Malta (Fig. la), knowing that also here there are differences between our findings on the one sıde and facts, assumptions, and descriptions in earlier studies on the other hand. However, we are convinced that the species we have at hand is identical with the single, damaged male Rebel described in 1919. “A single male specimen, only partly in a good condi- tion, of this very small new species was collected by Fig. 1. S. melitensis male. a. Malta, Adolf Andres, a war prisoner on the Island of Malta. < REE He a OP This probably happened in February 1915 and was mentioned (Andres 1916), already under the name suggested by me (1.1.)”. (This historical note was writ- ten in the German language by Rebel). Sciopetris melitensis species has been listed with little or no additional information by other authors in various Lepidoptera faunistic lists (Sieder 1959; De Lucca 1965: 514; Sammut 1984a: 65; 1984b: 10; 1985: 304; Sammut & Valletta 1989: 98; Sauter & Hättenschwiler 1999: 77). Recently Sammut (1989: 129; 2000: 42) translated to Maltese all the known information regarding this species. However, all these references regard the only two captures of S. melitensis known at that time. Since then many pro- fessional and amateur entomologists have been searching for this little species without success. It took over 50 years until one of us, Michael Zerafa, took two male specimens and found a 6-8 mm long bag that contained eggs. He took it home and after a few days, very small larvae were crawling out of the little bag and started to build a mini- ature bag of their own. He reared them successfully through the summer and in January and February of the next year adults, males and females, were hatching out of the bags and were mating and laying their eggs. From these eggs more specimens were reared by us and the life history could be observed and the species was studied. In the fol- lowing lines we give a better and more complete description of the male, female, egg, larva, pupa, bag, and life history. | The genus Sciopetris belongs to the Taleporiini Tutt, 1900 (Psychidae: Taleporiinae). Taleporiini can be recognized by the short epiphysis and 6 veins from discoidal cell in hindwing. Characteristics for Sciopetris are: (1) forewing with 8 veins off the discoidal Nota lepid. 30 (2): 397-406 399 Figs 2-3. 2. Wing venation of male S. melitensis. 3. Male genitalia of S. melitensis, a. lateral view, b. ventral view. cell and without accessory cell; (2) males with ocelli (but also present in Bankesia, Pseudobankesia, Taleporia); (3) antenna with an antero-ventral half-circle of bristles (Fig. 1b) (Sauter & Hattenschwiler 1999). Redescription of Sciopetris melitensis Rebel, 1919 from Malta Material. The following description is based on 21 males, 9 females, 6 larvae, some of which were mounted on slides, and approximately 50 bags, all from the same location: Malta, Bormla, leg. M. Zerafa, reared ex. ova, or males collected in the field. One bag was collected at Manikata, also by Zerafa, and from it the parasite emerged. The specimens are deposited in the collections of the authors. Description. Male. Wings narrow and long. Average wingspan 9.6 mm (n= 21; smallest 8.2 mm, largest 11.2 mm; wingspan of S. hartigi and S. amseli 11 mm, S. technica: 10-11 mm). Bred specimens are on an average slightly smaller than wild collected males. Forewing with 8 veins off the discoidal cell, without accessory cell (Fig. 2). Colour whitish with brown scales forming variable dots and small brown areas (Fig. la). Hindwing with 6 veins off the cell, with an intercalated cell, veins m2+m3 stalked, no connection between radial ramus and sub-costal (the original description gives incomplete data on the wings due to the strong damage on the holotype; Sieder 1959 also did not remove the wing scales of the only available specimen when de- scribing S. amseli and therefore the actual venation of this species remains unknown; Hättenschwiler 1996 studied and published the wing venation of S. karsholti, and it differs in some aspects from that of S. melitensis). Scales of forewings wide, class 4-5 (Sauter 1956), on hindwings class I-3, whitish in colour. Head without ocelli. Labial palps with 3 segments, bent forward. Antenna long, % to %4 of wing length, the 27-36 segments with scattered ventral bristles which stand at base of segments nearer together and forming kind of a ventral half-ring (Fig. Ib). Scapus and 3-4 basal seg- ments coated with whitish scales. Eyes round and large; distance between eyes nearly twice that of eye diameter. Head and body covered with long whitish hairs. Forelegs with epiphysis; midlegs with one pair and hindlegs with two pairs of tibial spurs. Male genitalia (Fig. 3): saccus triangular. Valva about as long as tegumen. Sacculus with pointed thorn. Phallus long, straight. 400 HÄTTENSCHWILER et al.: On Sciopetris melitensis from Malta Fig. 4. Female of S. melitensis, a. genitalia in ventral view, b. side view, c. enlarged head part, d. 1 foreleg, 2 midleg, 3 hindleg enlarged. Female (Fig. 4). Wingless, 3-4 mm long (excluding ovipositor), and about 1.5 mm in diameter, cylindrical. Head and thoracic segments dorsally and laterally sclerotized, dark brown. Head small, eyes miniature, antenna reduced. Legs long, tarsi mostly with 4 segments, first about as long as 2-4 together (Fig. 4d); some specimens only have three tarsal segments, 2 short and | long. Abdominal segments 1-7 whitish to pale yellow, with light brown dorsal plates (Fig. 4b). Segment 8 with full circle of golden abdominal hair tuft; genital opening ventral, between segments 8 and 9 (Fig. 4a); ovi- positor telescopic. Egg. Yellowish when fresh, slightly oval, 0.35-0.45 mm long, without any markings or sculpture. The colour changes with the life history of the larva to dark grey. In two of the bags opened we counted 65 and 71 eggs respectively. The body of one dead female contained 49 fully developed eggs. Larva (Fig. 5a). Whitish to pale yellow; head dark; thoracic segments 1-3 with dark brown markings and stripes; length 3.5-4.5 mm, diameter approx. | mm. (Fig. 5b). Ventral side of labrum with four pairs of setae. Tineidae have 3 pairs only (Davis 1978). Bag (Figs 6a-b). Triangular in cross section. Female bags range from 6.6-8.2 mm in length and male bags from 5.3-6.8 mm. About 2 mm wide. Opening of female bags decorated with debris of dried leaves and lichens. Silk structure with coating of sand and miniature stones; often partly, or even completely covered with organic matter like bits of plant and wood tissue and also minute parts of dead insects. Pupa (Fig. 7). Male pupa 3-4 mm long, 0.8-1.2 mm in diameter. Sclerotized exuvia very delicate and fragile; light brown skeleton thinner than in other genera of Taleporiini. Abdominal segments (Fig. 7a) equipped with row of thorns facing backwards on each segment anteriorly. Sheaths of appendices arranged as usual; long antennal sheaths reaching rear end of pupa. Thorax-head plate (Fig. 7c) with two pairs of bristles. Nota lepid. 30 (2): 397-406 401 {Rated eS ST 1 LEE eier Tr A ER RT LUS Visa ee N, pat LE Figs 5-7. Sciopetris melitensis. 5. Larva, a. lateral view, b. ventral side of labrum with 4 pairs of setae. 6. Bags, a. bag with only silk and sand, b. bag, upper side completely covered with plant material, only the ventral side showing the silken basic construction. 7. Male pupa after hatching, a. lateral view, b. ventral view, c. details of thorax-head plate, and d. abdomen with dorsal rows of thorns facing backwards, apical two thorns facing forward. Female exuvia even thinner than in male; reduced to soft skın which ıs pushed back- wards rather than being crawled out of. After hatching, exuvia remaining as flat pack- age in bottom of bag. Life history. Larva. The young larvae hatch from after 2—4 weeks in their mother’s bag and hurry to get out. Their first action is to build a miniature bag with silk and of- ten with minute plant material taken from the mother’s bag or elsewhere. During their whole life the larvae maintain, repair, and enlarge their bag to accommodate their in- creasing size, but they never leave it. The bag is kept long enough and wide enough so that the larva can turn around within it. To enlarge the bag, the larva bites it open along one of the three selvages and inserts a small section of silk and also little bits of foreign matter. This procedure is done along one selvage after the other until the inside room is again wide enough for the grown larva to turn around inside the bag. The larvae mainly feed on the moss Tortula muralis Hedwig, 1801, but we observed also that dead insects, found on the moss, are eaten. 402 HÄTTENSCHWILER et al.: On Sciopetris melitensis from Malta Fig. 8. Biotope; a shady street with moss covered rocks and walls (photo by P. Sammut). During the warm and dry summer a diapause is observed for some 2-3 months. In au- tumn this diapause ends and the larvae eat again for some weeks and prepare for pupa- tion. For most of the time the larvae hide in the moss, where many also pupate. Pupa. The dorsal abdominal thorns would help the pupa to work itself out of the bag by bending the abdomen up and down and finding hold ın the wall of the bag. This behavior is normal with both sexes in all genera of the Naryciinae and Taleporiinae. However, according to our observations S. melitensis does not partly work the pupa out of the bag prior to hatching and the rows of thorns on the pupae can be interpreted as a relict (Fig. 7a, d), The two thorns on the last segment facing forward probably help the pupa in preventing it from falling out of the bag when preparing for hatching. Adult. Mating occurs early in the morning. The female hatches at dawn and waits outside of her bag with the ovipositor widely extended, calling for a partner with her pheromone. The male hatches at dusk and dries its wings. It remains on or near the bag until the next morning. When the pheromone of a female reaches the male it follows the scent against the airflow to find the waiting female. Mating lasts only a few min- utes. Immediately after, the female starts laying about 50-70 eggs into the bag beside the pupal exuvia and rubs off her abdominal hairs to place them between the eggs for cushioning and insulation. When finished, the female “walks” away and dies. The sex ratio is balanced. The species has one generation per year which begins approximately with egg laying in January or February, and lasts a full year with a diapause of 2 to 3 months in the summer. Parasites. The only parasitoid known emerged from a bag collected in Manikata. It was a male wasp of the genus Gelis Thunberg, 1827 (Ichneumonidae, Cryptinae). The biomass of the larva is practically identical with that of the wasp. Nota lepid. 30 (2): 397-406 403 € x 7 ” Mt ar LÉ LL PER A Pr BIN it > Fig. 8. Biotope; a shady street with moss covered rocks and walls (photo by P. Sammut). Habitat (Figs 8-9). This species is known only from Malta, and it is the only Sciopteris known on the Maltese archipelago, where it was found on mossy rocks and stone walls colonized mostly by Tortula muralis Hedwig, 1801 (Pottiaceae). Besides the type lo- cality, it is also known from Gharghur (Amsel 1953) and Manikata. The situation of the collecting localities is shown on the map (Fig. 10). Etymology: The name melitensis, meaning “Maltese” or “of Malta” is derived from the old name of the Island of Malta — Melita. Discussion With the exception of S. melitensis, from none of the other five species in the genus are all stages known and many identification criteria remain unknown. In the following table the published data are compiled. One can observe important criteria, such as the presence or absence of ocelli and the presence or absence of an intercalated cell in the hindwing. Such important criteria often serve to separate genera, but in the case of the genus Sciopetris, too many questions remain unanswered. For this reason we do not split the genus and accept it in its present, heterogeneous state until more is known of the species in question. In fact, the species S. melitensis, re-described in this paper, dif- fers from the general description of the genus by the absence of ocelli and the presence of an intercalated cell in the hindwing. However, we are convinced that the species reviewed here is the species found by Adolf Andres in 1915 and partly described by Rebel in 1919. HÄTTENSCHWILER et al.: On Sciopetris melitensis from Malta 404 2CT 75 | i eee SUISSTIU }SBO9 Bas JJO WY Ce “eISTUNT, uondııss>p [RUISIIO 9661 PAYSWNEH 1JOYSADY ST JOJOUURIP Ç'7 SU] (4) x z'xoidde ge “xoidde BIUIPIeS yorepy— Areniqay („) odAyesed Jo Apnys ‘UONdIIOSap JeUTSLIO 9L6T “19Pats 1851.10, Te eget [AN uondLII9sap JeuIsLIıo (ZL8I *UOJUIEJS) DSONaid 8-59 JEW -8°9—-E'¢ grew {rejnZuenn re sua] juosaid SUISSIUU (wu) Sue] Seq (wUI) 9ZIS 2[PUIS} 39[9107 uo sısAydıda ysıay 249 : aoueJSIp Saka SJUQUIB9S PUUQJUE SUIMOIO] (OS6I ones) €] UO ueyJ I9MOLIeU soyeos rey Soyeos SUIMPUIU poyeys EU+ QU [[99 payepesiggur JIM ‘9 Dan ose | M | ous au | m | me W (OST EAUSEM ‘UBJSIULUSJY a [TAod IY ‘PLIS Y AVITBOO]T à powers [199 [Bploosip u9JO CUI+ZTW ‘9 JJO SUIDA SUIMpUIy (9561 Ines) SO[BOS SUIMOIO]J [[99 TEPTOSSTP JJO SUI9A SUIMOIO] (wur) uedssurm dA uosvas SUIAY judy I € UMOUY Sa[eu J0 Joquinu SOIPNJS Jenyse UONdII9S9p [eUISLIO uONdLIISSP [EUTSLIO OJUI JO 291N0S 6161 ‘PA 6561 “I2P9IS 1681 sısuogyau asup SHOLIAOIN 2211190] * UMOUYUN,, AJPOIPUI svar JUPIG ‘satoads $242d012S [fe 107 vp umouy Jo Areunung “T° qeL Nota lepid. 30 (2): 397-406 405 en DLL Tw Cominotto @St.Paul's Is. . Manikata Gharghur : Naxxar- , - Rabat SPantelleria Linosa EN Gozo MALTA > = © Malta Lampedusa © © Filfla Is. Fig. 10. Map with the collecting localities of S. melitensis on the island Malta and position of the Maltese Archipelago in the Mediterranean Sea. Acknowledgements We are thankful for the help we obtained from various people, Prof. Dr. Willi Sauter for checking and discussing the manuscript, Dr. Martin Schwarz for his help in determining the parasitoid wasp, Dr. Gaden Robinson for his help in gathering the required literature, and not least, our thanks also go to Ruth Hättenschwiler for taking care of the nursery of these most interesting larvae. References Amsel, G. 1955. Uber mediterrane Microlepidopteren und einige transcaspische Arten. — Bulletin de l’Ins- titut royal des Sciences naturelles de Belgique 31 (83): 1-64. Andres, von A. 1916. Verzeichnis der wahrend meiner Kriegsgefangeschaft von mir auf Malta gesammel- ten Lepidoptera. — Entomologische Rundschau 33 (9): 43-45, (10): 48-49, (11): 50. Davis, D. R. 1978. The North American moths of the genera Phaeoses, Opogona and Oiniophila, with a discussion of their supergeneric affinities. - Smithsonian Contributions to Zoology 282. DeLucca, C. 1965. The place of the Lepidoptera in the Zoogeography of the Maltese Islands. — Extrait des rapports et procès-verbaux des réunions de la C.I.E.S.M.M. 18 (2): 511-515. Hattenschwiler, P. 1996. Sciopetris karsholti, eine neue Psychide aus Tunesien. — Nota lepidopterologica 19 (1/2): 107-112. Meyrick, E. 1891. A fortnight in Algeria, with description of new Lepidoptera. — Entomological Monthly Magazine 27: 58. 406 HÄTTENSCHWILER et al.: On Sciopetris melitensis from Malta Rebel, H. 1919. Zur Kenntnis palaearktischer Talaeporiiden. — Deutsche Entomologische Zeitschrift Iris, Dresden 32 (1918): 95-112. Sammut, P. 1984 a. A systematic and synonymic list of the Lepidoptera of the Maltese Islands. — Neue Entomologische Nachrichten 13: 1-124. Sammut, P. 1984 b. The present status of the endemic Lepidoptera of the Maltese Islands. — Potamon 13: 65-70. Sammut, P. 1985. Further additions to the Lepidoptera of the Maltese Islands. - SHILAP Revista de Lepi- dopterologia 13 (52): 304-306. Sammut, P. 1989. Il-Lepidoptera Maltija. — /n: T. Cortis (ed.), L-Identità Kulturali ta’ Malta. — Kungress Nazzjonali: 117-139. Sammut, P. & A. Valletta 1989. Lepidoptera. — Jn: P. J Schembri & J. Sultana (eds), Red Data Book for the Maltese Islands. — Department of Information, Valletta, pp. 97-104. Sammut, P. 2000. Kullana Kulturali 12. — Il-Lepidoptera. Pubblikazzjonijiet Indipendenza: 1—246. Malta. Sauter, W. 1956. Morphologie und Systematik der schweizerischen Solenobia-Arten. — Revue suisse de zoologie 63 (3): 451-550. Sauter, W. &. P. Hättenschwiler 1999. Zum System der palaearktischen Psychiden 2. Teil Bestimmungs- schlüssel für die Gattungen. — Nota lepidopterologica 22 (4): 262-295. Sieder, L. 1959. Neue Palaearktische Psychiden. — Zeitschrift der Wiener Entomologischen Gesellschaft 44: 145-150. Sieder, L. 1976. Eine neue Psychide aus Sardinien, Sciopetris hartigi spec. nov. — Entomologische Be- richten 36: 43-45 Stainton, H. T. 1872. List of Tortricina and Tinea collected in north-west Morocco by Mr Blackmoore, in 1870-71. — Entomologist’s Monthly Magazine 8: 233. Nota lepid. 30 (2): 407-410 407 Prays peregrina sp. n. (Yponomeutidae) a presumed adventive species in Greater London Davip AGASSIZ The Natural History Museum, Cromwell Road, London SW7 SBD; e-mail: d.agassiz@nhm.ac.uk Resume. Prays pergrina sp. n. is described from scattered localities in Greater London. The possible origin of this species is discussed. Introduction In the autumn of 2005 two British lepidopterists: Colin Plant and Philip Sterling con- tacted me about a mystery yponomeutid of which a specimen had been taken in West London. Shortly afterwards a second specimen was reported which appeared to belong to the same species. Over the following months further specimens were reported. I searched through the collections of the Natural History Museum (BMNH) but was un- able to find an exact match for the specimens. A male was dissected which seemed to confirm that the species belonged to the genus Prays. Pictures of adult and genitalia were circulated to entomologists in Europe and Australia, but none could suggest an identity. Moriuti (1977) described six new species from Japan but it resembles none of these. When a seventh specimen was reported in 2006 I made a further search of the BMNH collections and found that P. curulis Meyrick seemed to be the nearest species, a male had been dissected by Moriuti but there were some differences in the genitalia. It therefore seemed necessary to describe the species as new. A summary of recorded specimens is as follows: Date 15.viii.2003 | Parliament Hill | TQ2785_ | 0 19.vi.2005 | Chelsea Physic Garden | TQ2778 | 9 6.ix.2005 | Wood Green | TQUSSS | © 5.x.2005 ER EE 28.vii.2006 6.ix.2006 ae ee BE Bez WE 9 | Mani Br BE 16.ix.2006 18.x.2006 24.vi.2007 10.vii.2007 24.viii.2007 5.ix.2007 8.ix.2007 14.ix.2007 David Macklin 2.x.2007 | Wimbledon | TQ2670 | à | ViadProklov | The distribution of these records is shown in Fig. 5 Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 408 Acassız: À new Prays species described from London Prays peregrina Sp. n. Figs 1-4 Material. Holotype: 9 ‘5" Oct. 2005 D. Howdon | Melville Ave, Greenford, | Middx.’ ‘BM genitalia slide No. 31468.’ a red circled holotype label and ‘Prays peregrina Agassiz det D. Agassiz, 2007. Deposited in BMNH. — Paratypes: 10° Alexandra Road, Middlesex, London N8, 5.1x.2005; BM genitalia slide No. 31467, BMNH; 10° Parliament Hill, London NW3, 15.v111.2003, R. Softly, coll. C. W. Plant; 19 London NS 28.v11.2006, M. Ashby, coll. R. Terry; 19 Chelsea Physic Garden, SW, 19.vi.2005, T. H. Freed, coll. T. H. Freed; 19 Barnet, 18.x.2006, R. Terry, coll. R. Terry. Description of the adult (Figs 1-2). Wingspan 14 mm. Head pale grey, labial palpus white, segment 3 longer than segment 2, tuft of white hairs arising from base; antenna just over half wing length, weakly annulate pale fuscous and white; pair of white tufts behind head. Thorax dark fuscous. Forewing white with scattering of dark fuscous dots, dark fuscous mark arising from middle of dorsum in shape of curved triangle, dark fus- cous tornal spot, terminal cilia concolorous with adjacent wing. Hindwing uniform pale grey, small hyaline patch above vein CuP near base. Underside of all wings fuscous. Legs white, forelegs fuscous above. Abdomen white. The species is similar to Prays curulis Meyrick, but differs in lacking partial fascia from middle of dorsum, and having more pronounced tornal spot. : Male genitalia (Figs 3, 3a) Soci strong with tips drawn out to a point, gnathos forming an arched rod, valva with sacculus forming a small point, saccus long and slender. Phallus curved with a cluster of 2 large and 3-4 smaller cornuti. Differs from P. curulis in the more uniform socii, longer and narrower saccus and the stronger and fewer cornuti in the phallus. Female genitalia (Fig. 4) Ostium wide, ductus bursae strong and straight, slight- ly longer than ostial chamber, corpus bursae ovate, signum small sclerotised plate with two prongs. Distribution. All known specimens from Greater London (Fig. 5). The native range of the species is unknown, suspected to be in Asia, specimens of the closely related P. cu- rulis, described by Meyrick (1914), in the Natural History Museum are from northern India and Nepal. Life History unknown. Etymology. The name is taken from the Latin word meaning “that comes from foreign parts” Discussion The origin of these specimens remains a matter of speculation. Either they must be breeding locally or else emerging from imported plant material or foodstuffs. If the former were the case one would expect a local population to build up, and therefore the scattered nature of the records suggests that they are repeatedly imported. The lack of records from any other city or country is surprising, as well as the fact that there is no mention of such a Prays species in the economic literature. It is also puzzling that there are no records prior to 2003. The genus Prays Hiibner contains over 40 described species predominantly from the Old World; there are 23 species from Asia, 8 from Australia, 4 from Europe, 3 from Africa, and the remainder from certain oceanic islands or South America. Foodplants used are Oleaceae (Fraxinus, Juglans, Ligustrum, Olea), Rutaceae (Citrus), Caprifoliaceae Nota lepid. 30 (2): 407-410 409 Figs 1-4. 1. Set specimen of P. peregrina (photo Rachel Terry). 2. Live specimen of P. peregrina (pho- to Martin Jordan). 3. Male genitalia, with phallus on the left (photo Rachel Terry). 4. Female genitalia, 4a signum enlarged. (Viburnum) and a few other families. Larvae are known to feed on fruits and in shoots of the host plant as well as the leaves. The specimen taken in Chelsea Physic Garden was a female and a spermatophore in the corpus bursae indicated that it had paired with a male, so several specimens must have emerged at about the same time. It seems most likely that the specimens have been imported as larvae or pupae in some foodstuff from Asia, but this is purely conjecture, the span of dates of capture from June to October suggests either that the species is continuously brooded and repeatedly imported, or it may have become locally established. Malumphy (2007) listed various arthropod species intercepted on Citrus hystrix which has only recently been imported from Indonesia, since this belongs to an appropriate family it should be explored as a possible host plant. 410 Acassız: A new Prays species described from London Fig. 5. Map of Greater London showing distribution of records of P. peregrina (Dmap). Acknowledgements The recorders of all specimens cited are thanked for their willing collaboration and assistance, Dr Klaus Sattler has given advice and encouragement. Thanks to Rachel Terry for making the preparation and pho- tographing the male genitalia, the distribution map was generated using Dmap. References Malumphy, C. 2007. Arthropods intercepted on fresh leaves of Citrus hystrix DC imported into England and Wales. — Entomologist’s Gazette 58: 39-50. Meyrick, E. 1914. Indian Micro-Lepidoptera. — Journal of the Bombay Natural History Society 23: 125-126. Moriuti, S. 1977. Fauna Japonica: Yponomeutidae s.lat. (Insecta: Lepidoptera). — 327pp. 96 pls, Tokyo. Nota lepid. 30 (2): 411-414 All Dyspessa aphrodite sp. n. from Greece (Cossidae) ROMAN V. YAKOVLEV! & THOMAS J. WITT? ' ul. Chkalova, 57-81, Barnaul, 656049, Russia; e-mail: yakovlev_r@mail.ru 2? Tengstr.33, 80796 München, Germany; e-mail: thomas @ witt-thomas.com Abstract. Dyspessa aphrodite Yakovlev & Witt, sp. n. is described from Greece and compared with Dyspessa emilia (Staudinger, 1878), with which it has been confused. Therefore, D. emilia must be re- moved from the European list of Cossidae. Introduction Our knowledge of European Cossidae is considered satisfactory. After the revisions of Daniel (1955—1965) several comprehensive papers and books have been published with new data on the distribution and systematics of the European carpenter moths (de Freina & Witt 1990; Karsholt & Razowski 1996; Yakovlev 2005). However, while examining material in the Thomas Witt entomological museum (MWM) the authors surprisingly found a series of an intriguing carpenter moth of the genus Dyspessa Hübner, 1820 originating from Greece (Peloponnes, Mega Spileon). These moths had been wrongly identified as Dyspessa emilia (Staudinger, 1878) and were mentioned as such for the first time for the fauna of Greece by de Freina & Witt (1990: 31-32, pl. 4 figs 10-12), and later by Karsholt & Razowski (1996). A detailed examination of these specimens revealed that they represent a species new to science and its description is given below. As a result, Dyspessa emilia (Staudinger, 1878) has to be deleted from the list of European Cossidae and must be replaced by Dyspessa aphrodite Yakovlev & Witt, sp. n. Dyspessa aphrodite sp. n. Figs 1-3, 7, 9 Material. Holotype-C: “Greece | Peloponnes | Mega Spileon | 6.vi.1981 | Mühle leg.”, “Holotypus | Dyspessa aphrodite | Yakovlev & Witt”, MWM. — Paratypes: 70, same data (GenPr. 11821, 11822) MWM. Add separators between lines of labels and add Holotype label and others if applicble. Description. Forewing length 9-10 mm. Antenna bipectinate, medially with processes becoming 1.5 times longer than segment diameter, space between processes as wide as processus breadth. Forewing pale yellow with small dark spot in discal zone in area of cubital veins, relatively large brown spot in postdiscal zone in radial area (at discal cell apex), and weakly expressed brown spot at apex. Two paratypes miss these spots. Fringe mostly yellow, but basaly dark at tips of radial veins. Hindwing greyish yellow with pale yellow fringe. Male genitalia (Fig. 9). Uncus triangular with a beak-like pointed apex. Gnathos arms thin and long, gnathos small. Valva trapezoid with almost rectangular distal mar- gin, also with small crest on costal margin; distal end membranous. Arms of transtilla forming wide triangles. Saccus small, rounded. Juxta small. Phallus curved, thin and tapering to apex; vesica opening occupying half of its length. Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 412 YAKOVLEV & Witt: Dyspessa aphrodite sp. n. from Greece ak Figs 1-3. Dyspessa aphrodite Yakovlev & Witt, sp. n. (MWM). 1. Holotype. 2-3. Paratypes. Figs 4-6. Dyspessa emilia (Staudinger, 1878), males from Turkey, (MWM). 4. Anatolia, 25 km south of Sivas, 1500 m, 24—26.vii.1978, leg. W. Thomas. 5-6. Ak.-Ch.-Tschiftlik, 19-20.v11.1928, coll. v. Bartha. Diagnosis. The new species is close to Dyspessa emilia (Staudinger, 1878) (Figs 4-6, 8, 10) from which it differs by the following characters: The antennal processus in D. emilia is much longer; it is 2.5—3 times longer than the anten- nal diameter while in D. aphrodite it is not longer than 1.5 times this diameter. The forewing is much wider than in D. emilia. The basal parts of the forewing fringe are dark only at the distal end of the radial veins in D. aphrodite, whereas it is dark at the distal end of all veins in Figs 7-8. Antennae of male Dyspessa specimens. 7. D. aphrodite Yakovlev & Witt, Sp. n., paratype. 8. D. emilia (Staudinger, 1878). Nota lepid. 30 (2): 411-414 413 Fig. 9-10. Male genitalia Dyspessa aphrodite. 9. Dyspessa aphrodite Yakovlev & Witt, sp. n. 10. Dyspessa emilia (Staudinger, 1878). D. emilia. In forewing pattern, the spots of the postdiscal row are approximately equal in sizein D. emilia. In the new species the spot in the radial area is larger than the oth- ers, which are shifted into the discal zone. The male genitalia practically do not differ, except for the somewhat more expressed processus on the costal margin of the valva in Dyspessa emilia. However, our analysis of very large material in genus Dyspessa showed very weak interspecific differences in genitalia structure. Etymology. In Greek mythology Aphrodite is the goddess of love and beauty. Distribution. Known only from the type locality. Remark. The absence of forewing dark spots on two paratypes may be due to wear or individual variation. References Daniel, F. 1955. Monographie der Cossidae I. Kritische Beurteilung der bisher dem Genus Srygia Latr. zugeteilten Arten. — Mitteilungen der Münchener Entomologischen Gesellschaft 44-45 (1954-1955): 159-181, pl. 1. Daniel, F 1956. Monographie der palaearktischen Cossidae H. Die Genera Cossus Fabr. und Lamello- cossus gen. n. (Lep.). — Mitteilungen der Münchner Entomologischen Gesellschaft 46: 243-289, pls 8-10. Daniel, F. 1959: Monographie der palaearktischen Cossidae III. Das Genus Holcocerus Stgr. — Mitteilungen der Münchener Entomologischen Gesellschaft 49: 102-160. Daniel, F. 1960: Monographie der palaearktischen Cossidae IV. Die Genera Cossulinus Kby., Dyspessa- cossus Dan. und /soceras Tti. (Lep.). — Mitteilungen der Münchener Entomologischen Gesellschaft 50: 93-118. Daniel, F. 1961. Monographie der palaearktischen Cossidae V. Die Genera Parahypopta g.n., Sinicossus Clench und Catopta Stgr. — Mitteilungen der Münchner Entomologischen Gesellschaft 51: 160-212, pls 16-17. Daniel, F. 1962: Monographie der palaearktischen Cossidae VI. Genus Dyspessa Hbn. Erster Teil. — Mitteilungen der Münchener Entomologischen Gesellschaft 52: 1-38. Daniel, F. 1964: Monographie der palaearktischen Cossidae VII. Genus Dyspessa Hbn. Zweiter Teil. Genus Paropta Stgr. — Mitteilungen der Münchener Entomologischen Gesellschaft 54: 181-236. 414 YAKOVLEV & Witt: Dyspessa aphrodite sp. n. from Greece Daniel, F. 1965. Monographie der palaearktischen Cossidae VIII. Nachträge und Register zur Sub- familie Cossinae. — Mitteilungen der Münchner Entomologischen Gesellschaft 56: 77-114, pls 2-3. Freina, J. J. de & T. J. Witt, 1990. Die Bombyces und Sphinges der Westpalaearktis (Insecta, Lepidoptera). Band 2. Cossoidea, Hepialoidea, Pyraloidea, Zygaenoidea. — Edition Forschung & Wissenschaft Verlag GmbH, Miinchen. 134 pp., 10 pls. Karsholt, O. & J. Razowski (eds.) 1996. The Lepidoptera of Europe. A distributional checklist. — Apollo Books, Stenstrup, 380 pp. Yakovlev, R.V. 2005. New data of distribution and systematic of Cossidae (Lepidoptera) of Europe and adjacent territories. — Eversmannia 3/4: 18-27. (in Russian) Nota lepid. 30 (2): 415-421 415 Taxonomic notes on Acossus Dyar and Parahypopta Daniel (Cossidae) ROMAN V. YAKOVLEV ul. Chkalova, 57-81, Barnaul, 656049, Russia; e-mail: yakovlev_r@mail.ru Abstract. While studying material of Cossidae from North America it was found that Acossus Dyar, 1905 is a synonym of Lamellocossus Daniel, 1956, syn. n. Thus, the following combinations are made: Acossus terebra ({Denis & Schiffermüller], 1775) comb. n. and Acossus viktor (Yakovlev, 2004) comb. n. Genus Acossus includes five species and is Holarctic in distribution. The type and topotypical specimens of Parahypopta caestrum radoti (Homberg, 1911) were investigated. Distinct features of pattern and mor- phology differentiate it from Parahypopta caestrum (Hübner, 1804). Parahypopta radoti (Homberg, 1911) stat. n. proves to be a distinct species. 1. Acossus Dyar, 1905 Type species: Cossus undosus Lintner, 1878 Genus Acossus was described by Dyar (1905) for the North American Cossus undo- sus Lintner, 1878 and since 1905 the name was used only for representatives of the Nearctic fauna. Ureta (1957) described from Chile Acossus comadioides from Chile, a name that was later synonymised with Rhizocossus munroei Clench, 1957 (Gentili 1985). Half a century after the description of Acossus, Daniel (1956) described the genus Lamellocossus, to exclude Bombyx terebra Denis et Schiffermiiller, 1775 from Cossus Fabricius, 1793 with which it had been associated. As the main diagnostic char- acter, Daniel mostly used the antennal structure of the male. Bombyx terebra clearly differs in this character from the type species Cossus cossus (Linnaeus, 1756), because of its bipectinate antenna. Later, Russian authors (Zagulyaev 1978; Chistyakov 1999; Yakovlev 2004) soundly stressed the differences between Cossus and Lamellocossus. A new species, Lamellocossus viktor Yakovlev, 2004 was described from the terebra- group; it inhabits the arid regions of South Tuva (the northern part of the Great Lake Hollow, at Lake Tere-Khol’ and the Tes Khem River) and should occur also in northern Mongolia. While studying material of the private collections by Thomas Witt (Munich) and Armin Hauenstein (Untermiinkheim) and after an assessment of the literature I found that Acossus undosus and A. centerensis (Lintner, 1877) specimens from Canada presented striking external similarities with Lamellocossus specimens. My examination of the genitalia of both North American species showed them to be nearly identical with those of Lamellocossus terebra. The external traits, antenna and male genitalia structure leave no doubt that L. terebra and L. viktor belong to Acossus, which is thus Holarctic in range. Hitherto, the only known Holarctic taxon of Cossidae was Zeuzera pyrina (Linnaeus, 1761) (Hodges, 1983), which, however, quite probably was introduced to North America by man. Diagnosis: Medium-sized, gray-colored moths. Antenna bipectinate, each segment with two long processes. Thorax and abdomen with dense pubescence of gray hairy scales. Fore wing gray, somewhat darker at base, with wavy streaky pattern through- Nota lepidopterologica, 15.11.2007, ISSN 0342-7536 416 YAKOVLEV: On Palaearctic Cossidae Lorch] Figs 1-4. Adults of Acossus. 1. A. undosus, male, Canada, British Columbia, Hazelton, 30 km N Kispioux River, Resort, 400 m, 3.08.-10.08.1999, leg. S. Ortner, Museum Witt, München. 2. A. centerensis, male, Canada, British Columbia, Interior Carıbou Mountains, Lake Canım, 900 m, 26.07.-3.08.1997 coll. Hauenstein, Untermünkheim. 3. A. terebra, male, Russia, W. Altai Mts. [Altaiskii Krai Region, Zmeinogorski district], 5 km south of Samarka, 800 m, 25-26.07.1993, leg. Z. Varga, Museum Witt, München. 4. A. viktor, holotype, Zoological Institute, St. Petersburg. out; fringe checkered. Hindwing somewhat lighter gray, often with an expressed wavy streaky pattern. Male genitalia. Uncus long, beak-like; gnathos arms long and quite thick, quite large at point of fusion, covered with fine spinules. Valva broad, cup-like, swollen, with convex costal margin. Arms of transtilla strongly sclerotised, hook-shaped, pointed. Juxta small, strongly sclerotised; saccus weakly expressed. Phallus weakly curved, relatively thick and long; vesica opening dorso-apical, at about one third of phallus length. Female genitalia. Oviscapt short. Apophyses posteriores very long; papillae anales elongate and smoothly rounded apically; anthrum immersed; ductus bursae wide membranous; bursa ellipsoid, without signa; ductus seminalis from base of bursa base near connection of ductus bursae. Diversity. The genus includes five species: Acossus undosus (Lintner, 1878: 243) (Cossus) (Figs 1, 5), occuring in Canada and the USA (Barnes & McDunnough 1911; Schoorl 1990). Acossus populi (Walker, 1856: 1515) (Cossus), occuring in Canada and the USA (Barnes & McDunnough 1911; Schoorl 1990). Nota lepid. 30 (2): 415-421 417 Figs 5-7. Genitalia of Acossus. 5. A. undosus, male. 6. A. terebra, male (Chistyakov 1999). 7. A. terebra, female (Chistyakov 1999). Acossus centerensis (Lintner, 1877: 129) (Cossus) (Fig. 2) occuring in Canada and the USA (Barnes & McDunnough 1911; Schoorl 1990). Acossus terebra (Denis & Schiffermüller, 1775: 60) (Bombyx) comb. n. (Figs 3, 6-7) occuring in Eurasia (Israel, Turkey, northern Spain, middle and southern Europe, southern Sweden, Finland, Baltic States, Ukraine, central part of European Russia, Caucasus, southern Siberia including the Altai and Sayan Mts. to S. Yakutia, southern part of Far East, Korea, Heilongjiang, Jilin, inner Mongolia (de Freina & Witt 1990; Hua et al. 1990; Yakovlev 2004). Acossus viktor (Yakovlev, 2004), (Lamellocossus) comb. n. (Fig. 4) occuring in Russia, southern Siberia, southern part of Tuva Republic. Note. Fania connectus Barnes et McDunnough, 1916, formerly treated in Acossus, has been transferred to Fania Barnes, McDunnough, 1911 (Schoorl 1990), which is followed here. Remarks. Acossus belongs to Cossinae Leach, 1815, is similar to the Palaearctic Cossus, Gobibatyr Yakovlev, 2004 (type species: Cossus colossus Staudinger, 1887) and the North American Prionoxystus Grote, 1882 (type species: Cossus robiniae Peck, 1818). 418 YAKOVLEV: On Palaearctic Cossidae Coll. À. Homberg 1% P.E.L. Viette det. 19 { Engl Ir |. Caen Fun var. | radoË Homers Bull. Soc. end FE 14/) 5 * £ À € £ . : Dov RaTieV Fig. 8. Parahypopta radoti, lectotype. Fig. 9. Lables of the lectotype of Parahypopta radoti (Homberg, 1911). 2. Parahypopta radoti (Homberg, 1911) stat. n. Figs 8—12 Hypopta radoti Homberg, 1911. Type locality: France, Alpes-Maritimes, Cannes. Hypopta caestrum radoti Homberg, 1911: 143-144. Type locality: France, Alpes-Maritimes, Cannes. Homberg (1911) briefly described Hypopta caestrum radoti, pointing out the dark pat- tern of the specimens, collected at Cannes (Alpes-Maritimes). Seitz (1912: 424) and Dalla Torre (1923: 22) considered this taxon as a subspecies of Parahypopta caestrum. Daniel (1961: 161-162) wrote that Parahypopta radoti is «eine Zustandsform oder Mutation, die im Bereich einer engen Population einmal die vorherrschende Form wer- den kann», thus not considering these moths as a valid taxon. Leraut (1980: 52) men- tioned Parahypopta radoti Homberg, 1911 as a junior synonym of Parahypopta caes- trum (Hübner, 1804). De Freina & Witt (1990: 22) adopted the opinion of the previous authors and noted that ““Neben den bei Cossiden üblichen Größendifferenzen einzelner Populationen zeigt sich ... auch schiefergrau verdunkelte Individuen, die innerhalb jeder Population mehr oder weniger häufig auftreten können: f. radoti Homberg, 1911”. This taxon was not been mentioned in the comprehensive list of the lepidopteran fauna of Europe by Karsholt & Razowski (1996). In the Muséum National d’Histoire Naturelle Paris (MNHN), I investigated the lectotype (designated below) of Hypopta caestrum radoti and several topotypical specimens, including syntypes. I found that this taxon is a valid species, clearly distinct from Parahypopta caestrum (Hübner, 1804) by external characters as well as the male genitalıa structure. Description. Male. Fore wing length 12-13 mm. Fore wing whitish-brown, with a row of small roundish brown spots at costal margin; there are several brown spots partly fused to each other; submarginal area somewhat lighter than the rest of wing area; entire central part of wing occupied by a dark-brown spot in postdiscal area with Nota lepid. 30 (2): 415-421 419 Figs 10-15. Parahypopta spp. 10. P. radoti, male, Cannes, MNHN. 11. P. radoti, female, Cannes, MNHN. 12. P. radoti, male genitalia. 13. P. caestrum, male, Hungary. 14. P. caestrum, female, Hungary. 15. P. caestrum, male genitalia. wavy transversal lines; wing basal area without pattern. Fringe checkered dark at veins and light between them. Hind wing brown without pattern; with a light-brown fringe. Male genitalia. Uncus triangular, wide, with a membranous zone on its upper surface. Tegumen wide. Gnathos arms very stout and robust, gnathos covered with small spinules, especially dense laterally. Valva broad, distal end membranous and mobile; costal margin with robust trapeze-shaped processus with slightly dentate margin. Arms of transtilla with wide bases and curved and pointed ends. Juxta carina-shaped with broad weakly sclerotized lateral processes, slightly widening apically. Saccus wide, 420 YAKOVLEV: On Palaearctic Cossidae semicircular. Phallus stout, slightly curved, with slantingly cut at distal end; vesica with opening dorso-apically and occupying about half of phallus length; without cornuti. Diagnosis. Clearly differs from P. caestrum (Figs 13-15) by the much darker wing coloration, the slightly shorter forewing with rounded apex, and detalis of the male genitalia structure: the somewhat wider juxta, the much more developed processus on the costal margin of the valva, the shorter and less curved arms of transtilla, the curved phallus. Taxonomical notes. Taking into account the complicated taxonomy within this group, it is necessary to fix the name Parahypopta radoti to a specific specimen. Many syn- types of Hypopta caestrum radoti are preserved in the Natural History Museum of the Humboldt University of Berlin, the Zoologische Staatssammlung des Bayerischen Staates of Munich and there is one specimen labelled as lectotype at the MNHN Paris. However, according to my own data the lectotype designation has not been published. Thus, to preserve the selection of this specimen, I hereby designate it as the lectotype. The specimen (Fig.8) bears the following labels (Fig.9): (1) Red, rectangular, with ‘Type’ printed typographically; (2) White, rectangular, with ‘Coll. R. Homberg | Muséum Paris’ printed typographically; (3) White, rectangular ‚with R. Homberg’s hand-writing in black ink ‘Env. Cannes | 1. Juillet 1910 | ex larva. as page IR. Homberg’; (4) White, rectangular, with R. Homberg’s hand-writing in black ink ‘Var. Radoti. Hmbg. | Type | Bull. Soc. Ent. France | 12 avril 1911’; (5) White, rectangular, with five lines of text of which the first is typographically printed while the rest are written in black ink by P. Viette’s hand as follows ‘P.E.L. Viette det. 19 | Hypopta L. T. | caestrum var. | radoti Homberg | Bull. Soc. Ent. Fr., | 1911, p. 143’; (6) White, circular, with black mar- gin, with typographically printed ‘LECTO- | TYPE’; (7) White, rectangular, with A. Kondratyev’s hand-writing in black ink ‘gen. prep. | Ne. 5. | Kondratiev’; (8) Red, rec- tangular, with the following hand-written text in black ink: ‘LECTOTYPE | Hypopta caestrum Hb. | var. Radoti Homberg, 1911| des. R. V. Yakovlev’. Distribution. Reliably known only from the type locality. A clarification of the range is planned in the course of a revision of the genus Parahypopta Daniel, 1961. Acknowledgements I am thankful to all who helped me to prepare this work: A. Hauenstein (Untermünkhein), O. Kosterin (Novosibirsk), J. Minet (Paris), T. J. Witt (Munich), and V. Zolotuhin (Ulyanovsk). References Barnes, W. & J. H. McDunnough. 1911. Revision of the Cossidae of North America. — Contributions to the natural history of the Lepidoptera of North America 1: 1-36, pls 1-7. Chistyakov, Yu. A. 1999. Family Cossidae. — In: P. A. Ler (ed.), Opredelitel’ nasekomykh Dal’nego Vosto- ka Rossii 5 (2). — Vladivostok. Dal’nauka: 309-319. (in Russian) Dalla Torre, K. W. von. 1923. Cossidae. — In: W. Junk (ed.), Lepidopterorum Catalogus 29. Berlin. 63 pp. Daniel, F. 1956. Monographie der palaearktischen Cossidae. II. Die Genera Cossus Fabr. und Lamellocos- sus gen. n. (Lep.). — Mitteilungen der Münchner Entomologischen Gesellschaft E. V. 46: 243-289, pls 8-10. Nota lepid. 30 (2): 415-421 42] Daniel, F. 1961. Monographie der palaearktischen Cossidae. V. Die Genera Parahypopta g.n., Sinicossus Clench und Catopta Stgr. — Mitteilungen der Münchner Entomologischen Gesellschaft 51: 160-212, pls 16-17. Denis, M. & I. Schiffermüller 1775. Ankündigung eines systematischen Werkes von den Schmetterlingen der Wienergegend. — Augustin Bernardi, Wien. Dyar, H. G. 1905. New genera of South American moths. — Proceedings of the United States national Museum 29: 173-178. Daniel, F. 1956. Monographie der palaearktischen Cossidae. II. Die Genera Cossus Fabr. und Lamellocos- sus gen. n. (Lep.). — Mitteilungen der Münchner Entomologischen Gesellschaft E. V. 46: 243-289, pls 8-10. Freina, J. J. de & T. J. Witt. 1990. Die Bombyces und Sphinges der Westpalaearktis (Insecta, Lepidoptera). Band 2. Cossoidea, Hepialoidea, Pyraloidea, Zygaenoidea. — Edition Forschung & Wissenschaft, Miinchen. 134 pp., 10 pls. Gentili, P. 1985. La familia Cossidae de Patagonia (Lep.: Ditrysia). — (Thesis of) Facultad de Cs. Natura- les y Museo-UNLP, Argentina. Hodges, R. W. 1983. Cossidae. p. 31. — In: R. W. Hodges (ed.), Check list of the Lepidoptera of America North of Mexico. — E. W. Classey Ltd. Homberg, R. 1911. Description d’une nouvelle variété de Hypopta caestrum Hb. [Lep. Cossidae]. — Bulletin de la Société entomologique de France 7: 143-144. Hua, B., I. Chou, D. Fang & S. Chen 1990. The Cossid fauna of China (lepidoptera, Cossidae). — Ilustrata] cinaj insect-faunoj H. Tianze Eldonejo. Yangling, Shaanxi, China. 147 pp., 7 pls. Karsholt, O. & J. Razowski (eds.). 1996. The Lepidoptera of Europe. A distribution Checklist. — Apollo Books. 380 pp. Leraut, P. 1980. Liste systématique et synonymique des Lépidopteres de France, Belgique et Corse. — Supplément a Alexanor et au Bulletin de la Société entomologique de France. 334 pp. Lintner, J. A. 1877. On a new species of Cossus. - Canadian Entomologist 9: 129-130. Lintner, J. A. 1878. Contribution to the Economical Entomology of the year 1876. — Report on the New York State Museum of Natural History 30: 236-243. Schoorl, J. W. 1990. A phylogenetic study on Cossidae (Lepidoptera: Ditrysia) based on external adult morphology. — Zoologische Verhandelingen 263: 295 pp, 1 pl. Seitz, A. 1912. Die Gross-Schmetterlinge der Erde. 2: Die Palaearktischen Spinner & Schwärmer. — Stutt- gart, 479 pp. Ureta, E. 1957. Revision de la familia Cossidae (Lep. Het) en Chile. — Bulletin Museum nacional History naturale Chile 27 (2): 129-153. Walker, F. 1856. List of the Specimens of Lepidopterous Insects in the Collection of the British Museum, London 7: 1509-1808. Yakovlev, R. V. 2004. Carpenter-moths (Lepidoptera, Cossidae) of Siberia. — Euroasian Entomological Journal 3 (2): 155-163, pl. 1. (in Russian) Zagulyaev, A. K. 1978. Fam. Cossidae. pp. 177-186. — Drevotochcy. — In: G. S. Medvedev (ed.), Opredeli- tel’ nasekomykh evropeiskoi chasti SSSR. T. IV Lepidoptera (1). — Leningrad. Nauka. (in Russian) 422 Instructions for authors NOTA LEPIDOPTEROLOGICA A journal focussed on Palaearctic and General Lepidopterology Published by the Societas Europaea Lepidopterologica e.V. http://www.soceurlep.org INSTRUCTIONS FOR AUTHORS Co-Editors: Dr Bernard Landry and Dr Matthias Nuss Manuscripts and all related correspondence should be sent to Dr Matthias Nuss Museum fiir Tierkunde, Königsbrücker Landstr. 159, 01109 Dresden, Germany e-mail: matthias.nuss@snsd.smwk.sachsen.de General. Nota lepidopterologica is a scientific journal published by Societas Europaea Lepidopterologica (SEL) on May 15 and November 15 each year. Because SEL is a registered non-profit organization, no royalties will be paid to authors of contributions. A pdf and twenty- five reprints of each paper will normally be supplied free of charge to the first author; additional copies may be ordered on a form enclosed with the proofs. However, no reprints (even for charge) will be available for book reviews. 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Der Mitgliedsbeitrag ist jährlich am Jahresanfang zu entrichten. Er beträgt für Einzelpersonen € 35,00 bzw. für Vereine € 45,00. Die Aufnahmegebühr beträgt € 2,50. Die Zahlung wird auf das SEL-Konto 19 56 50 507 bei der Postbank Köln (BLZ 370 100 50) erbeten (IBAN: DE63 3701 0050 0195 6505 07; BIC: PBNKDEFF). Mitteilungen in Beitragsangelegenheiten werden an den Schatzmeister Dr. Robert Trusch, Staatliches Museum für Naturkunde, Erbprinzenstr. 13, D-76133 Karlsruhe; e-mail: trusch@smnk.de erbeten. Adressenänderungen sollten umgehend dem Mitgliedssekretär oder dem Schatzmeister mitgeteilt werden. Published by the Societas Europaea Lepidopterologica (SEL), Nota lepidopterologica is a scientific journal that members of SEL receive as part of their membership. Authors who would like to submit papers for publication in Nota lepidopterologica are asked to take into consideration the relevant instructions for authors available on the SEL homepage at http://www.soceurlep.org. The sales of single and back issues of Nota lepidopterologica as well as sales to non-members of SEL are under the responsibility of Apollo Books, Kirkeby Sand 19, DK-5771 Stenstrup; e-mail: apolio- books@vip.cybercity.dk. The membership is open to individuals and associations as provided for by the statutes of SEL. Applications for membership are to be addressed to the Membership Secretary Willy De Prins, Dorpstraat 401 B, B-3061 Leefdaal, Belgium; e-mail: willy.de.prins@telenet.be. The application form is available on the SEL homepage. The annual subscription is to be paid at the beginning of the year. It is 35.00 € for individuals or 45.00 € for associations. The admission fee is 2.50 €. Dues should be paid to SEL account no. 19 56 50 507 at Postbank Köln [Cologne] (bank code 370 100 50; IBAN: DE63 3701 0050 0195 6505 07; BIC: PBNKDEFF) or to local treasures as mentioned on the website. Communications related to membership contributions should be sent to the Treasurer Dr Robert Trusch, Staatliches Museum fiir Naturkunde, Erbprinzenstr. 13, D-76133 Karlsruhe; e-mail: trusch@smnk.de. Changes of addresses should be immediately communicated to the Membership Secretary or the Treasurer. Publié par la Societas Europaea Lepidopterologica (SEL), Nota lepidopterologica est un périodique scientifique envoyé à tous les membres de la SEL. Les auteurs qui désirent publier des manuscrits dans la revue sont priés de tenir compte des Instructions aux auteurs disponibles sur le site Web de la SEL: http://www.soceurlep.org. Les ventes de numéros supplémentaires ou d'anciens numéros de Nota lepidopterologica, ainsi que les ventes de numéros aux personnes n'étant pas membres de la SEL sont sous la responsabilité de Apollo Books, Kirkeby Sand 19, DK-5771 Stenstrup; courriel: apollobooks@vip.cybercity.dk. Tel que prévu dans ses statuts, les individus de même que les associations peuvent devenir membres de la SEL. Les demandes d'adhésion doivent être envoyées au Secrétaire responsable des adhésions, Willy De Prins, Dorpstraat 401 B, B-3061 Leefdaal, Belgique; courriel: willy.de.prins @telenet.be. Le formulaire d'adhésion est disponible sur le site Web de la SEL. L'adhésion se paie au début de l'année. Elle est de 35 € pour les individus et de 45 € pour les associations. Les frais d'admission sont de 2,50 €. Les paiements peuvent être envoyés au compte de la SEL: no. 19 56 50 507, Postbank Köln [Cologne] (code bancaire 370 100 50; IBAN: DE63 3701 0050 0195 6505 07; BIC: PBNKDEFF) ou au trésorier local tel que mentionné sur le site Web.. Toute question en rapport avec l'adhésion doit être envoyée au Trésorier, Dr. Robert Trusch, Staatliches Museum für Naturkunde, Erbprinzenstr. 13, D-76133 Karlsruhe; courriel: trusch@smnk.de. Tout changement d'adresse doit être mentionné immédiate- ment au Secrétaire responsable des adhésions ou au Trésorier. "SMITHSONIAN INSTITUTION LIBRARIES AMUN 3 9088 01388 3012 |