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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

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(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

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.

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.

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.

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),

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.

1.0 619 A

ale 656

O 627

0.6 625 O

628 654

0.4 618 EH 644 O

m O0 D 4623 630b

0.2 622 O

O649

> SE? 621

< ae on 657b

04 653 Se O

À 624b

-0.6 655

=. 624a

-1.0 629

12 643

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. 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. clarissima

P. droshica

P. esquilinus

P. gilgitica

P. lehana

P. turkestana

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.

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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-

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,

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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

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

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

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 <1 0 <p < 0.05 OBkir

Bükk

9Bzs

Figs 2-3. 2. Distribution of alleles with different frequencies in the populations of the two regions in the

two years of collection. Agg99: Karst samples collected in 1999; Agg00: Karst samples collected in 2000;

Bükk99: samples collected in the Bükk plateau in 1999. 3. UPGMA dendrogram of the A. artaxerxes

issekutzi samples constructed on the basis of Nei’s distances.

the two regions and between the two years. Three frequency categories were set: rare

alleles (p<0.05), common alleles with a frequency of 0.05<p<1 and fixed alleles (p=1).

The differences between the two regions in their allele distribution were significant

(x°,=8.04; 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

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)

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.

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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

| | 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

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

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 ).

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.

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.

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.

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:

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.

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.

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

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.

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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

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

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.

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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.

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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

00 05 19 15 20 25 30 0,24 026 0,28

Number of spines per valve Ocelli position

[ex]

Frequency

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.

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.

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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

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

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 ®

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 | ~

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

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

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

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

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

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.

FWD SUM

Bucegi

Itai

—

ofan

atra

Ibula

ioe)

ee)

Ne) D |W|\O BIN N |W} | WIN IN Teele

SLO TN INI DIN | D | NHN] oO u D ni = | OR oo I

NS)

Penninic.N

Pyrenees.E

auphiné

SS)

ie)

ernina

Ne)

OU)

ernese

Ötztal

Penninic.S

Pt.S.Bernh

damello

mberg

Un | Go

wlus0|Z|5/Zl0l0/z)> 2 \w | |O olrlel3lzlole 2!

= © ba O oO |= 0 &

& = = ate À 5

= = NE =

SQ [Où

OO |

Un [Un

(SY)

erlitzen

rtler.N

orway.S

olom.Mar

orway.N

olom.Lat

Grim

No)

ey)

Blolalian iN > —_ IS) un = > IR

AIRJBEIR[OIKRIKR [Dp lolatie OININ/OlEIMIn!lwl]D alo oO

co | aS U ON On DIRIRIDN i) os =

OU)

No)

—

Sweden.N

—

(oe)

—

RP [OS ID ID WIN [W]e Jw OS ID | mm | m = |N WIN IRIS |

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.

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

ern

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).

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

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

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-

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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

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

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)

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.

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)

‘Amata sp. n. | nahe sintensis <sic> 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 <sic> 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

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.

& 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.

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).

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

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,

Daten wie Holotypus (Gen.Präp. deFR 2006/50, Fig. 19, 20 24). 6. Paratypus 9, Fundort wie Fig. 2, je-

Mts., 900 m, 25.03.-05.4.2000, leg. Siniaev & Plutenko (Gen.Präp. deFr 2006/58). 8. Paratypus 9. Iran,

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.

19, 20. S Genital mit Phallus (vergrößert), N-Oman, Westl. Hajar-Gebirge, Oase (Wadi) Al Abyadah

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-

2006/58).

384 DE FREINA: Creatonotos omanirana sp. n.

(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.

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Oman Studies, Special Report 2: 187-216.

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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).

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

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Sesiidae). — Chinese Journal of Entomology 18: 141-165.

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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-

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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.

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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

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

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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

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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

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Nota lepid. 30 (2): 397-406 405

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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

5.x.2005 ER EE

28.vii.2006

6.ix.2006

ae ee

Bez

9 | Mani

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

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

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

€

: 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.

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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.

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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.

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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

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Nota lepidopterologica wird als wissenschaftliche Zeitschrift von der Societas Europaea

Lepidopterologica (SEL) herausgegeben und den Mitgliedern der SEL zugesandt. Autoren, die

Manuskripte für die Publikation in der Nota lepidopterologica einreichen möchten, finden die

jeweils gültigen Autorenrichtlinien auf der Homepage der SEL unter http://www.soceurlep.org.

Der Verkauf von Einzelheften und älteren Jahrgängen von Nota lepidopterologica sowie der Verkauf

der Zeitschrift an Nichtmitglieder erfolgt durch Apollo Books, Kirkeby Sand 19, DK-5771

Stenstrup; e-mail: apollobooks@vip.cybercity.dk. Die Mitgliedschaft bei der SEL steht

Einzelpersonen und Vereinen nach Maßgabe der Satzung offen. Der Aufnahmeantrag ist an den

Mitgliedssekretär Willy De Prins, Dorpstraat 401 B, B-3061 Leefdaal, Belgien; e-mail:

willy.de.prins@telenet.be zu richten. Das Antragsformular ist im Internet auf der Homepage der

SEL unter http://www.soceurlep.org erhältlich. 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.