ISSN 2367-5365 (online) ISSN 0342-7536 (print)

PENSOFT

Volume 38 • Number 2 • 2015 http://nl.pensoft.net

NOTA

LEPIDOPTEROLOGICA

Su

5-4-/

gNf

SOCIETAS EUROPAEA LEPIDOPTEROLOGICA e.V.

http:/ / nl.pensoft.net

Honorary Members

Günter Ebert (DE), Peder Skou (DK), Pamela Gilbert (GB), Barry Goater (GB),

Peter Hëttenschwiler (CH), Rienk de Jong (NL), Willy De Prins (BE), and Zoltân S. Varga (HU)

Council

A journal focused on Palaearctic and General Lepidopterology

Published by Pensoft Publishers on behalf of the Societas Europaea Lepidopterologica e.V.

Editors, Jadranka Rota and Carlos Lopez Vaamonde

Linguistic Editor. Adrian Spalding

Editorial Board. Sven Erlacher (subject editor; Chemnitz, DE), Thomas Fartmann (subject edi-

tor; Münster, DE), Zdenëk F. Fric (subject editor; Ceské Budëjovice, CZ), Axel Hausmann (sub-

ject editor; Munich, DE), Peter Huemer (subject editor; Innsbruck, AT), Lauri Kaila (subject edi-

tor; Helsinki, FI), Ole Karsholt (Copenhagen, DK), Bernard Landry (subject editor; Genève, CH),

Carlos Lopez Vaamonde (subject editor; Orléans, FR), Vazrick Nazari (subject editor; Ottawa,

CA), Erik J. van Nieukerken (subject editor; Leiden, NL), Matthias Nuss (Dresden, DE), Thomas

Schmitt (subject editor; Trier, DE), Wolfgang Speidel (Bonn, DE), Roger Vila (subject editor; Bar-

celona, ES), and Alberto Zilli (subject editor; London, GB).

Cover image: The chalk-hill blue (Polyommatus coridon, Lycaenidae) - the insect of the year for Germany,

Austria, and Switzerland. See T. Schmitt's article on page 107.

President:

Vice-President:

General Secretary:

Treasurer:

Membership Secretary:

Ordinary Council Members:

Erik van Nieukerken (NL)

Thomas Simonsen (DK)

Carlos Lopez Vaamonde (FR)

Robert Trusch (DE)

Hossein Rajaei (DE)

Ximo Baixeras (ES), Andrea Grill (AT), Iva Mihoci (HR),

Pasquale Trematerra (IT), and Boyan Zlatkov (B G)

Nota Lepidopterologica

© Societas Europaea Lepidopterologica (SEL)

ISSN 0342-7536

Printed by Druckhaus Dresden GmbH

All content is Open Access distributed under the terms of the Creative Commons Attribution License,

which permits unrestricted use, distribution, and reproduction in any medium, provided the original author

and source are credited. Authors are thus encouraged to post the pdf files of published papers on their

homepages or elsewhere to expedite distribution.

NOTA LEPIDOPTEROLOGICA

Volume 38 No. 2 • Sofia, 15.12.2015 • ISSN 0342-7536

Thomas Schmitt. Biology and biogeography of the chalk-hill blue

Polyommatus coridon - insect of the year 2015 for Germany,

Austria and Switzerland 107-126

Thomas Sobczyk. Transfer of Pygmaeotinea crisostomella Amsel,

1957 from Tineidae to Psychidae and its taxonomic status

(Lepidoptera) 127-131

Hans Christof Zeller, Peter Huemer. A new species of Micropterix

Hübner, 1825 from the Orobian Alps (Italy) (Lepidoptera,

Micropterigidae) 133-146

John A.M. van Roosmalen, Camiel Doorenweerd. Coleophora

gryphipennella (Hübner, 1796) (Lepidoptera, Coleophoridae) on

Fragaria vesca L. (Rosaceae), a novel host, in the coastal dunes

of The Netherlands 147-155

Joaquin Baixeras. Book Review: Eucosma Hübner of the Contiguous

United States and Canada (Lepidoptera: Tortricidae: Eucosmini)..

157-158

David Agassiz. Book Review: The Notodontidae of South Africa

159-160

Konstantin A. Efetov, Gerhard M. Tarmann, Teodora B. Toshova,

Mitko A. Subchev. Enantiomers of 2-butyl 7Z-dodecenoate are

sex attractants for males of Ads cita mannii (Lederer, 1853),

A. geryon (Hübner, 1813), and Jordanita notata (Zeller, 1847)

(Lepidoptera: Zygaenidae, Procridinae) in Italy 161-169

Nota Lepi. 38(2) 2015: 107-126 | DOI 10.3897/nl.38.4977

Biology and biogeography of the chalk-hill blue Polyommatus coridon -

insect of the year 2015 for Germany, Austria and Switzerland

Thomas Schmitt12

1 Senckenberg German Entomological Institute, Eberswalder Straße 90, D-l 5374 Müncheberg

2 Entomology, Department of Zoology, Institute of Biology, Faculty of Natural Sciences I, Martin Luther University Halle-

Wittenberg, D - 06099 Halle (Saale); Thomas.Schmitt@Senckenberg.de

http://zoobank.org/90176804-4354-45CF-BF91-D768F3B133A3

Received 25 March 2015; accepted 16 May 2015; published: 24 July 2015

Subject Editor: Zdenek Frie.

Abstract. The chalk-hill blue was nominated insect of the year 20 1 5 for Germany, Austria and Switzerland.

The species is strongly associated with base-rich short-turfed swards; the caterpillars feed mainly on horse-

shoe vetch Hippocrepis comosa and show myrmecophilous behaviour. The species is restricted to Europe,

where it is widely distributed but missing from the northern parts of the continent. Polyommatus coridon

survived the last ice age in Mediterranean réfugia in Italy and the Balkan Peninsula from where it colonised

more northern regions postglacially.

The chalk-hill blue -insect of the year 2015 for Germany, Austria and Switzerland

The insect of the year for Germany, Austria and Switzerland has been nominated by a curatorium

every year since 1999. This panel is composed of experts representing a variety of scientific societ-

ies and institutions in these three countries. The goal is to advertise insects in general to a broader

public by focusing on one charismatic species every year. However, the aesthetic appeal of the

selected species is not the most important aspect of the insect of the year. Beauty in this case is just

a vehicle to stimulate interest in the highly remarkable ecology and behaviour of the species. In

particular, the nomination of an insect of the year is intended to enhance awareness of the general

importance of insects and the necessity of their conservation. After nomination of the brimstone

Gonepteryx rhamni in 2002 and the bumet Zygaena carniolica in 2008, the chalk-hill blue Poly-

ommatus coridon (Poda, 1761) (Figure 1) became the third lepidopteran species to be nominated

insect of the year, in 2015. Against this background, an overview of the biology and biogeography

of this butterfly species is presented.

Portrait of the species

The wing pattern of the chalk-hill blue is remarkably dimorphic. The colour of the upperside of the

wings is light blue in the male, often with a silvery hue. The margin of the forewings is brown, but

the width of this varies regionally. A pattern of white circles can be observed in this margin. How-

ever, this patterning is much less frequent in eastern than in western European populations (Schmitt

et al. 2005). On the hindwing, this margin is mostly dissected into brown spots with white margins.

108

Thomas Schmitt: Biology and biogeography of the chalk-hill blue

Figure 1. A freshly emerged male of the chalk-hill blue with open wings sitting on the dry inflorescence of a

knapweed. Spiazzi, Monte Baldo, northern Italy, 02.IX.2013. Photo: Thomas Schmitt.

The dominating colour of the wing upperside of the females is brown. The hindwings in most

cases show orange spots at the margin. These spots are less pronounced on the forewing and fade

out towards the apex. The darker central spot on the forewing is often bordered by a narrow white

line (Figure 2).

The underside of both wings has a characteristic spot pattern, which is similar in males and

females. The dark spots are always surrounded by white lines. The colour of the underside of the

wings is always lighter in males than in females. In males this is a slightly yellowish light brown

or pale grey, whereas in females it is a considerably darker brown (Figure 3).

The larvae have a typically lycaenid shape. Their dominant colour is greenish, but with a some-

what dirty aspect. Two broken yellow lines run dorsally along the entire body. Other yellow mark-

ings can be found laterally (Figure 4).

Closely related species

In central Europe, the male of the chalk-hill blue can hardly be mixed up with any other lycaenid.

A somewhat similar blue is only observed in Polyommatus damon and Polyommatus daphnis ,

but these two species show considerably different wing patterning (Settele et al. 2009). However,

the situation becomes much more complicated in south-western Europe. Polyommatus hispana

is widely distributed in eastern Spain, southern France and a geographically rather limited area

Nota Lepi. 38(2): 107-126

109

Figure 2. Copula of the chalk-hill blue with open wings sitting on the inflorescence of a blue-flowering Er-

yngium species. The differences in wing pattern between female (left) and male (right) are easily recognised.

Spiazzi, Monte Baldo, northern Italy, 02.IX.2013. Photo: Thomas Schmitt.

in north-western Italy. This sibling species is morphologically mostly similar to P coridon, but

allozyme polymorphisms strongly supports it status as a distinct species (Schmitt et al. 2005).

Also Polyommatus albicans , widely distributed in central Iberia, but also in the Atlas Mountains of

Morocco, is difficult to distinguish if relying on its morphology alone; however, the colour of the

wings in general is more whitish than in P coridon.

The species status of Polyommatus philippi , which is restricted to a limited region in north-east-

ern Greece, is highly controversial and it has often been synonymised with P. coridon. Also the

morphologically differing P. coridon caelestissima, which is endemic to a restricted area in eastern

Spain geographically separated from the continuous distribution of the species, has a doubtful tax-

onomic status. In this case, the males have a sky-blue and not a light silvery blue wing colour; it

is still debated whether this taxon represents a well differentiated subspecies or a separate species

(Femândez-Rubio 1991, Tolman and Lewington 1998, Tshikolovets 2011).

The populations in the mountain areas of the islands of Corsica and Sardinia were only discov-

ered in the 1970s and 1980s, respectively, and were described as P. coridon nufrellensis and P.

coridon gennargenti. While the former is mostly accepted as being just a subspecies of P. coridon ,

the latter is frequently assumed to be a good species, as supported for example by genetic analysis

and rearing experiments (Marchi et al. 1996, Jutzeler et al. 2003). Polyommatus corydonius is

110

Thomas Schmitt: Biology and biogeography of the chalk-hill blue

Figure 3. Copula of the chalk-hill blue with closed wings. The differences in wing pattern between female

(left) and male (right) are easily recognised. Csâkvâr, Vertes mountains, western Hungary, 08.VIII.20 14.

Photo: Thomas Schmitt.

another morphologically similar species distributed in Turkey and the Caucasus region, but which

never occurs in sympatry with P. coridon (Tshikolovets 2011).

The female of P. coridon is much more easily misidentified than the male as it is rather similar

to several other species. Even females of Polyommatus bellargus , which is well differentiated in

males by the sky-blue wing colour, are not easy to distinguish. One of the clearest characters of the

female P. coridon is the distribution of the intensity of the brown colour on the underside of the

forewing: the relatively dark brown at the margins becomes gradually lighter to the centre. How-

ever, particularly in old and hence worn females, this cannot be assessed without doubt remaining

in some cases (Tolman and Lewington 1998, Tshikolovets 201 1).

Due to the close relatedness within the genus Polyommatus , natural interspecific hybrids are

frequently observed, as for example between P. coridon and P. bellargus (e.g. de Lesse 1969a).

These hybrids have an intermediate pattern and colouring of the wings and are named Polyomma-

tus x polonus.

Life cycle

The chalk-hill blue is strictly univoltine over most of its range. Males in most regions start emerg-

ing by mid- July. A peak of male emergence can often be observed around 20 July. Early males may

Nota Lepi. 38(2): 107-126

111

Figure 4. The caterpillar of the chalk-hill blue shortly before pupation, on its host plant horse-shoe vetch,

being visited by an ant. Photo: Albert Krebs (E-Pics ETH Zürich). Publication with permission of the copy-

right holders.

occur in the first half of July or even in late June, but are relatively rare (Ebert and Rennwald 1991,

Haag and Eller 2007, Trampenau 2007, Pfeuffer 2013); exceptionally early sightings were record-

ed for Bavaria where butterflies were observed in the wild as early as 1 5 May (Pfeuffer 2013). Such

early males might be predestined for hybridisation with females of the first generation of P bel-

largus. However, even in rather warm and dry years, the normal flight season of P coridon is only

marginally earlier than in normal years. Females mostly emerge one week later than males, hence

showing protandry (Thiel and Meyer 2007), as in many other butterflies and insects in general. The

flight season of the chalk-hill blue is relatively long, but the number of individuals starts to decline

around 20 August in most years. Nevertheless, the species is frequently still to be observed in early

September, but mostly in relatively small numbers. Observations in the second half of September

have only been made in some years. Sightings in early October exist, but are exceptions.

Truly bivoltine populations are only known in a geographically restricted area of south-western

Slovakia in the Vah valley. They were even described as a separate species, Polyommatus slovacus

(Vit’az et al. 1997). However, analyses of allozyme polymorphisms clearly demonstrated that these

bivoltine populations show no genetic differentiation from the nearest univoltine populations; further-

more, no indication of a genetic bottleneck could be detected; therefore, their species status has to be

rejected (Schmitt et al. 2005). In these bivoltine populations of south-western Slovakia, a first genera-

tion by the end of May and in June composed of relatively small numbers of individuals is followed by

a second generation, which normally is much more numerous than the first generation and on the wing

during the normal flight period of univoltine populations of adjoining regions (Schmitt et al. 2005).

112

Thomas Schmitt: Biology and biogeography of the chalk-hill blue

The closely related P. hispana in eastern Spain and southern France (Kudma et al. 2011) also has two

generations a year, but the first one is on the wing earlier than in the Vah valley, while the second one

tends to be later than P. coridon (Tolman & Lewington, 1998). All other closely related species only

have a single generation per year, with adults flying in mid-summer (Tshikolovets 2011).

Populations of P. coridon other than that in the Vah valley are strictly univoltine. Schurian ( 1 989)

reports that it is not possible to obtain the next generation of butterflies without diapause, albeit

under artificial breeding conditions. However, Comont et al. (2009) obtained spontaneous develop-

ment of British provenances, with adults reared in a green house but under conditions similar to the

normal climatic environmental conditions emerging by mid-November. A similar phenomenon also

might explain the few exceptionally early butterflies observed in Bavaria (see above).

The females lay their eggs after being fertilised. Often the eggs are not laid directly on the larval

host plant, but on dry grass or moss nearby. If eggs are laid directly on the larval host plant, this

is not on living parts of the plant like shoots and flowers, but on dry parts (Ebert and Rennwald

1991, Pfeuffer 2000). The young larva, when ready to hatch, remains in the egg shell for the entire

winter (but see Comont et al. 2009). It only hatches in the following spring when the sun heats its

surroundings to an adequate temperature. Hatching thus takes place from mid-March onwards. The

caterpillars then develop relatively quickly, but considerably more slowly than related lycaenid spe-

cies with more than one generation per year. Pupation takes place close to the ground, close to the

host plants, in early June or later. The pupal stage lasts about one month, so that the imagoes hatch

punctually by mid-July, thereby completing one complete life cycle (Ebert and Rennwald 1991).

Habitats

The chalk-hill blue has relatively demanding habitat requirements (Brereton et al. 2008). The most

typical habitats in central Europe are semi-natural calcareous grasslands such as Mesobrometum

(Figure 5) and Xerobrometum, but also the Coelerietum, often in the form of pastured Juniperus

heathlands (Figure 6) (Ebert and Rennwald 1991, Haag and Eller 2007, Pfeuffer 2013). Addition-

ally, the sparse vegetation which rapidly becomes established in abandoned limestone queries and

is often remarkably similar structurally to the above mentioned grasslands can be suitable habitats

too (Lotzing 1990, Benes and Konvicka 2002, Höttinger et al. 2013). At some places, as for exam-

ple in the Nahe valley (Rhineland-Palatinate, Germany) or in northern Bohemia, P. coridon is also

present on base-rich soils over vulcanitic rocks, especially on rocky slopes with sparse vegetation

and on Stipa grasslands. Occurrences on acid or neutral soils are really rare in western Europe (e.g.

the Rotenfels in the Nahe valley, Rhineland-Palatinate, Germany). Occurrences on such soils are

somewhat more common, but still rare, in eastern and south-eastern Europe. This difference might

be also due to the regional differences in the level of larval host plant specialisation with western

European populations being more specific than eastern ones (see below).

In some regions, in particular in eastern Brandenburg (e.g. on the slopes along the river Oder),

large populations are also found on base-rich sandy soils, where the preferred habitats are Stipa

grasslands. However, occurrences on sandy soils are rather rare elsewhere. One of these exceptions

is the Mainzer Sand in Rhineland-Palatinate (Germany).

Large populations of the chalk-hill blue can also be observed in the southern part of the species

range. However, here it retreats from the hot plains and valleys and inhabits the somewhat cooler

hills and mountains. The karst regions of the Balkan Peninsula (Figure 7) and the Apennines in

Italy are regions of southern Europe with a particularly high number of suitable habitats.

NotaLepi. 38(2): 107-126

113

Figure 5. The Mesobrometum represents a characteristic habitat of the chalk-hill blue. The photos show a

typical habitat in spring when the caterpillars are shortly before pupation. Nature Reserve Perfeist near Was-

serliesch (Rhineland-Palatinate, Germany), 10.V.2009. Photos: Thomas Schmitt.

Figure 6. Flower-rich Juniperus heathlands, which belong to the plant association Coelerietum, are among

the best habitats for the chalk-hill blue. The photos show the summer aspect when the butterflies are on the

wing. Bucovica, Durmitor, Montenegro, 31. VII. 2014. Photos: Thomas Schmitt.

In calcareous mountain ranges, the butterflies can be found on highly inaccessible rocky slopes

(Fig. 8) where they inhabit small patches of grassland on steep slopes and within rocky fields. Here,

the imagoes use the existing flowers for nectaring and the larvae feed on their host plants, which

can be found between stones. However, the population densities are generally lower in these hab-

itats than they are in the most suitable ones.

Although the chalk-hill blue is considered to be a moderately thermophilic species, individu-

al-rich populations are observed in the Alps as high as 2000 m asl. However, the numbers of indi-

viduals decreases considerably above 2000 m asl., with only occasional observations documented

from above 2500 m asl. (Schweizerischer Bund fur Naturschutz 1987, Huemer 2004, Stettmer et

al. 2007). It is remarkable that individuals in the Alps are also found in regions with acidic ground

rock where this is intermixed with other more base-rich rock or even limestone. This phenomenon

can for example be observed in the Großglockner and the Matterhorn regions.

114

Thomas Schmitt: Biology and biogeography of the chalk-hill blue

Figure 7. The karst landscapes in the southern European mountain regions have many habitats suitable for the

chalk-hill blue. The habitat here is in the border region of Bulgaria and Greece at an altitude of about 1500 m

asl. Trigrad, southern Bulgaria, 04.VIII.2010. Photo: Thomas Schmitt.

Ecology of the larvae

The caterpillars preferably feed on the horse-shoe vetch Hippocrepis comosa (Figure 9). This spe-

cies apparently is the exclusive larval host plant in the western part of the range. In more eastern

regions from the Balkan Peninsula in the South to Brandenburg and Poland in the North, the lar-

vae also feed on the purple crown vetch Securigera varia (Abb. 10) (Benes and Konvicka 2002,

Höttinger et al. 2013). In some regions without populations of horse-shoe vetch, as in Poland,

eastern Brandenburg and Saxony, purple crown vetch can even be the only host plant of the cat-

erpillars (Trampenau 2007, Buszko and Maslowski 2008, Settele et al. 2009). Some authors also

mention the liquorice milkvetch Astragalus glycyphyllos (Schweizerischer Bund fur Naturschutz

1987, Höttinger et al. 2013); however, this host plant might be generally of minor importance.

Tshikolovets (2011) additionally mentions Hippocrepis glauca, a species restricted to the Mediter-

ranean region. For the populations from Corsica, Hippocrepis conradiae , endemic to this island,

was mentioned as a host plant (Parmentier and Zinszner 2013). Searching for the caterpillars is

relatively easy, as they can frequently be found beneath larger individuals of their host plants, often

in the moss layer (Ebert and Rennwald 1991).

The caterpillars are often accompanied by ants, with which they live in symbiosis (e.g. Schurian

1989, Ebert and Rennwald 1991, Fiedler et al. 1992, Pfeuffer 2000, Asher et al. 2001). The chalk-hill

NotaLepi. 38(2): 107-126

115

Figure 8. The chalk-hill blue is also found on rocky cliffs with limited accessibility. The butterflies use

the small grassy habitat patches, which are scattered over the steep slopes. Bade Herculane, Cema valley,

south-western Romania, 29.VII.2010. Photo: Thomas Schmitt.

blue is hence a myrmecophilous butterfly species, as are many lycaenids (Figure 4). Both partners

benefit from this association (but see Malicky 1969, 1970 for an alternative opinion). The caterpillars

have special glands from which they secret a liquid rich in sugar and amino acids, which is taken up

by the ants (Maschwitz et al. 1975, Daniels et al. 2005). Fiedler and Maschwitz (1988) were even

able to show that the amount of honeydew-like secret production is sufficiently high to “contribute

significantly to the nutrition of the attending ants”.

In return, the ants defend “their” caterpillars against enemies. Thus, they hinder for example

parasitoids such as parasitic wasps and flies from laying their eggs on the larvae. However, the presence

of the ants might also be a general protection against other, more opportunistic, predators. Hence, the

carnivorous ants protect an otherwise suitable prey. However, this protection is far from being perfect;

many caterpillars of the chalk-hill blue are still infested by parasitoids. Nevertheless, even the pupae

are frequently found close to ant nests (Pfeuffer 2000, Asher et al. 2001). This symbiosis has already

been observed for different ant species. In central Europe, relationships with P. coridon are known to

involve the genera Lasius, Tetramorium and Myrmica (e.g. Fiedler 1987, Maschwitz and Fiedler 1988,

Schurian 1989, Ebert and Rennwald 1991, Fiedler et al. 1992, Pfeuffer 2000, 2013).

In general, myrmecophilous behaviour has frequently been observed in many lycaenid species,

with many ant species involved all around the world (e.g. Fiedler et al. 1991, New 1993). The

116

T HOMAS Schmitt: Biology and biogeography of the chalk-hill blue

Figure 9. The horse-shoe vetch Hippocrepis comosa is the most important larval host plant of the chalk-

hill blue. In the western part of its distribution, this plant species is even the only host plant. An inflores-

cence is shown to the left, the typical pinnate leaves to the right. Gant, Vertes mountains, western Hungary,

21. VII. 20 14. Photos: Thomas Schmitt.

Figure 10. The purple crown vetch Securigera varia is an important additional larval host plant from the

Balkan Peninsula to eastern Brandenburg and Poland. In some regions, e.g. Brandenburg, this plant is the sole

larval host plant. Strausberg, eastern Brandenburg, 17.VI.2014. Photo: Thomas Schmitt.

Nota Lepi. 38(2): 107-126

117

protective benefits for the larvae of lycaenids have already been demonstrated in the wild for Glau-

copsyche lygdamus (Pierce and Mead 1981, Pierce and Easteal 1986). Furthermore, higher growth

rates of lycaenid larvae with than without ant attendance have been demonstrated (e.g. Fiedler and

Saam 1994, Wagner and del Rio 1997).

Ecology of the imagoes

The butterflies are much less choosy in the selection of their nectar sources than the caterpillars

are with their host plants. However, the imagoes tend to prefer classic butterfly flowers, mostly of

the plant families Lamiaceae, Asteraceae and Caprifoliaceae, although species of Leguminosae

are also frequently visited for nectaring. The most visited plant genera are apparently Origanum ,

Scabiosa , Knautia and Centaurea (Weidemann 1986, Ebert and Rennwald 1991, Lörtscher et al.

1995, Pfeuffer 2013). The distribution of nectar sources strongly influences the microdistribution

of the butterflies within the habitat (Lörtscher et al. 1995). It seems that butterflies give preference

to violet flower heads, but visits to white and yellow flowers can also frequently be observed (Fig.

11). However, the butterflies also gather at muddy places to take up water (e.g. Pfeuffer 2013) and

even visit excrements and carcasses (e.g. Jones 2000). Like many other lycaenids, the chalk-hill

blue aggregates in sleeping groups in the evening (Figure 12) (e.g. Weidemann 1986). After sunset,

they mostly descend into the more closed vegetation to spend the night.

On sunny days, it is mainly the male individuals that can be observed flying around in the hab-

itat. Females fly less and invest more time in nectaring or just sitting in the vegetation. Therefore,

one might get the impression that fewer females than males are present. However, this impression

is misleading. Rearing of more than a hundred larvae randomly collected in the wild in Oik (south-

ern Eifel, Rhineland-Palatinate, Germany) resulted in a nearly equal number of both sexes (Ashoff

and Schmitt 2014).

The intensive flight activity within habitats leads one intuitively to overestimate the actual dis-

tance of translocations within and between habitats. Thus, a mark-release-recapture experiment

with 2,211 marked butterflies in the Keuperscharren area south-west of Bitburg (southern Eifel,

Rhineland-Palatinate, Germany) demonstrated that the exchange between habitat patches is much

less than one would expect from the high flight activity of the butterflies. Only four individuals

were detected to have moved between two patches separated by a distance of 600 m containing ar-

able fields and intensive grassland with few flowers. Just one butterfly was found to have travelled

a greater distance of 3.7 km (Schmitt et al. 2006). Similar experiments in Hampshire (southern

Britain) showed that only 1 to 2% of the population exchanged between populations one to two km

apart (Asher 2001). In another study in southern Britain, only 12 out of 1,797 marked butterflies

moved between the three analysed habitats which were 350 to 1100 m distant from each other;

however, five individuals even crossed a motorway while changing their habitat (Adey and Wilson

2010). In Luxembourg, none of 304 recaptured individuals (out of 2,085 marked ones) moved

between three sites that were 4.2 to 1 1.2 km apart from each other (Thiel and Meyer 2007). For

P. gennargenti, which might or might not be conspecific with P. coridon (see above), mark-re-

lease-recapture in four habitat patches with a maximum distance of 350 m between them showed

that, with the exception of one rather small patch, emigration rates ranged from 3 to 11% (Casula

et al. 2004).

In Britain, however, P. coridon adults were also found 1 0 to 20 km from known colonies, thus

supporting the idea that a rather small (but ecologically highly important) proportion of the individ-

118

Thomas Schmitt: Biology and biogeography of the chalk-hill blue

Figure 11. The imagoes of the chalk-hill blue visit flowers of different plant species for nectaring, here the

cream scabious Scabiosa ocholeuca (left: Csâkvar, Vértes mountains, western Hungary, 21.VII.2014) and

a yellow-flowering Fabaceae (right: Luka nad Vahom, south-western Slovakia, 09.VIII.2014; note that the

depicted individual is a member of the second generation of a bivoltine population). Photos: Thomas Schmitt.

uals is quite mobile (Asher et al. 2001). Similar findings are also known for Baden-Württemberg

(Ebert and Rennwald 1991). One such particular case is described in more detail by Leverton (2014).

Even within a seemingly homogeneous habitat, the individuals did not mix randomly. Thus in

the study performed in the Keuperscharren near Bitburg, one of the patches with a size of 6.7 ha

was divided into two parts similar in size. No discontinuity in habitat separated these two subplots.

Only 15 individuals out of 703 marked here could be recaptured. However, 13 of these were re-

captured in the sector where they were first marked, and only two changed over to the respective

other subplot (Schmitt et al. 2006). A quite similar finding was made in Luxembourg where only

12.5% of the recaptured individuals within one major calcareous grassland area were detected on

a different part of this habitat (Thiel and Meyer 2007). This clearly demonstrates that individuals

living in a larger habitat plot only use a relatively small fraction of the available habitat for their

daily activities. Lörtscher et al. (1997) also supported this point of view by demonstrating in their

mark-release-recapture experiment in Alpe di Poma (Ticino, Switzerland) that males on average

moved 135 m between two capture events and did not mix randomly within one habitat. This was

even more pronounced in females, which moved significantly less (on average 90 m) than males

and changed between different parts of the habitat less frequently.

Further unpublished studies by the author in the nature reserve Badstube near Mimbach (Blies-

gau, Saarland, Germany) in the year 1998 also support these data. The results of this study, based on

more than 3,000 marked individuals, also showed that no random mixing took place on a seemingly

homogeneous grassland area of some few hectares. Furthermore, a strip of deciduous forest within a

deeply incised valley of approximately 100 m width reduced exchange to some very few individuals.

However, it seems to be a common behavioural pattern of butterflies not to use larger continuous hab-

itats entirely, but only a fraction of these, even if they do not exhibit territorial behaviour. A further

example of this phenomenon is the lesser marbled fritillary Brenthis ino , studied in an assay in which

all individuals were marked individually and positioned by a GPS device (Weyer and Schmitt 2013).

Despite these low exchange rates among habitats, mobility in general seems to be sufficient to

counteract noticeable differentiation among populations, as was demonstrated by genetic analyses;

Nota Lepi. 38(2): 107-126

119

Figure 12. The imagoes of the chalk-hill blue often congregate in sleeping groups in the evening. Many

individuals can assemble at exposed places, to utilise the last sunshine of the day. Gant, Vértes mountains,

western Hungary, 08.VIII.20 14. Photo: Thomas Schmitt.

also see the rare long distance movements cited above (Ebert and Rennwald 1991, Asher et al.

2001). One study of allozyme polymorphisms at 20 different loci included 874 individuals from

22 populations sampled in Rhineland- Palatinate and the Saarland. The Bliesgau, a region with a

remarkable density of viable populations (Schmitt 2002), had no significant genetic differentiation

between the analysed populations. The amount of exchange in this region seems to be sufficient

to completely hinder genetic differentiation between these populations. The calcareous regions of

the western Saarland, the southern Eifel and the central Eifel, which all have a less dense network

of populations than the Bliesgau, also had low rates of genetic differentiation among populations,

and the mean of genetic variance between populations was less than 2% of the entire variance in

all three of them. Hence, genetic differentiation between populations is apparently far from being

critical in terms of conservation. The limited exchange rates seem to be sufficient to bolster against

such a differentiation. Nevertheless, the larger populations expressed higher genetic diversity than

the small ones (Schmitt and Seitz 2002a). Similar genetic findings with low differentiation be-

tween populations are also known for the Göttingen region. Here, it was demonstrated that the

Leine valley with its intensive agriculture genetically separated the populations to the East and

West of it (Krauss et al. 2004).

120

Thomas Schmitt: Biology and biogeography of the chalk-hill blue

These results on population genetics are mostly supported by a classical ecological study from

the Göttingen region which demonstrated that the population density of P. coridon is mainly de-

pendent on the quantity of its larval host plant, in this case H. comosa, but not on the effect of hab-

itat isolation and habitat quality (Krauss et al. 2005). Quite similar results were obtained by Rosin

et al. (201 1) southwest of Krakow (southern Poland); they showed that the best predictors for a

potential habitat being occupied or not are its size and the percentage cover by the host plant, in this

case S. varia. Hence, the chalk-hill blue is mostly dependent on the preservation of large habitats.

Another study from the Göttingen area yielded somewhat different results. No impact of the

habitat area on the butterfly was found, but the importance of habitat connectivity was revealed.

However, connectivity neither impacted the larval host plant occurrence nor the infection rate by

parasitoids (Brückmann et al. 2011). This is also supported by population genetic data from this

region showing that the expected heterozygosity of allozymes decreased with distance to other

populations (Krauss et al. 2004).

Distribution and biogeography

The chalk-hill blue is mostly restricted to Europe (Kudma et al. 2011), and the species is only found

in a very restricted part of western Asia north of the Caspian Sea (Anikin et al. 1 993, Lukhtanov and

Lukhtanov 1994); it is missing in Turkey apart from a record of a single individual (Hesselbarth et

al. 1995). The border of its south-western distribution is located in northern Spain (Garcia-Barros

et al. 2004). In the North-West, P. coridon is found up to south-eastern England (Emmet and Heath

1990; Asher et al. 2001). In Italy, the species can be found throughout the peninsula, but mostly

in the Apennines at higher altitudes. Similarly, the species is widespread in the Balkan Peninsula

where it is found as far south as the Peloponnese, but predominantly in mountain ranges (Pamperis

1997). The northern limit of distribution stretches along the northern margin of the German middle

mountains (Bink 1992), and in the Netherlands it is only found in the extreme South of the country

(Wynhoff et al. 1992). In eastern Germany, the chalk-hill blue is found along the Oder almost as far

north as the Baltic Sea, which is reached in Poland (Buszko 1997, Buszko and Maslowski 2008).

The species is completely missing in Scandinavia (Henriksen and Kreutzer 1982).

Studies of allozyme polymorphisms of several thousands of individuals of the chalk-hill blue al-

lowed the reconstruction of the distribution dynamics over time. In this context, two major genetic

lineages could be distinguished. A western lineage is found in Italy, France, the western and central

Alps and major parts of Germany. An eastern lineage is distributed from the Balkan Peninsula, stretch-

ing over the Carpathian Basin to Brandenburg and Poland (Schmitt and Seitz 2001). Sequencing of

mitochondrial genes also supports the differentiation into at least two major lineages, an eastern and a

western one (Talavera et al. 2013). De Lesse (1969b) distinguished two major groups by their number

of chromosomes, a western group with 87 or 88 chromosomes and an eastern group with 90 to 92

chromosomes. The distribution of these two groups almost perfectly matches the distribution of the

two major allozyme groups, which also can be distinguished by morphological features (see above).

Furthermore, it is only the eastern lineage which commonly uses S. varia as its larval host plant.

Along the contact zone between the lineages, intensive hybridisation was detected in some

regions of the eastern Alps based on the allozyme data set. However, hybrid populations are rath-

er rare north of the Alps, but for example were also found at two localities in Sachsen-Anhalt

(Schmitt and Zimmermann 2012).

NotaLepi. 38(2): 107-126

121

Within the western lineage, populations from the Pyrenees are well distinguished from all oth-

ers. Furthermore, the populations from southern Germany (Baden-Württemberg to southern Thur-

ingia) showed a genetic make-up that distinguished them as an individual group from all other

populations (Schmitt et al. 2002). However, this group also includes populations from the Alps, as

indicated by still unpublished data. Remarkable differences in genetic diversity can be observed

between neighbouring regions in the western lineage, as for example between north-eastern France

and western Germany (Schmitt et al. 2002). A continuous loss of genetic diversity from western

Hungary to Brandenburg was observed for the eastern lineage. After performing a linear regression

for the number of allozyme alleles for these populations, a highly significant correlation (p < 0.00 1 )

was obtained which explained 78% of the regional variation (Schmitt and Seitz 2002b).

These genetic patterns strongly support the existence of ice age réfugia in Italy and in the

Balkan Peninsula at least during the last glacial period. Here, these lineages evolved in allopa-

try. However, the process of evolution was not necessarily restricted to a single glacial period,

but might have taken place during repeated periods of glacial isolation in these réfugia. Ice age

survival in these Mediterranean réfugia has long been postulated (de Lattin 1949) and has been

supported by numerous phylogeographic analyses since then (cf. Schmitt 2007). However, the

differentiation of the populations in the Pyrenees as well as in southern Germany and parts of the

Alps calls for additional refuge areas not resembling the pattern of the classical Mediterranean

refuge areas. Additional glacial réfugia therefore could have existed south of the Pyrenees and

Alps. This assumption is also supported by still unpublished data based on sequences of two mi-

tochondrial loci. Hence, the chalk-hill blue shows biogeographical traits of a species that is much

more cold-tolerant than previously thought and which was thus able, at least additionally, to sur-

vive glacial periods in so called extra-Mediterranean réfugia north of the classical Mediterranean

réfugia (cf. Schmitt and Varga 2012).

These genetic analyses also allow a relatively detailed reconstruction of the postglacial range ex-

pansion. The Adriato-Mediterranean lineage evolving in the Italian peninsula most probably had a

north-western distribution limit during the last ice age located in north-western Italy or south-eastern

France. Starting here, this lineage could colonise the regions northwards to Lorraine without genetic

erosion, with the Rhone valley most probably representing an important expansion corridor. During

the subsequent expansion into the western German region, remarkable genetic impoverishment has

taken place. This might be explained by the, if compared to north-eastern France, considerably less

favourable environmental conditions for this species in western Germany resulting in remarkable

genetic erosions in the wake of the colonisation of this region (Schmitt et al. 2002).

During glacial conditions, the Ponto-Mediterranean lineage which survived the last ice age in

the Balkan Peninsula most probably had its north-western distribution edge in the region of the

northern Dalmatian Coast. Starting there, postglacial range expansion reached as far north as east-

ern Brandenburg. During a first advance, the species was able to colonise to the forelands of the

eastern Alps. Here, the route of further expansion bifurcated. An eastern branch ran along the

Hungarian middle mountains to eastern Slovakia. A western branch reached Moravia via the Porta

Hungarica (lowland area between the north-eastern Alps and the south-western foothills of the

Tatra Mountains). By a westwards advance, the species colonised the limestone regions of the

Czech Republic. This advance was stopped by the mountain ranges between the Czech Republic

and Germany, which were too cold and where soils were too acid to permit the survival of the spe-

cies; on the other side of these mountains, all populations derive from the Adriato-Mediterranean

122

Thomas Schmitt: Biology and biogeography of the chalk-hill blue

region (Schmitt and Zimmermann 2012). The further expansion out of Moravia northwards most

probably first followed the river Vistula, and then in a westerly direction along the Torun-Eber-

swald glacial valley finally reaching eastern Brandenburg; most parts of the Odra region in Poland

have rather few suitable habitats available for P. coridon so that this putative expansion corridor

has to be considered less likely. The linear decline of the number of alleles from western Hungary

to eastern Brandenburg implies a constant loss of genetic diversity in the Ponto-Mediterranean

lineage during its postglacial range expansion (Schmitt and Seitz 2002b).

Acknowledgement

I thank Andrew Liston (Senckenberg German Entomological Institute, Müncheberg) for correcting my Eng-

lish. The constructive comments of Vladimir Hula (Mendel University, Bmo) and another anonymous referee

on a previous version of the manuscript are greatly acknowledged.

References

Adey JP, Wilson SF (2010) The impact of the M40 motorway on populations of Chalkhill Blue Lysandra

coridon and Silver-spotted skipper Hesperia comma at Aston Rowant National Nature Reserve. British

Journal of Entomology and Natural History 23: 7-19.

Anikin W, Sachkov SA, Zolotuhin VV (1993) „Fauna lepidopteriologica Volgo-Uraliensis“ 150 years later:

changes and additions, part 1. Rhopalocera (Insecta, Lepidoptera). Atalanta 24: 89-120.

Asher J, Warren M, Fox R, Harding P, Jeffcoate G, Jeffcoate S (2001) The millennium atlas of butterflies in

Britain and Ireland. Oxford University Press, Oxford, 433 pp.

Ashoff M, Schmitt T (2014) Are different allozyme genotypes of the butterfly Polyommatus coridon adapted

to resist cold and heat shocks? Annales Zoologici Fennici 51: 413^122. doi: 10.5735/086.051.0502

Benes J, Konvicka M (2002) Butterflies of the Czech Republic: Distribution and conservation I. Vydal SOM,

Praha, 478 pp.

Bink FA (1992) Ecologische Atlas van de Dagvlinders van Noordwest-Europa. Schuyt Co. Uitgevers en Im-

porteurs, Haarlem, 512 pp.

Brereton TM, Warren MS, Roy DB, Stewart K (2008) The changing status of the Chalkhill Blue butterfly

Polyommatus coridon in the UK: the impacts of conservation policies and environmental factors. Journal

of Insect Conservation 12: 629-638. doi: 10. 1007/s 1084 1-007-9099-0

Brückmann SV, Krauss J, van Achterberg C, Steffan-Dewenter I (2011) The impact of habitat fragmentation

on trophic interactions of the monophagous butterfly Polyommatus coridon. Journal of Insect Conserva-

tion 15: 707-714. doi: 10.1 007/s 1 084 1 -0 1 0-9370-7

Buszko J (1997) Atlas rozmieszczenia motyli dziennych w Polsce (Lepidoptera: Papilionidae, Hesperiidae).

Edycja Turpress, Torun, 170 pp.

Buszko J, Maslowski J (2008) Motyle dzienne Polski. Wydawnictwo Koliber, Nowy S^cz, 274 pp.

Casula P, Scanu D, Cmjar R, Grill A, Marchi A (2004) The fragmented population structure of the Sardi-

nian chalk hill blue butterfly (Lepidoptera, Lycaenidae). Journal for Nature Conservation 12: 77-83. doi:

10.101 6/j .jnc.2003 .09.00 1

Comont RF, McCracken M, Hesketh H, Hails RS, Roy HE (2009) Bivoltine blues: a second generation in the

British chalkhill blue Polyommatus coridon (Lepidoptera: Lycaenidae). British Journal of Entomology

and Natural History 22: 71-72.

Daniels H, Gottsberger G, Fiedler K (2005) Nutrient composition of larval nectar secretions from three species of

myrmecophilous butterflies. Journal of Chemical Ecology 31: 2805-2821. doi: 10.1007/sl0886-005-8395-y

Nota Lepi. 38(2): 107-126

123

De Lattin G (1949) Beiträge zur Zoogeographie des Mittelmeergebietes. Verhandlungen der deutschen Zoo-

logischen Gesellschaft, Kiel, 143-1 5 1 .

De Lesse H (1969a) Les hybrides naturels entre Lysandra coridon Poda et L. bellargus Rott. Alexanor 6: 73-82.

De Lesse H (1969b) Les nombre de chromosomes dans le groupe de Lysandra coridon [Lep. Lycaenidae].

Annales de la Société Entomologique de France N.S. 5: 469-522.

Ebert G, Rennwald E (Eds) (1991) Die Schmetterlinge Baden- Württembergs, Volume 2. Eugen Ulmer, Stutt-

gart, 535 pp.

Emmet AM, Heath J (Eds) (1990) The butterflies of Great Britain and Ireland. - Vol 7 of ’’The moths and

butterflies of Great Britain and Ireland”. Harley Books, Colchester, 370 pp.

Femândez-Rubio F (1991) Guia de mariposas diumas de la Peninsular Ibérica, Baléares, Canarias, Azores

y Madeira (Libytheidae, Nymphalidae, Riodinidae y Lycaenidae). Ediciones Pirâmide, Madrid, 418 pp.

Fiedler K (1987) Tetramorium caespitum (LINNEAEUS, 1758) (Hymenoptera: Formicidae), eine weitere

mit Polyommatus coridon (PODA, 1761 ) (Lepidoptera: Lycaenidae) vergesellschaftete Ameisenart. Nach-

richten des entomologischen Verein Apollo N.S. 8: 60.

Fiedler K (1991) Zoogeographische Aspekte der myrmekophilen Beziehungen der Bläulinge (Lepidoptera:

Lycaenidae). Verhandlungen der deutschen zoologischen Gesellschaft 84: 502.

Fiedler K, Maschwitz U (1988) Functional analysis of the myrmecophilous relationships between ants (Hy-

menoptera: Formicidae) and lycaenids (Lepidoptera: Lycaenidae). Lycaenid larvae as trophobiotic part-

ners of ants - a quantitative approach. Oecologia 75: 204-206. doi: 10.1007/BF00378598

Fiedler K, Saarn C (1994) Does ant-attendance influence development in 5 European Lycaenidae butterfly

species? (Lepidoptera). Nota lepidopterologica 17: 5-24

Fiedler K, Schurian KG, Seufert P (1992) Neue Beobachtungen zu Ameisenassoziationen europäischer Bläu-

lingsraupen (Lepidoptera: Lycaenidae). Mitteilungen internationaler entomologischer Verein 17: 121-130.

Garcia-Barros E, Munguira ML, Martin Cano J, Romo Benito H, Garcia-Pereira P, Maravalhas ES (2004)

Atlas de las mariposas diumas de la Peninsula Ibérica e islas Baléares (Lepidoptera: Papilionoidea Hespe-

rioidea). Monografias Sociedad Entomolögica Aragonesa 11, 228 pp.

Haag M, Eller O (2007) Silbergrüner Bläuling - Polyommatus coridon (Poda, 1761). In: Schulte T, Eller O,

Niehuis M, Rennwald E (Eds) Die Tagfalter der Pfalz. GNOR-Eigenverlag, Landau, 377-383.

Henriksen HJ, Kreutzer IB (1982) The butterflies of Scandinavia in Nature. Skandinavisk Bogforlag, Odense,

215 pp.

Hesselbarth G, van Oorschot H, Wagener S (1995) Die Tagfalter der Türkei und angrenzender Länder, Band

1 . Selbstverlag Sigbert Wagener, Bocholt, 754 pp.

Höttinger H, Pendl M, Wiemers M, Pospisil A (2013) Insekten in Wien Tagfalter. Österreichische Gesell-

schaft für Entomofaunistik, Wien, 349 pp.

Huemer P (2004) Die Tagfalter Südtirols. Folio Verlag, Wien, 232 pp.

Kudma O, Harpke A, Lux K, Pennersdorfer J, Schweiger O, Settele J, Wiemers M (2011) Distribution atlas

of butterflies in Europe. Gesellschaft für Schmetterlingsschutz, Halle, 576 pp.

Krauss J, Schmitt T, Seitz A, Steffan-Dewenter I, Tschamtke T (2004) Effects of habitat fragmentation on

the genetic structure of the monophagous butterfly Polyommatus coridon along its northern range margin.

Molecular Ecology 13: 311-320. doi: 10.1046/j.l365-294X.2003.02072.x

Krauss J, Steffan-Dewenter I, Müller CB, Tschamtke T (2005) Relative importance of resource quantity, iso-

lation and habitat quality for landscape distribution of a monophagous butterfly. Ecography 28: 465—474.

doi: 10. 1 1 1 l/j.0906-7590.2005.04201 .x

Leverton R (2014) Dispersive behavior by male chalk hill blue Polyommatus coridon (Poda) (Lep.: Lycaenidae).

Entomologist's Record and Journal of Variation 126: 48.

Lörtscher M, Erhardt A, Zettel J (1995) Microdistribution of butterflies in a mosaic-like habitat - the role of

nectar sources. Ecography 18: 15-26. doi: 1 0.1 1 1 1/j.l 600-0587. 1995.tb00 11 5.x

124

Thomas Schmitt: Biology and biogeography of the chalk-hill blue

Lörtscher M, Erhardt A, Zettel J (1997) Local movement patterns of three common grassland butterflies in

a traditionally managed landscape. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 70:

43-55.

Lotzing K (1990) Massenvorkommen von Ly sandra coridon PODA in einem aufgelassenen Kalksteintagebau

im Kreis Staßfurt (Lep., Lycaenidae). Entomologische Nachrichten und Berichte 34: 40.

Lukhtanov V, Lukhtanov A (1994) Die Tagfalter Nordwestasiens. Herbipoliana 3, Verlag Ulf Eitschberger,

Marktleuthen, 440 pp.

Jutzeler D, Casula P, Gascoigne-Pees M, Grill A, Leigheb G (2003) Confirmation du statut spécifique de

Polyommatus gennargenti (Leigheb, 1987) de Sardaigne comparé à Polymmatus coridon (Poda, 1761)

de la région de Schaffhouse (CH) par élevage parallèle (Lepidoptera: Lycaenidae). Linneana Belgica 19:

109-118.

Malicky H (1969) Versuch einer Analyse der ökologischen Beziehungen zwischen Lycaeniden (Lepidoptera)

und Formiciden (Hymenoptera). Tijdschrift voor Entomologie 1 12: 213-298.

Malicky H (1970) New aspects of the association between lycaenid larvae (Lycaenidae) and ants (Formicidae,

Hymenoptera). Journal of the Lepidopterists’ Society 24: 190-202.

Marchi A, Addis G, Hermosa V, Cmjar RE (1996) Genetic divergence and evolution of Polyommatus coridon

gennargenti (Lepidoptera, Lycaenidae) in Sardinia. Heredity 77: 16-22. doi: 10.1038/hdy.l996.103

Maschwitz U, Fiedler K (1988) Koexistenz, Symbiose, Parasitismus Erfolgsstrategie der Bläulinge. Spektrum

der Wissenschaften 5/1988: 56-66.

Maschwitz U, Wüst M, Schurian K (1975) Bläulingsraupen als Zuckerlieferanten für Ameisen. Oecologia 18:

17-21. doi: 10.1007/BF00350631

New TR (Ed.) (1993) Conservation biology of Lycaenidae (Butterflies). International Union for Conservation of

Nature and Natural Resources (IUCN), Gland, 173 pp.

Pamperis L (1997) The butterflies of Greece. Bastas-Plessas Publications, Athens, 766 pp.

Parmentier L, Zinszner E (2013) Contribution to the knowledge of two endemic Corsican butterflies, Poly-

ommatus coridon nufrellensis and Plebejus bellieri (Lepidoptera: Lycaenidae) in relation to Hippocrepis

conradiae : first evidence as hostplant, discovery of a new locality and update on distribution, biology and

conservation. Phegea 41: 26^41 .

Pfeuffer E (2000) Zur Ökologie der präimaginalstadien des Himmelblauen Bläulings (Lysandra bellargus

Rottemburg 1775) und des Silbergrünen Bläulings (Lysandra coridon Poda 1761), unter besonderer

Berücksichtigung der Myrmekophilie. Berichte des Naturwissenschaftlichen Vereins für Schwaben e. V.

104: 72-98.

Pfeuffer E (2013) Silbergrüner Bläuling Polyommatus coridon (Poda, 1761). In: Bräu M, Bolz R, Kolbeck H,

Nummer A, Voith J, Wolf W (Eds) Tagfalter in Bayern. Eugen Ulmer, Stuttgart, 319-321.

Pierce NE, Mead PS (1981) Parasitoids as selective agents in the symbiosis between lycaenid butterfly larvae

and ants. Science 211:11 85-1 1 87. doi: 1 0. 1 126/science.2 1 1 .4487. 1 1 85

Pierce NE, Easteal S (1986) The selective advantage of attendant ants for the larvae of a lycaenid butterfly,

Glaucopsyche lygdamus. The Journal of Animal Ecology 55: 451-462. doi: 10.2307/4730

Rosin ZM, Skorka P, Lenda M, Moron D, Sparks TH, Tryjanowski P (2011) Increasing patch area, proximity

of human settlement and larval food plants positively affect the occurrence and local population size of

the habitat specialist butterfly Polyommatus coridon (Lepidoptera: Lycaenidae) in fragmented calcareous

grasslands. European Journal of Entomology 108: 99-106. doi: 1 0. 1441 l/eje.201 1.01 3

Schmitt T (2002) Biogeographie und regionale populationsgenetische Strukturierung von Polyommatus co-

ridon (Lycaenidae: Lepidoptera) im Saarland und in Rheinland-Pfalz. Abhandlungen der Delattinia 28:

117-130.

Schmitt T (2007) Molecular Biogeography of Europe: Pleistocene cycles and postglacial trends. Frontiers in

Zoology 4: 11. doi: 10.1186/1742-9994-4-11

Nota Lepi. 38(2): 107-126

125

Schmitt T, Seitz A (2001 ) Allozyme variation in Polyommatus coridon (Lepidoptera: Lycaenidae): identifica-

tion of ice-age réfugia and reconstruction of post-glacial expansion. Journal of Biogeography 28: 1 1 29—

1136. doi: 10.1046/j. 1365-2699.200 1.0062 1.x

Schmitt T, Seitz A (2002a) Influence of habitat fragmentation on the genetic structure of Polyommatus coridon

(Lepidoptera: Lycaenidae): implications for conservation. Biological Conservation 107: 291-297. doi:

10.101 6/S0006-3207(02)00066-6

Schmitt T, Seitz A (2002b) Postglacial distribution area expansion of Polyommatus coridon (Lepidoptera:

Lycaenidae) from its Ponto-Mediterranean glacial refugium. Heredity 89: 20-26.

Schmitt T, Varga Z (2012) Extra-Mediterranean réfugia: The rule and not the exception? Frontiers in Zoology 9:

22. doi: 10.1186/1742-9994-9-22

Schmitt T, Zimmermann M (2012) To hybridize or not to hybridize: what separates two genetic lineages of

the Chalk-hill Blue Polyommatus coridon (Lycaenidae, Lepidoptera) along their secondary contact zone

throughout eastern Central Europe? Journal of Zoological Systematics and Evolutionary Research 50:

106-115. doi: 10. 1 1 1 1/j. 1439-0469.20 1 1 .00644.x

Schmitt T, Gießl A, Seitz A (2002) Postglacial colonisation of western Central Europe by Polyommatus

coridon (Poda 1761) (Lepidoptera: Lycaenidae): evidence from population genetics. Heredity 88: 26-34.

doi: 1 0. 1 03 8/sj .hdy.6800003

Schmitt T, Varga Z, Seitz A (2005) Are Polyommatus hispana and Polyommatus slovacus bivoltine Polyom-

matus coridon (Lepidoptera: Lycaenidae)? The discriminatory value of genetics in taxonomy. Organisms,

Diversity Evolution 5: 297-307. doi: 10.1016/j.ode.2005.01.001

Schmitt T, Habel JC, Besold J, Becker T, Johnen L, Knolle M, Rzepecki A, Schultze J, Zapp A (2006) The

Chalk-hill Blue Polyommatus coridon (Lycaenidae, Lepidoptera) in a highly fragmented landscape: How

sedentary is a sedentary butterfly? Journal of Insect Conservation 10: 311-316. doi: 10.1007/sl0841-006-

9000-6

Schurian KG (1989) Revision der Lysandra-Gmppe des Genus Polyommatus LATR. (Lepidoptera: Lycaenidae).

Neue Entomologische Nachrichten 24: 1-181.

Schweizerischer Bund für Naturschutz (Ed.) (1987) Tagfalter und ihre Lebensräume. Arten • Gefährdung •

Schutz. Schweiz und angrenzende Gebiete. Volume 1. Fotorotar, Egg, 516 pp.

Settele J, Steiner R, Reinhardt R, Feldmann R, Hermann G (2009) Schmetterlinge Die Tagfalter Deutschlands.

2. aktualisierte Auflage. Ulmer, Stuttgart, 256 pp.

Stettmer C, Bräu M, Gros P, Wanninger O (2007) Die Tagfalter Bayerns und Österreichs. 2nd edition. Bayerische

Akademie für Naturschutz und Landschaftspflege, Laufen, 248 pp.

Talavera G, Lukhtanov VA, Pieppel L, Pierce NE, Vila R (2013) In the shadow of phylogenetic uncertainty:

The recent diversification of Lysandra butterflies through chromosomal change. Molecular Phylogenetics

and Evolution 69: 469—478. doi: 10. 101 6/j.ympev.201 3.08.004

Thiel G, Meyer M (2007) Suivi des populations de Polyommatus coridon (Poda, 1761) sur trois pelouses

sèches calcicoles au Luxembourg (Lepidoptera, Lycaenidae). Bulletin de la Société des naturalistes luxem-

bourgeois 108: 55-62.

Tolman T, Lewington R (1998) Die Tagfalter Europas und Nordwestafrikas. Franckh-Kosmos, Stuttgart.

319 pp.

Trampenau M (2007) Polyommatus coridon (Poda, 1761) Silbergrüner Bläuling. In: Reinhardt R, Sbieschne

H, Settele J, Fischer U, Fiedler G (Eds) Tagfalter von Sachsen. Beiträge zur Insektenfauna Sachsens, Band

6, Dresden, 308-313.

Tshikolovets V (2011) Butterflies of Europe and the Mediterranean area. Vadim Tshikolovets, Pardubice,

544 pp.

Vit’az L, Bâlint Z, Zitnan D (1997) Polyommatus slovacus sp. nov (Lepidoptera, Lycaenidae): the bivoltine

relative of Polyommatus coridon in Slovakia. Entomological Problems 28: 1-8.

126

Thomas Schmitt: Biology and biogeography of the chalk-hill blue

Wagner D, del Rio CM (1997) Experimental tests of the mechanism for ant-enhanced growth in an ant-tended

lycaenid butterfly. Oecologia 1 12: 424-429. doi: 10.1007/s004420050328

Weidemann H-J (1986) Tagfalter. Band 1. Verlag J. Neumann-Neudamm, Melsungen, 288 pp.

Weyer J, Schmitt T (2013) Knowing the way home: strong philopatry of a highly mobile insect species, Brenthis

ino. Journal of Insect Conservation 17: 1 197-1208. doi: 10. 1007/s 1084 1-0 13-960 1-9

Wynhoff I, van Swaay C, van der Made J (1992) Dagvlinders van de Benelux, 2nd edition. Stichting Uitgeverij

Koninklijke Nederlandse Natuurhistorische Vereinigung De Vlinderstichting, 187 pp.

Nota Lepi. 38(2) 2015: 127-131 | DOI 10.3897/nl.38.5090

Transfer of Pygmaeotinea crisostomella Amsel, 1957 from Tineidae to

Psychidae and its taxonomie status (Lepidoptera)

Thomas Sobczyk1

1 Diesterwegstraße 28, D-02977 Hoyerswerda, Germany, email: ThomasSobczyk@aol.com

http://zoobank.org/9E0B96CC-B486-4B01-849D-D560DF146DFE

Received 14 April 2015; accepted 13 July 2015; published: 7 August 2015

Subject Editor: Jadranka Rota.

Abstract. Pygmaeotinea Amsel, 1957, syn. n. is transferred from Tineidae to Psychidae and regarded as

congeneric with Eumasia Chrétien, 1904. The type species of Pygmaeotinea is combined with Eumasia as

Eumasia crisostomella Amsel, 1957, comb. n. Adult male and female of this species are redescribed and illus-

trated for the first time. Eumasia crisostomella Amsel, 1957 is only known from the type locality in Portugal

at Singeverga and one of three Eumasia species from Iberian Peninsula.

Zusammenfassung. Pygmaeotinea Amsel, 1957, syn. n. wird von den Tineidae zu den Psychidae transferriert

und ist congenerisch mit Eumasia Chrétien, 1904. Die Typusart von Pygmaeotinea wird mit Eumasia als

Eumasia crisostomella Amsel, 1957, comb. n. kombiniert. Adulte Männchen und Weibchen dieser Art werden

erstmals abgebildet und die Beschreibung durch weitere Merkmale ergänzt. Eumasia crisostomella Amsel,

1957 ist nur von der Typenlokalität bekannt und eine von drei Eumasia- Arten der Iberischen Halbinsel.

Introduction

The monotypic Pygmaeotinea Amsel, 1957 was described within Tineidae. Its type-species,

P. crisostomella Amsel, 1957 is only known from the type specimens and no further specimens

have been reported thus far.

Amsel (1957) noted that the relationship of Pygmaeotinea crisostomella to other Tineidae is

unknown. Karsholt and Razowski (1996) placed the genera Eumasia and Pygmaeotinea together

with Apterona Millière, 1857 within Apteronini (Psychidae, Oiketicinae). Sauter and Hättenschwiler

(1999) argued that this placement was done by the editors and does not reflect the opinion of the

authors. They do not consider the first genus as belonging to Apteronini. Unfortunately, the original

description does not allow us to judge the systematic placement of Pygmaeotinea , except that the in-

formation available suggests that it does not belong to Psychidae (Sauter and Hättenschwiler 1999).

Recently, Pygmaeotinea was re-transferred to Tineidae by Sobczyk (2011), but still provision-

ally. Even more recently, Gaedike (Bonn) found the type-series at the Staatliches Museum für

Naturkunde Karlsruhe (SMNK). As an expert of Tineidae, after having examined the specimens,

he came to the conclusion that they do not belong to Tineidae. Subsequently, he kindly arranged

the loan of the type specimens to me.

128

Thomas Sobczyk: Transfer of Pygmaeotinea crisostomella Amsel, 1957...

Systematic part

Eumasia crisostomella Amsel, 1957, comb. n.

Figs 1-6

Examined material. Holotype S “vi [1]950 / Singeverga” back cover (handwritten Amsel) “Portugal / coll. Monteiro”,

“GU 3199”, red label “ Pygmaetinea crisostomella”, “coll. SMNK”, genital slide with same information and additional

“Holotypus”; Paratype 1 $ “Singeverga / vi.1953” back cover (handwritten Amsel) “Portugal / coll. Monteiro”, genital

preparation 3198, red label “Allotypus Ç, leg. H. Amsel”, “coll. SMNK”.

Citation of original description by Amsel.

(. Pygmaeotinea )

Fühler des S kurz bewimpert, bis etwa V2 Costa reichend. Kopfhaare abstehend. Vflg.-Geäder:

Zelle offen, nur 3 Radialadern: r2+r3 und r3+r4 zu je einer Ader zusammengefallen; mit ml und

r4+r5 gestielt; m2 fehlend; ax ohne Wurzelschlinge. Hflg. -Geäder: rr und ml gestielt, die Spitze

umgreifend, m2 fehlend. Axillaradern stark reduziert. Genitalapparat des f: Uncus abgerundet,

Gnathos fehlend, Vinculum (Saccus) zugespitzt, Valven in einen basalen und einen terminalen Teil

gegliedert, ohne sonstige Strukturen.

( crisostomella )

Spw. 7 mm. Kopfhaare gelblich, Palpen bräunlich. Fühler des $ bis V2 Costa reichend, sehr kurz

bewimpert, fast pubscent. Zwischen oberen Außenrand und der Fühlerwurzel ein großer quecksil-

berartiger Fleck. Vfgl. Gelblich. Ein großer bräunlicher Fleck am Innenrand bei V2, weitere Flecke

im apikalen Flügelteil. Die Art ist durch den großen, fast viereckigen Fleck am Innenrand bei V2 gut

charakterisiert. An der Costa stehen einige bräunliche Schuppen, im äußeren Drittel der Flügel

häufen sich diese zu Flecken. Fransen graubraun. Hflg. Nur wenig schmaler als die Vflg. Fransen

etwas länger als der Flügel breit ist. Beine ungeringelt. Genitalapparat des f: Uncus abgerun-

det, Gnathos fehlend. Vinculum ziemlich lang, zugespitzt. Valven aus zwei Teilen bestehend, einen

äußeren, nicht strukturierten und einen inneren, der deutlich gegenüber dem Außenteil abgesetzt

ist. Aedoeagus leicht gebogen, ziemlich lang, ohne Cornuti. Genitalpräparat 3199 (...) Einige

Stücke wurden aus abgeflachten Säcken gezüchtet, die 7 mm lang mit Sandkörnchen besetzt sind.

Redescription crisostomella. Forewing length (with fringes) 3.5-4 mm, wingspan 7-8

mm. Labial palpi tripartite, median segment of double width and equal length as distal seg-

ment. Both segments covered with brown, ventrally directed hair-like scales. Antennal seg-

ments round, covered with short setae, dorsally with a semi-circular fan-like arrangement

of broad, dark brown scales. The angle of this fan is 3CM0° (see description of measurement

by Hättenschwiler 1998). Forelegs with bristle brush (without epiphysis). Forewing sca-

les long-oval, rounded distally, usually four- to six-pointed. Forewing ground colour yello-

wish, with a distinct dark brown spot at V2 of posterior margin and partly the anterior margin

with some very narrow dark brown spots; distal third significantly spotted and converging par-

tially to transverse lines. Fringe gray, hair-like, bi- or tricuspid. Flindwings uniformly dark gray.

Male genitalia (genital preparation no. 3199 by Amsel): Total length 0.5 mm, very wea-

kly sclerotized (perhaps an artifact due to excessive maceration). In ventral view, almost

three times as long as wide. Valva extending beyond posterior margin of tegumen, wea-

kly sclerotized distally with fine setae. Phallus 0.4 mm long, tubular, almost straight.

Nota Lepi. 38(2): 127-131

129

Figures 1-6. Eumasia crisostomella Amsel, 1957. 1. S Holotype, Portugal, Singeverga, vi. [1]950. 2. Holotype

genital preparation 3199. 3. Detail antennae. 4. Foreleg. 5. $ Paratype $ same data, but vi.1953. 6. $ Paratype

genital preparation 3198. Scale bar: 1 mm (1, 5), 0.1 mm (2, 3), 0.5 mm (4, 6).

130

Thomas Sobczyk: Transfer of Pygmaeotinea crisostomella Amsel, 1957...

Female (differences to males). Antenna filiform, wingspan 8 mm. Wing pattern as in the male, but

missing dark spots at the anterior margin of basal 2/3 of the forewings. Abdominal segment eight

with dense, wavy hairs. Female genitalia (genital preparation no. 3198 by Amsel): 2.0 mm total

length. Oviscapt with fine setae distally, supported by three pairs of apophyses. Posterior apophy-

ses with a length of 1.9 mm, almost as long as the entire genitalia, anterior apophyses 0.85 mm

long, forked distally, distal edge of antevaginal plate concave, covered with fine, distally directed

spines. Ovipositor also with a third pair of 0.5-mm-long apophyses.

Diagnosis. E. crisostomella Amsel, 1957, comb. n. differs from E. parietariella (Heydenreich,

1851) from Central and South Europe and Eumasia brunella Hättenschwiler, 1998 from the Iberian

Peninsula by the forewing pattern. Forewings of the latter two species are also covered with dark

spots basally, but the dark spots of E. crisostomella are concentrated on distal third. On the basal

2/3 there are two or three small partial dark spots on the anterior margin. Most conspicuous is a

large, almost square dark spot in the middle of the posterior margin. Hindwings of E. parietariella

are light cream, of E. brunella brown and of E. crisostomella grey.

Remarks. The original description of Pygmaeotinea neither contains information about a dif-

ferentiation from other genera nor why the description of a new genus was considered necessary.

The venation (figure 5, Amsel 1957) and antennal and genital structures are clearly consistent

with typical structures of Eumasia. A diagnosis for the genus is given by Hättenschwiler (1998).

Particularly noteworthy is the scalation of the male antennae with a dorsally semi-circular fan-

like arrangement of broad scales. Each antennal segment has a dorsal compartment of broader

scales. Thus, Pygmaeotinea Amsel, 1957, syn. n. is identical to Eumasia Chrétien, 1904 and is

synonymized here with it and its type species transferred to Eumasia , as Eumasia crisostomella

Amsel, 1957, comb. n.

The label data do not completely match with information provided in the original description.

Holotype $ denotes VI. [1]950 as date. There was no label on the pinned specimen that indicated

it as the holotype. That information was only present on the genital slide belonging to this

specimen and the slide number 3199 is present on both, the genitalia slide and on the pinned

holotype specimen.

In the description the date for the female (allotype) is listed as vi.1950; however, the specimen

label reads ‘vi.1953’.

Acknowledgements

I thank Reinhard Gaedike (Bonn), who inspired the study, and Peter Hättenschwiler, who provided important

information. I thank Robert Trusch and Michael Falkenberg (both State Museum of Natural History, Karlsru-

he, Germany) for the loan of the type material of E. crisostomella. Last but not least, I thank Matthias Nuss

(Senckenberg Museum of Zoology Dresden) for comments on the manuscript.

References

Amsel HG (1957) Neue Microtineiden aus Portugal (Lepidoptera, Tineidae). Beiträge zur naturkundlichen

Forschung in Südwestdeutschland 16: 30-33.

Hättenschwiler P (1998) Neue Eumasia Arten aus Mittelspanien und den Malediven und einige Ergänzungen

zur Kenntnis der Gattung Eumasia (Psychidae). Nota lepidopterologica 21(4): 264-282.

Nota Lepi. 38(2): 127-131

131

Sauter W, Hättenschwiler P ( 1 99 1 ) Zum System der palaearkti sehen Psychiden (Lep., Psychidae) 1 . Teil: Liste

der palaearktischen Arten. Nota lepidopterologica 14 (1): 69-89.

Sauter W, Hättenschwiler P (1996) Psychidae. In: Karsholt O, Nielsen ES. The Lepidoptera of Europe. A

distributional Checklist. Apollo Books, Stenstrup, 39-46.

Sauter W, Hättenschwiler P (1999) Zum System der palaearktischen Psychiden (Lep., Psychidae). 2. Teil:

Bestimmungsschlüssel für die Gattungen. Nota lepidopterologica 22(4): 262-195.

Sobczyk T (2011) Psychidae. World Catalogue of Insects 10. Apollo Books Stenstrup, 467 pp.

Nota Lepi. 38(2) 2015: 133-146 | DOI 10.3897/nl.38.5058

A new species of Micropterix Hübner, 1825 from the Orobian Alps (Italy)

(Lepidoptera, Micropterigidae)

Hans Christof Zeller1, Peter Huemer2

1 Forsthubfeld 14, A-5303 Thalgau, Austria; christof.zeller@gmx.net

2 Tiroler Landesmuseen Betriebsges. m.b.H., Naturwissenschaftliche Sammlungen, Feldstraße 11a, 6020 Innsbruck;

p. huemer@tiroler-landesmuseen. at.

http://zoobank.org/4D0DEFC4-83DC-4309-8D65-1529E71BDEFB

Received 8 April 2015; accepted 18 August 2015; published: 22 October 2015

Subject Editor: Lauri Kaila.

Abstract. Micropterix gaudiella Zeller & Huemer, sp. n. is described from the southern part of the Orobian

Alps (Piedmont, Italy) and compared with its likely closest relatives Micropterix isobasella Staudinger, 1871

and Micropterix stuebneri Zeller, Wemo & Kurz, 2013. The new species is well characterized by its wing

pattern and colour and by structures of the male genitalia. The species status is furthermore supported by mo-

lecular data of the DNA barcode region. The distance to its nearest neighbour Micropterix schaefferi Heath,

1975 is 2.65%. M. gaudiella is the seventh species of the genus Micropterix Hübner, 1825 probably endemic

to the Alps.

Introduction

The European fauna of Micropterigidae has recently gained increasing attention, reflected by

faunistic reviews and several taxonomic papers (Corley 2007; Zeller-Lukashort et al. 2007;

Thierry and Nel 2012; Zeller-Lukashort et al. 2013). The actual species inventory now seems well

advanced as new species are rarely found and usually originate from insufficiently explored Medi-

terranean countries, with only two remarkable exceptions from the Central European Alps within

the last four decades (Heath and Kaltenbach 1 984; Kurz et al. 2004). Sampling of an unidentified

Micropterix in the Italian Alps (Pizzo Arero, Orobian Alps, Piedmont, Italy) by PH and colleagues

in June and July 2013 and 2014 thus came as a surprise. Although only few females were available

in first hand, phenotypic appearance and the DNA barcode did not match any hitherto described

species from the region. It was therefore decided to search for additional samples and we finally

succeeded in collecting a large series of the species in summer 2014 including the male sex.

Subsequent analysis of male genitalic characters supported the recognition of a new species, which

is described here.

134

Hans Christof Zeller & Peter Huemer: A new species of Micropterix from Italy

Material and methods

Our study is based on almost 277 specimens of the new Micropterix species and uncounted material

of all European congeners. The type material is only partially set whereas several samples were only

spread and dried immediately after collecting to ensure sufficient quality of DNA samples. Genitalia

preparations followed standard techniques used for the family Micropterigidae (Zeller-Lukashort et al.

2007). Photographs of the adults were taken with an Olympus SZX 10 binocular microscope and an

Olympus E-3 digital camera and developed using the software HELICON FOCUS 4.3, ADOBE PHO-

TOSHOP CS4 and LIGHTROOM 2.3. DNA barcode sequences are based on a 658 base-pair long

segment of the mitochondrial COI gene (cytochrome c oxidase 1). DNA samples (dried legs) were

prepared according to the prescribed standards (deWaard et al. 2008). Present authors and associated

colleagues tried to obtain DNA barcodes of the majority of European Micropterigidae. Legs from 379

specimens belonging to 57 species of Micropterix have so far been processed at the Canadian Centre

for DNA Barcoding (CCDB, Biodiversity Institute of Ontario, University of Guelph) to obtain DNA

barcodes (BOLD 2015) using the standard high-throughput protocol described in Ivanova et al. (2006)

and deWaard et al. (2008). DNA sequencing resulted in a barcode fragment of 658 bp for a total of 149

specimens and 29 species (BOLD 2015), partially published earlier by Lees et al. (2010). Ninety-one

sequences belonging to 25 species are treated in this study and enable delimitation of the new species.

Details of successfully sequenced voucher specimens including complete voucher data and images can

be accessed in the Barcode of Life Data Systems: public dataset “Lepidoptera of the Alps - Micropterix

[DS-LEALMIC]” (BOLD 2015; Ratnasingham and Hebert 2007). Degrees of intra- and interspecific

variation in the DNA barcode fragment were calculated under Kimura 2-parameter (K2P) model of

nucleotide substitution using analytical tools in BOLD Systems v3.0 (BOLD 2015). A neighbour-join-

ing tree of DNA barcode data of European taxa was constructed using MEGA 5 (Tamura et al. 201 1)

under the K2P model for nucleotide substitutions.

The morphology of the new species is compared with similar species from the Alps and also

from other regions of Europe (Kurz and Kurz 2015). We consequently build on the important

identification treatments by Heath (1987), Kozlov (1989, 1990a, b) and Zeller-Lukashort et al.

(2007). For more information about collection sites, preparation techniques and data archive of

Micropterix spp. see Zeller-Lukashort et al. (2007).

Abbreviations of private and institutional collections:

MBCG Italy, Bergamo, Museo di Scienze Naturali “Enrico Caffi”;

RCTM Research Collection Toni Mayr, Feldkirch, Austria;

RCCZ Research Collection Christof Zeller, Thalgau, Austria;

RCNP Research Collection Norbert Poll, Bad Ischl, Austria;

RCSO Research Collection Siegfried Ortner, Bad Ischl, Austria;

TLMF Tiroler Landesmuseum Ferdinandeum, Innsbruck, Austria.

Results

Checklist of European Micropterix

The species listed below occur within Europe (Karsholt 2015, Kurz and Kurz 2015), presented in

an order considered to reflect morphological relationships among the species as suggested by Kurz

Nota Lepi. 38(2): 133-146

135

et al. (2015). Many of the morphological characters of Micropterix have been found to be more

or less similar in smaller or greater groups of species and it has been assumed that similarities or

synapomorphies of the male genitalia indicate a closer relationship. Based on a specific character

matrix several characters have been selected and attributed as plesiomorphic or apomorphic (but

see Kurz et al. 2015).

Micropterix mansuetella Zeller, 1 844

Micropterix amsella Heath, 1975

Micropterix calthella (Linnaeus, 1761)

Micropterix isobasella Staudinger, 1871

Micropterix stuebneri Zeller, Wemo & Kurz, 2013

Micropterix gaudiella sp. n.

Micropterix granatensis Heath, 1981

Micropterix aglaella (Duponchel, 1838)

Micropterix wockei Staudinger, 1870

Micropterix aureatella (Scopoli, 1763)

Micropterix herminiella Corley, 2007

Micropterix aruncella (Scopoli, 1763)

Micropterix corcyrella Walsingham, 1919

Micropterix lakoniensis Heath, 1985

Micropterix kardamylensis Rebel, 1903

Micropterix igaloensis Amsel, 1 95 1

Micropterix cassinella Kurz, Kurz & Zeller, 20 1 0

Micropterix klimeschi Heath, 1973

Micropterix completella Staudinger, 1871

Micropterix tunbergella (Fabricius, 1787)

Micropterix sicanella Zeller, 1 847

Micropterix cypriensis Heath, 1985

Micropterix aureoviridella (Höfner, 1898)

Micropterix maschukella Alphéraky, 1876

Micropterix facetella Zeller, 1850

Micropterix jeanneli Viette, 1949

Micropterix renatae Kurz, Kurz & Zeller-Lukashort, 1997

Micropterix minimella Heath, 1973

Micropterix italica Heath, 1981

Micropterix erctella Walsingham, 1919

Micropterix uxoria Walsingham, 1919

Micropterix paykullella (Fabricius, 1794)

Micropterix garganoensis Heath, 1 960

Micropterix imperfectella Staudinger, 1859

Micropterix tuscaniensis Heath, 1960

Micropterix hartigi Heath, 1981

Micropterix allionella (Fabricius, 1794)

Micropterix trifasciella Heath, 1 965

136

Hans Christof Zeller & Peter Huemer: A new species o/ Micropterix from Italy

Micropterix rothenbachii Frey, 1856

Micropterix huemeri Kurz, Kurz & Zeller-Lukashort, 2004

Micropterix ibericella Caradja, 1920

Micropterix zangheriella Heath, 1963

Micropterix schaefferi Heath, 1975

Micropterix emiliensis Viette, 1950

Micropterix osthelderi Heath, 1975

Micropterix trinakriella Kurz, Zeller-Lukashort & Kurz, 1997

Micropterix vulturensis Heath, 1981

Micropterix rablensis Zeller, 1 868

Micropterix myrtetella Zeller, 1850

Micropterix croatica Heath & Kaltenbach, 1984

Micropterix fenestrellensis Heath & Kaltenbach, 1984

Taxonomic part

Micropterix gaudiella Zeller & Huemer, sp. n.

http://zoobank.org/4AA658ED-DED7-46A7-908E-231CBC52A8A7

Material. Holotype f: Italia sept., Bergamo, Alpi Orobie Rifugio Ca d'Arera 1600 m 9°47,8’E, 45°55,07’N 25.vi.2014,

leg. Huemer TLMF 2014-006 (TLMF), label with identification numbers CZ-Z30577, label “DNA BARCODE TLMF

Lep 14851” and red label “HOLOTYPE of Micropterix gaudiella Zeller & Huemer”. - Paratypes: 4 Ç, same locality as

holotype, but 16.vii.2013, leg. Massaro M. (MBCG), labels with identification numbers CZ-Z30771, CZ-Z30774-CZ-

Z30776; 3 S, 2$, same locality as holotype, but 17.vii.2014, leg. Siegfried Ortner (RCSO), labels with identification num-

bers CZ-Z29417, CZ-Z29459-CZ-Z29462; 40 S, 22 $, same data as holotype, but leg. Norbert Poll (RCNP), labels with

identification numbers CZ-Z29463-CZ-Z29524; 69 <$, 26 Ç, same data as holotype, but leg. Mayr Toni (RCTM), labels

with identification numbers CZ-Z29360-CZ-Z29369, CZ-Z29371-CZ-Z29416, CZ-Z29418-CZ-Z29445, CZ-Z29446 and

AP: MK-1072, CZ-Z29448-CZ-Z29457; 76 15 Ç, same data as holotype (TLMF), labels with identification numbers

CZ-Z29330-CZ-Z29347, CZ-Z30561-CZ-Z30576, CZ-Z30578-CZ-Z30580, CZ-Z30582-CZ-Z30623, CZ-Z30625-CZ-

Z30631, CZ-Z30633-CZ-Z30634, CZ-Z30636-CZ-Z30638; 16 è, same locality as holotype, but 23.vi.2014, labels with

identification numbers CZ-Z29314-CZ-Z29327, CZ-Z29328 and AP-Nr 1/2014, CZ-Z29329; 1 S, same data as holotype

(TMLF), label with identification number CZ-Z30581, label “DNA BARCODING TMLF Lep 14853”; 1 »same data as

holotype (TMLF), label with identification number CZ-Z30624, label “DNA BARCODING TMLF Lepl4852”; 1 Ç, same

locality as holotype, but 25.vi.2013, leg. Huemer (TLMF), label with identification number CZ-Z29359, label “DNA BAR-

CODING TMLF Lep09987”. All 276 paratypes bear red label “PARATYPE of Micropterix gaudiella Zeller & Huemer”.

Description. Adult (Figs 1, 2). Forewing length 3. 5-3. 9 mm (c?, n=206), 4. 0-4.4 mm (Ç, n=71).

Head black-brown; vestiture of hair-like scales on the head, yellow to dirty yellow; antennae dark

brown, respectively 3/4 (c?) and 1/2 (Ç) of forewing length; thorax and tegulae violet with golden

reflection; forewing dull bluish with conspicuously broad and mostly bronzy golden markings: a

trapeziform fascia at 1/4, at 1/2 a moderately broad straight fascia across the whole wing width,

inwardly oblique, at 3/4 a broad, variably shaped, inwardly convex fascia; fringe bronzy golden,

basally bluish violet; hindwing bronzy golden, apically tinged purplish; fringe bronzy golden; legs

and abdomen brown, golden shining.

Nota Lepi. 38(2): 133-146

137

Figures

1-2: Adults of M. gaudiella sp. n. 1: 2:?.

There is some variation in the ground coloration from dull bluish to purple-violet. Sometimes

there is also a small spot on the costa at 3/5. The wing pattern differs between the sexes only a little

in the broadness of the fasciae, as is typical for the genus.

$ Genitalia (Fig. 3). Uncus long, slender, with a broad rounded tip; a paired association of hair-

like setae ventrally beyond the uncus; accessory claspers moderately long, nearly keel-shaped;

along distal margin, a row of about 10-12 moderately long distally hook-like modified thickened

setae oriented in slightly ventral direction; at inner surface proximally another irregular row of five

or six shorter, finer setae; valvae moderately long, stout, distal third enlarged and strongly bent

dorsally, constricted at the point of inflection; on the inner surface of the valvae a group of shorter

138

Hans Christof Zeller & Peter Huemer: A new species of Micropteri xfrom Italy |

Figures 3-4: Genitalia of M. gaudiella sp. n.. 3: <$. 4. $.

setae postbasally; on the inner surface of the lower margin two to three irregular rows of shorter

spinous setae and some longer setae on the distal fourth part.

Ç Genitalia (Fig. 4). Tergite IX missing; stemite IX reduced, constricted medially, with strongly

fringed lateral margins, without diagnostic features. Terminal papillae with two sclerotized plates

forming an undiagnostic band; receptaculum seminis long and slender, constricted in the first third,

with transverse striation typical for the genus.

Diagnosis. M. trifasciella from Piedmont (Italy) shows somewhat similar but conspicuously

narrower fasciae and differs in the ground colour. M. rablensis from Friuli (Italy), Carinthia (Aus-

tria) and western Slovenia shows a somewhat similar wing pattern but is predominantly smaller

and differs in the reddish bronzy golden to purple ground coloration of the forewings. M. comple-

tella , endemic to Sardinia and Corsica (Thierry and Nel 2012), has usually lighter purple- violet

forewings with comparably broad fasciae and differs in the shape of the outer fascia. From all

these mentioned species, the new species is furthermore clearly separated by its male genitalia,

e.g. by the distinct shape of uncus and accessory claspers as well as by the orientation and form

of the 10-12 long setae on the accessory claspers (Kurz and Kurz 2015). In the male genitalia, the

new species closely resembles M. isobasella from Valais (Switzerland) (Fig. 5) and the recently

described M. stuebneri from Sierra Nevada (Spain), being distinguished from M. isobasella mainly

Nota Lepi. 38(2): 133-146

139

Figure 5. Male genitalia of M. isobasella.

by its longer uncus and the partly hook-like modified thickened setae at the rounded distal margin

of the accessory claspers and from M. stuebneri mainly by its distinctly shorter and smaller acces-

sory claspers (Zeller- Lukashort et al. 2013). M. isobasella can also easily be distinguished by its

unicolorous golden forewings without any significant markings. In the female genitalia the new

species differs from M. rablensis by the shape of stemite IX, from M. trifasciella and M. comple-

tella by its strongly fringed lateral margins of stemite IX. There are no differences in the female

genitalia between M. isobasella and the new species.

Based on morphological characters (Kurz et al. 2015), the new species is considered to belong to

a species-complex together with M. calthella, M. isobasella and the recently described M. stuebneri.

Molecular data. The intraspecific divergence of the barcode region of M. gaudiella sp. n. is

moderate with a mean distance of 0.46% and a maximum distance of 0.77% (n=4) (Tab. 1). The

minimum distance to the nearest neighbour M. schaefferi on the contrary is much higher with

2.65% (mean dist. 3.18%, max. dist. 3.76%; n=9). The minimum distance to the morphologically

closest M. isobasella is 3.67% (mean dist. 3.85%, max. dist. 4.18%; n=l) and to M. stuebneri

4.73% (mean dist. 4.99%, max. dist. 5.22%; n=l).

Distribution. The new species is only known from the mountain Pizzo Arero (Alpi Orobie,

Piedmont, Italy) from an elevation of about 1600 m.

140

Hans Christof Zeller & Peter Huemer: A new species o/ Micropterix from Italy

Table 1. Intraspecific distance and interspecific divergence to the nearest neighbour in the genus

Micropterix , based on 25 European species. Source: DNA Barcode data from BOLD (Barcode of

Life Database, cf. Ratnasingham and Hebert 2007).

Life history. The early stages are unknown. The new species was observed from the end of June

to mid- July near the border of a montane-subalpine beech forest with tall herbaceous vegetation

and bushes (Ligs 6, 7), congregating on Rosa sp. feeding on its pollen. Together with the new spe-

cies, M. aruncella (Scopoli, 1763) was recognized on flowers of Rosa sp. too (Fig. 8). At the same

locality M. rothenbachii Frey, 1856 was also found. Several specimens of the new species were

swept from flowers of Helianthemum spp. in adjacent subalpine grassland. The habitat is southern

exposed slopes on limestone.

Etymology. It was a great pleasure to find this unexpected new species from the Italian Alps.

Therefore the new species is called “ gaudiella ”, derived from the Latin word “gaudium”, which

means “fun, pleasure, happiness”.

Remark. The labels used by PH, Melania Massaro, Toni Mayr, Norbert Poll and Siegfried

Ortner differ in the usage of local names of the type locality but all refer to the same spot centred

on 9°47,8’E; 45°55,07’N (DDM). According to Melania Massaro there are additional specimens

collected at the type locality and deposited in MBCG, which are not examined and therefore not

included in the type series.

Nota Lepi. 38(2): 133-146

141

Figure 6. Biotope of M. gaudiella and the lucky collectors Marlies Mayr, Toni Mayr and Norbert Pöll.

Figure 7. M. gaudiella sp. n., resting on a flower of Rosa sp..

142

Hans Christof Zeller & Peter Huemer: A new species of Micropterix from Italy

Molecular analysis

Sequencing of European species of Micropterix resulted in barcode fragments for 6 1 specimens,

plus 30 unpublished or public records from external projects (e.g. Barcoding Fauna Bavarica and

Finnish Barcode of Life), covering altogether 25 species, supplementing data of extra-European

species that have been published by Lees et al. (2010). 74 full barcode sequences of 658 bp, 13

sequences longer than 600 bp and four sequences of ca. 400 bp were included in the analysis. The

smaller fragments were single sequence only (M. amsella, M. mansuetella , and M. rablensis).

Barcode variation is insufficiently known for a considerable portion of species due to lack of

successfully sequenced samples (Table 1). Intraspecific distance is low and ranges from 0% to

2.99% (mean 0.92%) but may include cases of overlooked cryptic diversity in species with high

divergence (i.e. M. aruncella, M. facetella). Interspecific divergence for the whole sample is mod-

erately high with mean divergence of 4.77% and maximum of 8.91%, and a mean and maximum

distance to the nearest neighbour of 2.84% and 5.35% respectively. The divergence is <1% only in

one species pair (M. schaefferi and M. vulturensis) (Table 1, Fig. 9).

Discussion

Alpha-taxonomy of European Micropterix seems quite well established with relatively few taxa de-

scribed during the last decades, so it is striking to find a new species in Central Europe. By contrast,

the distribution of the majority of Micropterix species is only moderately well documented with few

Figure 8. M. aruncella and M. gaudiella sp. n. feeding on Rosa sp.

Nota Lepi. 38(2): 133-146

143

I Micropterix aureatella (n=14; AT, Fl, IT, SW)

Micropterix erctella (n=1; IT)

Micropterix hartigi (n=1; IT)

Micropterix rothenbachii (n=3; AT, IT)

^ Micropterix allionella (n=7; AT, FR, IT)

Micropterix rablensis (n=1; IT)

^ Micropterix aureoviridella (n=5; AT, IT)

Micropterix osthelderi (n=2; AT)

Micropterix mansuetella (n=1; Fl)

Micropterix amsella (n=1; MN)

Micropterix wockei (n=1 ; GR)

Micropterix jeanneli (n=1; MN)

Micropterix facetella (n=3; MN)

Micropterix klimeschi (n=1; GR)

Micropterix aruncella (n=17; AT, DE, Fl, IT, MK)

Micropterix stuebneri (n=1; ES)

Micropterix isobasella (n=1; CH)

Micropterix vulturensis (n=1; IT)

Micropterix schaefferi (n=9; AT, DE, IT)

Micropterix myrtetella (n=1; IT)

-4 Micropterix tunbergella (n=6; AT, CH, DE, LV)

Micropterix igaloensis (n=1; MN)

Micropterix sp. (n=2; IT)

Micropterix gaudiella (n=4; IT)

Micropterix sicanella (n=1; IT)

Micropterix calthella (n=5; Fl, UK)

o.oi

Figure 9. Neighbour-joining tree (Kimura 2-parameter, built with MEGA 5; cf. Tamura et al. 201 1). Note: the

scale bar only applies to internal branches between species. The width of the triangles represents the sample

size, and the depth the relative genetic variation within the cluster (2x scale bar). Source: DNA Barcode data

from BOLD (Barcode of Life Database, cf. Ratnasingham and Hebert 2007).

faunistic studies, particularly as concerns the Mediterranean region. Despite this shortcoming, it al-

ready appears evident from published sources that Micropterix exhibits remarkable levels of narrow

range endemicity. Indeed about three-quarters of the European taxa can be clustered into the follow-

ing disjunct endemism groups (Karsholt 2015; Zeller-Lukashort et al. 2013; Kurz and Kurz 2015).

144

Hans Christof Zeller & Peter Huemer: A new species o/ Micropterix from Italy

Apennines: 9 out of 20 species are endemic.

Alps: 7 out of 19 species are endemic.

Balkan Peninsula: 9 out of 17 species are endemic.

Iberian Peninsula: 5 out of 8 species are endemic.

Mediterranean islands: 8 species are endemic.

Black Sea area: 1 species is endemic.

Also, only 12 species (about one-quarter of European Micropterix) are widely distributed across

Europe, and only one species (M aureatelld) reaches Japan (Kurz and Kurz 2015). M. aruncella,

M. aureatella , M. calthella , M. mansuetella and M. tunbergella are known to range across from the

mainland to geographically well separated islands (although a few micropterigids occur within an

island group, such as Sabatinca occurring in both North and South Island of New Zealand: Gibbs

1983; Gibbs and Lees 2014; see also Imada et al. 2011). Endemism in the Lepidoptera fauna of the

Alps in general and Micropterix in particular probably largely results from the history of glaciation,

which is reflected in nine biogeographic zones defined by vegetation (Ozenda 1988). These zones

have been partially considered as areas of endemism for Lepidoptera (Huemer, 1988). The majority

of endemic Micropterix occur in the Western Alps and are restricted to the so-called “Inner Alps”

(i.e. M. isobasella, M. paykullella , M. trifasciella and M. fenestrellensis) whereas M. huemeri is

restricted to the adjacent “Pre-Ligurian” biogeographic zone. However, the highest diversity of en-

demic Lepidoptera in general is present in the so-called “Gardesan-Illyrian zone” of the south-east-

ern Alps which is defined by limestone massifs at the southern border of the Alps reaching from

Lake Como in the west to the Karawanks in the east. More than 30 endemic Lepidoptera species

are known from this area (Huemer 1998). M. gaudiella is the first species of the genus Micropterix

probably endemic to this zone, while a further species, M. rablensis , is more widespread and also

occurs in the northern part of the Eastern Alps. The Orobian Alps in particular seem to have attract-

ed insufficient attention among lepidopterists so far and still host a considerable number of cryptic

species belonging to various groups. A new species of Kessleria (Huemer & Mutanen, 2015) and

a distinct subspecies of Colostygia (Huemer & Mayr, 2015) are striking examples. It seems not

unlikely that further undescribed endemic Micropterix may be found in other remote and under-

collected areas of Europe, e.g. on the Balkan and Iberian Peninsulas. Outside Europe, despite the

efforts of John Heath, there are probably a number of undetected species in North Africa, while the

Himalayas as far as China are virtually unexplored for the genus (but see Lees et al. 2010).

Acknowledgements

We are particularly grateful to Paul Hebert and his team at the Canadian Centre for DNA Barcoding

(Guelph, Canada), whose sequencing work was enabled by funding from the Government of Can-

ada to Genome Canada through the Ontario Genomics Institute. We are also grateful to the Ontario

Ministry of Research and Innovation and to NSERC for their support of the BOLD informatics

platform. We are furthermore indebted to the Promotion of Educational Policies, University and

Research Department of the Autonomous Province of Bolzano - South Tyrol for helping to fund

the project “Genetic biodiversity archive - DNA barcoding of Lepidoptera of the central Alpine

region (South, East and North Tyrol)”, and to the Austrian Federal Ministry of Science, Research

and Economics for funds received in the framework of ABOL (Austrian Barcode of Life).

Nota Lepi. 38(2): 133-146

145

Dr. Marko Mutanen (Oulu) and Dr. Andreas Segerer (Munich) kindly gave us access to un-

published barcode sequences. We are furthermore grateful to Toni Mayr, Siegfried Ortner and

Norbert Poll for the loan of their material and provision of useful photos, to Mag. Michael Kurz

for help regarding the preparation of the female genitalia. Dr. David Lees (London) and Martin

Corley (Faringdon, Oxford) kindly improved our English and gave us useful hints. Stefan Heim

(Tiroler Landesmuseen, Innsbruck) kindly developed figures of set adults and genitalia. Finally we

acknowledge the kind support received from Dr. Marco Valle, Dr. Paolo Pantini and Dr. Melania

Massaro (Museo Civico di Science Naturali “E. Caffi”, Bergamo) during our stay in Italy and sup-

plementing samples of the new species.

References

BOLD (2015) BOLD: The Barcode of Life Data System (www.barcodinglife.org). http://www.boldsystems.org

Corley M (2007) A brief review of the Micropterigidae of Portugal, with description of a new species of Mi-

cropterix Hübner. Nota lepidopterologica 30(1): 71-78.

deWaard JR, Ivanova NV, Hajibabaei M, Hebert PDN (2008) Assembling DNA Barcodes: Analytical

Protocols. In: Cristofre M (Ed.) Methods in Molecular Biology: Environmental Genetics. Humana Press

Inc., Totowa, USA, 275-293. doi: 10.1 007/978- 1-59745-548-0J 5

Heath J, Kaltenbach T (1984) New species of Micropterix Hübner (Lepidoptera: Zeugloptera: Micropterigidae)

from Italy and Yugoslavia. Entomologist’s Gazette 35: 21-23.

Gibbs GW (1983) Evolution of Micropterigidae (Lepidoptera) in the SW Pacific. GeoJoumal 7.6: 505-510.

doi: 10.1007/bf002 18523

Gibbs GW, Lees DC (2014) New Caledonia as an evolutionary cradle: a re-appraisal of the jaw-moth genus

Sabatinca (Lepidoptera: Micropterigidae) and its significance for assessing the antiquity of the island’s

fauna. In: Guilbert É, Robillard T, Jourdan H, Grandcolas P (Eds) Zoologia Neocaledonica 8. Biodiversity

studies in New Caledonia. Muséum national d’Histoire naturelle, Paris, 239-266. [Mémoires du Muséum

national d’Histoire naturelle; 206]

Heath J (1987) A check list of the genus Micropterix Hübner [1825] (Lepidoptera: Zeugloptera, Micropterigidae).

Entomologist’s Gazette 38: 205-207.

Huemer P (1998) Endemische Schmetterlinge der Alpen - ein Überblick (Lepidoptera). Stapfia 1998(55):

229-256.

Huemer P, Mayr T (2015) Eine neue Unterart von Colostygia kitschelti aus den Bergamasker-Alpen, mit Be-

merkungen zum Artkonzept im C. austriacaria-Komplex (Lepidoptera: Geometridae). Wissenschaftliches

Jahrbuch der Tiroler Landesmuseen (in press).

Huemer P, Mutanen M (2015) Alpha taxonomy of the genus Kessleria Nowicki, 1864, revisited in times of

DNA-barcoding (Lepidoptera, Yponomeutidae). ZooKeys 503: 89-133. doi: 10.3897/zookeys.503.9590

Imada Y, Kawakita A, Kato M (2011) Allopatric distribution and diversification without niche shift in a

bryophyte-feeding basal moth lineage (Lepidoptera: Micropterigidae). Proceedings of the Royal Society B

278: 3026-3033. doi: 10.1 098/rspb.20 11.0134

Ivanova NV, deWaard JR, Hebert PDN (2006) An inexpensive, automation-friendly protocol for recovering

high-quality DNA. Molecular Ecology Notes 6: 998-1002.

Kozlov MV (1989) Short review and key for determination of Micropterix Hbn. (Lepidoptera, Micropterigidae)

species of Palaearctic. 2. Key for determination (pt. 1). Vestnik zoologii 1989(6): 26-31. [In Russian]

Kozlov MV (1990a) Short review and key for determination of Micropterix Hbn. (Lepidoptera, Micropterigi-

dae) species of Palaearctic. 3. Key for determination (pt. 2). Vestnik zoologii 1990(2): 21-26. [In Russian]

146

Hans Christof Zeller & Peter Huemer: A new species of Micropterix from Italy

Kozlov MV (1990b) Short review and key for determination of Micropterix Hbn. (Lepidoptera, Micropterigi-

dae) species of Palaearctic. 4. Results of investigation of the type species. Vestnik zoologii 1990 (3): 28-33.

[In Russian]

Karsholt O (2015) Fauna Europaea: Lepidoptera, Micropterigidae. Fauna Europaea version 2.6.2, http://

www.faunaeur.org

Kurz MA, Kurz ME (2015) Naturkundliches Informationssystem, http://www.nkis.info

Kurz MA, Kurz ME, Zeller-Lukashort HC (2004) A new Micropterix species from southern France (Microp-

terigidae). Nota lepidopterologica 26(3/4): 111-114.

Kurz MA, Kurz ME, Zeller HC (in press) Morphological characters as indicators for a hypothetical arrange-

ment of species in the genus Micropterix Hübner. Mitteilungen der Naturkundlichen Gesellschaft.

Lees DC, Rougerie R, Zeller-Lukashort C, Kristensen NP (2010) DNA mini-barcodes in taxonomie assign-

ment: a morphologically unique new homoneurous moth clade from the Indian Himalayas described in

Micropterix (Lepidoptera, Micropterigidae). Zoologica Scripta 39(6): 642-661. doi: 10. 1 1 1 1/j. 1463-

6409.2010.00447.x

Ratnasingham S, Hebert PDN (2007) BOLD: The Barcode of Life Data System (http://www.barcodinglife.

org). Molecular Ecology Notes 7: 355-364. doi: 10.1 1 1 l/j. 1471-8286.2007.01678.X

Ozenda P (1988) Die Vegetation der Alpen im europäischen Gebirgsraum. Gustav Fischer Verlag, Stuttgart,

New York, 353 pp.

Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S (201 1) MEGA5: Molecular Evolutionary Ge-

netics Analysis using Maximum Likelihood, Evolutionary Distance, and Maximum Parsimony Methods.

Molecular Biology and Evolution 28: 2731-2739. doi: 10.1093/molbev/msrl21

Thierry V, Nel J (2012) Microlépidoptères nouveaux ou peu connus de la faune de France (Lep. Micropterigidae,

Blastobasidae, Cosmopterigidae, Gelechiidae). Oreina 20: 17-18.

Zeller-Lukashort HC, Kurz ME, Lees DC, Kurz MA (2007) A review of Micropterix Hübner, 1825 from

northern and central Europe (Micropterigidae). Nota lepidopterologica 30(2): 235-298.

Zeller-Lukashort HC, Wemo A, Kurz MA (2013) A new species of Micropterix Hübner, [1825] from southern

Spain (Lepidoptera: Micropterigidae). SHILAP Revista de Lepidopterologia 41(164): 489^194.

Nota Lepi. 38(2) 2015: 147-155 | DOI 10.3897/nl.38.6289

Coleophora gryphipennella (Hübner, 1796) (Lepidoptera, Coleophoridae)

on Fragaria vesca L. (Rosaceae), a novel host, in the coastal dunes

of The Netherlands

John A.M. van Roosmalen1, Camiel Doorenweerd2

1 Buizerdweg 16, 1826 GG Alkmaar, The Netherlands; j.roosmalen6@upcmail.nl

2 Department of Terrestrial Zoology, Natur alls Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands;

camiel. doorenweerd@naturalis. nl

http://zoobank.org/E8B0465E-D133-4ACF-AAll-ABCE86DDE594

Received 21 August 2015; accepted 19 October 2015; published: 6 November 2015

Subject Editor: Bernard Landry.

Abstract. During regular surveys of Lepidoptera in the coastal dune North Holland Dune Reserve, we ob-

served larval cases and feeding traces typical for Coleophoridae on wild strawberry {Fragaria vesca L.,

Rosaceae). The spatulate or pistol shape of the cases excluded Coleophora violacea (Ström, 1783) and C.

potentillae Elisha, 1885. According to the literature, C. albicostella (Duponchel, 1842), very rare in The

Netherlands, was the only Coleophora species known to create spatulate cases on this host. We collected

larval cases for DNA analysis on the larvae and for rearing, which revealed that none of the collected larvae

belong to the Coleophoridae previously recorded feeding on this host, but a mixture of three other Coleophora

species. We found that early instar larvae of C. lutipennella (Zeller, 1838) and C. flavipennella (Duponchel,

1843), normally feeding on oaks {Quercus spp., Fagaceae) only, may be found feeding on F vesca in the fall.

We also found that C. gryphipennella (Hübner, 1796), abundant on Rose {Rosa spp., Rosaceae) in the coastal

dunes of The Netherlands, regularly feeds on F vesca and rearing experiments proved that it can complete

its larval stage on F vesca. We therefore conclude that Fragaria is a new host genus for C. gryphipennella.

After reviewing all the C. albicostella records from The Netherlands, we conclude that it is a very rare species,

likely restricted to the southernmost provinces. None of the confirmed records are from reared specimens.

The host range of C. albicostella in literature is possibly overestimated and may not even include Fragaria.

Introduction

The coastal dunes of the province North Holland constitute a region rich in Lepidoptera. The North

Holland Dune Reserve, a 53 km2 dune area roughly between the cities of Wijk aan Zee and Bergen,

is a particularly rich area. Here, 1,449 species of moths (macro- and microlepidoptera) have been

recorded in the Dutch national observation database “NOCTUA” (Ellis 2015a), 1,353 of which

were recorded from year 2000 onwards. In total in this database approximately 2,400 different

species of Lepidoptera have been recorded for The Netherlands. The North Holland Dune Reserve

is protected by Natura 2000 laws and is managed by the private water company PWN. The first

author is one of the volunteers who inventory Lepidoptera in this reserve. During day-time collect-

ing, he encountered larval cases and feeding traces typical for Coleophoridae on wild strawberry

{Fragaria vesca L., Rosaceae).

148

Roosmalen & Doorenweerd: Coleophora gryphipennella (Hübner, 1796)...

Many Coleophoridae are leaf miners as larvae, although, unlike most other leafmining groups,

they can arbitrarily change their host during the larval life stage and do not stay in the leafmines con-

tinuously. The larvae build a portable case that is constructed from silk and plant tissue, usually the

epidermal layers of a plant leaf (Emmet et al. 1996). The cases often have a distinctive shape and sev-

eral types of cases have been designated by different authors and are included in identification keys

(Hering 1951; Toll 1953; 1962; Patzak 1974; Emmet et al. 1996). However, the shape of the case can

change significantly between early and later instars. Furthermore, some species shift between leaf

mining to seed mining or flower or flower-bud mining, for example Coleophora salinella Stainton,

1859, C. tricolor Walsingham, 1899, and C. bernoulliella (Goeze, 1783) (Emmet et al. 1996). None-

theless, most species are monophagous and the host plant is an important character for identification.

Three species of Coleophoridae were known to feed on Fragaria in Europe according to the

literature (Hering 1957; Klimesch 1958). Two of these create lobe type cases, viz. Coleophora

violacea (Ström, 1783) and C. potentillae Elisha, 1885 and can easily be recognized (Emmet et al.

1996; Ellis 2015b). The third species that had been reported to feed on Fragaria is C. albicostella

(Duponchel, 1842), which creates a spatulate case, a more common case type amongst Coleop-

horidae. C. albicostella is a very rare species in The Netherlands with about a dozen registered

observations or collected specimens since 1880 (Küchlein and Donner 1993; Willem Ellis, pers.

comm.; RMNH and ZMA collections [Naturalis Biodiversity Center, Leiden, The Netherlands]).

To find out which species we found feeding on Fragaria with spatulate or pistol shaped cases, we

photographed, collected, and reared them and analysed their DNA.

Material and methods

Localities

The North Holland Dune Reserve stretches roughly from 52.487°N, 4.590°E to 52.682°N, 4.69 1°E

and has an area of roughly 53 km2. We did most of our observations in the area just south of the

village Bergen aan Zee at two main localities. One is a small patch of woodland surrounded by an

open dune area relatively close to the inland border of the North Holland Dune Reserve, with En-

glish Oak ( Quercus robur L., Fagaceae) as the main tree species and F vesca dominant in the herb

layer, further indicated as ‘Ll Oak’. The other locality is a small patch of woodland surrounded

by an open dune area relatively close to the North Sea with Birch (Betula sp., Betulaceae) as the

main tree species and again F vesca dominant in the herb layer, further indicated as ‘L2_Birch’. In

addition, we carried out some observations in the surrounding area. An overview of all localities

and dates is provided in Table 1. JvR took the photographs, with a Canon EOS 7D camera with

a Canon EF 100mm f/2.8L Macro IS USM lens, except for the photographs in figures 15 and 16,

which were taken with a Nikon D80 camera with a Micro-Nikkor AF 60mm f/2.8 D lens.

Rearing

On 29.X.2013 and 19.xi.2013, we collected spatulate cases on Fragaria vesca at the L2_Birch local-

ity. We kept the collected larval cases in a small transparent plastic jar and put them in a garden shed

to hibernate, along with small pieces of the host plant. In March 2014 we transferred the cases to

fresh Fragaria leaves in the garden. We first observed fresh feeding traces on 1 l.iv.2014, the active

larvae still with spatulate cases. On 05.V.2014 we observed that the most active larva had stopped

feeding and that the case had moved to the petiole of the leaf it had been feeding on. The case now

Nota Lepi. 38(2): 147-155

149

Table 1. Localities of the Coleophora observations of this study in chronological order.

*The case types following Emmet et al. (1996)

had a trivalved appearance. We transferred the case into a small plastic jar and kept it indoors until

emergence. We found the second case lying on a leaf rather than being attached to the underside and

we assumed that the larva had died. We obtained another rearing result from a still fresh spatulate

case on F. vesca on 07.V.2014 in the dunes near Egmond (GPS coordinates 52.637°N, 4.649°E) by

collecting the case together with fresh leaves that were taken indoors until emergence.

DNA barcoding

We selected ten specimens from different localities and hosts for DNA analysis, see Table 2. We

pulled the larvae from their case with forceps, damaging the case as little as possible. The cases are

stored as vouchers in the RMNH collection. For DNA analysis we used the mitochondrial COI-bar-

code gene region (Hebert et al. 2003). DNA extraction and PCR amplification followed the meth-

ods described in van Nieukerken et al. (2012). We added the collecting and sequence data of the

specimens to the Barcode of Life Datasystems (BOLD; Ratnashingham and Hebert 2007), under

their unique RMNH registration number as sample ID. Sufficient reference material of all potential

Coleophora species that feed on Fragaria was available in BOLD and we made identifications of

the larvae by examining the distance to the nearest neighbour. There was a 1 00% match for most

samples, and also the monophyly criterion was met, when we evaluated whether our sequence

fitted within a monophyletic DNA barcode species cluster.

Abbreviations

RMNH Rijks Museum voor Natuurlijke Historie collection, housed at Naturalis Biodiversity Center,

Leiden, The Netherlands.

ZMA Zoologisch Museum Amsterdam collection, housed at Naturalis Biodiversity Center, Leiden,

The Netherlands.

Results

Field observations

In the spring of 2011 at locality ‘Ll Oak’, during an intensive search for Tinagma perdicella

Zeller, 1839 (Douglasiidae), which flies during daytime and can be found close to Fragaria , JvR

150

Roosmalen & Doorenweerd: Coleophora gryphipennella (Hübner, 1796)...

noticed occupied Coleophoridae larval cases and feeding damage on Fragaria (Figs 1, 2). These

spatulate cases at first impression had a shape similar to cases of Coleophora albicostella, and

given the host plant, led to the assumption that this was the rare C. albicostella. However, rearing

was unsuccessful, leaving the identifications unconfirmed. In the autumn of that same year, again

spatulate cases were found, this time at locality ‘L2_Birch\ At that time rearing was not attempted.

It was assumed to be easier to look for larval cases in the spring and attempt to rear those, allowing

the larvae to hibernate under natural conditions. However, in the spring of 2012 JvR encountered

no cases and only in the next autumn, JvR observed cases again.

In the autumn of 2012 at locality ‘L2_Birch’ we stumbled upon a small Rosa sp. bush with

almost each leaf occupied by a Coleophora case, and most leaves were eaten out. At the same

locality, a few meters further, we discovered a Fragaria vesca plant also with typical Coleophora

feeding traces and with cases indistinguishable from the ones on the Rosa sp. bush. Both even

showed the small, approximately three mm long, rectangular shaped first stage cases still present

on the plants (Figs 3-6). Only three cases were found on F. vesca and JvR decided to return in the

spring to collect cases rather than to attempt to hibernate and rear these cases.

However, in the spring of 2013 again JvR found no larval cases on Fragaria. It took until

October 2013 until finally JvR could collect cases. At this point collaboration started with CD to

use DNA barcode analysis to identify the species that creates these cases on Fragaria and Rosa.

To collect fresh material, we intensified the search for larval cases and this yielded two cases

from Rosa spp. as well as from Fragaria vesca from different localities (Table 1). During this

collecting effort, we found a second type of feeding trace and larval Coleophora case on F. vesca :

tiny fleck mines created by larvae in small pistol-shaped cases (Figs 7-8) at locality ‘Ll Oak’.

We sequenced a total of ten larvae (see Figs 8-10) from the different localities on 19.X.2013 and

29.X.2013 (Table 2).

DNA barcode analysis

Nine out of the ten barcoded specimens yielded a DNA barcode. All identifications were unambig-

uous (Table 2). None of the specimens proved to be Coleophora albicostella. Instead, a mixture of

three other species, none of which had been recorded to feed on Fragaria , was found: C. gryphipen-

nella (Hübner, 1796), C. lutipennella (Zeller, 1838), and C. flavipennella (Duponchel, 1843). The

Table 2. Material collected for DNA barcoding and subsequent molecular identification.

Nota Lepi. 38(2): 147-155

151

Figures 1-8. 1. Spatulate case found on Fragaria 20.V.201 1. To the right of the case feeding damage can be

seen. 2. Spatulate case found on Fragaria 20.V.201 1, with larva. 3. First stage case on Fragaria , showing also

the place where the leaf was cut for a second stage case, 02.X.2012. 4. Spatulate case on Fragaria, 02.X.2012.

5. Fragaria leaf from Figure 3 showing feeding damage, first stage case and ‘leaf cut’, 02.x. 2012. 6. First

stage cases and spatulate cases on Rosa, 02.x. 2012. 7. Tiny fleck mines on Fragaria, 29.x. 2013. 8. Small

pistol case on Fragaria, 29.X.2013.

152

Roosmalen & Doorenweerd: Coleophora gryphipennella (Hübner, 1796)...

larger cases, i.e. >5 mm long, were all of the spatulate type and were identified as C. gryphipennella.

The smaller cases were pistol shaped. These were either C. lutipennella or C. flavipennella , species

that normally feed on oaks ( Quercus spp.). We did not find larger pistol shaped cases.

Rearing

We collected a few autumn cases from Fragaria vesca in November 2013 for rearing. They hi-

bernated in a garden shed and began feeding on Fragaria again in the spring (Figs 11-12). On

21.V.2014 an adult emerged. Based on the external characters we identified it as Coleophora gry-

phipennella, which can be readily distinguished from C. albicostella by the lack of a white stripe

along the costa (see Figs 13-14). In addition, a fresh case collected by Luc Knijnsberg on F vesca

on 7.V.2014 (Fig. 15) at ‘Ll Oak’ also successfully yielded an imago of C. gryphipennella on

10.vi.2014 (Fig. 16).

Discussion

Host range and distribution of Coleophora albicostella

We initially expected to have found the rare Coleophora albicostella in the North Holland Dune

Reserve, but none of the specimens that we reared or analysed for DNA proved to be C. albicostel-

la. Although this does not completely exclude the option that C. albicostella occurs on Fragaria

vesca in the coastal dunes of The Netherlands, further investigation of the host records of this

species made it even more unlikely. Fragaria spp. is first mentioned as a host plant for C. albi-

costella by Hering (1957), along with Potentilla spp. and Rubus spp. The status and source of

these observations are unclear, as they are mentioned without reference to the stage (e.g. larval

or adult stage) of the studied material or whether rearing was attempted. Hering (1957) does pro-

vide a drawing of the larval case, which appears as a rather general looking spatulate case that

cannot be distinguished from, for example, C. gryphipennella. Furthermore, the case may include

slight differences based on the host plant that was included in its construction. Huemer (1988)

only mentions Potentilla spp. and occasionally also Comarum spp., Filipendula spp., Rubus spp.,

and Sanguisorba spp. as hosts (all Rosaceae). Rearing has been documented from Potentilla spp.

(Bryner 2015; Richter 2015) and Geum spp. (Hugo van der Wolf, pers. comm., in collection). We

believe that these records represent the predominant hosts. Even if C. albicostella may feed on the

other genera, it is likely only so in areas where Potentilla or Geum may be found. In any case, also

in the light of our findings here, C. albicostella cannot readily be reported when spatulate larval

cases are encountered on Fragaria and should always be reared, or DNA barcoded, to efficiently

identify such larvae.

Fragaria vesca as a new host for Coleophora gryphipennella

We demonstrated both by DNA analysis on the larvae from spatulate shaped larval cases collected

on Fragaria vesca in the fall as well as by successfully rearing adults from larvae that have been

feeding on F vesca both in the fall and in the spring that F. vesca is a novel host for Coleophora

gryphipennella. At least in the study area, this seems unlikely to be due to a shortage of Rosa ,

especially Rosa pimpinellifolia L. but also larger species such as Rosa rubiginosa L. are abundant

and C. gryphipennella can be found in many localities feeding on Rosa throughout the habitat.

The adaptation to another host plant for C. gryphipennella could be a local habit, only occurring in

Nota Lepi. 38(2): 147-155

153

Figures 9-16. 9. Spatulate leaf case on Fragaria collected for DNA analysis, 29.X.2013. 10. Spatulate leaf

case on Rosa collected for DNA analysis, 29.X.2013. 11. Final, trivalved case of the reared larva after hiberna-

tion and after feeding ended on 05.V.2014. 12. Fleck mines produced by the reared larva after hibernation and

after feeding ended on 05.V.2014. 13. Emerged imago Coleophora gryphipennella reared from a hibernating

larva. 14. Emerged imago Coleophora gryphipennella reared from a hibernating larva. 15. Fresh green case

on Fragaria, 07.V.2014, leg. and photograph Luc Knijnsberg. 16. Emerged imago from the case found on

07.V.2014, photograph Luc Knijnsberg.

154

Roosmalen & Doorenweerd: Coleophora gryphipennella (Hübner, 1796)...

the dunes of the North Holland Dune Reserve. On the other hand, it could also be a more general,

widespread habit that can be found throughout its distribution in Europe. More evidence to sup-

port this came from a recent find of a C. gryphipennella larva on F vesca near Durbuy, Belgium

(Steve Wullaert, pers. comm.), as confirmed by DNA analysis (RMNH.INS.30422 on BOLD). C.

gryphipennella is distributed throughout most of Europe, Turkey, and Central and Eastern Siberia

(Baldizzone et al. 2006). C. gryphipennella is mostly described as monophagous on Rosa spp. Our

observations include feeding signs and larval cases on Rosa pimpinellifolia and Rosa rubiginosa.

Rubus sp. and Rubus corylifolius Sm. (Rosaceae) are mentioned as incidental host plants by Her-

ing (1957). Hering describes that in the fall the cases are somewhat pressed together sideways and

are terminated with a two-sided valve and that in the spring they become more cylindrical, ending

with a three-sided valve. Our observations are that after hibernation, the cases are still flat with a

two-sided valve. Before pupation they turn more cylindrical with a three-sided valve.

Status of Coleophora albicostella and C. gryphipennella in The Netherlands

Our findings indicate that Coleophora albicostella is rare in The Netherlands, even more rare

than it appeared before we started working on this manuscript, and we have had to conclude that

some of the records in the national database “NOCTUA” (Ellis 2015a) involved different species.

In some cases this was due to identifications based on larval cases, but sometimes also because

of a confusion with the similarly named C. albicosta (Haworth, 1828), which feeds on Ulex spp.

(Fabaceae). Only a handful of verified C. albicostella records remain, all from the southernmost

part of the country, the south of the province of Limburg. Outside this area, two doubtful records

are reported, one from the southwest and one from the east of the country. C. gryphipennella on

the other hand is abundant in The Netherlands with records throughout the country in all provinces

(Küchlein and Donner 1993; Muus 2015). Most records are reported from the coastal area where

Rosa pimpinellifolia is common, which is also reported to be the main host plant (Hering 1957).

Identification of spatulate-type Coleophora cases on Fragaria vesca

Spatulate cases on Fragaria in the spring or the larger cases in the autumn are most likely all Coleop-

hora gryphipennella , although we cannot completely exclude C. albicostella. It appears that the final

cases in the spring are trivalved (see Fig. 11) and from this stage the adults emerge about a month later

(Fig. 15). Small pistol cases (i.e. <5 mm) that may be encountered in the autumn on Fragaria are C.

lutipennella or C. flavipennella , and can most likely be found in the vicinity of oaks ( Quercus spp.), the

common host for these species (Hering 1957). C. lutipennella and C. flavipennella are in the same spe-

cies group as C. gryphipennella {sensu Emmet et al. 1996), but we find it unlikely that they can com-

plete their larval stage without oaks. It is more likely that, when they come down from the oaks in the

autumn to find a safe place to hibernate, they consume small amounts of Fragaria opportunistically.

Conclusions

Fragaria is a new host genus for Coleophora gryphipennella , which was previously only reported

to feed on Rosa , or occasionally on Rubus. C. gryphipennella creates spatulate type cases that may

be found in the fall and the spring and the species can complete its larval life on Fragaria vesca.

Small pistol shaped cases that may be found on Fragaria in the fall belong to the Quercus feeding

species C. flavipennella or C. lutipennella. The actual host range of C. albicostella , which was

Nota Lepi. 38(2): 147-155

155

initially known to be the only species creating spatulate cases on Fragaria, remains unclear, but

there are no confirmed rearing records of this species from Fragaria.

Acknowledgements

We would like to thank Hugo van der Wolf, Tymo Muus, Erik van Nieukerken, Willem Ellis, and Jacques

Wolschrijn for useful comments. Special appreciation goes to Luc Knijnsberg for contributing his rearing

results and photos and to Steve Wullaert for contributing his Belgian find. We would like to thank Giorgio

Baldizzone, Bernard Landry and Jadranka Rota for reviewing and editing and their comments that further

improved the manuscript.

References

Baldizzone G, van der Wolf H, Landry J-F (2006) Coleophoridae, Coleophorinae (Lepidoptera). In: Landry B

(Ed.) World Catalogue of Insects, Vol. 8. Apollo Books, Stenstrup.

Bryner R (2015) Coleophora albicostella. http://lepiforum.de/lepiwiki.pl7Coleophora_Albicostella [accessed

26.iv.20 15]

Ellis WN (2015a) NOCTUA, the database of The Netherland’s Lepidoptera, maintained by the Working

Group Lepidoptera Faunistics and Dutch Butterfly Conservation, [accessed 05.V.2015]

Ellis WN (2015b) Bladmineerders van Europa / Leafminers of Europe, http://www.bladmineerders.nl/index.

htm [accessed 26.iv.20 15]

Emmet A, Fletcher D, Harley B, Langmaid J, Robinson GS, Skinner B, Sokoloff P, Tremewan W, Heath J (Eds)

(1996) The moths and butterflies of Great Britain and Ireland. Harley Books, Essex, England, 452 pp.

Hebert PDN, Cywinska A, Ball SL, DeWaard JR (2003) Biological identifications through DNA barcodes.

Proceedings of the Royal Society of London Series B-Biological Sciences 270: 313-321. doi: 10.1098/

rspb.2002.2218

Hering EM (1951) Biology of the Leaf Miners. Dr W. Junk, ’s-Gravenhage, 419 pp.

Hering EM (1957) Blattminen von Europa. Dr W. Junk, ’s-Gravenhage, Band I, 648 pp.

Huemer P (1988) Kleinschmetterlinge an Rosaceae unter besonderer Berücksichtigung ihrer Vertikalverbreit-

ung (excl. Hepialidae, Cossidae, Zygaenidae, Psychidae und Sesiidae). Neue Entomologische Nachrichten

20: 1-376.

Klimesh J (1958) Beiträge zur Kenntnis der Lepidopteren-Fauna der Wachau in Niederösterreich (Microlepi-

doptera). Zeitschrift der wiener entomologischen Gesellschaft 43: 17-22, 43^14, 76-77, 91-97.

Küchlein JH, Donner JH (1993) De kleine vlinders: handboek voor de faunistiek van de Nederlandse Micro-

lepidoptera. Pudoc, Wageningen, 715 pp.

Muus TST (Ed.) (2015) Microlepidoptera.nl Atlas van de kleinere vlinders van Nederland, www.microlepidoptera.nl

Nieukerken EJ van, Doorenweerd C, Stokvis FR, Groenenberg DSJ (2012) DNA barcoding of the leaf-mining

moth subgenus Ectoedemia s. str. (Lepidoptera: Nepticulidae) with COI and EFl-a: two are better than one

in recognising cryptic species. Contributions to Zoology 81 : 1-24.

Patzak H (1974) Beiträge zur Insektenfauna der DDR: Lepidoptera - Coleophoridae. Beiträge zur Entomologie

24: 153-278.

Ratnashingham S, Hebert PDN (2007) BOLD: The Barcode of Life Data System (http://www.barcodinglife.org).

Molecular Ecology Notes 7: 355-364. doi: 10.1 1 ll/j.l471-8286.2007.01678.x

Richter I (2015) Coleophora albicostella. http://www.coleophoridae.bluefile.cz/?p=3418 [accessed 26.iv.20 15]

Toll S (1953) Rodzina Eupistidae polski. Documenta Physiographica Poloniae 32: 103.

Toll S (1962) Materialien zur Kenntnis des paläarktischen Arten der familie Coleophoridae (Lepidoptera).

Acta Zoologica Cracoviensia 7: 557-719.

Nota Lepi. 38(2) 2015: 157-158 | DOI 10.3897/nl.38.6909

Book Review: Eucosma Hübner of the Contiguous United States and

Canada (Lepidoptera: Tortricidae: Eucosmini)

Joaquin Baixeras1

1 Cavanilles Institute of Biodiversity and Evolutionary Biology, University of Valencia, C/Catedràtic José Beltran 2, 46980

Paterna (Valencia), Spain; joaquin.baixeras@uv.es

Received 20 October 2015; accepted 23 October 2015; published: 6 November 2015

Subject Editor: Jadranka Rota.

Donald Wright and Todd Gilligan (2015): Eucosma Hübner of the Contiguous United States and

Canada (Lepidoptera: Tortricidae: Eucosmini). Wedge Entomological Research Foundation. 256

pp. ISBN 978-0-933003-16-3. Price: €90 or £65.'

Donald Wright is Professor Emeritus at the University of Cincinnati and Todd Gilligan is Research

Scientist at Colorado State University. Both are renowned tortricid experts and they have already

co-authored several publications on the taxonomy and phylogeny of the tortricid subfamily Ole-

threutinae, with special reference to the tribe Eucosmini and more concretely of its type genus

Eucosma Hübner.

The genus Eucosma , with more than 230 species,

is one of the most species-rich genera of tortricids.

Mostly Holarctic in distribution it is especially di-

verse in the Nearctic. It has been considered as one

of the most taxonomically recalcitrant genera, full of

difficulties and misidentifications. Most North Amer-

ican species of Eucosma have been historically placed

in the genus Phaneta and the limits between these

two genera and Pelochrista have remained obscure.

The recent research by these same authors on this ex-

tremely difficult taxonomic area has been absolutely

critical and the present classification of the group can

only be understood through their contribution. In the

words of the authors themselves, “This volume is, in

part, a culmination of that study”. No comprehensive

treatment of the North American Eucosma had been

attempted since Heinrich’s (1923) monograph on the

“Eucosminae” and so this publication comes to fill

an important gap in the lepidopterological literature

Figure 1. The cover of the book. (Fig- 1)-

Published by the Wedge Entomological Research Foundation (New Mexico, USA), the book is distributed in

Europe through Antiquariat Goecke & Evers (Germany) as well as Pemberley Natural History Books (UK).

158

Baixeras: Book Review: Book Review: Eucosma Hübner...

Figure 2. One of the adult habitus plates from the

book.

Wright and Gilligan concentrate their knowl-

edge of North American Eucosma in a mono-

graph of 256 pages, 115 of them beautifully il-

lustrated. The layout of the book is impeccable.

An introduction to the genus provides a useful

historical background. Those interested in de-

scribing or interpreting the morphology of Eu-

cosmini will find especially useful the method-

ological section that includes an account of the

morphology with an interesting character coding.

The 133 species - including nine new species -

studied are distributed into 1 8 groups plus a mis-

cellaneous group of non-assigned species. Every

group is briefly introduced. Each species is then

examined in detail including a list of synonyms,

records and misidentifications, type deposition,

diagnostic comments, and distributional and bi-

ological data when available. Habitus images of

the adults (right side) are illustrated in 29 full col-

our plates (Fig. 2). The head is also photographed

when distinctive characters are relevant. Male

and female genitalia drawings are compiled in

49 plates where Wright feels free to demonstrate

his outstanding qualities as illustrator (Fig. 3).

The sterigma- sternum 7-ductus bursae complex

is figured separately at a useful scale, allowing

accurate details of the female genitalia. Unlike

other faunal works, the variability receives spe-

cial attention and the male genitalia of differ-

ent specimens are illustrated when necessary,

completing a total of 450 adult images and 629

genitalia drawings. The usual appendices (food

plants, taxon names. . .) include a novel and inter-

esting comparative biometric table.

The book is essential for those interested in

Tortricidae. But even if focused on the North

American fauna and a single genus, anyone in-

terested in Lepidoptera will find this publication

attractive, not only for the information provided,

but also as a benchmark to which other authors

can aspire.

Nota Lepi. 38(2) 2015: 159-160 | DOI 10.3897/nl.38.7012

Book Review: The Notodontidae of South Africa

David Agassiz1

1 Department of Life Sciences, The Natural History Museum, London SW7 5 BD; agassiz@btinternet.com

Received 23 September 2015; accepted 19 October 2015; published: 6 November 2015

Subject Editor: Jadranka Rota.

Alexander Schintlmeister and Thomas J. Witt 2015: The Notodontidae of South Africa in-

cluding Swaziland and Lesotho (Lepidoptera: Notodontidae). Proceedings of the Museum

Witt, Volume 2, Munich and Vilnius. 104 distribution maps, 37 colour plates, 42 plates with

genitalia figures, 288 pages. ISBN: 978-3-940732-19-4. Price €89 plus additional postage.1

Figure 1 . The cover of the book “The Notodontidae

of South Africa including Swaziland and Lesotho

(Lepidoptera: Notodontidae)”.

It came as something of a surprise to see this

book. Well-illustrated taxonomic works on the

Lepidoptera of sub-Saharan Africa are seldom

seen. It is nicely bound in A4 format; often such

publications are only available on the Internet,

which has the advantage of availability at low

cost, but a book still has much appeal if it can

be afforded. More of a surprise, when reading

the introduction, was that it appears to have been

conceived only in 2013. A colossal amount of

work must have been put in preparing text and

photographs to a high standard.

The Introduction sets out clearly the way

in which species are to be described, then fol-

lows an historical account of the study of these

moths in South Africa with biographical details

of the chief contributors. Much of the material

described is in the Ditsong National Museum of

Natural History (formerly the Transvaal Muse-

um) in Pretoria.

A checklist of the 99 species treated follows

with 44 names reduced to synonymy which is as

important as the twelve new species and three

genera which are described.

1 The book can be ordered online from the Museum Witt Munich website (http://www.insecta-web.org/

MWM/htmls/museum_proceedlings_en.html).

160

Agassiz: Book Review: The Notodontidae of South Africa

The taxonomic part, including description or redescription of each species, fills the main part of

the book. Each species has the original description cited and is then described under the headings:

diagnosis, bionomics, and distribution. A dot distribution map is included in each case, which in-

cludes all the countries of Southern Africa north to the Zambezi River. In critical cases the descrip-

tions are augmented with illustrations pointing out the differences between closely related species.

For newly described species the label data of the type series is given in full. There follows a full

list of references and a list of the taxonomic changes introduced.

The genitalia figures, occupying 8 1 pages, are photographs in monochrome, male and female of

the same species alongside each other; in many cases the posterior segments of the male abdomen

are also illustrated. The colour plates of adults show life size specimens on a uniform pale blue-

grey background in most cases and are of high quality. For some species there are also photographs

of blown larvae; the data relating to each specimen illustrated are cited in full.

Colour photographs of live specimens and of larvae of some species follow filling the next six

plates, then there are photographs of many of the habitats referred to in the text, indicated also by

adjoining maps.

This work has been well researched, the taxonomic treatment is thorough and well documented

and sensible choices have been made about what information should be included.

For anyone working in Southern Africa this book will be a huge asset and it should also be of

interest to those farther afield who are interested in this family. I commend it wholeheartedly.

Nota Lepi. 38(2)2015: 161-169 | DPI 10.3897/nl.38.6312

Enantiomers of 2-butyl 7Z-dodecenoate are sex attractants for males of

Adscita mannii (Lederer, 1853), Æ geryon (Hübner, 1813), and Jordanita

notata (Zeller, 1847) (Lepidoptera: Zygaenidae, Procridinae) in Italy

Konstantin A. Efetov1, Gerhard M. Tarmann2, Teodora B. Toshova3,

Mitko A. Subchev3

1 Crimean Federal University, Department of Biological Chemistry and Laboratory of Biotechnology, 295006

Simferopol, Crimea; efetov.konst@gmail.com

2 Tiroler Landesmuseen, Ferdinandeum, Naturwissenschaftliche Abteilung, Feldstrasse lia, A-6020 Innsbruck, Austria;

g. tarmann@tiroler-landesmuseen. at

3 Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Str., 1113 Sofia, Bulgaria;

teodoraJoshova@yahoo. com; subchev@yahoo. com

http: //zoobank. org/0D9 7E7B1-EA 1D-4E2D-B6B2-5E62525 77D43

Received 23 August 2015; accepted 30 October 2015; published: 10 November 2015

Subject Editor: Thomas Fartmann.

Abstract. The R- and S-enantiomers of 2-butyl (7Z)-dodecenoate (alone or in mixtures), recently identified

in the natural extracts of Illiberis rotundata pheromone glands, were used as lures in sticky traps to study the

occurrence of Procridinae species in Italy in 14 localities during 2010 and 2011. Three species were attracted

during the study - Adscita mannii (Lederer, 1853), A. geryon (Hübner, 1813), and Jordanita notata (Zeller,

1847). The most numerous species was A. mannii. Lures with (25)-butyl (7Z)-dodecenoate attracted males

of Adscita mannii and A. geryon , while those containing (2i?)-butyl (7Z)-dodecenoate attracted males of

Jordanita notata.

Introduction

Four pheromone compounds were identified in extracts of female sex pheromone glands of Illiberis

(. Primilliberis ) rotundata Jordan, 1907: (2Æ)-butyl (7Z)-dodecenoate [R-7-12], (2*S)-butyl (7Z)-do-

decenoate [S-7-12], (27?)-butyl (9Z)-tetradecenoate [R-9-14] and (2S)-butyl (9Z)-tetradecenoate

[S-9-14] (Subchev et al. 2009). Mixtures of two of them, R-7-12 and R-9-14, were found to be

attractive for the males of I. rotundata and /. (. P. ) pruni Dyar, 1905 (Subchev et al. 2012; 2013).

Our experience with the use of all four components in Bulgaria, Crimea, Hungary, Armenia, Tur-

key, and Afghanistan (Efetov et al. 2008, 2010a, 2010b, 2011; Efetov et al. 2014; Subchev 2014;

Subchev et al. 2010) showed that only R-7-12 and S-7-12 and their mixtures were attractive for dif-

ferent species of the genera Rhagades Wallengren, 1863, Zygaenoprocris Hampson, 1900, Adscita

Retzius, 1783, and Jordanita Verity, 1946. Thus, during our field trips in Italy in 2010 and 2011 we

used only the last two mentioned substances.

Zygaenidae fauna of Italy is represented by 45 species, of which 14 belong to the subfamily Pro-

cridinae (Bertaccini and Fiumi 1999; Efetov 1994, 2004; Efetov and Tarmann 1999, 2000, 2014;

Efetov et al. 2011). The aim of our work was to check the attractiveness of R-7-12 and S-7-12 and

their mixtures for Italian species of Procridinae.

162

Efetov et al. : Enantiomers of 2-butyl 7Z-dodecenoate...

Materials and methods

The pheromone compounds were synthesized at the Institute of Organic Chemistry, Hamburg

University, and pheromone baits and traps were prepared at the Institute of Zoology, Bulgarian

Academy of Sciences. For pheromone baits we used penicillin vial caps of grey rubber on which

the synthetic pheromone compounds were applied as hexane solutions. After evaporation of the

solvent, the caps were wrapped in aluminium foil and kept in a refrigerator at 5 °C until ready for

use. In most cases, sticky Delta traps were used. The removable sticky layers were covered with

Tanglefoot® insect glue. In addition to the sticky traps we also used commercially available traps

(plastic cylinders) for obtaining living material. Moreover, in some localities we also collected

attracted specimens by netting them.

Traps baited with the synthetic Procridinae sex pheromone compounds R-7-12 and S-7-12 alone

and in mixtures were placed and inspected in the field in 14 habitats located from northern Ita-

ly (Alps) to the southern part of the country (Calabria) during the periods 9.vi-18.vi.2010 and

28.vi-8.vii.201 1 . During these two periods the first two authors travelled from the Alps to southern

Italy and placed traps in position during the southwards trip and checked them on the return trip

northwards (Efetov et al. 2012).

List of studied localities in Italy (Figs 1-3)

2010

Province L’Aquila, Rocca di Mezzo, 1322 m, traps placed 9.vi.2010, traps inspected 9.vi. and

17.vi.2010.

Province L’Aquila, Sperone, 1212 m, placed 9.vi.2010, inspected 9.vi., 17.vi. and 18.vi.2010.

Province Latina, Lenola, 534 m, placed 10.vi.2010, inspected lO.vi. and 16.vi.2010.

Province Napoli, Monte Faito, 843 m, placed ll.vi.2010, inspected 1 l.vi.2010.

Province Potenza, Roccarossa, 1370 m, placed 12.vi.2010, inspected 12.vi., 13.vi. and 15.vi.2010.

Province Potenza, Monte Pollino, 1206 m, placed 13.vi.2010, inspected 13.vi., 14.vi. and

15.vi.2010.

Province Cosenza, Lago Ampollino, 1308 m, placed 15. vi. 2010, inspected 15.vi.2010.

2011

Province Verona, Monte, 316 m, placed 28.vi.2011, inspected 8.VÜ.2011.

Province Bologna, Loiano NNW, 248 m, placed 28.vi.2011, inspected 8.VÜ.2011.

Province Potenza, Roccarossa, 1370 m, placed 30.vi.2011, inspected 3.vii., 4.vii. and 5.VÜ.2011.

Province Cosenza, Lago Ampollino, 1308 m, placed 30.vi.2011, inspected l.vii., 2.vii. and

3.VÜ.2011.

Province Potenza, Lagonegro NE, 1340 m, placed 4.vii.201 1, inspected 4.vii. and 5.vii.201 1 .

Province Chieti, Castiglione Messer Marino N, 873 m, placed 5.VÜ.201 1, inspected 6.VÜ.201 1.

Province L’Aquila, Capistrello (1), 757 m, placed 6.VÜ.2011, inspected 7.VÜ.201 1 .

Province L’Aquila, Capistrello (2), 911 m, placed 6.VÜ.2011, inspected 7.VÜ.2011.

Province Roma, Cervara di Roma, 1127 m, locality visited on 7.vii.201 1.

Province Roma, Jenne, 936 m, locality visited on 7.VÜ.201 1 .

Province Trento (Trentino), Monte Bondone, 967 m, trap placed 9.VÜ.2011, inspected 9.VÜ.2011.

NotaLepi. 38(2): 161-169

163

Figure 1. Distribution of Jordanita notata in Italy (blue dots) and studied localities with attracted specimens

(red dots).

Figure 2. Distribution of Adscita geryon in Italy (blue dots) and studied localities with attracted specimens

(red dots).

Figure 3. Distribution of Adscita mannii in Italy (blue dots) and studied localities with attracted specimens

(red dots).

164

Efetov et al.: Enantiomers of 2-butyl 7Z-dodecenoate...

Results

Three Procridinae species were attracted to R-7-12 and S-7-12 and their mixtures (Figs 1-3), viz.

Adscita ( Tarmannita ) mannii (Lederer, 1853), A. (Adscita) geryon (Hübner, 1813), and Jordanita

( Tremewania ) notata (Zeller, 1 847). Jordanita notata males (9 specimens) were attracted to R-7-12

or mixtures containing this compound in two habitats in the province of L’ Aquila and one habitat

in the province of Potenza (Fig. 4, Table 1). Adscita geryon males (4 specimens) were attracted in

two habitats in the province of L’ Aquila (Table 2). A. mannii males (136 specimens) were attracted

to S-7-12 or mixtures containing this compound in the provinces of Trentino, Bologna, L’ Aquila,

Roma, Latina, Chieti, Napoli, Potenza, and Cosenza (Fig. 5, Table 3). Twelve specimens of A.

mannii in the habitats Sperone (2010), Cervara di Roma (2011), and Jenne (2011) were attracted to

the box containing all substances immediately after opening the bag, which is why they were not

included in Table 3.

When all three variants of the attractant were present in the habitat (S-7-12, R-7-12 and their

mixture), A. mannii came mainly to S-7-12 (up to 38 males were attracted to one trap over a period

of three days at Roccarossa). When we had only R-7-12 and the mixture of R-7-12 and S-7-12,

some specimens were also in traps containing the mixture. Furthermore, in Monte Pollino (2010)

and Loiano (201 1) where A. mannii was abundant and where we placed only one trap baited with

R-7-12, we found one and five males respectively in the traps (Table 3).

Figure 4. Sticky trap baited with R-7-12 with two males of Jordanita notata , Roccarossa, 15.vi.2010.

Nota Lepi. 38(2): 161-169

165

1 66 Efetov et al. : Enantiomers of 2-butyl 7Z-dodecenoate...

Nota Lepi. 38(2): 161-169

167

Figure 5. Sticky trap baited with S-7-12 with 38 males of Adscita mannii , Roccarossa, 3.VÜ.2011.

Discussion

A. mannii is distributed from north-eastern Spain and south-western France through the southern

parts of central Europe, Italy (including Sicily) and the Balkans to eastern Romania, Greek islands

(except Crete), and north-western Turkey. A. geryon is known from the Iberian Peninsula and

Great Britain (England, Wales) through most of central and southern Europe to Moldavia, south

of European Russia and north-western Turkey. J. notata is distributed in western and central Eu-

rope, northern Mediterranean (including Sicily and Crete) to Ukraine, Crimea, Northern Caucasus,

Transcaucasia, Turkey, and north-western Iran (Efetov and Tarmann 1999; Efetov 2004). The dis-

tributions of these three species in Italy are shown in Figs 1-3 (blue dots).

Our results confirm the data obtained earlier in Bulgaria and the Crimea that R-7-12 is an at-

tractant for Jordanita notata , while S-7-12 attracts Adscita geryon and A. mannii (Subchev et al.

2010) . It is interesting to note that the presence of S-7-12 does not inhibit the attractiveness of

R-7-12 for J. notata and the presence of R-7-12 also does not influence the attractiveness of S-7-12

for A. geryon and A. mannii. In July 2007, during an expedition to Armenia, K. A. Efetov and V.

M. Kiselev obtained the opposite result with Zygaenoprocris taftana (Alberti, 1939) (Efetov et al.

2011) , a species from the subgenus Molletia Efetov, 2001, which was attracted to R-7-12, while

168

Efetov et al: Enantiomers of 2-butyl 7Z-dodecenoate...

the presence of S-7-12 completely cancelled the attractiveness of R-7-12. This means that S-7-12

is an inhibitor of R-7-12 for Z. taftana. The genus Zygaenoprocris is represented by four subge-

nera (Efetov 2001a; 2001b). It looks as though the same situation as found in Molletia is present

in two species of another subgenus, viz. Zygaenoprocris Hampson, 1900. During an expedition

to Afghanistan in July 2011 A. Hofmann attracted Zygaenoprocris {Zygaenoprocris) eberti (Al-

berti, 1968) and Z (Z.) chalcochlora Hampson, 1900, to R-7-12 and a mixture of R-7-12+R-9-14

(Subchev 2014; Efetov, Hofmann and Tarmann 2014).

Jordanita notata belongs to the subgenus Tremewania Efetov & Tarmann, 1 999, Adscita geryon to

the subgenus Adscita Retzius, 1783, and A. mannii to the subgenus Tarmannita Efetov, 2000 (Efetov

and Tarmann 1999; Efetov 2004). Additional investigations are necessary to find attractants for repre-

sentatives of the three other subgenera of Procridinae that inhabit Italy, viz. Roccia Alberti, 1954 (one

species), Jordanita Verity, 1946 (three species), and Solaniterna Efetov, 2004 (one species).

It seems that the same attractants can be active for different species of the subgenus, but the

attractiveness for different species can depend on the ratio of the components in the mixture. For

example, the subgenus Primilliberis Alberti, 1954, of the genus Illiberis Walker, 1854, includes

four species (Efetov 1997; Efetov and Tarmann 2012), and in two of them, viz. I. (P) rotundata

and I. {P.) pruni , the males were attracted by different ratios of R-7-12 and R-9-14 (Subchev et

al. 2012; 2013). It is possible that a similar situation can be present in Adscita {Adscita), Adscita

{Tarmannita) and Jordanita {Tremewania). The confirmation of this needs further investigations.

Acknowledgements

We thank our friend Dr W. G. Tremewan (Great Britain) for his company and help during our field trips in Italy in 2010 and

2011 and for editing the English text. This research was partially supported by a Grant D002-244/2008 from the Bulgarian

National Scientific Fund.

References

Alberti B (1954) Über die stammesgeschichtliche Gliederung der Zygaenidae nebst Revision einiger Grup-

pen (Insecta, Lepidoptera). Mitteilungen aus dem zoologischen Museum der Humboldt-Universität Berlin

30: 1 1 5 — 480. doi: 10.1002/mmnz. 19540300202

Alberti B (1968) Zur Kenntnis des Genus Procris F. in Afghanistan (Lepidoptera, Zygaenidae). Reichenba-

chia 10: 249-253.

Bertaccini E, Fiumi G (1999) Bombici e Sfingi d’Italia (Lepidoptera Zygaenidae). Natura, Giuliano Russo

Editore, Monterenzio, 160 pp.

Efetov KA ( 1 994) Jordanita ( Gregorita ) algirica (Rothschild, 1917) (Lepidoptera, Zygaenidae) - a new for-

ester for the fauna of Europe. Vestnik zoologii 1 : 36.

Efetov KA (1997) Three new species of the genus Illiberis Walker, 1854, from Taiwan and Vietnam (Lepido-

ptera: Zygaenidae, Procridinae). Entomologist’s Gazette 48: 231-244.

Efetov KA (2001a) On the systematic position of Zygaenoprocris Hampson, 1900 (Lepidoptera: Zygaenidae,

Procridinae) and the erection of two new subgenera. Entomologist’s Gazette 52: 41^48.

Efetov KA (2001b) An annotated check-list of Forester moths (Lepidoptera: Zygaenidae, Procridinae). Ento-

mologist’s Gazette 52: 153-162.

Efetov KA (2004) Forester and Bumet Moths (Lepidoptera: Zygaenidae). The genera Theresimima Strand,

1917, Rhagades Wallengren, 1863, Zygaenoprocris Hampson, 1900, Adscita Retzius, 1783, Jordanita

Verity, 1946 (Procridinae), and Zygaena Fabricius, 1775 (Zygaeninae). CSMU Press, Simferopol, 272 pp.

Nota Lepi. 38(2): 161-169

169

Efetov KA, Can F, Toshova TB, Francke W, Subchev MA (2010a) Catches of Procridinae (Lepidoptera: Zy-

gaenidae) by pheromone traps in Hatay, Turkey (2009). In: Can F (Ed.) XII International Symposium on

Zygaenidae, Hatay, Turkey, 5-9 May 2010. Hatay, 14.

Efetov KA, Can F, Toshova T, Subchev M (2010b) New sex attractant for Jordanita anatolica (Naufock)

(Lepidoptera: Zygaenidae: Procridinae). Acta zoologica bulgarica 62: 315-319.

Efetov KA, Hofmann A, Tarmann GM (2014) Application of two molecular approaches (use of sex attractants

and DNA barcoding) allowed to rediscover Zygaenoprocris eberti (Alberti, 1968) (Lepidoptera, Zygaeni-

dae, Procridinae), hitherto known only from the female holotype. Nota Lepidopterologica 37: 151-160.

doi: 10.3897/nl.37.7871

Efetov KA, Kiselev VM, Subchev M, Toshova TB, Francke W (2008) Catches of Procridinae (Lepidoptera:

Zygaenidae) by pheromone traps in Armenia. XI International Symposium on Zygaenidae, Sofia, Bulgaria,

17-21 September 2008. Sofia, 15.

Efetov KA, Subchev MA, Toshova TB, Kiselev VM (2011) Attraction of Zygaenoprocris taftana (Alberti,

1939) and Jordanita horni (Alberti, 1937) (Lepidoptera: Zygaenidae, Procridinae) by synthetic sex phero-

mones in Armenia. Entomologist’s Gazette 62: 113-121.

Efetov KA, Tarmann GM (1999) Forester Moths. The genera Theresimima Strand, 1917, Rhagades Wallen-

gren, 1863, Jordanita Verity, 1946, and Ads cita Retzius, 1783 (Lepidoptera: Zygaenidae, Procridinae).

Apollo Books, Stenstrup, 192 pp.

Efetov KA, Tarmann GM (2000) On the systematic position of Procris alpina italica Alberti, 1937 and Proc-

ris storaiae Tarmann, 1977 (Lepidoptera: Zygaenidae, Procridinae). Tavricheskiy mediko-biologicheskiy

Vestnik 3(1-2): 161-167.

Efetov KA, Tarmann GM, Tremewan WG (2011) Zygaena nevadensis Rambur, 1858 (Lepidoptera: Zygaeni-

dae, Zygaeninae) newly recorded from the southern tip of the Penisola Appenninica (Apennine Peninsula),

Italy. Entomologist’s Gazette 62: 123-129.

Efetov KA, Tarmann GM (2012) A checklist of the Palaearctic Procridinae (Lepidoptera: Zygaenidae). CSMU

Press, Simferopol - Innsbruck. 108 pp.

Efetov KA, Tarmann GM (2014) A new European species, Adscita dujardini sp. nov. (Lepidoptera: Zygaeni-

dae, Procridinae) confirmed by DNA analysis. Entomologist’s Gazette 65: 179-200.

Efetov KA, Tarmann GM, Toshova T, Subchev MA (2012) Attraction of Adscita mannii (Lederer, 1853),

A. geryon (Hübner, 1813) and Jordanita notata (Zeller, 1847) (Lepidoptera: Zygaenidae, Procridinae) to

2-butyl 7Z-dodecenoate in Italy. In: Tarmann GM, Tremewan WG, Young MR (Eds) XIII International

Symposium on Zygaenidae, Innsbruck, Tirol, Austria, 16-23 September 2012. Innsbruck, 15.

Subchev M (2014) Sex pheromone communication in the family Zygaenidae (Insecta: Lepidoptera): a review.

Acta zoologica bulgarica 66: 147-158.

Subchev M, Efetov KA, Toshova T, Parshkova EV, Toth M, Francke W (2010) New sex attractants for species of

the zygaenid subfamily Procridinae (Lepidoptera: Zygaenidae). Entomologia Generalis (Stuttgart) 32: 243-250.

Subchev MA, Koshio C, Toshova TB, Efetov KA (2012) Illiberis ( Primilliberis ) rotundata Jordan (Le-

pidoptera: Zygaenidae: Procridinae) male sex attractant: Optimization and use for seasonal monitoring.

Entomological Science 15: 137-139. doi: 1 0. 1 1 1 l/j. 1 479-8298.20 1 1 .00485.x

Subchev M, Koshio C, Toshova T, Efetov KA, Francke W (2013) (2R)-butyl (7Z)-dodecenoate, a main sex

pheromone component of Illiberis ( Primilliberis ) pruni Dyar (Lepidoptera: Zygaenidae: Procridinae)?

Acta zoologica bulgarica 65: 391-396.

Subchev M, Toshova T, Koshio C, Franke S, Tröger A, Twele R, Francke W, Pickett JA, Wadhams LJ,

Woodcock CM (2009) Identification and biological activity of sex pheromone components from fe-

males of the plum moth Illiberis rotundata Jordan (Lepidoptera: Zygaenidae: Procridinae). Chemo-

ecology 19: 47-54. doi: 10.1007/s00049-009-0008-8

SOCIETAS EUROPAEA LEPIDOPTEROLOGICA e.V.

Nota Lepidopterologica wird als wissenschaftliche Zeitschrift von der Societas Europaea Lepidopterologica

(SEL) herausgegeben und den Mitgliedern der SEL zugesandt. Autoren, die Manuskripte fur die Publika-

tion in der Nota Lepidopterologica einreichen möchten, finden die jeweils gültigen Autorenrichtlinien auf

der Homepage unter http://nl.pensoft.net. Der Verkauf von Einzelheften und älteren Jahrgängen von Nota

lepidopterologica erfolgt durch das Antiquariat Goecke & Evers (Inh. Erich Bauer), Sportplatzweg 5, 75210

Keltern, Deutschland, E-Mail: books@insecta.de (www.insecta.de, www.goeckeevers.de); der Buchhan-

delspreis beträgt € 100,00 pro Band. Die Mitgliedschaft bei der SEL steht Einzelpersonen und Vereinen

nach Maßgabe der Satzung offen. Der Aufnahmeantrag ist an den Mit-gliedssekretar Willy De Prins, Dorp-

straat 401 B, 3061 Leefdaal, Belgien; e-mail: willy.de.prins@telenet.be zu richten. Das Antragsformular

ist im Internet auf der Homepage der SEL 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) er-

beten (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,

76133 Karlsruhe, Germany; 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 pub-

lication in Nota lepidopterologica are asked to take into consideration the relevant instructions for authors

available on the homepage at http://nl.pensoft.net. Single and back issues of Nota lepidopterologica can be

obtained from Antiquariat Goecke & Evers (Prop. Erich Bauer), Sportplatzweg 5, 75210 Keltern, Germa-

ny, E-Mail: books@insecta.de (www.insecta.de, www.goeckeevers.de); price € 100,00 per volume. 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, 3061 Leef-

daal, 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 für Naturkunde, Erbprinzenstr. 13, 76133

Karlsruhe, Germany; 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 sci-

entifique 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 Nota: http://

nl.pensoft.net. 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 Antiquariat Goecke & Evers (Erich Bauer, prop.), Sportplatzweg 5,75210 Keltern, Allemagne, cour-

riel: books@insecta.de (www.insecta.de, www.goeckeevers.de); en librairie, le prix est de € 100,00 le

volume. Tel que prévu dans ses statuts, les individus de même que les associations peuvent devenir mem-

bres de la SEL. Les demandes d’adhésion doivent être envoyées au Secrétaire responsable des adhésions,

Willy De Prins, Dorpstraat 401 B, 3061 Leefdaal, Belgique; courriel: willy.de.prins@telenet.be. Le for-

mulaire 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 indi vidus 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; IB AN: 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, 76133 Karlsruhe, Germany;

courriel: trusch@smnk.de. Tout changement d’adresse doit être mentionné immédiatement au Secrétaire

responsable des adhésions ou au Trésorier.

Contents

SMITHSONIAN LIBRARIES

3 9088 01854 6861

mai

Biology and biogeography of the chalk-hill blue Polyommatus coridon —

insect of the year 2015 for Germany, Austria and Switzerland

Thomas Schmitt

pgil|

'' ••A-vl'-S

KHHf

\ " , ’

Transfer of Pygmaeotinea crisostomella Amsel, 1957 from Tineidae to Psychidae

and its taxonomic status (Lepidoptera)

Thomas Sobczyk

A new species of Micropterix Hübner, 1825 from the Orobian Alps (Italy)

(Lepidoptera, Micropterigidae)

Hans Christof Zeller, Peter Huemer

Coleophora gryphipennella (Hübner, 1796) (Lepidoptera, Coleophoridae) on

Fragaria vesca L. (Rosaceae), a novel host, in the coastal dunes of The Netherlands

John A.M. van Roosmalen, Camiel Doorenweerd

WÊSÊ

itilll

Book Review: Eucosma Hübner of the Contiguous United States and Canada

(Lepidoptera: Tortricidae: Eucosmini)

Joaquin Baixeras

Book Review: The Notodontidae of South Africa

David Agassiz

Enantiomers of 2-butyl 7^-dodecenoate are sex attractants for males of Adscita

mannii (Lederer, 1853), A geryon (Hübner, 1813), and Jordanita notata (Zeller,

1847) (Lepidoptera: Zygaenidae, Procridinae) in Italy

Konstantin A. Efetov, Gerhard M. Tarmann, Teodora B. Toshova, Mitko A. Subchev

http://nl.pensoft.net