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Volume 80 - Number 2 - 2007




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NFO 11 2007


A journal focussed on Palaearctic and General Lepidopterology Published by the Societas Europaea Lepidopterologica e.V.



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

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Dr Enrique Garcia-Barros (Madrid, E), Prof. Dr Roger L. H. Dennis (Wilmslow, UK),

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Volume 30 No. 2 - Dresden. 15.11.2007 - ISSN 0342-7536


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


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



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.


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.


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.


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’ <difficult to read: this could be Boro Khoro Mts, E. Turkestan, 44.20N, 83.00 E; further location data is illegible>, 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



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’, <dry genitalia preparation glued to card>, ‘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’, <dry genitalia preparation glued to card>, ‘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’.


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


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.


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


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?


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.


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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-