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NOTA LEPIDOPTEROLOGICA

A journal focussed on Palaearctic and General Lepidopterology

Published by the Societas Europaea Lepidopterologica e.V.

Editor. Jadranka Rota

Associate Editor. Adrian Spalding

Editorial Board. Franziska Bauer (Dresden, D), Sven Erlacher (subject editor; Chemnitz, D),

Thomas Fartmann (subject editor; Münster, D), Zdenék F. Fric (subject editor; Ceské Budéjo-

vice, CZ), Axel Hausmann (subject editor; Munich, D), Peter Huemer (subject editor; Inns-

bruck, A), Lauri Kaila (subject editor; Helsinki, FI), Ole Karsholt (Copenhagen, DK), Bernard

Landry (subject editor, Genève, CH), Carlos Lopez-Vaamonde (subject editor; Orléans, F),

Vazrick Nazari (subject editor: Ottawa, CA), Erik J. van Nieukerken (subject editor; Leiden, NL),

Matthias Nuss (Dresden, D), Thomas Schmitt (subject editor; Trier, D), Wolfgang Speidel (Bonn, D),

Alberto Zilli (subject editor; Rome, I).

ISSN 0342-7536

Type setting: blattwerk | dd

Printed by Druckhaus Dresden GmbH

mechanical including photocopying, recording or any other information storage and retrieval system, without written

permission from the publisher. Authors are responsible for the contents of their papers.

NOTA LEPIDOP'TEROLOGICA

Volume 36 No.1 + Dresden, 17.06.2013 - ISSN 0342-7536

David Agassiz. Obituary: Paul Sokoloff (1946-2012) "ss 3

Wolfgang Wagner. Observations on the preimaginal ecology of Rhynchina canariensis

Pinker, 1962 (Erebidae: Hypeninae) and Abrostola canariensis Hampson, 1913

(Noctuidae: Plusiinae) on the Canary island of La Gomera .............cccccccceeesneeeceeeeesteeeeeeens 3

Ole Karsholt. Monochroa bronzella sp. n. from the southwestern Alps (Gelechiidae) .......... 13

Ivan N. Bolotov, Mikhail Yu. Gofarov, Alexander M. Rykov, Artyom A. Frolov &

Yaroslava E. Kogut. Northern boundary of the range of the Clouded Apollo butterfly

Parnassius mnemosyne (L.) (Papilionidae): climate influence or degradation

olelarvalhost plants Ye nee seen re ee nalen isn 19

Adrian Spalding, Iva Fukova & Richard H. Ffrench-Constant. The genetics

of Luperina nickerlii Freyer, 1845 in Europe (Noctuidae) .......uueeeeeeeeeeeesssenennnnnnneeneneeenenn 35

Stanislav K. Korb. The status of Satyrus abramovi var. korlana Staudinger, 1901

CN ne ers ee ee 47

Frans Groenen & Joaquin Baixeras. The “Omnivorous Leafroller”, Platynota stultana

Walsingham, 1884 (Tortricidae: Sparganothini), a new moth for Europe 53

Ozge Ozden. Habitat preferences of butterflies (Papilionoidea)

Hight IG arpaz DERINSUI OV DEUST ee ton eue 57

Ivan N. Bolotov, Mikhail Yu. Gofarov, Yulia S. Kolosova & Artyom A. Frolov.

Occurrence of Borearctia menetriesii (Eversmann, 1846) (Erebidae: Arctiinae)

in Northern European Russia: a new locality in a disjunct species range ...........n. 65

Oleksiy V. Bidzilya & Ole Karsholt. Two little-known species of Gelechiidae

HER Et UU @ [YAMA Fauna ware nennen sae sbivzardessaincs sotet sd pao ystcdivunscetledese sete 77

DO TEE Te TE eo 12, 85, 87

SMITHSON] a=

JUL 18 2013

LIBRARIES

Nota lepid. 36 (1): 3-4 3

Paul A. Sokoloff

1946 - 2012

Like many amateur entomologists Paul Sokoloff experienced a tension between his

entomology and his professional life. During his early career, even though working

towards further qualifications, he found time for fieldwork, especially near his home

in southeast London. He was interested in all Lepidoptera in Britain, especially the

Gelechiidae. He became active in entomological societies, being elected President

of the British Entomological & Natural History Society for 1984. He had published

an illustrated paper on the genera Teleiodes and Teleiopsis, he also updated a publi-

cation of the Amateur Entomologists’ Society Practical hints for collecting and stu-

dying Microlepidoptera in 1980 and produced a handbook Breeding the British and

European Hawk-moths in 1984. In 1985 he took over as editor of the Entomologist s

Record. His interest in literature and his ability with words were put to good use in

this role. After ten years he resigned when he took on more demanding professional

duties in a leading UK examinations board. In retirement he resumed his entomologi-

cal activity and became a member of Nota lepidopterologica’s editorial team. He met

and collaborated with other editors even though he had never managed to attend one

of the SEL Congresses. His editorial skill and the effort that he put into improving ma-

Nota lepidopterologica, 17.06.2013, ISSN 0342-7536

- Obituary: Paul A. Sokoloff (1946-2012)

nuscripts in English published in Nofa was greatly appreciated by the scientific editors,

as well as the numerous authors whom he helped. Regrettably he was diagnosed with

cancer in June 2012 and died in November. Our sympathy is extended to his widow

Linda and their son and daughter.

Davip AGASSIZ

Nota lepid. 36 (1): 5-11 5

Observations on the preimaginal ecology of

Rhynchina canariensis Pinker, 1962 (Erebidae: Hypeninae)

and Abrostola canariensis Hampson, 1913 (Noctuidae: Plusiinae)

on the Canary island of La Gomera

WOLFGANG WAGNER

Anton-Hohl-Str. 21a, D-87758 Kronburg, www.pyrgus.de; wolfgang@pyrgus.de

Received 18 June 2012; reviews returned 4 September 2012; accepted 24 September 2012.

Subject Editor: Alberto Zilli.

Abstract. In this work some information (including photos) is provided on larvae and preimaginal ecology

of two Canarian endemics Abrostola canariensis Hampson, 1913 and Rhynchina canariensis Pinker, 1962

from La Gomera. Larvae of R. canariensis were observed in Vallehermoso on Lotus emeroides R. P. Murray

(Fabaceae). They inhabit stony, semidry slopes with Juniperus turbinata Guss. (Cupressaceae) where there

are stands of L. emeroides on more or less open ground. The brownish, elongate larvae resemble those of

Zekelita antiqualis (Hübner, 1809). Eggs and larvae of Abrostola canariensis were found on Parietaria

Judaica L. (Urticaceae) on not too dry or partially shaded rocky slopes and especially stone walls made

of natural stone in cultivated or abandoned areas. Parietaria L. spp. should be the main host plants of this

species and Urtica L. spp. are likely to be used only occasionally.

Introduction

The Canary Islands are famous for their high rate of endemic plants and insects. While

the species composition is relatively well known, the preimaginal stages and bionomics

of many species are still in need of detailed study.

Rhynchina canariensis Pinker, 1962 (Erebidae: Hypeninae) and Abrostola cana-

riensis Hampson, 1913 (Noctuidae: Plusiinae) are both endemic to the Canary Islands.

While the former is known from Tenerife eastwards, the latter inhabits all islands of the

archipelago (Baez 1998; Hacker & Schmitz 1996). The larva and relevant life habits

were fully unknown in the case of R. canariensis and poorly known in that of A. cana-

riensis. The latter is said to use Urtica urens L. as host plant (e.g., Hacker & Schmitz

1996), but no reliable field observations have been published so far.

During a trip to La Gomera in December 6—19, 2011 the author had the chance to

find eggs and larvae of A. canariensis and larvae of R. canariensis, the latter being new

to La Gomera.

Material and methods

Eighteen larvae of Rhynchina canariensis were found on December 8 in Vallehermoso

(La Gomera, Canary Islands, Spain) at about 400 m above sea level by careful investi-

gation of Lotus emeroides R. P. Murray stands. As to Abrostola canariensis, five larvae

and two eggs were found in several localities (Vallehermoso, Agulo) on La Gomera

between December 8-15, 2011 by searching Parietaria judaica L. stands. The lar-

vae were successfully reared in small glass containers with perforated caps to avoid

Nota lepidopterologica, 17.06.2013, ISSN 0342-7536

6 WAGNER: Preimaginal ecology of Rhynchina canariensis and Abrostola canariensis on La Gomera

Fig. 2. Larval habitat of Rhynchina canariensis:

slopes with partially open ground in Vallehermoso

(La Gomera, December 2011).

excessive moisture at room temperatures

(18-20°C), and taxonomic identifications

were confirmed after they attained the im-

aginal stage.

Additionally, an ex-ovo rearing of

A. canariensis after oviposition of a female

(Fig. 1) from Valle Gran Rey (found at an

illuminated building) has been carried out

Fig. 1. Imago of Abrostola canariensis (La Gome- under the same conditions as mentioned

ra, Valle Gran Rey, December 2011). above.

Results

Bionomics. Larvae of Rhynchina canariensis inhabit dry to semidry, stony or rocky

slopes with partially exposed soil (Fig. 2) where procumbent shoots of Lotus emeroides

grow on mostly open ground in the “succulent” belt between the sea level and approxi-

mately 600-700 m above sea level. On La Gomera the species is obviously restricted to

the Juniperus turbinata Guss. (Cupressaceae) dominated slopes between Vallehermoso

and Hermigua where the observed host plant Lotus emeroides grows. This plant species

is endemic to La Gomera. Larvae of R. canariensis had been found already in December

2009 in the same locality, though nearly at the sea level, but rearing had failed so that

they could not have been identified. The larvae hide by day, stretching themselves along

the lower parts of the procumbent shoots (Fig. 3) of the host plants and in later instars

they feed preferentially at night. In captivity the moths emerged after 14 to 18 days of

pupal phase.

Larvae of Abrostola canariensis were observed on Parietaria judaica which grows

on walls bordering roads and fields (Fig. 4) or on rocks. The young, whitish green lar-

vae rest on the lower side of the leaves while in the last instar they tend to hide at the

base of the plant during daytime. The eggs (Fig. 5) were found singly on the lower side

Nota lepid. 36 (1): 5-11 7

Fig. 3. Larval habitat of Rhynchina canariensis: Fig. 4. Larval habitat of Abrostola canariensis at

Lotus emeroides on partially open ground at a slope Agulo (La Gomera, December 2011): rocks and

in Vallehermoso (La Gomera, December 2011). walls with Parietaria judaica.

of the leaves. The occupied plants were mostly growing isolatedly in rock and stone

niches on at least partially sunny ground. Pupation took place in captivity between the

end of December and January; all pupae (n = 12) entered dormancy and moths did not

emerge until late April and May 2012.

Searches for larvae of A. canariensis on Urtica urens in Fuerteventura (Pico de

la Zarza and above Cofete) in February 2011 did not result in any specimens except

for those of Vanessa vulcanica (Godart, 1819), which is rare on this eastern island.

Another examination of Urtica morifolia Poir. on La Gomera was also not successful

and resulted only in larvae of Vanessa vulcanica and Mniotype schumacheri (Rebel,

1917).

Habitus. The larvae (Figs 6—11) of Rhynchina canariensis are brownish, the first two

pairs of prolegs are reduced. They bear a variably broad (viz. not parallel-sided) darker

dorsal field which is bordered by a slightly white and then dark area. The ventral side

is light coloured, almost whitish. The head shows a darker finely reticulated pattern

and especially two large dark spots. The pupa (Fig. 12) is light yellowish to reddish

brown.

Young larvae of Abrostola canariensis are whitish green (Figs 13-14) and thus

well matching the lower sides of Parietaria leaves. In the last instar their colour ranges

from greenish yellow to light brown (Figs 15—16) with several small whitish marks

and speckles. The larva is similar to that of A. triplasia (Linnaeus, 1758), but, for ex-

ample, the dorsal markings on the fourth and fifth segments are different: dark triangles

point towards the head in A. triplasia whereas there are oppositely oriented subtriangu-

lar markings in A. canariensis. Additionally, the number of white spots and their size

and arrangement is different (e.g., two larger spots at the sides of the triangle of the

fourth segment in A. friplasia).

Typical traits of Abrostola Ochsenheimer, 1816 are well expressed: prolegs on ab-

dominal segments 3 —6, transverse, semicircular flecks on the dorsal zone of abdominal

segments 1, 2 and 8. The pupa (Fig. 17) is brown and does not differ significantly from

those of its European congeners.

8 WAGNER: Preimaginal ecology of Rhynchina canariensis and Abrostola canariensis on La Gomera

ES me “ Ed

Fig. 5. Egg of Abrostola canariensis (La Gomera, Fig. 6. Young larva of Rhynchina canariensis (La

December 2011). Gomera, Vallehermoso, December 2011)

Fig. 7. Larva of Rhynchina canariensis in the last Fig. 8. Larva of Rhynchina canariensis in the last

instar (lateral view). instar (dorsal view).

Discussion

Rhynchina canariensis (Fig. 18) is a xerothermophilous species of lower and middle

elevations, as shown by the localities where adults have been captured, mainly at light

(e.g., Hacker & Schmitz 1998; Pinker 1962). The species is not restricted to slopes, but

can also be found in drier coastal plains. On islands other than La Gomera the moth

must evidently rely on other Lotus spp. such as Lotus lancerottensis Webb et Berth.,

Lotus glaucus Dryand. in Aiton, Lotus glinoides Delile or Lotus campylocladus Webb

et Berth., which are locally abundant in biotopes where R. canariensis occurs (e.g., in

the low hills and valleys around Betancuria on Fuerteventura). It is questionable but

it should be examined whether R. canariensis is able to develop on other genera of

Fabaceae as well. Last instar larvae supplied in captivity with Onobrychis viciifolia

Scop. (Fabaceae) did not accept this plant. The larvae resemble in both external appear-

ance and behaviour those of Zekelita antiqualis (Hübner, 1809) (cf. Beck 1999), which

belongs to a closely related genus within the subfamily Hypeninae (Mayerl & Lédl

1997). For example, the larval head markings (Fig. 11) are very similar to each other.

Interestingly, larvae of Rhynchina (and Zekelita) show some characters commonly ob-

Nota lepid. 36 (1): 5-11

Fig. 9. Fully grown (some days prior to pupation) last Fig. 10. Fully grown last instar larva of Rhynchina

instar larva of Rhynchina canariensis (dorsal view). canariensis (lateral view).

Fig. 11. Head of larva of Rhynchina canariensis in Fig. 12. Pupa of Rhynchina canariensis (ventral view,

the last instar. cocoon removed)

Fig. 13. Larva of Abrostola canariensis in penulti- Fig. 14. Larva of Abrostola canariensis in penulti-

mate instar (La Gomera, Vallehermoso, December mate instar, dorsal view (La Gomera, Vallehermoso,

2011). December 2011).

served within the subfamily Catocalinae, e.g., the non-parallel sided darker dorsal field,

the overall shape, and their behaviour. The higher classification of Noctuoidea has been

in great flux recently and the closer affinity of some subfamilies formerly assigned

10 WAaGneR: Preimaginal ecology of Rhynchina canariensis and Abrostola canariensis on La Gomera

Fig. 15. Last instar larva of Abrostola canariensis Fig. 16. Last instar Larva of Abrostola canariensis

(lateral view). (dorsal view).

Fig. 17. Pupa of Abrostola canariensis (cocoon re- Fig. 18. Adult female of Rhynchina canariensis, ex

moved). larva, Vallehermoso, December 2011.

to Noctuidae in the old sense such as Hypeninae and Catocalinae is reflected by their

placement in the newly established family Erebidae (cf. Lafontaine & Fibiger 2006;

Zahiri et al. 2011).

Abrostola canariensis is also an inhabitant of semidry, rocky slopes of the “suc-

culent” belt and cultivated areas, and secondarily of stone walls along roads or between

fields. As Parietaria judaica is relatively widespread on the islands, it should be the

most important host plant for this species. In the literature there are hints and espe-

cially presumptions of Urtica being the host plant of A. canariensis. However, my

own examination of Urtica urens on Fuerteventura did not yield any larvae, but as the

larvae did accept Urtica dioica L. in captivity, it is likely that Urtica urens is a host

plant in nature, too. Urtica morifolia, as an endemic member of the genus Urtica L.,

which grows especially in the so called “Laurisilva”, is probably not suited because

of the cool microclimate prevailing in the humid areas where such wood formations

usually occur. Probably the moth also uses other Parietaria spp. such as the endemic

Parietaria filamentosa Webb & Berth. Rearing results indicate that this species is able

to survive the dry summer period in pupal dormancy in the same way as its Central

Nota lepid. 36 (1): 5-11 pi

European allies do during the cold winters. Sometimes there are hints about the occur-

rence of Abrostola canariensis on the Ilhas Selvagens which are located between the

Canaries and Madeira and belong to Portugal (e.g., the Fauna Europaea project), but

the species is not mentioned in the cited paper (Aguiar & Karsholt 2006).

References

Aguiar, A. M. F. & O. Karsholt 2006. Systematic catalogue of the entomofauna from the Madeira archipe-

lago and Selvages Islands. Lepidoptera. — Boletim do Museu Municipal do Funchal, Suppl. 9: 5— 139.

Baez, M. 1998. Mariposas de Canarias. — Editorial Rueda, Alcorcon (Madrid). 216 pp.

Beck, H. 1999. Die Larven der europäischen Noctuidae — Revision der Systematik der Noctuidae. — Her-

bipoliana 5: Vol. 1-4. 2160 pp.

Hacker, H. & W. Schmitz 1996. Fauna und Biogeographie der Noctuidae des makaronesischen Archipels

(Lepidoptera). — Esperiana 4: 167-221.

Lafontaine, J. D. & M. Fibiger 2006. Revised higher classification of the Noctuidae (Lepidoptera). — Ca-

nadian Entomologist 138: 610—635.

Mayerl, B. & M. Lédl 1997. Checkliste aller Arten der Gattungen Rhynchina Guenée, 1854 und Zekelita

Walker, 1863 der Paläarktischen und Indoaustralischen Region (Lepidoptera: Noctuidae: Hypeninae). —

Annalen des Naturhistorischen Museums in Wien 99B: 377-386.

Pinker, R. 1962. Interessante und neue Funde und Erkenntnisse fiir die Lepidopterenfauna der Kanaren.

I. Fortsetzung und Schluß. — Zeitschrift der Wiener Entomologischen Gesellschaft 47 (11): 169-179.

Zahiri, R., I. J. Kitching, J. D. Lafontaine, M. Mutanen, L. Kaila, J. D. Holloway & N. Wahlberg 2011. A

new molecular phylogeny offers hope for a stable family-level classification of the Noctuoidea (Lepi-

doptera). — Zoologica Scripta 40: 158-173.

12 Book review

Objectiu Natura — Associacié de Fotögrafs de Natura de Catalunya (ed.) 2012.

Mariposas por la vida. Guia visual de las mariposas ibéricas diurnas. — Objectiu Natura,

Barcelona, Spain, 255 pp. ISBN 978-84-616-1072-3. Price: 29.95 € plus shipping costs

(order information can be obtained at http://www.mariposasporlavida.org).

Over one hundred, mainly Spanish, photographers have contributed more than 1,000 photos.

Like all the other people involved, they have freely created a book that is special in many ways.

With “mariposas por la vida” (“butterflies for life”) a field guide based on photos, or simply

a “visual guide”, was published in November 2012, covering all the butterfly species of the

Iberian Peninsula and the Balearic Islands. Two hundred and twenty-nine species of the families

Papilionidae, Pieridae, Nymphalidae, Lycaenidae, Riodinidae and Hesperiidae are included. In

general, the book dedicates one page to each species, most of them with three coloured photos,

usually showing the upper- and the underside of the animals. All photos were taken in the field,

showing living specimens in their natural habitat. The information on each species is completed

by illustrations on distribution, flight time, butterfly size and degree of threat (according to

IUCN) as well as — sometimes a bit sparse — information on the larval food plants. Five ad-

ditional pages show pictures by photographers whose photos have not been published on the

pages dedicated to the species.

Being a “visual guide”, as the Spanish subtitle says, supplementary texts were deliberately

avoided. The taxonomy is up to date, the photos are of very good quality and aesthetically

impressive. For that reason alone the purchase of this book, which is moderately priced, can

be recommended. Some readers may regret that the butterflies of the Canary Islands are not

included, but this is not a book about Spanish but Iberian butterflies.

Yet another aspect makes this book so special: It is dedicated to Gabino Martin Toral, a

Spanish nature photographer and butterfly lover, who died much too early as a victim of the

insidious disease amyotrophic lateral sclerosis (ALS), a debilitating motor neuron disease. All

sales revenue of this book will be given to the foundation “Fundacion Miquel Valls”, to benefit

the care of ALS patients. So “mariposas por la vida” is a tribute to life, too.

TORSTEN VAN DER HEYDEN

Nota lepid. 36 (1): 13-18 13

Monochroa bronzella sp. n. from the southwestern Alps

(Lepidoptera: Gelechiidae)

OLE KARSHOLT !, JACQUES NEL’, FRANÇOIS FOURNIER *, THIERRY V ARENNE *

& PETER HUEMER?

! Natural History Museum of Denmark, University of Copenhagen, Universitetsparken 15,

DK-2100 Copenhagen, Denmark; okarsholt@snm.ku.dk

2 8 avenue Fernand Gassion, F-13600 La Ciotat, France; lucienne.nel(@orange.fr

3 25 rue de la Treille, F-65000 Clermont-Ferrand, France; ffournier63(@sfr.fr

* Thierry Varenne, 70 avenue Henry Dunant, F-06100 Nice, France; thierry.varenne@laposte.net

° Naturwissenschaftliche Abteilung, Tiroler Landesmuseen Betriebsgesellschaft m.b.H., Feldstrasse 11a,

A-6020 Innsbruck, Austria; p.huemer@tiroler-landesmuseen.at

Received 18 September 2012; reviews returned 9 October 2012; accepted 9 October 2012.

Subject Editor: Lauri Kaila.

Abstract. Monochroa bronzella sp. n. is described from the southwestern Alps (France, Italy). It is closely

related to M. nomadella (Zeller, 1868), with which it was hitherto confused. Literature records of M. no-

madella from France and northwestern Italy refer to M. bronzella sp. n. The two species are most clearly

distinguishable in the signa of the female genitalia. Females of both species have reduced wings, most

pronounced in M. nomadella. The new species is found in mountain areas at altitudes from around 800 to

2000 m. Adults and male and female genitalia of these two species are figured.

Resume. Monochroa bronzella sp. n. est décrit du sud-ouest des Alpes (France, Italie). Il est voisin de

Monochroa nomadella (Zeller, 1868) avec lequel il a été parfois confondu. Les signalisations de M. noma-

della de France et du nord-ouest de l’Italie concernent en réalité M. bronzella sp. n. Les deux espèces se

distinguent facilement par le signum des genitalia femelles. Les femelles des deux espèces ont les ailes

réduites, caractère plus prononcé chez M. nomadella. La nouvelle espèce vole dans des zones monta-

gneuses entre 800 et 2000 m. Imagos mâles et femelles des deux espèces sont figurés.

Introduction

Monochroa is a species-rich genus of Gelechiidae with altogether 30 species known

from Europe, including the Canary Islands (Huemer & Karsholt 2010; Karsholt 2011).

Similarly to other genera of the family a complete review on a continental scale is

lacking, though Elsner et al. (1999) give an overview of the central European taxa.

Monochroa species can be divided into species groups based on host-plant relation-

ships and genitalia characters (Gregersen & Karsholt, unpublished). The species dealt

with here belongs to the M. ferrea-group, which is characterised by having the vincu-

lum of the male genitalia with medial oval sclerotisation and the phallus cylindrical

with numerous small spines in the vesica, and larvae (as far as known) feeding on

Carex (Cyperaceae).

Most species of Monochroa are restricted to wetland habitats and only a few taxa

occur in mountain areas. Species of the M. ferrea-group inhabit sandy or rocky areas

from lowlands to high altitudes. Below we describe a new species which was hitherto

confused with M. nomadella (Zeller, 1868) and compare it with its closest relative.

Nota lepidopterologica, 17.06.2013, ISSN 0342-7536

14 KARSHOLT et al.: Monochroa bronzella sp. n. from the southwestern Alps

Abbreviations

BALD Collection of Giorgio Baldizzone, Asti, Italy

BAS Collection of Graziano Bassi, Avigliana, Italy

FOUR Collection of Francois Fournier, Clermont-Ferrand, France

MHNL Musée d’Histoire Naturelle de Lyon, France

TLMF Tiroler Landesmuseum Ferdinandeum, Innsbruck, Austria

VAR Collection of Thierry Varenne, Nice, France

ZMUC Zoologisk Museum, Natural History Museum of Denmark, Copenhagen, Denmark

Taxonomic part

Monochroa bronzella sp. n. Figs 1, 2, 5, 7, 9, 11

Material. Holotype ©, ‘route du Col de TENDE Alpes Maritimes 4.v111.2007 uv/vm 1360 m’ ‘Mo-

nochroa bronzella n. sp. © Th. Varenne leg.’ ‘HOLOTYPE’ ‘P. Huemer GEL 1176 ©’ (TLMF). —

Paratypes: France, 29, same data as holotype, genitalia slide Nel 21929 (VAR); same data as holo-

type, 19, 16.vii.2009, genitalia slide Fournier 798 (FOUR); Vaucluse, Saint-Christol, 20°, 8.vii.1992,

leg. Moulignier, genitalia slide Nel 1592, 1606 (MHNL, TLFM). Italy, Piemonte, Valsusa, Bussoleno,

Pian Cervetto, 1400 m, 19, 7.v1.1989, leg. Bassi (BAS); ibid, but Rocci Amelone, 19, 27.v.1990, leg.

Baldizzone, genitalia slide Hendriksen 1404 (ZMUC); ibid, but Mompantero, 1200 m, 30, 18.v1.1993,

leg. Bassi, genitalia prep. Elsner 882 (in tube) (BAS, ZMUC); ibid, but Mompantero, Mt. Rocciamelone,

800 m, 10°, 30.v.1998, leg. Bassi (BAS); ibid, but 1100 m, 10°, 24.v.2011, leg. Baldizzone, genitalia slide

Nel 25346 (BALD); Piemonte (CN), Parco Natur. Reg. Alpi Maritime, S. Giacomo di Entracque, Valle

della Rovina, 1537-1800 m, 10°, 14.vii.1996, leg. Baldizzone (BALD); ibid, S. Giacomo di Entracque,

sopra Lago della Rovina (Rocca Barbis), 1550-1850 m, 19, 20.vii.1997, leg. Baldizzone (BALD); ibid,

but 1850-2000 m, 10°, 26.v11.1997, leg. Baldizzone (BALD); ibid, but Entraque, Trinita, Vallone Grande,

1400 m, 20, 15.vii.1996, leg. Baldizzone, genitalia slide Hendriksen 2036 (BALD, ZMUC); ibid, but,

Mt. Ray, 1500-1800 m, 169, 39, 20. & 24.vii.1999, leg. Baldizzone, genitalia slides Hendriksen 2500,

2544, Huemer 12/1330 (BALD, ZMUC); ibid, but 60, 39, 18. & 20.vi1.2000, leg. Baldizzone (BALD);

ibid, Entracque, Trinita, Sentiero per Colle della Garbella, 1600—2000 m, 10°, 16.vii.2000, leg. Baldizzone

(BALD); ibid, 2000 m, 20°, 21.vii.2000, leg. Baldizzone (BALD); Prov. Cuneo, Colle della Lombarda,

1750 m, 60", 17.v11.2012, leg. Huemer (TLMF).

Description. Adult (Figs 1,2). Male (Fig. 1): Wingspan male 13-16 mm. Labial

palp slender; segment 2 slightly shorter than segment 3, cream-coloured, overlaid with

fuscous on lower and outer surface; segment 3 fuscous. Antenna dark brown; a few

paler rings near tip. Head, thorax and tegula shining fuscous. Forewing dark bronze

fuscous; an indistinct dark spot at apical part of the cell; fringe grey; no fringe line

present. Hindwing grey, with greyish fringe. Female (Fig. 2): Similar to male but small-

er (wingspan 9 mm) and with head shining metallic-fuscous and forewings shining

bronze-coloured, slightly darker towards apex, without any markings.

Remarks. The examined specimens show only little variation. Worn specimens be-

come paler, with more metallic shine.

Male genitalia (Figs 5, 7). Uncus digitate, short, with four long setae; gnathos

absent; valva heavily sclerotised, broad and weakly curved, distal part gradually ta-

pered towards apex, apex with sclerotised wall; sacculus broad, sub-oval, with weakly

concave outer and convex inner margin; vinculum with paired posteromedial ridge;

saccus short and broad, rounded; phallus massive, straight, distal third tapered; in situ

apical part of vesica with granular surface, medial part with separate group of about 50

small spines. Segment VIII with pair of short coremata in intersegmental membrane.

Nota lepid. 36 (1): 13-18 15

Figs 1-4. Monochroa spp., adults. 1. M. bronzella sp. n., ©’, holotype; 2. ibid, 9, paratype, Italy; 3. M. no-

madella (Zeller), ©, Italy; 4. ibid, 9, Russia.

Female genitalia (Figs 9, 11). Papillae anales elongate; apophyses posteriores

and anteriores slender, rod-like, about the same length; segment VIII smooth, ventral

part largely membranous, sclerotised subgenital plate semi-oval, longitudinal, covered

with numerous microtrichia; antrum and posterior part of ductus bursae membranous,

posteromedial part with long sclerotised plate; inception of ductus seminalis anteriorly

followed by short granular section; corpus bursae oval, covered with microtrichia; sig-

num a large irregularly shaped sub-oval plate, medially slightly constricted, anterior

and posterior part with about 4 teeth.

Diagnosis. M. bronzella sp. n. resembles the closely related, frequently slightly smaller

M. nomadella (wingspan of males 12—14 mm, females 8 mm) (Figs 3, 4), which has

a black streak in the fold and often also a black spot at 4/5 of the forewing, as well

as one at the apical part of the cell. Females of M. bronzella have shinier bronze-

coloured forewings than the dark greyish brown (and slightly brachypterous) female

of M. nomadella. The closely related M. ferrea (Frey, 1870) has darker, metallic grey

forewings with similar markings as in M. nomadella, and should not be confused with

M. bronzella sp. n. Its genitalia are figured by Sattler (1974) and Elsner et al. (1999)

(see also remarks). Eulamprotes unicolorella (Duponchel, 1843) is similar to females

of M. bronzella in having unicolorous, metallic shiny forewings, but those, as well as

the labial palps, head, thorax and tegulae, are distinctly darker.

16 KarsHOLT et al.: Monochroa bronzella sp. n. from the southwestern Alps

Figs 5-8. Monochroa spp., S-genitalia. 5. M. bronzella sp. n., slide Huemer GEL 1176; 6. M. nomadella

(Zeller), slide Huemer GEL 1177; 7. M. bronzella sp. n., phallus, slide Huemer GEL 1176; 8. M. nomadella

(Zeller), phallus, slide Huemer GEL 1177.

In the male genitalia the new species

differs from the most closely related M.

nomadella and M. ferrea by the distinctly

broader and not sickle-shaped or apically

pointed valva and the shape of the saccu-

lus, and from M. nomadella by having a

field of spines on the phallus (Figs 5-8;

Elsner et al. 1999: pl. 8, Fig. 67). The fe-

male genitalia are easily distinguished from

all other Monochroa by the large signum

of unique shape (Figs 9, 11) which is com-

pletely different, for example, in the exter-

nally similar M. nomadella (Figs 10, 12);

furthermore the long sclerite of the ductus

bursae is characteristic.

Distribution. Only known from the south-

western Alps of France and Italy.

Ecology/Habitat. Larval host plant and

early stages are unknown. Adults have

been observed from late May to late July

and they have been collected during night

Figs 9, 10. Monochroa spp., Q-genitalia. 9. M. bron-

zella sp. n., slide Huemer GU 12/1330; 10. M. no-

madella (Zeller), slide Huemer GU 12/1331.

Nota lepid. 36 (1): 13-18 17

1330; 12. M. nomadella (Zeller), slide Huemer GU 12/1331.

at light. It remains unclear if the female is able to fly. Elsner (in litt.) found hundreds

of males of M. nomadella coming to the UV light, whereas a single female specimen

was collected by sweeping grass and various vegetation. The larva of the related M. fer-

rea (Frey, 1870) has been reared from Carex ericetorum Pollich. (Cyperaceae) (Kaitila

1996). The habitats of M. bronzella sp. n. are steppic and xerothermic slopes. The spe-

cies seems to be restricted to siliceous soil whereas the related M. nomadella prefers

calcareous habitats (Elsner et al. 1999).

Etymology. The name of the new species refers to its uniformly bronze-coloured forewings.

General Remarks. Similar to descriptive taxonomy of other genera of Gelechiidae, our

description of male genitalia is based “on unrolled” slide preparations (Huemer 1987;

Pitkin 1986). According to such slides, the homology of the sacculus in Monochroa

seems doubtful, since this structure is articulated at the vinculum and should rather be

called the vincular process.

Literature records of M. nomadella from France (Fournier 2010) and northwestern

Italy (Karsholt 2004) refer to M. bronzella sp. n. and M. nomadella should be deleted

from the list of Lepidoptera found in France. Confirmed Italian records exist from

South Tyrol to northeastern Italy (Prov. Pordenone). As pointed out by Junnilainen et

al. (2010) the figure of the female genitalia of M. nomadella in the widely used book on

Central European Gelechiidae (Elsner et al. 1999) is erroneous. In fact two figures have

been inadvertently transposed on pl. 49 and thus fig. 67 refers to M. nomadella whereas

fig. 68 depicts M. ferrea (Elsner in litt.).

18 KarsHOLT et al.: Monochroa bronzella sp. n. from the southwestern Alps

Discussion

Monochroa is one of the morphologically particularly difficult genera of Gelechiidae

and identification from external appearance may cause serious problems. Dissection

of genitalia is thus often inevitable for a safe identification. Due to the overall similar-

ity of the adults and the frequently hidden living habits, the species inventory of the

European fauna is still incomplete. New taxa have been described more or less regu-

larly during the last decades, even from well explored areas such as Great Britain and

Scandinavia, but also from the Alps (Huemer & Karsholt 2010; Svensson 1992; Uffen

1991). However, considering the lack of a thorough generic revision, the description of

new taxa must be done with due care and is only possible within species-groups with

resolved taxonomy. The species group of M. nomadella and M. ferrea is not yet fully

resolved and according to preliminary results of DNA-barcoding may include further

cryptic species. However, the genitalia characters of M. bronzella sp. n. are unmistak-

able and it was obviously only by chance that the species was not recognised earlier.

As with several other recently described taxa (see Huemer & Karsholt 2010), this re-

cord reinforces the importance of the southwestern Alps for overlooked species, most

of which are restricted to this part of the Alps.

Acknowledgements

We thank Giorgio Baldizzone (Asti, Italy), Graziano Bassi (Avigliana, Italy) and Gustav Elsner (Prague,

Czech Republic) for providing specimens and information used in this study. The photographs were kindly

taken by Stefan Heim (Tiroler Landesmuseen), Innsbruck, Austria.

References

Elsner, G., P. Huemer & Z. Tokar 1999. Gelechiidae Mitteleuropas. — Verlag F. Slamka, Bratislava. 208 pp.

Fournier, F. 2010. Monochroa nomadella (Zeller, 1868) espèce nouvelle pour la faune de France (Lep.,

Gelechiidae). — Bulletin de la Societé entomologique de France 115: 192.

Huemer, P. 1987. Eine modifizierte Genitalpräparationstechnik für die Gattung Caryocolum.— Mitteilungen

der Schweizerischen entomologischen Gesellschaft 60: 207-211.

Huemer, P. & O. Karsholt 2010. A new endemic species of Monochroa from the south-western Alps

(Lepidoptera: Gelechiidae). — Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 62:

81-86.

Junnilainen, J., O. Karsholt, K. Nupponen, J.-P. Kaitila, T. Nupponen & V. Olschwang 2010. The gelechiid

fauna of the southern Ural Mountains, part. II: list of recorded species with taxonomic notes (Lepi-

doptera: Gelechiidae). — Zootaxa 2367: 1—68.

Kaitila, J.-P. 1996. Suomen jäytäjäkoiden (Gelechiidae) elintavat. — Baptria 21: 81-105.

Karsholt, ©. 2004. Gelechiidae. Pp. 112-141. — Jn: G. Baldizzone, I Microlepidotteri del Parco Naturale

Alpi Marittime (Italia, Piermonte) (Lepidoptera). — Bollettino del Museo Regionale di Scienze

Naturali, Torino 22: 1—318.

Karsholt, O. 2011. Gelechiidae. — Jn: O. Karsholt & E. J. van Nieukerken (eds), Lepidoptera, Fauna Euro-

paea, version 2.4, http://www.faunaeur.org [accessed 3.5.2012].

Pitkin, L. M. 1986. A technique for the preparation of complex male genitalia in Microlepidoptera. — En-

tomologist’s Gazette 37: 173-179.

Sattler, K. 1974. On Monochroa ferrea (Frey, 1870) and M. conspersella (Herrich-Schäffer, 1854) (Lepi-

doptera, Gelechiidae). — Entomologist’s Gazette 25: 177-282.

Svensson, I. 1992. Monochroa inflexella n. sp. (Lepidoptera, Gelechiidae). — Entomologisk Tidskrift 113:

47-51.

Uffen, R. W. J. 1991. Monochroa moyses sp. n., a new Gelechiid moth mining the leaves of Scirpus mari-

timus L. — British Journal of Entomology and Natural History 4: 1-6.

Nota lepid. 36 (1): 19-33 19

Northern boundary of the range of the Clouded

Apollo butterfly Parnassius mnemosyne (L.) (Papilionidae):

climate influence or degradation of larval host plants?

Ivan N. Bototov!*, MIKHAIL Yu. GOFAROV |, ALEXANDER M. Rykov’?,

ARTYOM A. FROLOV !, YAROSLAVA E. KOGUT!

! Institute of Ecological Problems of the North, Ural Branch of the Russian Academy of Sciences,

Northern Dvina Emb., 23, 163000 Arkhangelsk, Russian Federation

? The Pinega State Nature Reserve, 164610 Pinega, Russian Federation

“ corresponding author; inepras@mail.ru

Received 10 January 2012; reviews returned 27 February 2012 (first round), 13 July 2012

(second round); accepted 4 October 2012.

Subject Editor: Thomas Schmitt.

Abstract. The present paper summarises data on the northern localities of Parnassius mnemosyne (L.)

(Papilionidae), which are mostly situated in the Russian Federation, and gives a thorough description

of the species’ northern range location. It is shown that the northernmost populations exist within the

karst landscapes in the north of White Sea-Kuloi Plateau (between 65° 35’ and 66° 03’ N) in the lower

valleys of the rivers Soyana and Kuloi and in the north of Timan Highland (66° 10’ N) along the shore of

Kosminskoe Lake (the Pechora river basin). The northern limits of the Clouded Apollo’s range appear to

be strongly determined by the distribution of its larval host plants (primarily Corydalis solida (L.) Clairv.,

Papaveraceae) and the role of climate and relief seems to be of minor importance.

Introduction

The Clouded Apollo butterfly Parnassius mnemosyne (Linnaeus, 1758) (Papilionidae)

is an endangered species in Europe (Van Swaay & Warren 1999; Van Swaay et al.

2010). Its decline has been attributed to the cessation of traditional management, graz-

ing and mowing of semi-natural grasslands and coppicing in woodlands (Luoto et

al. 2001; Väisänen & Somerma 1985). The distribution of the species in European

countries is known with a high certainty, but it remains less thoroughly known in the

Russıan Federation (Kudrna et al. 2011; Weiss 1999) due to the less intensive recording

in northern Russia. In the meantime, a few individuals from Northern Russia have been

described as separate subspecies or other morphological forms (e.g., Eisner & Sedych

1964; Kreuzberg 1989; and others). Some predictive models have been published on

the distribution of P mnemosyne which reveal its possible change under biotic (larval

host plants) interactions, climate conditions (Araujo & Luoto 2007; Settele et al. 2008),

and habitats (Heikkinen et al. 2007), based on West European data.

The habitat preferences of the Clouded Apollo in European countries and the south-

ern regions of European Russia are well studied, but the northern part of Russia has not

yet been surveyed (Gorbach & Kabanen 2010; Lyvovsky & Morgun 2007; Weideman

1986; etc.). As populations of this species inhabit heterogeneous environments, their

structure generally conforms to the metapopulation model in which a landscape is di-

vided into suitable patches and unsuitable matrix (Gorbach & Kabanen 2010; Luoto et

Nota lepidopterologica, 17.06.2013, ISSN 0342-7536

20 BoLoTov et al.: Northern range boundary of Parnassius mnemosyne

northern localities of P mnemosyne

--—-- limits of the species range (by our data)

limits of the species range (by Weiss 1999)

| distribution of the species (by Weiss 1999)

E74 uncertain distribution of the species (by

Fig. 1. Distribution of the Clouded Apollo butterfly in Northern Europe and Western Siberia and the north-

ern boundary of the species range. The Russian localities — according to data from Table 1 (locality num-

bers on the map correspond to numbers in the Table); the Western European localities — according to

Settele et al. (2008) and Somerma & Yakovlev (1998).

al. 2001). For example, in Fennoscandia the Clouded Apollo inhabits a dense network

of semi-natural grasslands (with mating sites and nectar sources) and deciduous forest

patches (with larval food plants) (Heikkinen et al. 2007). Migration routes of individu-

als can extend onto the meadows and shrubs of open spaces in forests (Gorbach &

Kabanen 2010; Konvicka et al. 2006; Meier et al. 2005; Valimaki & Itamies 2003).

The Clouded Apollo butterfly is a specific K-strategist; females can mate only once

and lay about 50 eggs dispersed over a large area (Meglecz et al. 1997; Weideman

1986). P mnemosyne is an oligophagous species and its larvae develop on various plant

species of the genus Corydalis DC. (Papaveraceae). In the north of Russia, Corydalis

solida (L.) Clairv. and C. capnoides (L.) Pers. have been recorded (Korshunov 2002;

Tatarinov & Dolgin 1999). Reports from other countries include Corydalis solida for

Finland (Luoto et al. 2001; Somerma 1997), C. intermedia (L.) Mérat and C. pumi-

la (Host) Rchb. for Sweden (Franzen & Imby 2008), and C. intermedia for Norway

(Aagaard & Hanssen 1989). Knowledge of the host-plant species is important to ex-

plain the local and landscape distribution of the Clouded Apollo butterfly (Heikkinen

et al. 2005; Luoto et al. 2001). Spatial structure of P mnemosyne metapopulations

is determined by the distribution of the Corydalis populations (Gorbach & Kabanen

2010).

This paper maps the northern boundary of the P mnemosyne’s range, summarising

the information about the peripheral northern localities of this species and discussing

the relative influence of climatic factors and host-plant availability upon the limits of

the species range.

Nota lepid. 36 (1): 19-33 21

Materials and methods

The survey of marginal northern P mnemosyne populations was conducted in Arkhan-

gelsk oblast (Russian Federation). A. M. Rykov studied the populations in the Pinega

State Nature Reserve annually during 1978—2011. Field studies on the Soyana, Kuloi,

Pinega and Yula Rivers were conducted between 2002 and 2007. In 2003, collector

L. P. Shoshin (Arkhangelsk) sampled a few specimens of the Clouded Apollo in the

Ivovik Stream Valley, located at the Winter Coast of the White Sea. Data on other

northern P mnemosyne localities were obtained from different research papers. The

arrangement of the localities was digitised and mapped. The species range data in this

map were added from Weiss’ (1999) book.

The distribution of Corydalis plants was obtained from a digitised version of

“Atlas Flora Europaeae” (AFE) (Lahti & Lampinen 1999) and from “Flora Sibiriae”

(Malyschev & Pechkova 1994). Additional data originated from regional botanic pub-

lications (Liden 2001; Puchnina et al. 2000; Schmidt 2005). All botanical data were

transferred to the AFE grid map (squares of ca. 50 km x 50 km, the Universal Transverse

Mercator (UTM) projection and the Military Grid Reference System (MGRS)) (Jalas

& Suominen 1972—1996). Meteorological data were obtained from the website of the

World Data Center for Meteorology, Asheville, North Carolina.

Northern localities of the Clouded Apollo butterfly

As shown in Fig. 1, the northern boundary of the range stretches from the Norwegian

coast in the West to the Irtysh river headstream in the East, about 4000 km in length.

Some northern localities of this species in Fennoscandia are highly populated (Aagaard

& Hanssen 1989; Luoto et al. 2001; Opheim 1983; Somerma 1997; Väisänen & So-

merma 1985). There is little data on regional expansions (Marttila et al. 2001; Meier et

al. 2005). Information about marginal northern localities of P mnemosyne in Russia is

compiled in Tab. 1.

Tyumen oblast. In 1987-1988 P. mnemosyne populations were discovered in the

Irtysh (near the city of Tobolsk) and Iska (Korshunov 2002; Kreuzberg 1989) river

valleys. The populations inhabit hay-harvested and grazed floodplain meadows, main-

tained in river valleys since the 19" century.

Komi Republic. The most northern localities of P mnemosyne were discovered

in the valleys of the Pechora river basin at the foothills of the Northern Urals and

Timan Highland (Tatarinov & Dolgin 1999, 2001). The cited authors have conducted

field studies there since the 1990s. The highest density of populations was detected

in the Pechoro-Ilychsky Nature Reserve (Tatarinov & Dolgin 1999). The habitats of

the populations were natural humid mixed-herb meadows in river valleys, which are

characterised by heterogeneity of species composition and density of vegetation. The

dominant species were Filipendula ulmaria (L.) Maxim. (Rosaceae), Crepis sibirica

L. (Asteraceae), Thalictrum sp., Trollius europaeus L. and Aconitum septentrionale

Koelle (Ranunculaceae), Valeriana wolgensis Kazak. (Valerianaceae), Geranium sylva-

ticum L. (Geraniaceae).

22 BoLoTov er al.: Northern range boundary of Parnassius mnemosyne

Fig. 2. The Clouded Apollo butterfly specimens from peripheral northern populations inhabited meadows in

Moseev Ravine in the White Sea-Kuloi Plateau, Arkhangelsk oblast, Northern European Russia. A: upper-

side; B: underside.

Arkhangelsk oblast. The populations are located within the frontiers of the northern

part of the Timan Highland (Tatarinov & Dolgin 1999), at the White Sea-Kuloi Plateau

(Belomorsko-Kuloiskoe Plato) and in the Pinega river basin. These are probably the

largest populations of P mnemosyne among the northern ones. In the Southeast of the

White Sea-Kuloi Plateau, in the Pinega State Nature Reserve, observations of P mne-

mosyne populations have been made since 1978. The populations were discovered in

three large karst ravines (Moseev, Vizgunov and Severny), belonging to the Sotka river

basin (tributary of the Kuloi river) (Figs 2a, b). The butterfly inhabited small patches

of natural humid mixed-herb meadows at the ravine bottoms (Fig. 3). The dominant

species of these meadows are Aconitum septentrionale Koelle and Thalictrum sp.

(Ranunculaceae), Anthriscus sylvestris (L.) Hoffm. (Apiaceae), Geranium sylvaticum

L. (Geraniaceae), Filipendula ulmaria (L.) Maxim. (Rosaceae), Cirsium oleraceum

(L.) Scop. (Asteraceae), Chamerion angustifolium (L.) Holub (Onagraceae), Paeonia

anomala L. (Paeoniaceae) and Elymus caninus (L.) L. (Poaceae). Here, the ravines are

surrounded by Siberian spruce (Picea abies ssp. obovata (Ledeb.) Domin, Pinaceae)

forests, with small inclusions of Siberian larch (Larix sibirica Ledeb., Pinaceae). These

meadows were formed in karst ravines about 2500—3500 years ago and existed hereaf-

ter owing to harsh local microclimates, which prevented forest expansion (Titova et al.

2011). The total area of the Clouded Apollo habitats is ~4 ha within Vizgunov ravine,

~10 ha within Moseev ravine and ~15 ha within Severny ravine. The flight period

of adult P mnemosyne continues from mid-June to the beginning of August (13.vi—

6.viii), and adult density varies highly from year to year (Bolotov 2004; Rykov 2009).

Imagines were observed annually in 1978—2011 in two of the three patches, but in

Vizgunov ravine they have not been recorded since 2000.

The P mnemosyne population inhabiting the karst areas of the Soyana river val-

ley was observed in the Northeast of the White Sea-Kuloi Plateau during 2002-2007.

Zonal vegetation is represented by spruce and larch forests. The butterflies inhabit a

river valley about 50 km long, as well as humid mixed-grass meadows, which are

typical of the river valley bottom and which form small patches about 1-3 ha in size,

Nota lepid. 36 (1): 19-33 23

Figs 3—4. Habitats of the Clouded Apollo butterfly in the White Sea-Kuloi Plateau, Arkhangelsk oblast,

Northern European Russia. 3. Meadow in the Moseev Ravine. 4. Meadow in the Soyana river valley.

divided by thin forests and shrubs (Fig. 4). The meadows form natural floodplains, and

they were used for hay production until the end of the 20" century. The dominant plant

species were similar to those of meadows in large karst ravines.

A population of P mnemosyne was found in the Ivovik Stream valley (the northwest

of the White Sea-Kuloi Plateau) in 2003. The stream has a deeply scarred valley, re-

stricted to places of Vendian (Ediacara) rocky outcrop on the Winter Coast of the White

Sea. The butterflies inhabit small patches of natural humid mixed-herb meadows. This

population is isolated from all other localities by continuous stretches of spruce forests.

Nenetsky autonomous district. Only a few specimens of Pl. mnemosyne were dis-

covered on the Kosminskoe Lake shore meadows (northern part of Timan Highlands)

(Tatarinov 2006).

Karelia Republic. The distribution of the species is localised around Onega Lake

and the eastern part of Ladoga Lake areas (Gorbach & Kabanen 2010; Gorbach &

Reznichenko 2009; Kaisila 1947; Somerma & Yakovlev 1998). Localities of P mne-

mosyne also exist on upland meadows on the islands of Bolyshoy Klimenetskiy and

Kizhi in Onega Lake, and at the flood-land meadows along the Koloda river shores in

the southeastern part of Russian Karelia (Humala 1998). The meadows were used as

hayfields until the beginning of the 21" century. The dominant species of the meadows

are Heracleum sphondylium ssp. sibiricum (L.) Simonk. (Apiaceae), Rumex acetosa

ssp. thyrsiflorus (Fingerh.) Hayek (Polygonaceae), Centaurea scabiosa L., Tanacetum

vulgare L. and Taraxacum officinale F.H. Wigg. (Asteraceae), Barbarea vulgaris W.T.

Aiton (Brassicaceae), Poa pratensis L. and Phleum pratense L. (Poaceae). In some

localities, high abundance of adults was observed (Gorbach & Kabanen 2010).

Northern boundary of the species range: the outcome of biotic interactions

and climate conditions

It was mentioned before that P mnemosyne is not usually found farther north than

63—64° N (Kudrna et al. 2011; Lyvovsky & Morgun 2007; Settele et al. 2008; Weiss

24 BoLoTov et al.: Northern range boundary of Parnassius mnemosyne

Distribution of Corydalis spp.

[_®_] native [_® ] introduction

Distribution of Parnassius mnemosyne

F-] limits of the species range (by our data)

F--—-J limits of the species range (by Weiss 1999)

[7] distribution of the species (by Weiss 1999)

EZ uncertain distribution of the species (by

Weiss 1999)

Figs 5-6. Distribution of the four species of Corydalis spp. and the northern boundary of the Clouded

Apollo butterfly range. 5. C. solida (L.) Clairv. 6. C. capnoides (L.) Pers.

1999). New data allow us to specify the northern limits of the species distribution.

The world’s northernmost populations have been registered at the north of the White

Sea-Kuloi Plateau (between 65° 35’ and 66° 03’ N) in the Soyana and Kuloi lower river

valleys and in the north of Timan Highland (66°10’ N) at the Kosminskoe Lake shore

(the Pechora river basin).

Nota lepid. 36 (1): 19-33 25

S’E 10°E 15°E 20°E 25°E 30°E 35°E 40°E

a ‘¢

% Ÿ h

AA / on

6 IVE

30°E 35°F 40°E 45°F 50°E 55°E 60°E

Figs 7-8. Distribution of the four species of Corydalis spp. and the northern boundary of the Clouded

Apollo butterfly range. 7. C. pumila (Host) Rchb. 8. C. intermedia (L.) Mérat.

Sedimentary Paleozoic bedrock and modern areas of active karst processes form

both of these territories (Gofarov et al. 2006; Shvartsman & Bolotov 2008). The karstic

rocks are represented by Carboniferous limestone in the Timan Highland and Permian

gypsums and anhydrites in the White Sea-Kuloi Plateau. The plateau region is known

as a refuge for different animal and plant species, some of which are northern postgla-

26 BoLoTov et al.: Northern range boundary of Parnassius mnemosyne

cial and some are southern Atlantic relicts. Dryads Dryas octopetala L. and D. o. ssp.

punctata (Juz.) Hultén (Rosaceae), osiers Salix myrsinites L. and S. reticulata L. (Sa-

licaceae) (Puchnina et al. 2000; Simacheva 1986), pond damselflies Coenagrion gla-

ciale (Selys, 1872) and C. hylas (Trybom, 1899) (Coenagrionidae) (Bernard & Daraz

2010), carabid beetles Prerostichus brevicornis (Kirby, 1837) and Bembidion yuko-

num Fall, 1926 (Carabidae) (Mokhnatkin et al. 2010), collembolans Desoria tshernovi

(Martynova, 1974) and D. inupikella Fjellberg, 1978 (Isotomidae) (Babenko 2008)

may be considered to be postglacial relict species. In Europe, these postglacial relicts

are representatives of a cold-stenothermal fauna that probably colonised the subcon-

tinent during the late Pleistocene and early Holocene in the period of the maximum

distribution of birch and pine (Bernard & Daraz 2010; Elina et al. 2005). Other Atlantic

relicts, besides P mnemosyne, include its larval host plants Corydalis solida and C.

capnoides, as well as the plants Ste/laria nemorum L. (Caryophyllaceae), Cypripedium

parviflorum Salısb. (Orchidaceae), Paeonia anomala L. (Paeoniaceae) (Puchnina et

al. 2000; Simacheva 1986), the blue butterflies Cupido alcetas (Hoffmannsegg, 1803),

C. minimus (Fuessly, 1775) and Aricia nicias (Meigen, 1829) (Lycaenidae) (Bolotov

2004), and the carabid beetles Calosoma investigator (Illiger, 1798), Lebia cruxminor

(Linnaeus, 1758) and Badister lacertosus Sturm, 1815 (Carabidae) (Mokhnatkin et al.

2010). They probably migrated to Northern Europe during the Atlantic period of the

Holocene. According to the studies of molecular markers, expansion of P mnemosyne

northern lineages took place during the postglacial warming period 5000—7000 years

ago (Gratton et al. 2008).

During the present period, populations of both groups of relict species remain

isolated in the same regions of the European taiga, particularly in karst landscapes

(Shvartsman & Bolotov 2008). The coexistence of such different relict species is pos-

sible due to the high heterogeneity of karst landscapes. Sites with highly contrasting

temperatures exist in such areas: very cold sites near caves with long-term ice alternat-

ing with well-heated patches in slopes of south exposure and wide karst ravines. These

sites can easily be located near each other. For example, upland herb meadows grow

at the bottom of Moseev ravine (inhabited by the Clouded Apollo and other southern

relicts), whereas small patches of mountain dryads tundra with Salix myrsinites and S.

reticulata occur on the nearby gypsums and anhydrites outcrops.

The altitude range of the northern localities of P mnemosyne is very broad (Tab.

1). Here populations exist under different climatic conditions (Tab. 2). Therefore, cli-

mate and relief cannot be considered as major limiting factors for the expansion of the

Clouded Apollo northward. Dot maps of the distribution area of different Corydalis

species (Figs 5—8) reveal that the northern boundary of P mnemosyne’s range almost

fully correlates with the distribution of only one larval food species — Corydalis solida.

Corydalis intermedia is widespread in Western Fennoscandia, but P mnemosyne is

found only in a few localities. However, Fennoscandian populations of P mnemosyne

mostly prefer Corydalis solida, which is represented usually by introduced individu-

als (Liden 2001). These differ from native populations in flower and bract constitution

details. This plant species is widely cultivated in parks and gardens and escapes from

cultivation frequently. Many naturalised populations of Corydalis solida exist in the

Nota lepid. 36 (1): 19-33 ah

south of Norway (primarily along the seacoast), southern Sweden and central Finland

(AFE Secretariat, A. Sennikov, pers. comm.).

Studies of regional differences in the Clouded Apollo larval food preference may

be enlightening. For example, in the European part of Russia (Penza oblast), P. mne-

mosyne larvae were registered feeding only on Corydalis solida, although C. cava (L.)

Schweigg. & Körte and C. cava ssp. marschalliana (Willd.) Hayek occur in the same

biotopes (Polumordvinov & Shibaev 2007). Also, models using larval host plants as

a predictor of variability of the studied species predicted the presence of the Clouded

Apollo when Corydalis solida was present and the absence of the Clouded Apollo

when C. solida was also absent; this was true even when other Corydalis species were

present (Aratjo & Luoto 2007).

Given that the distribution of P mnemosyne in the north mostly correlates with

the presence of Corydalis populations, and populations of the butterfly inhabit natural

meadows, it is difficult to forecast significant future changes in the northern boundary

of the species range. Geographically, the majority of peripheral northern localities of

this species is concentrated in sparsely populated areas in the Russian Federation, in

predominantly non-disturbed taiga landscapes with difficult access, which do not seem

to be threatened by human activities. Many Russian populations inhabit the state nature

reserve territories: “Kizgi Scerries” Reserve (Karelia Republic), Pinega and Soyansky

Reserves (Arkhangelsk oblast), Pechoro-Ilychsky and “Belaja Kedva” Reserves (Komi

Republic).

P. mnemosyne occupies habitats with optimal ecological conditions in different bi-

omes, therefore this species has a zonal replacement of habitat preferences (Bei-Bienko

1966; Chernov 2008). In the north, it behaves like a typical mesophilous species, which

prefers open intrazonal habitats with medium humidity and solar heat (in different types

of open meadows). Southern populations prefer mostly humid and less warm habitats.

In Central Europe, including southern regions of European Russia, the Clouded Apollo

is a woodland species and inhabits forest steppes, sparse deciduous forests and forest

clearings where the larval host plants grow (Konviëka & Kuras 1999; Konvicka et al.

2006; Meglecz et al. 1997; Polumordvinov & Shibaev 2007; Weidemann 1986). In the

south of Europe, its populations avoid lowlands, where the environment is too hot and

dry, and reside primarily in the humid and cool habitats of mountain-subalpine belts

(Descimon & Napolitano 1993; Lyvovsky & Morgun 2007; Napolitano et al. 1988;

Napolitano & Descimon 1994). Hence, the distribution of the Clouded Apollo in the

North is limited principally by the distribution of its larval food plants. However, it

should be stated that the latitudinal change of landscape-habitat occupancy also de-

pends on regional climatic conditions (temperature and humidity). The results of this

paper agree with the importance of biotic interactions for modelling individual species

distribution at the macroecological scale under climate change (Araüjo & Luoto 2007).

Acknowledgements

The authors are grateful to AFE secretary A. Sennikov for providing data on Coryalis solida in Fennoscandia,

as well as N. Larionov, M. Podbolotskaya & M. Yartzeva for constructive remarks on the manuscript. Prof.

T. Schmitt, Dr. Z. Varga and Dr. M. Konvicka provided insightful comments and advice during revision

28 Bototov er al.: Northern range boundary of Parnassius mnemosyne

of the manuscript. The study has been supported by grants of the Russian Foundation for Basic Research

(Grant no. 10—04—008970), the President of Russia (no. MD-4164.2011.5), the Ural Branch of Russian

Academy of Sciences & the Ministry of Science and Education of the Russia.

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Nota lepid. 36 (1): 35-46 3)

The genetics of Luperina nickerlii Freyer, 1845

in Europe (Noctuidae)

ADRIAN SPALDING!, [va FUKOVA? and RRICHARD H. FFRENCH-CONSTANT ?

! Tremayne Farm Cottage, Praze, Camborne, Cornwall, TR14 9PH UK;

a.spalding@spaldingassociates.co.uk

2 Biosciences, University of Exeter in Cornwall, Penryn, TR10 9EZ, UK

Received 14 November 2012; reviews returned 7 January 2013; accepted 18 January 2013.

Subject Editor: Vazrick Nazari.

Abstract. We use mitochondrial markers to examine the genetic status of European subpopulations of Lu-

perina nickerlii Freyer, 1845 (Noctuidae) in Britain, Ireland, Spain and the Czech Republic. We show that

all the populations sampled belong to the same species Luperina nickerlii, despite considerable differences

in appearance, ecology and population isolation. Neighbour-joining tree based on mitochondrial markers

showed only three populations as separate clusters: gueneei, nickerlii and knilli. We show that subspecies

leechi, albarracina and demuthi are genetically close to each other and that both /eechi and gueneei show

significantly lower heterozygosity than the other subspecies sampled. L. n. albarracina and knilli show

high genetic variability. Isolation by distance was not supported in this study, suggesting populations were

probably linked to each other in the recent past.

Introduction

Luperina nickerlii Freyer, 1845 (Noctuidae) 1s widespread in mainland Europe occur-

ring on xerothermic slopes where the larvae feed on different grasses (Ganev 1982;

Hacker 1989; Karsholt & Razowski 1996; Robineau 2007; Steiner & Ebert 1998),

although in Britain and Ireland they are entirely coastal and certain subspecies are of

conservation concern (Goater & Skinner 1995). Eight subspecies of this moth have been

described (Tab. 1), based largely on phenotypes such as wing colouration, although

the taxonomy of the genus is in constant flux. Thus, L. n. leechi was described as a

new subspecies in 1976 (Goater 1976), whereas L. n. graslini and L. n. tardenota

were originally described as separate species but have since been synonymised and

are now regarded as subspecies of L. nickerlii (Zilli et al. 2005). L. n. knilli has also

been proposed to warrant full species status (De Worms 1978; Haggett 1980) but is

now considered a subspecies (Skinner 2008). L. n. albarracina was described as a new

subspecies in 1962 (von Zerny 1962) on wing colour and shape, but some authorities

now consider it merely a form (Zilli et al. 2005). L. n. graslini is similar in appearance

to L. n. gueneei but otherwise the subspecies all look different from each other in wing

colour and are generally easy to distinguish by eye. L. n. demuthi is the most variable

of all the subspecies and occasional specimens may look similar to those of subspecies

L. n. leechi, L. n. knilli and L. n. gueneei, but the majority of specimens are readily

separated from the other subspecies.

The European mainland subspecies all have similar ecologies, feeding on the same

food plants and occupying similar biotopes. However, the subspecies in Britain show

dramatic differences in their ecology and life styles and are prone to producing small

Nota lepidopterologica, 17.06.2013, ISSN 0342-7536

36 SPALDING et al.: The genetics of Luperina nickerlii in Europe

isolated colonies (Tab. 1). In Britain at least, the sparse fragmented distribution of this

moth over a large area may reflect a collapse in a former larger range size following

an initial increase at the end of the last glacial maximum when temperatures attained

levels similar to those of today, and as a result leaving behind small refuge populations,

or alternatively multiple post-glacial colonisation events from continental populations

or recent changes in habitat preference or behaviour (e.g., a host-plant switch or a

reluctance to fly). The populations of the British and Irish subspecies are at least 300

km apart from each other and a minimum of 320 km from the nearest known mainland

population near Paris (which is very small and possibly endangered), 850 km from

Spanish populations and 950 km from populations at Prague, with little possibility of

regular interchange. However, in captivity, L. n. gueneei and L. n. leechi (which have

similar ecologies) can pair and produce moths similar to L. n. leechi (Haggett 1980;

A. Myers, personal communication). These, at least, are the same species (Mayr 1942)

and have similar ecologies. The definition of a subspecies (Lincoln et al. 1985) as

isolated natural populations differing taxonomically and genetically from other groups

within the species supports the splitting of L. nickerlii into subspecies as differing

taxonomically.

The single isolated population of L. n. leechi has perhaps been studied in more

detail than the other subspecies (e.g., Spalding 1991a, b). It is restricted to a small

area of a shingle beach, 500 metres x 240 metres, where its larval food plant Elytrigia

Juncea (L.) Nevski (Poaceae) occurs (Spalding 1997; Spalding et al. 2012). It is listed

in a Biodiversity Action Plan listing as a unique subspecies currently under threat and

declining, and is thus of high conservation value (JNCC 2007). The population has

been studied by one of us (AS) since 1987. The population size of this iconic species is

very small, with the annual Index of Abundance (the sum of the weekly means based

on transect counts) between 1994 and 2009 ranging between 5 to 78 adult moths (mean

18.99) (Spalding 1997; Spalding & Young 201 1a).

Here we use mitochondrial DNA markers to ask several questions fundamental to

the origin and conservation of L. nickerlii subspecies. First, given the different ecologies

and appearance of the UK and Irish subspecies, do they belong to the same species

as the mainland populations? Secondly, we investigate the genetic variation of these

subspecies. Thirdly, we ask which populations should be the focus of conservation

concern.

Material and Methods

Sampling. Adult specimens of Luperina nickerlii from four different populations in

UK and Ireland and four mainland populations from the Czech Republic and Spain

were sampled (Tab. 2 and Fig. 1). Actinic light traps with chloroform were used

to collect the moths except for L. n. leechi, which was collected using torch light.

Specimens were kept alive until either snap frozen in liquid nitrogen (L. n. leechi, L. n.

gueneei, L. n. knilli and L. n. demuthi) or preserved in pure ethanol (Czech populations

of L. nickerlii) for genomic DNA extraction except for pinned and dried specimens

(Spanish populations of L. n. albarracina).

Nota lepid. 36 (1): 35-46 37

Tab. 1. Luperina nickerlii subspecies in Europe and their food plants and habitat

Subspecies Habitat Countries | Number of | Within site

sites abundance

demuthi Goater | Saltmarsh Puccinellia maritima England Few Abundant

& Skinner, 1995 (Huds.) Parl.

graslini Hot dry slopes | Festuca ovina L. and France Many Abundant

Oberthiir, 1908 other grasses (south)

gueneei Sand dune Elytrigia juncea England; Few Abundant

Doubleday, 1864 (Viviani) Runemark ex | Wales

Melderis

knilli Boursin, Coastal cliff Festuca rubra L.

1964

leechi Goater, Shingle beach | Elytrigia juncea England One Rare

1976 (Viviani) Runemark ex

Melderis

nickerlii Freyer, | Sparse open Festuca ovina L. and Germany; Many Abundant

1845 grassland; sandy | other grasses Czech

heaths Republic;

Bulgaria

tardenota Hot dry slopes | Festuca ovina L.and France Few Rare

Joannis, 1925 other grasses (central)

albarracina Hot dry slopes | Festuca ovina L. and Spain; Many Abundant

Schwingen- other grasses Portugal

schuss, 1962

Tab. 2. Luperina nickerlii sampling locations and mitochondrial COI haplotypes

Date Subspecies | Locality Coordinates COI* | Sample

size**

8 Sep 2008 Strood, UK | 51°47/51.53"N_ 0°55'09.23" E

19 Aug 2008 knilli Inch, 52°08'37.41" N 9°59'14.65" W |A,F,G

Ireland

19-20 Aug 2008 E

Food plant

Trabeg, 52° 07'16.44" N 10° 12'25.82" W 10

Ireland

5 Sep 2009 nickerlii Maslovice, |50°12'23.80" N 14°23'16.85” E |H,I, J

5 Sep 2009 Praha, CR |50°02'53.11” N 14°24'16.68" E

15 Sep 2009 albarracina 402353897 N 5°57. 187 W |A.R,O

Spain

16 Sep 2009 Paramos, | 42°35'17.95"N 3°43'56.36"W |A,K,L,

Spain M, N

* cytochrome c oxidase subunit I gene (COI) haplotypes sampled in the L. nickerlii populations.

** Number of specimens collected.

Genomic DNA isolation. Genomic DNA was extracted from the thorax or abdomen of

preserved individuals using either standard phenol/chloroform method (Blin & Stafford

1976) or a Genomic DNA Purification Kit (Fermentas, Burlington, Canada) according

to the manufacturer’s instructions. DNA concentration was estimated by NanoVue (GE

Healthcare, Buckinghamshire, UK) and adjusted to 500 ng/ul.

38 SPALDING et al.: The genetics of Luperina nickerlii in Europe

Yj"

"go

kilometres

Fig. 1. Map of Luperina nickerlii sampling localities and haplotype structure of each sampled population

based on the sequence of mitochondrial cytochrome c oxidase subunit I gene (COI). Each population is

represented by a circle which is proportional to the number of specimens used for the analysis. Regions

within each circle correspond to the proportion of individual COI haplotypes. Abbreviations: dem, L. n.

demuthi, Strood, UK; gue, L. n. gueneei, Gronant, UK; knl, L. n. knilli, Inch, Ireland; knT, L. n. knilli,

Trabeg, Ireland; lee, L. n. leechi, Loe Bar, UK; mas, L. n. nickerlii, Maslovice, Czech Republic; pro,

L. n. nickerlii, Praha, Czech Republic; par, L. n. albarracina Paramos de Masa, Spain; and ama, L. n.

albarracina, Amavida, Spain. Sample sizes are given in Table 2. L. n nickerlii also occurs elsewhere, e.g.,

in France and Portgual, but was not sampled there.

Genetic marker sequencing. In order to characterise the genetic distance and variability

of different populations of L. nickerlii we selected hybrid primers for mitochondrial

cytochrome c oxidase subunit I gene (COI). PCR was set up using a commercially

available master mix provided by Qiagen (Hilden, Germany) containing the reagents

12.5 ul dH,O, 2 ul 10x buffer, 2 ul MgCl, Primer F & R 2 x 1 ul, 0.4 wl I dNTP, 0.1 ul

Taq polymerase, | ul of DNA extracts, for a total of 20 ul PCR reactions, following the

manufacturer’s recommendations.

An initial denaturation at 94°C for 3 min was followed by 35 cycles of 30 sec at

94°C, 30 sec at the annealing temperature of 50°C, and 1 min 30 sec at 72°C, and

by a final extension step of 7 min at 72°C. Quality and amount of PCR products

were checked on a 1% agarose gel. In total 1,212 bp of mitochondrial sequence were

obtained. Sequences can be retrieved from GenBank under the following accession

numbers: GU903504—582 and HM068967-79.

Nota lepid. 36 (1): 35-46 39

Tab. 3. Population pairwise F,, values. The calculations were based on mitochondial COI sequences.

albarracina 0.04 NS |0.54*** |0.44** |0.63*** |0.12** |0.35*** 0.57*** |0.10 NS

Paramos

Significance level: NS = not significant; * P< 0.05; ** P< 0.01; *** P< 0.001

Sequence analyses. Mitochondrial sequences were aligned using the program MUSCLE

(Edgar 2004) implemented in Geneious (Biomatters Ltd., Auckland, New Zealand).

Heterozygotes were detected by Heterozygote plug-in in Geneious and manually

corrected. Out of 1,212 bp of mitochondrial sequence, 12 sites were informative.

Neighbor-joining trees were constructed by Geneious under Jukes-Cantor genetic

distance model (Jukes & Cantor 1969; Saitou & Nei 1987). Bootstrap was performed

with 1,000 replicates and branches with less than 50% support were collapsed.

Population structure analysis. Analysis of Molecular Variance (AMOVA), F-statistics

(fixation indices) and population pairwise differences (Excoffier et al. 1992; Weir 1996;

Weir & Cockerham 1984) were calculated using Arlequin version 3.1 (Excoffier et al.

2005). Heterozygosity was computed as described in Nei (1987). Correlation of genetic

and geographic distance was tested by Mantel test performed on matrices of pairwise

geographic distances (given as In km) and linearised pairwise F4. values (F</(1-Fsr))

(Mantel 1967; Slatkin 1995). Hardy-Weinberg equilibrium (Guo & Thompson 1992;

Levene 1949) and exact test of differentiation (Goudet et al. 1996; Raymond & Rousset

1995) were computed as implemented in Arlequin v3.1. Statistical parsimony network

(Templeton et al. 1992) was constructed using TCS v1.21 (Clement et al. 2000).

Results

Fsr values are summarised in Tab. 3. Despite our expectations and although the habitat

and phenotype of L. n. leechi is closest to L. n. gueneei, this subspecies is genetically

closer to L. n. demuthi and L. n. albarracina. Pairwise F,, value between L. n. leechi and

L. n. gueneei was 0.93 compared to the F,; between L. n. leechi and L. n. demuthi which

was 0.13. High genetic differentiation was shown between L. n. gueneei and L. n. knilli.

Neighbour-joining tree based on mitochondrial marker showed only three popula-

tions as separate clusters: the population of L. n. gueneei was well separated (Fig. 2);

L. n. nickerlii, L. n. knilli and L. n. albarracina individuals formed three separate

SPALDING et al.: The genetics of Luperina nickerlii in Europe

nickerlii

Czech Rep.

Fig. 2. Unrooted neighbor-joining tree calculated from mitochondrial COI haplotypes sampled among 91

L. nickerlii individuals from 9 populations. Each haplotype is represented by a letter code (for distribution

and frequency see Tab. 1 and Fig. 1). Numbers above branches indicate percentage of bootstrap support out

of 1,000 repetitions. Shaded rectangles mark unique haplotypes belonging to a particular population. Both

L. n. knilli and L. n. albarracina (Spain) marked by an empty rectangle and dashed line contain haplotype

A beside the enclosed ones.

demuthi

nickerli

Czech Rep.

knilli

albarracina

Spain

Fig. 3. Parsimony network constructed from sequence of mitochondrial cytochrome c oxidase subunit I

gene (COI) of 91 L. nickerlii individuals from 9 populations. (L. nickerlii occurs elsewhere in Europe but

these populations were not sampled). Total number of 15 different haplotypes were sampled (A-O listed in

Tab. 2; for their prevalence see Fig. 1). Elipse areas are proportional to the haplotype frequencies. Haplotype

connections were parsimonious at the 95% level. Haplotype with the highest outgroup probability is

displayed as a square (A). L. nickerlii subspecies are marked with shaded rounded squares. Shape overlaps

denote sharing of the haplotype A between different populations. Subspecies /eechi consists of a single

haplotype (A) and is not highlighted in the figure. Numbers along lines are the nucleotide positions in the

sequence that changed. Empty nodes represent missing unsampled intermediate haplotypes.

Nota lepid. 36 (1): 35-46 4]

Tab. 4. Genetic diversity indices based on mt COI gene.

Gene diversity (h) Average no. of pairwise differences (x)

Population

demuthi 0.49 +/- 0.18 1.02 +/— 0.74

gueneei 0.22 +/- 0.17 0.22 +/- 0.29

knilli Inch 0.67 +/- 0.31 1.33 +/— 1.10

knilli Trabeg 0.00 +/— 0.00 0.00 +/— 0.00

leechi

nickerlii Mäslovice

nickerlii Praha

albarracina Amavida

albarracina Paramos 0.72 +/- 0.16

Tab. 5. Analysis of molecular variance (AMOVA). Mitochondrial sequence (COI) was used for the analy-

sis. For abbreviations see legend to Fig. 1.

= among populations | 053 | 63.09 | 0 | |

= within populations | 03728 | 36.91 | | Fe = 0.6309

= within populations | 03728 | 3598 | 0 | Fe = 0.6402

57.62

0 | Fes

|=within populations 0.3728 | 3426 | 0

60.54 | 0.0004 | Fer= 0.6054

clusters (Fig. 1). Topology of the remaining samples was not resolved. The populations

of L. n. leechi and L. n. gueneei were very homogeneous. On the other hand, high varia-

bility was revealed between populations of L. n. albarracina and L. n. knilli (Fig. 1).

Values of expected heterozygosity in populations sampled are low when compared with

published results in other species of Lepidoptera (Neve 2009). The estimates, however,

strongly depend on the markers used (e.g., microsatellites, allozymes) and many of

the studies listed by Neve (2009) have used allozymes. Among populations studied

here, L. n. leechi showed significantly lower heterozygosity (Tab. 4). Finer structure of

population genealogy was achieved by haplotype network construction (Fig. 3).

The AMOVA analysis showed the populations are structured but in contrast to our

prior expectations, L. n. leechi is genetically closer to L. n. demuthi, L. n. knilli and

L. n. albarracina (they widely share one haplotype) rather than to L. n. gueneei (Tab. 5).

The Mantel test for isolation by distance was not significant (r? = 0.014; p = 0.78).

4? SPALDING ef al.: The genetics of Luperina nickerlii in Europe

Discussion

We used mitochondrial markers to look at the current genetic composition of L. nickerlii

moths in Britain and Ireland in order to understand where they came from and in particular

why they are prone to splitting-off genetically identifiable groups that are isolated from

other groups. Taken together the genetic markers support the null hypothesis that all the

populations sampled belong to the same species Luperina nickerlii, despite differences

in appearance and population isolation — and some differences in ecology between UK,

Irish and mainland populations, although the alternative hypothesis that evolutionary

divergence is too recent to be fully reflected genetically could also be considered.

More detailed analyses of the subspecies populations reveals a complex pattern of

within species population differentiation. Population structure for Lepidoptera depends

on the spatial distance of habitats, the dispersal abilities of the species (Nève 2009) and

population origin (e.g., Hewitt 1996). Luperina nickerlii appears to show low dispersal

abilities, perhaps due to population isolation (Spalding & Young 2011b), although

occasionally singletons are found at some distance from known populations (Goater

1974; Wedd 1991), indicating dispersal activity. Results from the COI data indicate that

there may have been some historical gene flow between the subspecies as isolation by

distance was not supported in this study, suggesting populations were probably linked

to each other in the recent past, either reflecting a collapse in a former larger pre-

glacial range size or multiple post-glacial colonisation events; the degree of genetic

differentiation between the British populations may suggest the second hypothesis as

otherwise greater similarity between populations might be expected (e.g., Dapporto et

al. 2011). Movement is likely between continental populations, e.g., in Spain and the

Czech Republic, where several populations occur in close proximity, and also in Wales

and south-east England where L. n. gueneei and L. n. demuthi exist in extensive dune

(for L. n. gueneei) and saltmarsh (for L. n. demuthi) habitat. In contrast, the results for

L. n. knilli indicate little interchange between populations despite occupying the same

extended coastal cliff habitat in south-west Ireland and L. n. leechi is isolated by at least

300 km from known L. nickerlii populations.

The origin of the British subspecies is unclear. Those occurring on the western

fringes may be part of an Atlantic Arc species assemblage that includes species such

as the Quimper Snail Elona quimperiana (Férussac) (de Beaulieu & Le Moigne 1991)

and Killarney Fern Trichomanes speciosum Willd (Page 1997). L. n. leechi shows

some genetic similarity to L. n. albarracina, L. n. knilli and L. n. demuthi, but not

(despite our expectations) L. n. gueneei. It is possible that L. n. leechi is a population

founded by a single stray L. n. demuthi or L. n. albarracina. If so, L. n. leechi would

be more likely to feed as larvae on Festuca rubra L. or Puccinellia maritima (Huds.)

Parl. (Poaceae). However, it would appear that L. n. leechi may have been present on

or near Loe Bar long enough to adapt to a different habitat and transfer from former

food plants to Elytrigia juncea; in fact Festuca rubra is abundant in that locality. Rapid

changes in larval host-plant preferences have been reported in butterflies (e.g., Asher et

al. 2001; Pratt 1986-1987; Thomas et al. 2001) and moths, e.g., Lithophane leautieri

(Boisduval, 1829) (Noctuidae) (Young 1997). Further research, perhaps involving L.

nickerlii specimens from France and Portugal, may reveal additional linkages between

the subspecies.

Nota lepid. 36 (1): 35-46 43

The origin of the extensive populations of L. n. gueneei remains a mystery; this

subspecies appears to show significantly lower heterozygosity despite forming extensive

populations on the north coast of Wales and the west coast of Lancashire, with at least

some linkage between populations. Despite similarities in ecology to L. n. leechi (both

species occurring on coastal dunes and beaches and both feeding on Elytrigia juncea),

there is no indication that these two species have a common origin.

Genetic factors are important when assessing threatening processes and devising

conservation plans for threatened species (Frankham & Ralls 1998). From a genetics

perspective, the primary conservation goals are to preserve as much genetic diversity

and variability as possible as well as the evolutionary processes responsible for this

diversity (Clarke & O’Dwyer 2000; Coates 2000; Crandall et al. 2000). It is perhaps

useful to rank populations on patch size, habitat quality and land tenure (Clarke &

O’ Dwyer 2000), variation in phenotype (Crandall et al. 2000) and host-plant performance

(Legge et al. 1996) in addition to genetic diversity. The continental subspecies appear

to have similar ecologies although there are some phenotypic and genetic differences;

the British and Irish subspecies show phenotypic and genetic differences as well as

having different host plants and habitats. Populations of L. n. tardenota, L. n. gueneei,

L. n. leechi and L. n. knilli appear to be small and possibly declining; we provisionally

suggest that key conservation effort should be directed to these subspecies.

However, not all subspecies should be considered equal (Ryder 1986). It is important

to take account of the evolutionary processes associated with current levels of species

diversity at the approriate geographical scale (e.g., Coates 2000). Low heterozygosity

combined with increased levels of inbreeding associated with a limited number of

individuals and a distorted sex ratio have been shown to decrease survival rates (e.g.,

Gerber 2006; Saccheri et al. 1998) and the small isolated population of L. n. leechi may

not survive for long. The phenotypic characters that have been used to differentiate L.

n. leechi as a subspecies are perhaps subject to environmental plasticity and may not

be under genetic control. In this case this population would no longer be considered

of conservation importance as L. n. leechi contains a single haplotype that is widely

shared with other subspecies (e.g., L. n. demuthi and L. n. albarracina). The lack of

genetic diversity possibly as a result of its recent origin and the small population size

suggests that this isolated subspecies may be less worthy of conservation than some of

the other subspecies. The case for L. n. gueneei is less clear as this subspecies possesses

a unique haplotype and forms extensive populations on the north coasts of Wales and

north-east Lancashire.

Acknowledgements

We would like to acknowledge insect collection by the following people: Jiti Skala (Prague, Czech

Republic), Jifi Darebnik (Holeëov, Czech Republic), Jaroslav Zäme£nik (Muzeum vychodnich Cech v

Hradci Krälové, Hradec Krälové, Czech Republic), Arcadi Cervellö (Societat Catalana de Lepidopterologia,

Barcelona, Spain) and Jordi Dantart (Museu de Ciéncies Naturals de Barcelona, Barcelona, Spain). We

would also like to thank Martin Honey of the Natural History Museum London for obtaining the reference

for the description of subspecies albarracina. Further, our thanks are due to Paul Wilkinson for sequencing

and to Martina Zurovcova (Institute of Entomology, Biology Centre ASCR, Ceské Budéjovice, Czech

Republic) and Alexie Papanicolaou (CSIRO Ecosystem Sciences, Canberra, Australia) for suggestions on

44 SPALDING et al.: The genetics of Luperina nickerlii in Europe

the data analysis. We are also grateful to the three anonymous referees for their comments which helped

very much improve this paper. This research was funded by a voucher from the Technology Transfer office

at the University of Exeter under the Smart-Solution initiative and by Spalding Associates (Environmental)

Ltd of Truro.

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Nota lepid. 36 (1): 47-52 47

The status of Satyrus abramovi var. korlana Staudinger, 1901

(Nymphalidae)

STANISLAV K. KORB

a/ya 97, Nizhny Novgorod. 603009 Russia; stanislavkorb@list.ru

Received 14 August 2012; reviews returned 9 October 2012; accepted 8 January 2013.

Subject Editor: Zdenék F. Fric.

Abstract. A new status for Satyrus abramovi var. korlana Staudinger, 1901 as a subspecies of Karanasa

regeli (Alphéraky, 1881) is proposed. The diagnostic characters of K. abramovi (Erschoff, 1884) and K. re-

geli (Alphéraky, 1881) in male genitalia are discussed. Lectotypes are designated for Satyrus abramovi var.

korlana Staudinger, 1901 and Satyrus regeli var. regulus Staudinger, 1887.

Pe310Me. VCTaHOBJIeHO, YTO TaKcoH Satyrus abramovi var. korlana Staudinger, 1901 aBıseTca NOABUOM

Karanasa regeli (Alphéraky, 1881). BpinesteHnpi xnarHocTuuecKkye npu3HakH K. abramovi (Erschoff, 1884)

u K. regeli (Alphéraky, 1881) B reHuTarınax camyosB. O603HayeHbI JIEKTOTUNBI Satyrus abramovi var. kor-

lana Staudinger, 1901 u Satyrus regeli var. regulus Staudinger, 1887.

Introduction

The taxonomy of some of the taxa of the genus Karanasa Moore, 1893 inhabiting

high mountainous Central Asıa remains unclear. One of these taxonomic problems

is the status of the species-group taxon Satyrus abramovi var. korlana, described by

O. Staudinger from “Korla” (eastern extensions of Tian-Shan in China near the city

of Korla). A. Avinoff & W. R. Sweadner (1951) listed this taxon as the separate spe-

cies Karanasa korlana (op. cit.: 101) and also as a subspecies of K. regeli (Alphéraky,

1881) (op. cit.: 191, 195). These two authors have been the last who revised the genus

Karanasa, but they did not use genitalia features in their revision. For clarification of

this problem, to resolve the status of the taxon kor/ana, I revised its type material as well

as the type material of other closely related taxa.

Abbreviations

SK S. K. Korb collection, housed in Nizhny Novgorod, Russia

ZMMU Zoological Museum of the Moscow University, Moscow, Russia

ZMHB Museum fiir Naturkunde an der Humboldt-Universitat, Berlin, Germany

Material and methods

The following name-bearing types have been studied: syntypes of Satyrus abramo-

vi var. korlana; lectotype of Satyrus regeli Alphéraky, 1881 (lectotype designated by

Korb 2012: 46); syntypes of Satyrus regeli var. regulus Staudinger, 1887 (all three taxa

are currently classified in Karanasa).

Nota lepidopterologica, 17.06.2013, ISSN 0342-7536

Kors: Status of Satyrus abramovi var. korlana

Figs 1-8. Karanasa Moore, 1893. 1, 2. topotype male, upperside (1) and underside (2), 12.vi11.2006,

Kyrgyzstan, Chatyr-Kul Lake environs, 2800 m. 3, 4. Karanasa regeli korlana (Staudinger, 1901), lec-

totype male, upperside (3), underside (4), Korla. 5,6. Karanasa abramovi regulus (Staudinger, 1887),

lectotype male, upperside (5), underside (6), Transalai. 7, 8. Karanasa regeli (Alphéraky, 1881), lectotype

male, upperside (7), underside (8), Tian Shan.

Nota lepid. 36 (1): 47-52 49

Ko rla

Figs 9, 10. Lectotype labels. 9. Karanasa regeli korlana (Staudinger, 1901). 10. Karanasa abramovi re-

gulus (Staudinger, 1887).

For nomenclatural stability and to fix the exact type locality I designate here the

lectotype of Satyrus abramovi var. korlana. For the same reason I designate here the

lectotype of Satyrus regeli var. regulus. Lectotypes are preserved in ZMHB.

Satyrus abramovi var. korlana Figs 3, 4, 9, 12, 15

Material. Lectotype ©, ‘Origin.’; ‘v. Korlana Stgr.’; ‘Korla’; ‘Abramovi | v.’; printed on red paper

‘LECTOTYPUS @| korlana | Stgr. | S. K. Korb design. 17.04.2012’. Paralectotype 19.

Satyrus regeli var. regulus Figs 5, 6, 10, 13, 16

‘LECTOTYPUS © | regulus | Stgr. | S. K. Korb design. 17.04.2012’. Paralectotypes 30, 19.

Karanasa regeli (Alphéraky, 1881) Figs 7-10, 12, 14, 15

Material. Kazakhstan: 120°, 39, Transilian Alatau Mts, Assy valley, 24.vii.2010, leg. P. Egorov, SK;

29, Transilian Alatau Mts, Koram, 2200 m, 14.vii1.1957, Panfilov leg., ZMMU. Kyrgyzstan: 20, 19,

Kungey Ala-Too Mts, Grigoryevskoye valley, 2500 m, 12.viii.2003, leg. S. K. Korb, SK; 19, Kungey

Ala-Too Mts, Temirovka, vi11.2010, local collector, SK; 19, 19, Kungey Ala-Too Mts, Toguzbulak, 2000

m, 7.vili.2003, leg. S. K. Korb, SK.

Karanasa abramovi (Erschoff, 1884) Figs 1-8, 11, 13, 16

Material. Kyrgyzstan: 20, Chatyr-Kul Lake environs, 2800 m, 12.viii.2006, leg. S. K. Korb, SK;

29, Kyrgyz Mts, Ala-Archa Nature Reserve, upper course of Ala-Archa river, 3000 m, 02.viii.2003, leg. S.

K. Korb, SK; 69, 29, West Tian-Shan <sic!>, Dolon Pass, 3000 m, 9.viii.1967, leg. A. Tsvetaev, ZMMU;

29, Transalai Mts, Aram-Kungei, unknown collector, SK.

50 Kors: Status of Satyrus abramovi var. korlana

Figs 11-14. Male genitalia. 11. Karanasa abramovi (Erschoff, 1884), topotype. 12.v111.2006, Kyrgyzstan,

Chatyr-Kul Lake environs, 2800 m. 12. Karanasa regeli korlana (Staudinger, 1901), lectotype. 13. Kara-

nasa abramovi regulus (Staudinger, 1887), lectotype. 14. Karanasa regeli (Alphéraky, 1881), paralecto-

type, Tian Shan.

Figs 15, 16. Vesica and distal part of phallus. 15. Karanasa regeli korlana (Staudinger, 1901), lectotype.

16. Karanasa abramovi regulus (Staudinger, 1887), lectotype.

Discussion and Conclusions

During the examination of the type material and additional specimens the following

differences between K. regeli and K. abramovi were found in the male genitalia (Figs

11—16). In X. abramovi the valva is elongated, with no extension in its distal part; in K.

regeli it is more massive, with an extension in its distal part. In K. abramovi the dorsal

Nota lepid. 36 (1): 47-52 51

Fig. 17. Variability of genitalia in Karanasa species. A-D. K. regeli (Alphéraky, 1881), Transili Alatau

Mts, Assy valley (A, B), Terskey Ala-Too Mts, Pokrovka environs (C, D). E-H. K. abramovi (Erschoff,

1884), Akshiyrak Mts, Dolon Pass.

32 Kors: Status of Satyrus abramovi var. korlana

side of the valva continues towards the apex in a more or less smooth line, whereas in

K. regeli it forms a bend at about one quarter from the apex. In K. abramovi the out-

growth on the dorsal side of the valva at about one third from the base is always pointed

and can be divided into two or three parts, whereas in K. regeli it is always somewhat

rounded and forms one whole. In K. abramovi the dorsal teeth on the valva, mostly pre-

sent apically, are always separated from this outgrowth with an area without teeth; in

K. regeli they start almost immediately after the outgrowth. In K. abramovi the vesica

has two small spine-like cornuti; in K. rege/i it has two quite large scale-like cornuti.

Due to the fact that genitalia features of the taxon korlana match much more closely

those of K. regeli than those of K. abramovi, korlana is now considered a subspe-

cies of K. regeli. It is its most widely distributed subspecies, distributed in Khalyktau,

Borokhotan, Narat, Borto-Ula, Kuruktag, Avral-Ula and the Uken Mountains in north-

western China. Both species (X. abramovi and K. regeli) are similar in wing pattern but

significantly different in genitalia (as described above).

Only one infrasubspecific taxon for this group is currently established: K. abramovi

ab. erschovi Avinov, 1910 (a yellow aberration of the female). Individual variation in

both taxa is only present in the colour and width of the light-coloured band on the upper-

side of the wing and in the size of the eyespots. Wing pattern variability in both species

is very large. Very high variation in genitalia structures is also present, but this does not

obscure the specific features as the variation only occurs in smaller details. This varia-

tion should be studied in more detail in the future (Fig. 17).

Acknowledgments

I thank Dr. A. V. Sviridov (Zoological Museum of the Moscow State University, Moscow, Russia) and

Dr W. Mey (Museum fiir Naturkunde, Berlin, Germany) for providing access to collections and type

specimens. I am very thankful also to the first anonymous referee of the manuscript, who worked hard to

improve this paper.

References

Avinoff, A. & W. R. Sweadner 1951. The Karanasa butterflies, a study in evolution. — Annals of the

Carnegie Museum 32 (1): I-II + 1-217.

Korb, S. K. 2012. Butterflies (Lepidoptera: Papilionoformes) of North Tian-Shan. Part 1. Hesperiidae,

Papilionidae, Pieridae, Libytheidae, Satyridae. — Eversmannia Suppl. 3: 1—84.

Nota lepid. 36 (1): 53-55 53

The “Omnivorous Leafroller”, Platynota stultana Walsingham,

1884 (Tortricidae: Sparganothini), a new moth for Europe

FRANS GROENEN ! & JOAQUIN BAIXERAS ?

! Dorpstraat 171, 5575 AG Luyksgestel, The Netherlands; groene.eyken@chello.nl

2 Institut Cavanilles de Biodiversitat i Biologia Evolutiva, Universitat de Valencia,

C/ Catedratic Jose Beltran 2, 46022 Valencia, Spain; joaquin.baixeras@uv.es

(corresponding author)

Received 2 February 2013; reviews returned 25 February 2013; accepted 6 March 2013.

Subject Editor: Jadranka Rota.

Abstract. Platynota stultana Walsingham, 1884, a polyphagous tortricid and economically important spe-

cies, is formally recorded for the first time for Europe.

Introduction

Platynota stultana Walsingham, 1884 is an invasive species of Tortricidae native to

Mexico and the southwestern United States, accidentally introduced to the Hawaiian

Islands (Miller 1995). Known in the entomological literature as the “omnivorous leaf-

roller”, its potential range of food plants includes more than 20 plant families including

relevant ornamental plants, agricultural crops, and even forest species (Powell & Brown

2012).

Its presence in Europe was first detected in 2009 by pest control services of the

provinces of Murcia and Almeria in Spain during routine monitoring of agricultural

areas, mostly on pepper crops (Capsicum sp., Solanaceae). Although there has been no

reaction in the entomological literature, several popular electronic agricultural journals

and leaflets have included information on this pest and provided details on its distribu-

tion and potential control in Spain (Hymenoptera 2011).

Parallel field work developed in Spain in the period 2005-2008 in the provinces

of Almeria, Alicante, and Granada by A. Cox and M. Delnoye rendered a good se-

ries of specimens of an unknown Sparganothini species that was finally identified by

A. Schreurs and the first author of this paper as belonging to P. stultana. Because of

the economic importance of the species and the limited attention that the entomologi-

cal literature has paid to this new pest introduction, it seems appropriate to publish this

note to formally record its presence in Spain.

Platynota stultana is a small moth, the wingspan of the male is 10—15 mm and of

the female 14—19 mm. As in most members of the tribe Sparganothini, the labial palpi

are long and frontally projected. This character is not found in the European fauna ex-

cept in the few species of the genus Sparganothis Hübner, reducing potential mistakes

in identification. Male forewings possess a small costal fold. The general upperside

ground colour is brown in the approximately basal half and golden brown in the distal

half (Fig. 1A). In the female the markings are less distinct (Fig. 1B). Some colour varia-

tion is common. Male and female genitalia include unmistakable features (Figs 1C, D).

Nota lepidopterologica, 17.06.2013, ISSN 0342-7536

54 GROENEN & BAIXERAS: Platynota stultana

Fig. 1. Platynota stultana. A: Male (Cabo de Gata, Almeria, Spain). B: Female (Granada, Spain). C: Male

genitalia (GS: FG2409). D: Female genitalia (GS: FG2408).

Detailed information on its morphology and biology is compiled by Powell & Brown

(2012):

In the United States the moth has 4—6 generations a year. The female lays a patch of

about 100 eggs. After hatching the larvae move to the top of the plant and feed within

a bud or between the two leaves. In greenhouse conditions the larvae are fully grown

within a period of 20-30 days. They hibernate between the third and fifth instar in

webbed nests. Pupation takes place in a rolled leaf.

Material. Spain, 29 specimens, Almeria, Aqua Dulce, x.2005, leg. AC; 3 specimens, Alicante, La Ma-

rina, leg. MD; 18 specimens, Almeria, Cabo de Gata, vi.2007; 5 specimens, same locality but dated x.2007;

82 specimens, Granada, Castillio de Banos, x.2008, genitalia slides FG21380, FG23549, FG2355¢0;

10 specimens, same locality but dated vi.2010; 1 specimen, xi.2011 [GNL, AS, AC].

Nota lepid. 36 (1): 53-55 55

Abbreviations

AS Collection A. Schreurs, Kerkrade, The Netherlands

AC Collection A. Cox, Mook, The Netherlands

GNL Collection F. Groenen, Luyksgestel, The Netherlands

MD Collection M. Delnoye, Susteren, The Netherlands

Acknowledgements

The authors want to express their gratitude to John Brown (USDA, Washington, USA), Tomas Cabello

(University of Almeria, Spain), Anton Cox and Martin Delnoye (The Netherlands), Ferran Garcia-Mari

(Polytechnic University of Valencia, Spain), Arnold Schreurs (The Netherlands), Marja von der Straten

(Plant Protection Service, The Netherlands), and Boyan Zlatkov (Sofia University, Bulgaria) for their col-

lections, information, and helpful comments.

References

Miller, S. E. 1995. Platynota stultana, the omnivorous leafroller, established in the Hawaiian Islands (Le-

pidoptera: Tortricidae). — Bishop Museum Occasional Papers 42: 36-39.

Powell, J. A. & J. W. Brown 2012. The Moths of North America. Fascicle 8.1, Tortricoidea, Tortricidae

(Part), Sparganothini and Atterini. - The Wedge Entomological Research Foundation, Washington,

229 pp.

Hymenoptera 2011. Platynota stultana, un nuevo lepidöptero plaga en el sudeste español. — Homo agri-

cola, 1: 33-38.

Nota lepid. 36 (1): 57-64 51

Habitat preferences of butterflies (Papilionoidea)

in the Karpaz Peninsula, Cyprus

ÖZGE ÖZDEN

Department of Landscape Architecture, Faculty of Architecture, Near East University, Nicosia,

North Cyprus Mersin 10 Turkey; ozgeozden77(@yahoo.com

Received 23 August 2012; reviews returned 1 November 2012; accepted 8 March 2013.

Subject Editor: Zdenék F. Fric.

Abstract. The Mediterranean region comprises of some of the world’s unique and biogeographically im-

portant areas, harbouring high levels of biological diversity. On the other hand, anthropogenic disturbances

are causing degradation of diverse ecosystems within the region. The aim of this study was to determine

the habitat preferences of butterfly species and the potential threats they may face within the Karpaz

Peninsula of Cyprus. To understand the importance of local vegetation characteristics of butterflies in the

Karpaz Peninsula, ‘Pollard Walk’ transect counts were used to assess the abundance and species richness

of butterflies. Butterflies resting on plants and those in flight were counted and identified. Preferred plant

species and habitat types (EUNIS and EU Habitats) of the butterflies are also identified. During the surveys

in 2006, eleven butterfly species were recorded. Two of them (Glaucopsyche paphos and Maniola cypri-

cola) are endemic to Cyprus. Construction developments and road improvements were recorded within

the region and have resulted in habitat loss and degradation. Our results provide valuable knowledge about

important habitats for Cypriot butterflies within the Karpaz Peninsula and additionally highlight the need

for their conservation in the face of large infrastructure developments and unregulated construction.

Introduction

The Mediterranean Basin is rich in biodiversity and in need of conservation (Myers

1990). Cyprus is the third largest island within the Mediterranean Basin, after Sicily and

Sardinia. It harbours a variety of ecosystems including pine forests, garrigue, maquis,

rocky areas, coastal rocky areas, coastal dunes, wetlands, and agricultural areas togeth-

er with a high number of threatened and endemic plant and animal species (Baier et al.

2009; Flint & Stewart 1992; Makris 2003; Tsintides 1998; USAID 2006). The northern

part of the island has been politically isolated for many years and as a result, develop-

ment has been relatively slow compared with similar regions in the Mediterranean. In

general, the northern part of the island offers a range of varied terrestrial habitats, such

as pine forests (both lowland and mountain), juniper shrubs, garrigue, phrygana, lime-

stone pavements and dune vegetations (Tsintides 1998; Viney 1994).

The Karpaz Peninsula is biologically one of the most important areas in Cyprus. It

is situated at the easterly point of the Five Finger Mountain Range (Besparmak Sira

Daglan). It consists of hill-like formations covered with maquis, pine forests, olive and

carob plantations, plains containing arable lands, semi-dry stream beds and a coastal

zone. The Karpaz Peninsula is particularly known for its unspoilt landscapes and its

interesting wildlife; therefore it is an important ecotourism area. The area has recently

received official legal protection as an important natural resource for the northern part

of the island and was declared a “Special Environmentally Protected Area” according

to Environment Law (21/97) article 11 by the Turkish Cypriot authorities. The Karpaz

Nota lepidopterologica, 17.06.2013, ISSN 0342-7536

58 OzpeNn: Habitat preferences of butterflies in the Karpaz Peninsula

Special Environmentally Protected Area has been selected due to the occurrence of in-

ternationally important habitats and species, including marine turtles Chelonia mydas

(Linnaeus, 1758), Caretta caretta (Linnaeus, 1758), Audouin’s Gull Larus audouinii

Payraudeau, 1826, and Mediterranean Monk Seal Monachus monachus (Hermann,

1779) (Godley & Broderick 1992; Haigh 2004; Iris & Gucel 2008). Karpaz is not only

known for important animal species; it also harbours many endemic and rare plant spe-

cies such as Cyprus Orchid (Ophrys kotschyi H. Fleischm. & So6) which is listed under

EU Annex II plant species (European Commission 2007a; Kreutz 2004).

Although the peninsula is declared as a “Special Environmentally Protected Area”

by local authorities, herbicide and pesticide use in agricultural fields is still allowed.

Agricultural cereal fields in the area are mainly wheat and barley production areas and

are open monocultures. These areas are not irrigated and farmers apply shallow plough-

ing in these monoculture fields. The field margins between the grassland fields are not

more than 1.5 m wide.

It is known that about one third (31%) of European butterflies has declined over

the last 10 years (van Swaay et al. 2010). There are many documented threats to but-

terflies in Europe, including the increasing use of agricultural herbicides and pesticides,

habitat loss, climate change, land management, agricultural conversion and fragmenta-

tion (Grill et al. 2005, Stefanescu et al. 2005; Wilson & Maclean 2011). Building and

infrastructure developments such as roads, quarries and housing are also strong drivers

of population declines affecting 80% of the threatened butterfly species within Europe

(van Swaay et al. 2009).

So far in Cyprus, 53 species and subspecies of butterflies have been recorded. They

include three endemic species (Maniola cypricola (Graves, 1928), Hipparchia cypri-

ensis Holik, 1949 and Glaucopsyche paphos Chapman, 1920) (Makris 2003). Two of

the endemic species, M. cypricola and G. paphos, are of European Conservation Con-

cern (van Swaay & Warren 1999). Here we used a case study of butterfly abundance

and behaviour within the Karpaz Peninsula in different habitat types. We compared

the butterfly abundance/activity and species richness between different habitat types

during the spring season. Understanding the response of different butterfly species to

different habitats is essential in order to design conservation management, especially

in Mediterranean mosaic landscapes (Pe’er et al. 2011). The aim of this study was to

determine the habitat preferences of butterfly species and the potential threats that they

may face within the Karpaz Peninsula of Cyprus.

Materials and Methods

This study was carried out in the Karpaz Peninsula of Cyprus between the beginning

of April until the end of May 2006. The vegetation in the area is mainly dominated by

Juniperus phoenicea L. (Cupressaceae), Olea europaea Linnaeus (Oleaceae), Ceratonia

siliqua L. (Fabaceae), Pistacia lentiscus L. (Anacardiaceae), Thymus capitatus (L.)

Hoffmanns. & Link (Lamiaceae), Sarcopoterium spinosum (L.) Spach (Rosaceae) and

Genista sphacelata Spach (Fabaceae). During the spring there are many wild flowers

in the area including several endemic plant species such as Anthemis tricolor Boiss.

Nota lepid. 36 (1): 57-64 59

34° 05' 34° 10' 34° 15' 34° 20' 34° 25' 34° 30'

(ind 2013 May 1 13:31:03 | seaturtle.org/maptool Projection: Mercator

Fig. 1. Numbered open circles (©) provide the location points for each butterfly-survey transect within the

Karpaz Peninsula.

(Asteraceae), Helianthemum obtusifolium Dunal (Cistaceae) and Ophrys kotschyi (Or-

chidaceae).

According to local records, the average annual rainfall in the area is 455—506 mm.

The highest rainfall occurs during December—January. The average temperature is

20°C in the region (Yeni Erenkoy Meterological Station).

The census procedure used for this research was the Pollard Walk method, as de-

scribed by Pollard (1977) and Royer et al. (1998). Five Pollard walk transects were es-

tablished across the Karpaz Peninsula during this project (Fig. 1.). Transects of a fixed

length (1 km) were walked and adult butterflies recorded. All transects were carried out

at the optimum time of the day for seeing butterflies (11.00-13.00 hours), on warm

sunny days at temperatures of 24—28°C, with little or no wind (Beaufort force 0-2)

and all transects were below 130 metres elevation. Transects were chosen to cover a

range of habitat types. Plant identifications were carried out during transect selection.

Butterflies nectaring on plants and those in flight were counted and identified. If the

exact identification of the species was not possible, a butterfly net was used to capture

those butterflies in question, in order to facilitate field identification using the avail-

able literature (Makris 2003; Tolman & Lewington 1997). After identification, captured

butterflies were released at their point of capture. In addition, butterflies nectaring on

plants were recorded together with the plant species.

60 Ozpen: Habitat preferences of butterflies in the Karpaz Peninsula

Tab. 1. Transects and habitat classifications for each transect

Transect | coordinates | EU EUNIS Common Habitat

number habitat type habitat type flowering plants | definition

35,59293 N J2 Low density | Onopordum Village Area

34,38642 E buildings cyprium,

Chrysanthemum

crononarium

35,59206 N | 5210 Arborescent Cistus spp., Arborescent

34,33646 E | matorral with Thymus capitatus | matorral

Juniperus spp.

3 35,46469 N E 2.6 Centaurea Agricultural area

34,14230 E Agriculturally- hyalolepis,

improved, Onopordum

re-seeded and cyprium

heavily fertilized

grassland,

including sports

fields and grass

lawns

35,60391 N | 5420 C2 Surface Few Cistus spp. |Phrygana

34,34566 E | Sarcopterium running waters and Onopordum

spinosum cyprium

phryganas

35,64884 N | 5210 Arborescent Cistus spp. Arborescent

34,46318 E | matorral with matorral

Juniperus spp.

Transect | was established within the Dipkarpaz village, and the transect was

walked along the house garden edges which were mainly dominated by Onopordum

cyprium Eig (Asteraceae), Chrysanthemum coronarium L. (Asteraceae), Cistus cre-

ticus L. (Cistaceae) and grassy patches. Transect 2 was established in the arborescent

matorral with Juniperus habitat. The habitat was dominated by Pistacia lentiscus,

Juniperus phoenicea and Cistus salviifolius L. (Cistaceae) with small grassland patch-

es. Transect 3 was established along the edge of the field crop (wheat and barley) grow-

ing area. The vegetation was dominated by Centaurea hyalolepis Boiss. (Asteraceae)

and Onopordum cyprium. Transect 4 was established along the Ronnas River and veg-

etation was dominated by Pistacia lentiscus. Transect 5 was established within the

arborescent matorral with Juniperus habitat. The habitat was dominated by Juniperus

phoenicea, Pistacia lentiscus and Calicotome villosa (Poir.) Link (Fabaceae) (Tab. 1,

Fig. 1).

The plant species were identified using Viney (1994) as a reference. In addition,

for each transect site the existing habitat type was also identified according to the

Interpretation Manual of European Union Habitats and EUNIS habitat classification

(Tab. 1) (Davies et al. 2004; European Commission 2007b).

Results and Discussion

Butterfly abundance and species richness were studied in different habitat types within

the Karpaz Peninsula. During the surveys, a total of 169 individual butterflies from

Nota lepid. 36 (1): 57-64 61

Tab. 2. Butterfly species and their total abundance observed in different habitats. T = Transect.

Species Name Village area | Arborescent | Agricultural | Phrygana Total number

(T 1) matorral area (T 3) (T 4)

(T 2 and T 5)

Pieris rapae

a Ce

Gee | 0 | 0° |

geo SD | of

Eiuaconiyehe pphos[ + | 9 |

mom 1 | © |

Rene fn un Je

En | | 0

ci OR ER CSSS

Éprausateon | 0 | + |

a RE PRE

Wanessa atatonta | 1 | + |

Total Abundance | 17 | 28 |

11 species were recorded across the five transects in the Karpaz Peninsula region of

Cyprus. Surprisingly, the highest number of butterflies (86) was recorded from agri-

cultural habitat, especially high abundance of the endemic species Maniola cypricola.

Most of the individuals recorded from farmland habitat were nectaring on Centaurea

hyalolepis along the field margins. Flower-rich field margins may be crucial for spring-

flying butterflies (Dover 1989), as nectar feeding increases individual longevity, fe-

male fecundity and patterns of oviposition in local populations (Erhardt & Mevi-Schiitz

2009; Stefanescu & Traveset 2009). An important factor behind butterfly losses is the

loss of flower-rich habitats from open farmlands (Nilsson et al. 2008).

The second highest number of butterflies (38) was recorded from Sarcopoterium spi-

nosum phrygana habitat (EU Annex I 5420) from the Ronnas River area. Sarcopoterium

phryganas are low thorny shrub-like formations within the thermo-Mediterranean zone

of Aegean islands, Greece, Coastal Anatolia and Cyprus. This habitat type harbours

many flowering and aromatic plant species such as Thymus capitatus, Cistus creticus,

Cistus salviifolius and Teucrium spp. (Lamiaceae) (EC 2007b). In particular Cistus spp.

and Teucrium spp. are important butterfly nectar sources in Cyprus (Ozden & Hodgson

2011). Species richness of butterflies was similar within four transects apart from the

species-poor arborescent matorral habitat transect (Tab. 2).

Maniola cypricola (60), Thymelicus acteon (Rottemburg, 1775) (27) and Vanessa

cardui (Linnaeus, 1758) (20) were the three most abundant species observed from dif-

ferent transects. The endemic M. cypricola was recorded from arborescent matorral,

phrygana habitats and agricultural farmlands but not from the village area. This obser-

vation was interesting, because M. cypricola is a very common species across Cyprus

(Ozden et al. 2008; Ozden & Hodgson 2011). Natural habitats and field margins may

provide suitable habitat for this species; however, this limited data cannot be consid-

ered conclusive. As expected, the generalist Pieris brassicae (Linneaus, 1758) was

recorded from all types of habitats (Tab. 2).

62 OzpeNn: Habitat preferences of butterflies in the Karpaz Peninsula

Tab. 3. Number of butterflies recorded while nectaring on different plant species. * — A total of 32 Maniola

cypricola was recorded on C. hyalolepis from agricultural area. ** — A total of 14 (all) Thymelicus acteon

was recorded on C. hyalolepis from agricultural area. *** — A total of 16 Vanessa cardui was recorded on

C. hyalolepis from agricultural area.

cyprium creticus hyalolepis sphaceleata

Creme | 0

[ Collas erocea | 0

| Femuenzm

RE ee.

RUES

ren

Fee

Peete:

Perez

BEE:

Vanessa cardui Be

Maniola cypricola

| we _

pi. 87 | ON

Vanessa atalanta

Total Numbers

Regarding nectaring records, the highest number of butterflies utilised Centaurea

hyalolepis (82 individuals) as a source of nectar. Thymelicus acteon was also re-

corded nectaring on Centaurea hyalolepis (Tab. 3). T. acteon is considered as a Near

Threatened species at the European level (van Swaay et al. 2010). Plants of the genus

Centaurea are widely regarded as providing a good source of nectar. The high nectar

yield of plants in this genus makes it very attractive to insects, especially butterflies

(Wackers et al. 2005). Centaurea hyalolepis is spreading annually or biennially reach-

ing a height of 60 cm and is much-branched; it is a common plant throughout northern

Cyprus, flowering from April to July.

Conclusion

In conclusion, the results presented in this paper provide valuable information on

Cypriot butterflies and their close relationship with different habitat types within the

Karpaz Peninsula. Further research is needed in order to discover which plant species

are preferred sources of nectar for butterflies over a longer time frame.

Observations of butterfly behaviour have revealed that the flora of the field mar-

gins provides rich nectar sources for butterflies in Karpaz. Therefore, local author-

ities should be made aware of the importance of field margins in cereal farmlands

and this information should be used when implementing the management plans of the

EU Habitats Directive within the agricultural farmland ecosystems of Karpaz Special

Protected Area. Also, for the future protection of special biodiversity-rich habitats, un-

necessary road improvements along with uncontrolled building constructions should be

excluded from this area.

Nota lepid. 36 (1): 57-64 63

Acknowledgements

This research was supported by United Nations Development Programme Partnership for the Future

(UNDP-PFF). I would like to thank Chris van Swaay (Dutch Butterfly Conservation) for his encourage-

ment to publish this research. Maptool, a program for analysis and graphics (product of Seaturtle.org,

www.seaturtle.org), was used in this paper.

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Nota lepid. 36 (1): 65-75 65

Occurrence of Borearctia menetriesii (Eversmann, 1846)

(Erebidae: Arctiinae) in Northern European Russia:

a new locality in a disjunct species range

Ivan N. BoLoTov”, MIKHAIL YU. GOFAROV, YULIA S. KOLOSOVA &

ARTYOM A. FROLOV

Institute of Ecological Problems of the North, Ural Branch of the Russian Academy of Sciences,

Northern Dvina Emb., 23, 163000 Arkhangelsk, Russian Federation

* corresponding author; inepras@mail.ru

Received 18 December 2012; reviews returned 7 February 2013; accepted 22 March 2013.

Subject Editor: Alberto Zilli.

Abstract. Disjunctive distribution is typical for the transpalaearctic tiger moth Borearctia menetriesii

(Eversmann, 1846) (Erebidae: Arctiinae) at the present time. The new discovery at the East European

(Russian) Plain modifies the previously known distribution pattern of this species. A specimen of this moth

was recorded on a small patch of a humid mixed-herb meadow on the forest karst landscape of the White

Sea-Kuloi Plateau. Its location is in the upper part of the Sotka River Valley and is surrounded by northern

primary forest of spruce with inclusions of larch patches. It is characterised by a cold microclimate. This

paper summarises data on previously known localities of B. menetriesii that are situated mainly within the

Russian Federation and analyses the species distribution using Bailey’s Ecoregions system.

Introduction

The Menetries’s Tiger Moth Borearctia menetriesii (Eversmann, 1846) (Erebidae: Arc-

tiinae) is one of the rarest species of Palaearctic tiger moths and disjunctive distribution

is typical for ıt (Dubatolov 1984, 2010; Kurentzov 1965, 1973). Contemporary data

include the following areas in this species range: Europe: Middle-Finland; European

Russia: Karelia and Ural Mountains; Siberia: lower Ob river, the Altai and Sayan

Mountains, Baikal, Transbaikalia, Evenkia and Yakutia; Kazakhstan: northeastern re-

gion; Far Eastern Russia: northern region of Amur basin, Sikhote-Alin Mountains and

Sakhalin Island (Dubatolov 1996, 2010; Dubatolov & Gordeeva 2005; Ermakov 2006;

Nupponen & Fibiger 2012; Saarenmaa 2012).

Single specimens were found in most of the listed localities (Dubatolov 1984,

2009, 2010; Koshkin 2010; Krogerus 1944; Marttila et al. 1996; Shodotova et al. 2007;

Silvonen 2010). Sometimes, the records are separated from each other by decades, for

example in Finland (Fabritius 1914; Krogerus 1944; Marttila et al. 1996; Lappi et al.

2004; Silvonen 2010) and Sakhalin Island (Hori 1926; Klitin 2009). In recent times

(i.e. 21st century) species records are known for Finland (Lappi et al. 2004; Silvonen

2010) and Russia: Yamal-Nenets District (Gorbunov & Olschwang 2012), North Ural

Mountains (Ermakov 2006; Nupponen & Fibiger 2012), Kuznetsky Alatau Mountains

(Sutchev & Skalon 2012), Transbaikalia (Saarenmaa 2012), Sikhote-Alin Mountains

(Silvonen 2010) and Sakhalin Island (Klitin 2009).

The largest gap in the species range is situated in the territory of north-eastern

Europe: in the West there are only a few localities in East Fennoscandia and in the East

Nota lepidopterologica, 17.06.2013, ISSN 0342-7536

66 Bototov et al.: A new locality for Borearctia menetriesii in Northern European Russia

we know only of North Ural and Western Siberian records (Dubatolov 2004, 2010;

Ermakov 2006; Nupponen & Fibiger 2012).

Larval development of B. menetriesii was described for the first time by Krogerus

(1944) and was recently illustrated in detail via in vitro observations (Saarenmaa 2012).

The aim of the present paper is to analyse the distribution range of B. menetriesii in the

light of the first species record from the White Sea-Kuloi Plateau, Northern European

Russia.

Material and methods

The White Sea-Kuloi plateau is the largest karst region of Northern European Russia,

with an area of approximately 24,000 km?. The average plateau altitude varies from 70

to 140 m.a.s.l. (Gofarov et al. 2006). Lower Permian gypsum and anhydrite rocks form

the eastern part of the plateau and intensive karst processes occur in this area.

The karst landscape environment on White Sea-Kuloi Plateau has specific ecologi-

cal conditions and the most important are the following: (1) high relief heterogeneity:

alternating karst craters and ravines, deep incised river valleys, rock outcrops, slide-

rocks and caves with perennial subterranean ice; (2) high levels of stream flow and

low levels of waterlogging; (3) significant microclimate variability, from very cold to

warm conditions, which is determined by a high complexity of vegetation cover; (4)

high mineralisation of ground waters and domination by carbonate-rich soils in the soil

coverage; (5) high activity of exogenous geological processes (karst formation) and

soil erosion (Puchnina et al. 2000; Shvartsman & Bolotov 2008).

Siberian spruce (Picea abies ssp. obovata (Ledeb.) Domin, Pinaceae) and larch

(Larix sibirica Ledeb., Pinaceae) primary forests dominate the vegetation cover.

According to data from Pinega meteorological station (for the period 1903-2003), the

annual mean air temperature is 0.1°C, annual precipitation is 554 mm, the mean air

temperature of the coldest month (January) is -15.0°C and of the warmest month (July)

is 15.5°C; the summarised daily means above 10°C equal 1216°C.

We were conducting our entomological research on the Sotka River shore (Kuloi

River drainage) during the periods 7—17.vi.2000, 13—-20.v11.2000, 21 -28.vin1.2000,

26.v11.2001, 26-31.v111.2004, 8—11.vii.2005, 21-22.v11.2007. We collected main-

ly butterflies (Papilionoidea), but several individual moths were caught selectively.

Butterfly nets were used for collecting.

Data on other B. menetriesii localities were obtained from different studies (see

Appendix). We included only reliable references where species identification was veri-

fied by specialists; for the majority of Russian localities identification was performed by

Dr. V. V. Dubatolov (Siberian Zoological Museum of the Institute of Animal Systematics

and Ecology, Siberian Branch of the Russian Academy of Science, Novosibirsk city).

The arrangement of the localities was digitised and mapped using ESRI ArcGIS 10. We

mapped only those data pertaining to collected specimens in order to avoid including

several visual and non-specific records (Appendix). The presumed error of determi-

nation of the locality coordinates is around + 1-2 km, because published records of

moths are usually ascribed to approximate locations (for example, near a certain vil-

Nota lepid. 36 (1): 65-75 67

Fig. 1. Borearctia menetriesii specimen (female) from the Sotka River valley, Arkhangelsk Oblast, North-

ern European Russia (photo by Yu. Kolosova).

lage). Two localities were digitised from the general range map by Dubatolov (2010)

and the obtained coordinates are highly approximated, probably by around + 5 km. We

used Bailey’s Ecoregions Map of the Continents (Bailey 1989) for generalised estima-

tion of the species preferences for ecosystem types. On this map, ecosystem units of re-

gional extent (ecoregions) are marked by climate and vegetation. An Arcview shapefile

containing ecoregions map data was obtained from the Global Ecosystem Data Base of

NOAAs National Geophysical Data Center, Boulder, Colorado, USA.

Results

The record of a female specimen of B. menetriesii was made on 9.v11.2005 (Fig. 1). The

moth flew at the top of tall mixed-herb vegetation and was caught during the flight (I. N.

Bolotov leg.). This specimen is deposited in the collection of the Biological Museum of

the Institute of Ecological Problems of the North, Ural Branch of the Russian Academy

of Sciences (Arkhangelsk city).

Locality description: Upper part of the Sotka River valley; 64°38'59” N, 43°04’ 08” E;

altitude 37 m.a.s.l; 16 km WSW from the Pinega settlement; karst landscape; a small

patch of natural humid mixed-herb meadow (Figs 2a, b). The dominant species of the

meadow were Aconitum septentrionale Koelle and Thalictrum sp. (Ranunculaceae),

Geranium sylvaticum L. (Geraniaceae), Filipendula ulmaria (L.) Maxim. (Rosaceae),

Cirsium oleraceum (L.) Scop. (Asteraceae), Chamerion angustifolium (L.) Holub (Ona-

graceae), Paeonia anomala L. (Paeoniaceae) and Elymus caninus (L.) L. (Poaceae).

The meadow is surrounded by the primary Siberian spruce forest on gypsum soils with

68 BoLoTov et al.: A new locality for Borearctia menetriesii in Northern European Russia

Fig. 2. Habitat of Borearctia menetriesii. A, the Sotka River valley with karst gypsum outcrop, Pinega

region, Arkhangelsk Oblast, Northern European Russia. B, forest humid mixed-herb meadow where speci-

men was collected (photos by Yu. Kolosova).

15°E 20°E 25°E 30°E 35°E 40°E 45°E 55‘°E 145°E 150’E 155°E 160°E

1 L 1 1 L 1 u ' 1 fi aD ate f fi 1 fi fi

i 250 500 1000

Fig. 3. Distribution range of Borearctia menetriesii. Species localities: 1 — our record, 2 — previously pub-

lished records (see Appendix). Species locality numbers on the map correspond to numbers in the appen-

dix.

inclusions of larch (Larix sibirica) patches. Large gypsum outcrops (20-30 m high)

are found near the location where the specimen was collected (Fig. 2a). Subterranean

solution cavities, caves with cold microclimate and perennial ice are situated in these

outcrops. Sparse larch forests and tundra communities with numerous Arctic and

Arctic-Alpine vascular plants occur in the outcrops: Dryas octopetala L. and D. o.

ssp. punctata (Juz.) Hultén (Rosaceae), Arctostaphylos alpina (L.) Spreng. (Ericaceae),

Hedysarum arcticum B. Fedtsch., Astragalus norvegicus Grauer and Oxytropis camp-

estris ssp. sordida (Willd.) C. Hartm. (Fabaceae), Salix myrsinites L., S. arbuscula L.,

S. recurvigemmis A. Skvorts. and S. reticulata L. (Salicaceae); etc. The study area is

situated in the centre of a large primary taiga forest massif, belonging to Pinega State

Nota lepid. 36 (1): 65-75 69

Fig. 4. Proportions of numbers of Borearctia menetriesii localities situated in different Bailey’s Ecoregions

(Bailey 1989). A total data of 39 localities was used (published data in the appendix and our own record).

Ecoregions divisions: TD — tundra division, SD — Subarctic division, SM — Subarctic regime mountains,

PM - prairie regime mountains, WM — warm continental regime mountains. Ecoregions provinces: / —

Arctic tundra, 2 — continental and extreme continental light deciduous taiga, 3 — continental dark evergreen

needle-leaf taiga, 4 — eastern oceanic taiga, 5 — moderate continental dark evergreen needle-leaf taiga,

6 — forest-creeping trees-tundra of extreme continental climate, 7 — forest-tundra of moderately and con-

tinental climate, & — open woodland-creeping trees-tundra, 9 — open woodland-tundra, /0 — continental

steppe-forest-tundra and steppe-forest-meadows, 11 — oceanic forest-creeping trees.

Nature Reserve. Anthropogenic activity on the reserve territory has been totally pro-

hibited since 1976 and works are allowed only for reserve staff and several authorised

scientists.

Discussion

The new record of B. menetriesii in north-eastern Europe significantly reduces the gap

between two Borearctia derivatives (Fig. 3). In the upper part of the Sotka River val-

ley the karst landscape produces cold microclimate (Puchnina et al. 2000; Shvartsman

& Bolotov 2008) by the action of two factors: the cold influence of karst groundwater

outpouring and the refrigerant effect of ice caves on the air of the river valley. Boreal

karst areas have a more continental climate in comparison to neighbouring territories

(Puchnina et al. 2000; Shvartsman & Bolotov 2008). This fact confirms the opinion

about the relative continentality of B. menetriesii (Kaisila 1947). Another important

fact is that larch forest patches are common in the karst landscapes of the White-Sea

Kuloı plateau, including the Sotka River valley (Puchnina et al. 2000; Shvartsman &

Bolotov 2008), because the Larix species are significant food plants in the majority of

the B. menetriesii range (Saarenmaa 2012).

Collected specimens of B. menetriesii inhabited only a few ecoregion types in spite

of very broad species range (Fig. 4 and Appendix). Most of the localities (19 sites, 49%

of the total known) were situated in two ecoregion provinces: 1) subarctic mountains

with forest-creeping trees-tundra of extreme continental climate (Transbaikalia and the

Russian Far East); 2) moderate-continental dark evergreen needle-leaf taiga (all north-

ern European localities). The northernmost species record was made at 71°52'N in

70 BoLoTov ef al.: A new locality for Borearctia menetriesii in Northern European Russia

Arctic tundra of the Yana-Indigirka Lowland of the Northern Yakutia (Appendix), but

scarce larch forests in the tundra of Yana-Indigirka can advance very far northwards

in the river valleys (including the Muksunuokha River valley where this specimen

was collected) and it is possible that B. menetriesii ranges as far north as larch forests.

The height of the Arsenyeva Mountain (1860 m, the Sikhote-Alin Mountains of the

Russian Far East) is the maximum altitude where a specimen of this species has been

collected.

According to the aforementioned, one can conclude that the distribution pattern

of B. menetriesii is primarily connected with ecosystems of primary taiga forests and

mountain forest-creeping trees-tundra with elfin forms of coniferous trees. These eco-

system types play the main role in vegetation cover of Northern Eurasia. In spite of

the exceptionally low abundance of B. menetriesii, we can assume that the scattered

transcontinental distribution of this species is determined by preference for specific

biomes.

Acknowledgements

The authors are grateful to Dr. A. Zilli and two anonymous reviewers for valuable comments on the manu-

script; and the employees of Pinega State Nature Reserve for great help in the field works. This study has

been supported by grants of the Russian Foundation for Basic Research (Grant no. 10—04—008970), the

President of Russia (no. MD—4164.2011.5), the Ural Branch of Russian Academy of Sciences; and the

Ministry of Science and Education of the Russia.

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BoLoTov et al.: A new locality for Borearctia menetriesii in Northern European Russia

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Nota lepid. 36 (1): 77-84 an

Two little-known species of Gelechiidae in the European fauna

OLexsıy V. BipziLyA |, OLE KARSHOLT ?

! Kiev National Taras Shevchenko University, Zoological Museum, Volodymyrska str., 60,

01601 MSP, Kiev, Ukraine; bidzilya@univ.kiev.ua

okarsholt@snm.ku.dk

Received 13 April 2013; reviews returned 22 April 2013; accepted 30 April 2013.

Subject Editor: Lauri Kaila.

Abstract. The identities of Doryphora orthogonella Staudinger, 1871 and Anacampsis azosterella Her-

rich-Schäffer, 1854 are discussed and they are confirmed as belonging to the genera Stomopteryx Heine-

mann, 1870 and Syncopacma Meyrick, 1925, respectively. Both species are redescribed based on types

and additional new material. The adults and the male genitalia of both species are illustrated. A lectotype

of Doryphora orthogonella is designated. Both species are new to the fauna of Ukraine, and Syncopacma

azosterella is shown to be relatively widely distributed in southern parts of central and eastern Europe and

in the Mediterranean.

Introduction

Gelechiidae is among the least known lepidopteran families in Europe (Bidzilya & Kars-

holt 2008). Although some progress has been made (e.g., Huemer & Karsholt 2010), there

are still numerous taxa awaiting revision. Here we deal with two such taxa. As a result of

study of material deposited at ZMKU two doubtful species of Gelechiidae belonging to

the genera Stomopteryx Heinemann, 1870 and Syncopacma Meyrick, 1925, currently in

the subfamily Anacampsinae (Karsholt et al. 2013), were found. Their identification ap-

peared problematic, and we therefore considered them worthy of detailed examination.

The first species, Stomopteryx orthogonella (Staudinger, 1871), was known only from

the type-series collected in Sarepta in Russia (nowadays a district of Volgograd) in 1871

and has not been recorded since its description. The discovery of two additional males of

this species from Ukraine encouraged us to re-examine the type material deposited in the

collection of ZMHU. As a result, S. orthogonella is redescribed here and its male genitalia

are described and illustrated for the first time.

The second species, Syncopacma azosterella (Herrich-Schäffer, 1854), has for a long

time been confused with other species of the genus Syncopacma. This was due to the fact

that its type material had not been previously revised. Recently the holotype of S. azoste-

rella was found by the second author, and this allowed us to clarify the status of this taxon

and link a number of regional records with this name.

Abbreviations of institutions

BMNH The Natural History Museum, London, U.K.

ZMHU Zoological Museum, Humboldt University, Berlin, Germany

ZMKU Zoological Museum, Kiev National Taras Shevchenko University, Kiev, Ukraine

ZMUC Zoological Museum, Natural History Museum of Denmark, Copenhagen, Denmark

Nota lepidopterologica, 17.06.2013, ISSN 0342-7536

Figs 1-3. Stomopteryx orthogonella (Staudinger). 1. Lectotype, Sarepta, wingspan 14.0 mm. 2. Paralecto-

type, Sarepta, wingspan 13.1 mm. 3. Male genitalia, lectotype.

Stomopteryx orthogonella (Staudinger, 1871) Figs 1-3

Doryphora orthogonella Staudinger, 1871: 307.

Aristotelia orthogonella (Staudinger); Meyrick, 1925: 42.

Stomopteryx orthogonella (Staudinger); Karsholt & Riedl, 1996: 118, 311.

19 1867, 10° 6.vili.1869, 10° 8.vili.1875 (all BMNH); Volgograd, 10° 18—24.v.1967, leg. V. Zouhar, geni-

talıa slide Karsholt 4076 (ZMUC). Ukraine, Dnepropetrovsk reg., Pavlograd distr., Bulakhovka, estuary,

saline: 29 15.vii.2011, leg. V. Afans’eva (ZMKU).

Redescription. Adult (Figs 1,2). Wingspan 12.5-14.5 mm. Head grey to light

brown; frons and lateral margins paler; labial palpus strongly upcurved, brown, inner

surface pale grey to cream; segment 3 paler, 1.5 times narrower and nearly as long as

segment 2, pointed; scapus brown, flagellum brown with white basal rings; thorax and

tegula as forewing. Forewing brown; a narrow black streak from the base along fold

to nearly half length of wing, with a few orange-brown scales; a diffuse black spot ın

middle of cell; a black dot in the cell corner; creamy spots at 3/4 of costa and dorsum;

subapical area mottled with grey scales; fringe grey, brown-tipped. Hindwing grey; a

narrow yellow-white line at border to grey fringe.

Variation. The black streak at the base of the forewing may be divided into two elon-

gated spots; the costal and dorsal cream-coloured spots may in some specimens be

reduced to a few scales or obsolete.

Male genitalia (Fig. 3). Uncus 3 times as long as broad, densely covered with long

strong setae, tip pointed, curved; gnathos weak, ring-shaped; tegumen prolonged, lateral

Nota lepid. 36 (1): 77-84 719

folds curved inwards; pedunculi short, tapered; valva evenly curved, slightly broadened

in middle, apex rounded, about as long as tegumen and uncus, covered with setae in

distal 2/3; sacculus subtriangular, densely covered with hairs; phallus basally swollen,

distal portion straight, gradually narrowed and pointed apically; lateral projection about

as broad as distal portion of the phallus at its base, with a prominent pointed tip.

Female genitalia. Unknown.

Biology. Early stages unknown. Adults have been collected in May, July and August,

and the species may thus be bivoltine. The habitat in Ukraine is a saline estuary.

Distribution. Russia: Lower Volga; Ukraine (new record).

Remarks. Doryphora orthogonella was described from two males collected at Sarepta

(now Krasnoarmeysk near Volgograd, 48°31’N, 44°34’E) by H. Christoph (Staudinger

1871). Although Staudinger compared it to Scrobipalpa acuminatella (Sircom, 1850)

he noted that its hindwings were similar to those of Gelechia detersella Zeller, 1847,

the type species of the genus Stomopteryx Heinemann, 1870. It was, however, only

combined with Stomopteryx in 1996 by Karsholt & Riedl.

The five specimens held in the collection of the BMNH have also most likely been

collected at the type locality, although they are not part of the type series. The few

specimens of S. orthogonella available from the type locality and two specimens col-

lected in Ukraine are identical both externally and in the male genitalia.

Externally S. orthogonella may resemble S. hungaricella Gozmany, 1957, but the

latter is usually nearly uniformly black rather than brown, and without any trace of

streaks or spots. The lateral projection that is placed near the right angle of the phallus

is the most prominent character; it separates S. orthogonella from most other species of

Stomopteryx, although a similar structure 1s found in S. basalis (Staudinger, 1876) and

related species, which are easily separated by the reddish-brown base of the forewing.

S. orthogonella is apparently a rare species. Since its description in 1871 it has not

been dealt with in the literature apart from checklists and catalogues (e.g., Meyrick

1925) until now. Anikin et al. (1999) considered it to be extinct from its type locality.

Syncopacma azosterella (Herrich-Schäffer, 1854) Figs 4-8

Anacampsis azosterella Herrich-Schäffer, 1854: 194

Syncopacma azosterella (Herrich-Schäffer, 1854) — Gozmäny, 1957: 121, fig. 4.1. (misidentification of

S. albifrontella Heinemann, 1870).

Syncopacma sp. 1 — Elsner et al. 1999: 52, Farbtaf. 25; Taf. 25, Abb. 306.

talpräparat No. 1042 det. J. Klimesch, Linz” | “Holotype” (ZMHU). France, Alpes-Maritimes, Domaine

de Maure Vieil: 10° 26.1v.1999, gen. slide Hendriksen 2462, 39 2-3.v1.2000, gen. slide Hendriksen 2680,

leg. H. Hendriksen (ZMUC). Greece: Lakonia, 5 km S Monemvasia, 29 8.vili.1979, 19 15.1x.1979, 19

6.v11.1980, 19 13.v1.1981, 19, 24.v1.1981, 19 18.vii1.1982, 19 26.v111.1982, 10° 29.v1.1984, leg. G. Chris-

tensen; 20° 4.v11.1984, leg. B. Skule, gen. slide Hendriksen 4430; 7 km SW Monemvasia, 150 m, 19

22.1x.1979, leg. G. Christensen, gen. slide Hendriksen 4431; 19 9.iv.1981, leg. B. Skule; 19 2.vii.1982,

leg. S. Langemark & B. Skule; Prov. Serra, 20 km E Sidirokastro, Kapnophyton, 450 m, 10° 18.vii.1990,

21-23.v.1994, leg. O. Karsholt, gen. slide Hendriksen 3615 (all ZMUC). Hungary, Veszprem County, 10

km N Veszprem: 47°10’N 17°58’E, 300 m, 20° 17.v11.2005, leg. C. Hviid, B. Skule & E. Vesterhede, gen.

slide Hendriksen 6209 (ZMUC). Morocco, High Atlas Mts, Asni area: 1100-1400 m, 30°, 39 la. 8-10.

80 BipzityaA & KarsHOLT: Two little-known gelechiids in the European fauna

iv.1989, Cytisus sp., leg. O. Karsholt, gen. slide Hendriksen 4708 (ZMUC). Romania, Carpatii orientali,

Cheile Bicazului, 1250 m: 19 11-12.v111.1988, leg. & coll. S. & Z. Kovacs. Slovenia, SW part, 11 km

above Kozina, Stavnik Mts, 45°32’N 13°58E, 950 m: 19 30.vi. 2003, C. Hviid & B. Skule, gen. slide

Hendriksen 4876 (Z MUC). Spain: Prov. Segovia, San Ildefonso, no date, 19, 29, gen. slide Karsholt

5006; Prov. Malaga: Camino de Istan, 400 m, 19 25.v1.1975, leg. E. Traugott-Olsen, gen. slide Hen-

driksen 4547; Sierra de Marbella, El Mirador, 700 m, 19 19.viii.1977, leg. E. Traugott-Olsen, gen. slide

Hendriksen 4549; Camino de Ronda, Urb. Madronal, Loma de Colmenas, 500 m, 19 23.v.1986, gen. slide

Hendriksen 5271; 10° 19.v111.1988, gen. slide Hendriksen 3749; 29 28.vii.1988; 30°, 19 30.viii.1988, gen.

slide Hendriksen 2390; 40°, 39 4.1x.1988; 20° 10.1x.1988; 19 13.1x.1988, leg. E. Traugott-Olsen; Prov.

Granada: 10 km NW Otivar, Lopera, 1200 m, 29 24.v11.2003, leg. P. Skou, gen. slide Hendriksen 4677;

25 km N Almunecar, Moscaril, 500 m, 10° 28.viii-9.1x.2004, leg. G. Jeppesen, gen. slide Hendriksen 5089

(all ZMUC). Ukraine: Lugansk reg., Melovoi distr., Strel’tsovskaya Step Nat. Res., 20° 19 10.vii.2002,

at light, leg. A. Bidzilya, gen. slide 5/12; Donetsk reg., Kemennye Mogily Nat. Res., 19 18.vii.1989, leg.

A. Zhakov (ZMKU).

Redescription. Adult (Figs 4-6). Wingspan 9.5—13.5 mm. Head black, frons slight-

ly lighter, dark-grey; labial palpus up-curved, segment 3 about 1.5 times narrower and

nearly as long as segment 2, acute, light grey to white, underside black, segment 2 light

grey; scapus black, flagellum black, underside white-ringed; thorax and tegulae black;

forewing black, subapical fascia white, narrow; cilia grey, black-tipped; hindwing grey.

For variation see below.

Male genitalia (Fig. 7). Uncus twice as long as broad, lateral folds densely se-

tose, apex weakly rounded; gnathos hook stout, curved at nearly right angle in middle,

apex tapered, curved; tegumen prolonged with well-developed lateral folds, pedunculi

short, slender; valva nearly of equal width, slightly constricted before apex, exceeding

apex of uncus; vinculum narrow, band-shaped, posterior-lateral margin bulging, bear-

ing long hair-like setae; vincular projections moderately broad, outer margin evenly

curved after half length, inner margin straight, apex weakly pointed; saccus sub-quad-

rangular, anterior margin weakly emarginated; phallus bulbous in basal half, distal half

tapered, apical quarter needle-shaped.

Female genitalia (Figs 8, 8a). Papillae anales subovate, twice as long as wide,

about as long as length of segment VIII, covered with short setae, a few long hair-like

setae at base. Apophyses anteriores nearly four times shorter than apophyses posteri-

ores and about 3 times shorter than segment VIII. Segment VIII slightly broader than

long, trapezoidal, posterior margin nearly straight, anterior margin weakly concave in

middle. Lateral folds of sternum VIII broadly separated by medial zone with well-

developed A-shaped sclerotisation that reaches nearly to the anteromedial corners of

these folds. Ostium opening near anterior margin of sternum VIII, posterior subostial

sclerite semicircular, anterolateral sclerites joining anteromedial corners of the folds

of sternum VIII. Antrum strongly sclerotised, posterior margin evenly concave with

tapered posteriolateral corners. Ductus bursae slender, membranous, slightly narrowed

near the entrance of corpus bursae. Corpus bursae weakly sclerotised, subovate, about

as long as ductus bursae. Signum absent.

Biology. Early stages have not been described. The species has been reared from

Adenocarpus intermedius DC. (Fabaceae) first by Mendes (1904) from Säo Fiel in Beira

Baixa in Portugal, but it was identified as Anacampsis vorticella (Scopoli). According

to Mendes (op cit.) the larva is common in March and April on Adenocarpus, tying the

leaves into a bud-like form (M. Corley in litt.). A small series of moths were bred from

Nota lepid. 36 (1): 77-84 81

Figs 4-8. Syncopacma azosterella (Herrich-Schäffer). 4. Holotype, Austria, wingspan 12.5 mm. 5. Spe-

cimen from Ukraine, wingspan 11.1 mm. 6. Specimen from Morocco, wingspan 12.2 mm. 7. Male geni-

talia, Ukraine, gen. slide Bidzilya 5/12. 8. Female genitalia, France, gen. slide Karsholt 5006; a: segment

VIII (enlarged).

82 BıpzıL ya & KarsHorr: Two little-known gelechiids in the European fauna

larvae feeding on broom (‘Cytisus sp.’, Fabaceae) in the High Atlas Mts of Morocco.

Adults have been collected from May to September, mostly at light, but in Bulgaria

flying around Genista sp. (Junnilainen et al. 2010). The species has probably one gen-

eration in central Europe and two or three generations in the Mediterranean lowlands.

Distribution. Austria, Bulgaria (Junnilainen et al. 2010), Czech Republic (Elsner et

al. 1999), France (new record), Greece (new record), Hungary, Morocco (new record),

Romania, southern Ural in Russia (Junnilainen et al. 2010), Slovenia (new record),

Spain (new record), Ukraine (new record). In Portugal (new record) S. azosterella is a

locally common species feeding on Adenocarpus. Occasionally larvae may be quite nu-

merous (M. Corley in litt.). Records from Switzerland (e.g., Gozmany 1957) are based

on misidentification (SwissLepTeam 2010: 187). Records from Poland (e.g., Karsholt

& Riedl 1996) probably date back to Rebel (1901: 154), but were not confirmed and are

probably due to misidentification.

Remarks. Anacampsis azosterella was described from a single specimen collected by

H. Lederer in Austria: Wien (Herrich-Schaffer 1854). The whereabouts of the holotype

were unknown for a long time (Hering 1952: 206; Wolff 1958: 258), which made it im-

possible to correctly apply this name to any taxon. We were able to locate the holotype

in the ZMHU but, unfortunately, the corresponding genitalia slide seems to be lost. The

type is in rather good condition, although somewhat faded. That may have been the

case already when Herrich-Schäffer described it, as he described the (white) subapi-

cal fascia as “etwas bräunlich” [somewhat brownish]. In the photograph (Fig. 4) the

holotype specimen looks more broad-winged than the specimens shown in Figs 5-6,

but that is because it does not have the wings spread to horizontal position. Comparing

it to other old, faded specimens of S. azosterella gave an exact match. After its descrip-

tion S. azosterella remained a poorly known taxon, and it was only mentioned in a few

publications. Gozmany (1957: 121) tried to solve the identity of S. azosterella, but

he mixed it with S. albifrontella (Heinemann, 1870) and its synonym S. ignobiliella

(Heinemann, 1870) (Wolff 1958: 258), causing further misidentifications of S. azoste-

rella in the literature. Based on a preliminary study of the holotype, Karsholt & Riedl

(1996: 119) reintroduced S. azosterella for the species dealt with here, although without

an explanation, and that probably caused Elsner et al. (1999) to doubt its identity and

treat the species as “Syncopacma sp. 1.”

We have considered whether the holotype of S. azosterella could belong to another

Syncopacma species, and one could argue that it might be either (a small) S. cinctella

(Clerck, 1759) or S. ochrofasciella (Toll, 1936). However, the species dealt with here

is a Surprisingly variable species (see below) so it is difficult to argue that the holotype

of S. azosterella does not belong here. We are, moreover, of the opinion that this solu-

tion serves the stability of nomenclature best. We have also considered the possibility

of treating Anacampsis azosterella Herrich-Schäffer as a species incertae sedis and to

give anew name to “Syncopacma sp. 1” of Elsner et al. (1999), but we prefer not to do

so as we find that a less satisfactory solution.

S. azosterella is a variable species. Specimens from southern Greece and southern

Spain are small (wingspan 7—11 mm), with the smallest specimens from the summer

and autumn generations. They have a clear white subapical band on the forewing and

Nota lepid. 36 (1): 77-84 83

no black spots, and they also look more slender-winged. Specimens from central Spain

(San Ildefonso) are larger (wingspan 11-13 mm) but otherwise similar. However, two

specimens from southern Spain, prov. Granada (wingspan 11 mm) differ from other

Spanish specimens examined in having almost black forewings with only a few lighter

scales at the costa near the apex and at the tornus; moreover, they have black spots in

the middle of the wing. These two specimens more closely resemble a series of reared

specimens from Morocco (wingspan 11-13 mm), which have black spots in the fold

and in the middle of the wing followed by a few light yellow scales, and a yellowish

white subapical fascia interrupted in the middle.

The examined slides of male genitalia show slight variation, not just in size, but in

the relative proportions of the length of the vincular projections and the saccus, and

also of the basal, swollen part and the apical, thin part of the phallus. Furthermore,

there are small differences in the shape of the phallus. However, we found no correla-

tion between these small differences and the differences in forewing pattern described

above. We therefore conclude that this variation is most probably due to differences in

preparing the studied genitalia slides, and especially in the pressure put on the genitalia

through the cover slip.

Most species of Syncopacma are more or less difficult to recognise from external

characters, and it is often necessary to examine the genitalia to reach a safe identifica-

tion. Fortunately, the male genitalia of most species exhibit characteristic differences

(e.g., Elsner et al. 1999; Wolff 1958). As no such differences could be found between

the more or less different looking populations studied by us, we here conclude, at least

tentatively, that they belong to one variable species. Studies of the DNA from differ-

ent populations may well contradict this, and especially specimens from Spain and

Morocco, having an interrupted subapical fascia and black spots in the forewing, may

well turn out to represent a distinct species.

S. azosterella may be confused externally with S. cinctella (Clerck, 1759) and other

Syncopacma species with narrow white subapical fascia. The male genitalia of S. azos-

terella most resemble those of S. suecicella (Wolff, 1958) and S. linella (Chrétien,

1904), but differ from the first mentioned in the basal portion of the phallus being

longer and in the absence of a prominent lateral vincular projection. S. linella differs in

the apically pointed rather than rounded posterior vinculum projections as in S. azos-

terella. A-shaped sclerotisation on sternum VIII and semicircular posterior subostial

sclerite are characteristic features of the female genitalia.

Acknowledgements

Yuriy I. Budashkin (Karadag Nature Reserve, Ukraine), Veronika O. Afanas’eva and Kyrylo K. Holo-

borod’ko (Dnepropetrovsk State University, Ukraine), and Zoltan Kovacs (Miercurea Ciuc, Romania)

provided material used in this study. Wolfram Mey assisted us during our work with the collection of

ZMHU. Peter Huemer (Tiroler Landesmuseum, Innsbruck, Austria), Klaus Sattler (BMNH, London, UK)

and Andreas Segerer (Zoologische Staatssammlung, Miinchen, Germany), helped with information. Leif

Aarvik (Zoologisk Museum, Oslo, Norway) and Martin Corley (Faringdon, UK), commented on the man-

uscript, and the latter also improved the English language. We also received suggestions for improving

the manuscript from Peter Huemer and an anonymous reviewer. Reinhard Sutter (Bitterfeld, Germany)

provided the photograph for Fig. 8. We are grateful to all for their help.

84 BipzityA & KarsHorr: Two little-known gelechiids in the European fauna

References

Anikin, V. V.,S. A. Sachkov & V. V. Zolotuhin 1999. “Fauna Lepidopterologica Volgo-Uralensis“ 150 years

later: changes and additions. Part 4. Coleophoridae, Gelechiidae, Symmocidae and Holcopogonidae

(Insecta, Lepidoptera). — Atalanta 29: 295-336.

Bidzilya, O. & O. Karsholt 2008. New data on Anomologini from Palaearctic Asia (Gelechiidae). — Nota

lepidopterologica 31: 199-213.

Elsner, G., P. Huemer & Z. Tokär 1999. Die Palpenmotten (Lepidoptera, Gelechiidae) Mitteleuropas. — Ver-

lag F. Slamka, Bratislava. 208 pp., 1-85, 1-28 pls.

Gozmäny, L. 1957. Notes on the generic group Stomopteryx Hein., and the description of some new Micro-

lepidoptera. — Acta Zoologica Academiae Scientarum Hungaricae 3: 107-135.

Hering, E. M. 1952. Generische Unterschiede zwischen Stomopteryx Hein. und Aproaerema Durr. (Lep.

Gelech.). — Opuscula Entomologica 17: 201-207.

Herrich-Schäffer, G. A. W. 1847-1855. Systematische Bearbeitung der Schmetterlinge von Europa. 5, 394

pp., 124 + 7+ 1 pls. Regensburg.

Huemer, P. & O. Karsholt 2010. Gelechiidae II (Gelechiinae: Gnorimoschemini). Pp. 1—586. — In: P.

Huemer, O. Karsholt & M. Nuss (eds), Microlepidoptra of Europe 6, Apollo Books, Stenstrup.

Junnilainen, J., O. Karsholt, K. Nupponen, J.-P. Kaitila, T. Nupponen & V. Olschwang 2010. The gelechiid

fauna of the southern Ural Mountains, part. II: list of recorded species with taxonomic notes (Lepido-

ptera: Gelechiidae). — Zootaxa 2367: 1—68.

Karsholt, O., M. Mutanen, S. Lee & L. Kaila. 2013. A molecular analysis of the Gelechiidae (Lepidoptera,

Gelechioidea) with an interpretative grouping of its taxa. — Systematic Entomology 38: 334-348.

Karsholt, O. & T. Riedl. 1996. Gelechiidae, excl. Gnorimoschemini. Pp. 103-113, 118-122, 310, 312. —

In: O. Karsholt & J. Razowski (eds), The Lepidoptera of Europe. A distributional checklist, Apollo

Books, Stenstrup, 380 pp.

Mendes, C. de Azevedo 1904. Lepidopteros de Portugal. II. Lepidopteros da regiäo de S. Fiel (Beira Bai-

xa). — Brotéria 3: 223-254.

Meyrick, E. 1925. Lepidoptera Heterocera. Fam. Gelechiadae. Pp. 1-290, 5 pls — In: P. Wytsman (ed.),

Genera Insectorum 184, Bruxelles.

Rebel, H. 1901. Famil. Pyralidae — Micropterygidae. Pp. 1-368. — Jn: O. Staudinger & H. Rebel (eds),

Catalog der Lepidopteren des Palaearctischen Faunengebietes 2, Berlin.

mologische Zeitschrift 14 (1870): 273-330.

SwissLepTeam 2010. Die Schmetterlinge (Lepidoptera) der Schweiz: Eine kommentierte systematisch-

faunistische Liste. — Fauna Helvetica 25: 1-349.

Wolff, N. L. 1958. Further Notes on the Stomopteryx Group. — Entomologiske Meddelelser 28: 224-281.

Book review 85

P. Huemer 2013. Studiohefte 12. Die Schmetterlinge Osterreichs (Lepidoptera).

Systematische und faunistische Checkliste. — Tiroler Landesmuseen-Betriebsgesellschaft

m. b. H., Innsbruck, Austria. 304 pp. ISBN 978-3-900083-42-7. Price: 14.80 € plus shipping

costs. Orders can be placed online at http://www.tiroler-landesmuseen.at/shop.php/de/druck-

werke alle /studiohefte

Austrian Lepidoptera sparked a great interest in many European lepidopterists who have spent

a great deal of their time hunting butterflies and moths in the picturesque alpine valleys and

high rocky mountains of the Alps. Austria is one of the most geologically and biogeographi-

cally interesting European countries, and therefore it comes as no surprise that the publica-

tions on different species of Lepidoptera of this country started in the 18" century, for example

the Gracillariidae from Austria being studied already by Fabricius (1798). After two hundred

years, this interest in Austrian butterflies and moths is still very much alive. New molecular

data, which recently became available, require correct species identification and accompanying

taxonomic information, which can be provided only by standardised checklists. This book rep-

resents an updated, re-written, and taxonomically improved edition of the Austrian catalogue of

Lepidoptera which was published two decades ago (Huemer & Tarmann 1993) and is adapted

to the present-day purposes. The novelty of the 2013 edition is summarized on p. 15. A total of

4071 species, presently recorded from Austria, are listed, and 119 species are indicated either as

false, ambiguous, or based on accidental records, so the community of lepidopterists is invited

to cautiously re-evaluate and re-study these interesting cases. In contrast to many taxonomic

catalogues, this systematic and faunistic checklist of Austrian Lepidoptera is the result of a huge

collecting effort of generations of lepidopterists and it particularly demonstrates the great field

experience of the author himself.

Furthermore, the faunistic data, presented in the form of a table divided per provinces of

Austria, are based on voucher specimens deposited in the collection of the Tiroler Landesmuseum

Ferdinandeum or associated collections in other museums, so these voucher specimens credit the

checklist as a highly reliable reference source. Two new synonymisation acts (in Oecophoridae

and Crambidae) are included in the book and one case of synonymy is revised (in Tortricidae).

Nine species belonging to the families Gracillariidae, Oecophoridae, Gelechiidae, Elachistidae,

Tortricidae, Geometriidae and Noctuidae are presented as new for the Austrian fauna.

The Checklist begins with colour plates containing 128 photographs of butterflies or moths

in nature in their resting position. Usually such systematic-faunistic checklists are quite dull

publications, so the inclusion of a subset of colourful and high-quality photographs makes this

publication visually attractive. The systematic-faunistic checklist, which occupies a major part of

the book (p. 32—203), follows van Nieukerken et al. (2011) for the classification of the lepidop-

teran families and Kaila et al. (2011) for the classification of the superfamily Gelechioidea and

hypothesised phylogenetic relationships. The short introductory list of suborders, infraorders,

clades and families is presented at the beginning of the chapter allowing the reader to easily spot

these higher taxa in the faunistic table. However, for species one needs to look at the index first

before trying to find them in the very long faunistic table. The species arrangement within the

genera is not as handy for use of this checklist as one could wish. Probably it would have been

easier for the reader to find species within the genera if they were arranged alphabetically. The

determination of the phylogenetic position of alpine species of Lepidoptera is far from com-

plete. We still lack a clear picture of relationships of species and numerous species complexes,

especially in microlepidoptera, and despite the truly rapid advances in molecular techniques, the

problems of specific relationships and species delimitations still fall on speculations in many

cases, so the alphabetical order of the species within the genera might have its own advantages

such as user-friendliness and easy finding of any species of interest.

Synonyms in the species group have been kept to a strict minimum and were subjectively

chosen for those cases in which these names have been used often in earlier literature or when

they were used as valid species names in the earlier version of the catalogue (Huemer & Tar-

86 Book review

mann 1993) either as species or subspecies. The author refers to the online world catalogues

for a complete synonymy. Synonyms in the genus-group have been kept to a strict minimum as

well. At the end of the book (p. 204—243), a comprehensive chapter Comments provides use-

ful and interesting information on the taxonomic and/or distributional peculiarities of certain

species indicated by the letter K in the systematic and faunistic checklist. The reference list (p.

244-261) is robust, non-abbreviated and gives a good overview of publications on the Austrian

lepidopteran fauna. The book ends with the highly needed index.

To summarise, the author should be cordially thanked for sharing his extraordinary taxo-

nomic and faunistic knowledge of Lepidoptera of his home country and congratulated for the

impressive results. This checklist should be in the library of any lepidopterist interested in

European Lepidoptera and especially in the libraries of those amateurs and professionals who

spend their holidays in this beautiful country or collecting in the SEL study area, which is just

across the Austrian border. The book is in German; however, I am very certain that this fact will

not hinder any lepidopterist from buying it and admiring this meticulous work.

References

Fabricius, J. C. 1798. Supplementum entomologiae systematicae. Hafniae, apud Proft et Storch: 480-572.

Huemer, P. & G. Tarmann 1993. Die Schmetterlinge Osterreichs (Lepidoptera). Systematisches Verzeich-

nis mit Verbreitungsangaben für die einzelnen Bundesländer. — Veröffentlichungen des Tiroler Landes-

museums Ferdinandeum 73: | —224.

Kaila, L., M. Mutanen & T. Nyman 2011. Phylogeny of the mega-diverse Gelechioidea (Lepidoptera):

adaptations and determinants of success. — Molecular Phylogenetics and Evolution 61: 801-809.

Nieukerken, E. J. van, L. Kaila, I. J. Kitching, N. P. Kristensen, D. C. Lees, J. Minet, C. Mitter, M. Mu-

tanen, J. C. Regier, T. J. Simonsen, N. Wahlberg, S.-H. Yen, R. Zahiri, D. Adamski, J. Baixeras, D.

Bartsch, B. A. Bengtsson, J. W. Brown, S. R. Bucheli, D. R. Davis, J. De Prins, W. De Prins, M. E.

Epstein, P. Gentili-Poole, C. Gielis, P. Hattenschwiler, A. Hausmann, J. D. Holloway, A. Kallies, O.

Karsholt, A. Kawahara, S. J. C. Koster, M. Kozlov, J. D. Lafontaine, G. Lamas, J.-F. Landry, S. Lee,

M. Nuss, K. T. Park, C. Penz, J. Rota, B. C. Schmidt, A. Schintlmeister, J. C. Sohn, M. A. Solis, G. M.

Tarmann, A. D. Warren, S. Weller, R. V. Yakovlev, V. V. Zolotuhin, & A. Zwick 2011. Order Lepido-

ptera Linnaeus, 1758. — Jn: Z.-Q. Zhang (ed.), Animal biodiversity: An outline of higher-level classifi-

cation and survey of taxonomic richness. — Zootaxa 3148: 212-221.

JURATE DE PRINS

Book review 87

P. Leraut 2012. Moths of Europe, volume 3, Zygaenids, Pyralids 1 and Brachodids.

N. A. P. Editions, Verrierres-le Buisson. ISBN 978-2-913688-15-5. Price: £79.99 plus

shipping costs.

This book is an English translation from French of volume 3 of Les Papillons de nuit d’Eu-

rope — Zygénes, Pyrales 1. The present review deals only with the family Zygaenidae, as we

understand that the section on pyralids is being dealt with by a specialist in that group.

Undoubtedly, as with volumes 1 and 2, this book is an excellent identification guide to the

moths of Europe, based on the colour illustrations. However, the text is unfortunately marred

by scientific error and misinformation, so that it would have benefited greatly, if it had been

peer-reviewed by relevant specialists before publication. Moreover, in this respect, members of

‘Groupe d’Information de Recherche et d’Animation sur les Zygaenidae — GIRAZ’, a society

formed by a dedicated team of French entomologists who specialise in making a specific study

of the French zygaenid fauna, appear not to have been consulted.

While much of the information provided in the text is compilatory, regarding the Procridinae

the now out-of-date Forester Moths (Efetov & Tarmann 1999) is the only publication that is

cited in the references. However, many publications devoted to the taxonomy of this group have

subsequently been published (Efetov 2001c, 2004, 2005; Efetov et al. 2003) but apparently are

not referred to.

The characterisation of the Zygaenidae, as defined in this book (p. 41), has several shortcom-

ings. For example, the Phaudinae is included as a subfamily, although quite recently (Nieukerken

et al. 2011) it was placed as a family within the Zygaenoidea. All species of Zygaenidae have

ocelli (not only in the Zygaeninae, as mentioned) and, together with the presence of the chae-

tosemata (not mentioned), are two of the most important characters of the family. The antenna

of Zygaenidae is bipectinate, biserrate or simple with a clubbed terminal end and not only

‘pectinate and club-shaped’. The labial palpi are prominently developed in the tribe Artonini

(subfamily Procridinae), of medium length in the tribe Procridini and only in the Chalcosiinae

and Zygaeninae are they ‘weakly developed’.

The wing venation representing the Zygaeninae (apparently of a Zygaena species) is figured

(p. 42, fig. 20) but, as there is no indication from which species the drawing was made, the impres-

sion is given that this character situation is constant in the Zygaeninae. This is incorrect as there

are strong differences in some of the Zygaeninae (e.g., Pryeria sinica or Epizygaenella caschmi-

rensis, see Alberti 1954: 445, pl. 44, figs 1 and 7, respectively). The same can be said about the

figure of the wing venation of the Procridinae (genus Jordanita) (p. 43, fig. 21), as there are some

important differences within this subfamily. In the case of Jordanita, for example in J. (Roccia)

naufocki, veins R, and R, are stalked or connate in the forewing, while in the closely related spe-

cies J. (R.) tianshanica (pl. 4, fig. 15) R, and R, arise separately from the cell (Efetov 1990: 11).

The description of the habitus of the Procridinae is misleading; it is incorrect to state that

the forewing is ‘usually narrow’ (p. 42) and that most Procridinae ‘have a uniform single-tone

colouring’. In fact the habitus of Procridinae is very diverse. In Europe most species do have a

uniform colouration with a submetallic sheen on the body and forewing upper side, but some

of the tropical species can be very colourful with yellow, red and white spots and stripes, with

green or blue metallic pattern, or even with almost completely translucent wings. The antennae

in Procridinae are bipectinate in the male, bipectinate or biserrate in the female and only in the

Central American genus Pseudoprocris do they consist of a simple flagellum without lateral

extensions, thus forming a clubbed antenna as in Zygaena.

On page 54 it is stated that Jordanita subsolana belongs to the subgenus Lucasiterna, but

Ino subsolana is the type-species of the subgenus Solaniterna; therefore the correct combina-

88 Book review

tion is Jordanita (Solaniterna) subsolana (Efetov 2004: 33, 119). Jordanita graeca sultana is

cited (p. 55) as a valid subspecies, but this is a synonym under J. graeca graeca (Efetov 2001b:

156). It is stated (p. 61) that the larval host plant of Adscita jordani is unknown, but the larva

feeds on Rumex species (Efetov & Tarmann 2003a, 2003c). It should have been mentioned on

pages 63 and 64 that Adscita bolivari and A. mannii belong to the subgenus Tarmannita (Efetov

2000: 169). The larval host plants of Adscita obscura belong not only to the family Cistaceae,

as mentioned on page 66, but also to the Rosaceae and Fabaceae (Tarmann & Tremewan 2001).

On page 66 it is considered that Adscita alpina has two valid subspecies, viz. A. alpina alpina

and A. alpina italica. However, Efetov & Tarmann (2000) have shown that A. alpina and A.

italica are two well-differentiated species that have strong differences in the female genitalia.

Adscita italica is found in central and southern Italy, whereas A. alpina is only found in the Alps,

viz. south-eastern France, southern and south-eastern Switzerland, western Austria and northern

Italy (Efetov & Tarmann 2000, 2003b). Adscita (Zygaenoprocris) taftana 1s briefly mentioned

on page 67 but following the revision of the genus Zygaenoprocris, the current placement of this

species 1s Zygaenoprocris (Molletia) taftana (Efetov 2001a: 45).

With regard to the distribution of Procridinae species, there are a number of errors. The map

on page 47 implies that Rhagades pruni inhabits the whole of Spain, but it is found only in a

very restricted area in the north-eastern part of the country (Efetov 2004: 14). Adscita mannii 1s

regarded as highly local (p. 65), but in Italy, for example, it is widely distributed and even mass

occurrences are sometimes found in many habitats. On page 66 it is stated that Adscita krymen-

sis was first described in the Crimea and also reported from Ukraine (p. 66), but the species is

known only from the Crimea (Efetov 2001c); moreover, the latter is part of southern Ukraine.

Of the 108 Zygaena species currently considered to be valid (Hofmann & Tremewan 2010),

63 are listed in the check-list on page 68, but it is rather puzzling that 36 of these are extra-

limital to Europe. The criterion for such a selection is not given and it remains unclear why

many European species are excluded, e.g., four European endemics (Z. romeo, Z. rhadaman-

thus, Z. oxytropis, Z. anthyllidis) and five species with a wide distribution in Europe, viz. Z.

osterodensis, Z. nevadensis, Z. filipendulae, Z. lonicerae and Z. ephialtes. Moreover, Z. mana

and Z. alpherakyi, two endemics to the Caucasus region and bio-elements of the fauna of the

Russian territory, are also excluded. Generally speaking, one can say that the check-list is very

poorly compiled, incomplete and inconsistent and without any systematic concept; moreover, it

does not reflect the relevant literature (Tremewan 1988; Hofmann & Tremewan 1996: 187-219,

2010).

The arrangement of the genitalia figures is puzzling and it is unclear as to what the author

is trying to do in this respect. For example, on page 71 the male genitalia of Z. exulans, Z.

minos and Z. purpuralis are compared (the first-mentioned not closely related to the two last-

mentioned species), while on page 73 the female genitalia of Z. purpuralis, Z. minos and Z.

youngi are figured (the last-mentioned species not closely related to the former two and placed

in a different subgenus).

With regard to the distribution of Zygaena species, the map on page 69 shows a single record

of Z. purpuralis from Sicily; presumably this follows Naumann et al. (1984: 96). However, there

are no authentic records of this species from the island and even Bertaccini & Fiumi (1999: 65)

refer to the distribution map in Naumann et al. According to the distribution map on page 85,

Z. trifolii occurs throughout Sicily but the species is only known from a few records from the

vicinity of Syracusia (Hofmann et al. 1994: 43; Hofmann & Tremewan, 1996: 183). On page 92

it is stated that the Isle of Skye is the sole locality in Scotland for Z. lonicerae, but the species

has spread during the last few years from northern England into the border counties of Scotland

(Bland 2001). It is stated (p. 95) that Z. nevadensis possibly occurs in Italy near the frontier with

Book review 89

France, but there are no records of this species from that region. However, it was recently dis-

covered in Calabria (Efetov et al. 2011), a record that has been overlooked in the book. Zygaena

exulans is said to occur from 1000—3000 m.a.s.l., depending on latitude (p. 120), but in Scotland

the species occurs at around 700-850 m, while in northern Scandinavia and the northern part

of European Russia it is found near the sea level. Pryeria sinica, described from Japan with

a distributional range from there to Taiwan, South Korea, China and the Far East of Russia,

has recently been reported from Europe (England, Spain); however, it is erroneously stated (p.

123) that the species was originally from western Asia. The distribution of Z. tamara is cited as

Turkey to Afghanistan but the most easterly known site for Z. tamara is in the vicinity of Semnan

in the Iranian Alborz mountains and no records are known from further east and, of course, from

Afghanistan (A. Hofmann, pers. obs.). For Z. cambysea it is stated ‘Iran’ but in fact this species

is also widely distributed in eastern Turkey and Armenia and recorded from Azerbaijan and Iraq

(Hofmann & Tremewan 1996). Although Z. rosinae (p. 250, pl. 13 fig. 9) is labelled ‘Téhéran’ (a

city of 15 million inhabitants), its distribution is cited as Turkey and Caucasus. As far as Turkey

is concerned, the distribution is peripheral and there are only a few records from Transcaucasia,

its main occurrence being throughout Iran (A. Hofmann, pers. obs.).

With reference to cyanogenesis and the toxic properties found in the Zygaenidae, it would

have been better to use the term ‘glucosides’ rather than heterosides (p. 44), the latter apparently

referring to such compounds found in plants. Moreover, the use of glucoside is well established

in the zygaenid literature, e.g., Franzl (1992). It is also stated that linamarin and lotaustralin are

biosynthesized by the larvae, which is correct, but these compounds can also be sequestered by

the larvae from their host-plants that contain them. On the same page it is stated that ‘... the

caterpillars feed without really hiding themselves’, which does apply to many species, but some

only feed at dusk and dawn, e.g., those of Z. transalpina in Europe, while those of many species

in the Middle East feed only at night (Hofmann & Tremewan pers. obs.).

In several places the word ‘adrets’, meaning ‘southerly facing slopes’, has been used, with

reference to habitats; while ‘adrets’ is a geographical term acceptable in both French and English,

it is rarely if ever used in the latter language and is not included in many English dictionaries.

Greater consistency in the botanical nomenclature would have been desirable. The host plant

for Z. angelicae is cited as Coronilla varia (p. 107), but five pages before it is stated that Z.

ephialtes lives on Securigera varia (the correct combination); one of these two names should

also have been mentioned as the host plant of ‘Z. hippocrepidis’ (p. 106), which in this book is

separated as a valid species from Z. transalpina, a placement that is not generally accepted by

most Zygaena specialists (Hofmann & Tremewan 1996). On page 93 it is stated that a larval host

plant for Z. romeo is Trifolium montanum, but there are no authentic records of the larva of this

species ever feeding on this plant or on any members of the genus Trifolium.

The flight period of Z. sedi is stated to be exclusively May (p. 85), but in the Crimea (Ukraine)

the species occurs from the end of May to the beginning of July (Efetov 2005: 170), in Greece it

flies from mid-June to the beginning of July and in Turkey from the end of June to mid-July (A.

Hofmann & W. G. Tremewan pers. obs.). It is considered that the flight period of Z. occitanica

is mainly in July (p. 112), but it emerges in many localities (e.g., eastern and southern Spain) in

mid-May and its flight period is already over before the end of June; moreover, in the vicinity of

Almeria it is even found at the end of April (A. Hofmann & W. G. Tremewan pers. obs.).

On a positive note, the reproduction of the photographs of most of the Zygaena adults is

good and the figures should enable anyone to identify any specimens (if correctly determined by

Leraut) that they might encounter except for those that need to be dissected. Even then, the speci-

mens are obviously figured at different scales, e.g., Z. zuleima (p. 248, pl. 12 fig. 14) is seemingly

larger than Z. truchmena and Z. persephone. The same can be said about Z. nevadensis (p. 265,

90 Book review

pl. 20 figs 6-13), for example, the figures being reproduced almost as large as Z. lavandulae or

Z. theryi. In those depicting the Procridinae, the shadow on the right hand side of the specimens

is somewhat distracting and gives not only the impression of an unfocused picture but also ex-

tends the proportions optically. In thıs respect, the line drawings of the genitalia should help, but

unfortunately these are so finely drawn and reproduced so small that many critical characters are

not readily visible. For example, those purporting to illustrate the lamina dorsalis of Z. minos and

Z. purpuralis (p. 71) are inadequate and do not show the diagnostic characters clearly.

This English translation has many typographical and/or translation errors. To give only a

few examples, ‘reticulum’ for retinaculum (p. 41), “Nedblstreif’ for Nebelstreif (p. 103), ‘Bade-

Wurtemberg’ for Baden-Wiirttemberg’ (p. 107); such shortcomings are also found in some of

the scientific names, e.g. Z. loyselis ‘unguemachi for Z. loyselis ungemachi (p. 248).

References

Alberti, B. 1954. Uber die stammesgeschichtliche Gliederung der Zygaenidae nebst Revision einiger Grup-

pen (Insecta, Lepidoptera). — Mitteilungen aus dem Zoologischen Museum der Humboldt-Universitat

Berlin 30: 115-480, pls 1-62.

Bertaccini, E. & G. Fiumi 1999. Bombici e Sfingi d’Italia 3 (Lepidoptera Zygaenidae). — Giuliano Russo

Editore, Monterenzio. 160 pp., 13 pls, text-figs, distr. maps.

Bland, K. P. 2001. A new site for Zygaena lonicerae latomarginata Tutt, 1899 (Lepidoptera: Zygaenidae)

in Scotland. — Entomologist’s Gazette 52: 70.

Efetov, K. A. 1990. A new species of the genus Adscita (Lepidoptera, Zygaenidae) from the Middle

Asia. — Vestnik Zoologii 1990 (4): 8-11, figs 1-4.

Efetov, K. A. 2000. A new subgenus of the genus Adscita Retzius, 1783 (Lepidoptera: Zygaenidae, Pro-

cridinae). — Tavricheskiy mediko-biologicheskiy Vestnik 3 (1-2): 168-174, figs 1-12.

Efetov, K. A. 200la. On the systematic position of Zygaenoprocris Hampson, 1900 (Lepidoptera:

Zygaenidae, Procridinae) and the erection of two new subgenera. — Entomologist’s Gazette 52: 41—48,

figs 1-15.

Efetov, K. A. 2001b. An annotated check-list of Forester moths (Lepidoptera: Zygaenidae, Procridinae). —

Entomologist’s Gazette 52: 153-162, figs 1-5.

Efetov, K. A. 2001c. A Review of the Western Palaearctic Procridinae (Lepidoptera: Zygaenidae). — Sim-

feropol. 328 pp., col. frontispiece, 98 text-figs, 44 monochrome, 29 col. pls.

Efetov, K. A. 2004. Forester and Burnet 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), 272 pp., col. frontis-

piece, 183 figs, 1 col. pl. Simferopol.

Efetov, K. A. 2005. The Zygaenidae (Lepidoptera) of the Crimea and other regions of Eurasia, 420 pp.,

col. frontispiece, 78 figs, 27 monochrome, 32 col. pls, distr. maps. Simferopol.

Efetov, K. A. & G. M. Tarmann 1999. Forester Moths: The genera Theresimima Strand, 1917, Rhagades

Wallengren, 1863, Jordanita Verity, 1946, and Adscita Retzius, 1783 (Lepidoptera: Zygaenidae,

Procridinae). — Apollo Books, Stenstrup. 192 pp., figs 1-415, 12 col. pls.

Efetov, K. A. & G. M. Tarmann 2000. On the systematic position of Procris alpina italica Alberti, 1937,

and Procris storaiae Tarmann, 1977 (Lepidoptera: Zygaenidae, Procridinae). — Tavricheskiy mediko-

biologicheskiy Vestnik 3 (1-2): 161-167, figs 1-8.

Efetov, K. A. & G. M. Tarmann 2003a. On the systematic position of Adscita bolivari (Agenjo, 1937) and

Adscita jordani (Naufock, 1921) (Lepidoptera: Zygaenidae, Procridinae). Pp. 6569, figs 1-5. — /n: K.

A. Efetov, W. G. Tremewan & G. M. Tarmann (eds), Proceedings of the 7th International Symposium

on Zygaenidae (Lepidoptera), Innsbruck (Austria), 4-8 September 2000: 360 pp., col. frontispiece,

text-figs. Simferopol.

Efetov, K. A. & G. M. Tarmann 2003b. On the biology and distribution of Adscita (Adscita) alpina (Alberti,

1937), A. (A.) italica italica (Alberti, 1937) and A. (A.) italica storaiae (Tarmann, 1977) (Lepidoptera:

Zygaenidae, Procridinae). — Jn: T. Keil (ed.), VIII International Symposium on Zygaenidae, Dresden,

10—14 September 2003: 1415.

Efetov, K. A. & G. M. Tarmann 2003c. New data on the biology of Adscita (Adscita) jordani (Naufock,

1921) (Lepidoptera: Zygaenidae, Procridinae). — Jn: T. Keil (ed.), VIII International Symposium on

Zygaenidae, Dresden, 10—14 September 2003: 16.

Efetov, K. A., G. M. Tarmann & W. G. Tremewan 2011. Zygaena nevadensis Rambur, 1858 (Lepidoptera:

Zygaenidae, Zygaeninae) newly recorded from the southern tip of the Penisola Appenninica (Apennine

Peninsula), Italy. - Entomologist’s Gazette 62: 123-129, figs 1-5.

Book review 9]

Efetov, K. A., W. G. Tremewan & G. M. Tarmann (eds) 2003. Proceedings of the 7th International Sym-

posium on Zygaenidae (Lepidoptera), Innsbruck (Austria), 4—8 September 2000: 360 pp., col. frontis-

piece, text-figs. — Simferopol.

Franzl, S. 1992. Synthesis, transport and storage of cyanogenic glucosides in larvae of Zygaena trifolii

(Esper, 1783) (Lepidoptera: Zygaenidae). Pp. 21-37, text-figs 1-6, pls 1-3. — Jn: C. Dutreix, C. M.

Naumann & W. G. Tremewan (eds), Proceedings of the 4th Symposium on Zygaenidae, Nantes 11-13

September 1987. Recent advances in burnet moth research (Lepidoptera: Zygaenidae). — Theses zoo-

logicae 19: 193 pp., 6 pls.

Hofmann, A., G. Reiss & W. G. Tremewan 1994. Preliminary notes on the Zygaena Fabricius, 1777, fauna

of Tunisia (Lepidoptera: Zygaenidae): part 2. - Entomologist’s Gazette 45: 39-51, figs 1—7.

Hofmann, A. & W. G. Tremewan 1996. A systematic Catalogue of the Zygaeninae (Lepidoptera: Zygae-

nidae). — Harley Books, Colchester. 251 pp.

Hofmann, A. & W. G. Tremewan 2010. A revised check-list of the genus Zygaena Fabricius, 1775 (Le-

pidoptera: Zygaenidae, Zygaeninae), based on the biospecies concept. — Entomologist’s Gazette 61:

119-131.

Naumann, C. M., R. Feist, G. Richter & U. Weber 1984. Verbreitungsatlas der Gattung Zygaena Fabricius,

1775 (Lepidoptera, Zygaenidae). — Theses zoologicae 5: 1-45, text-fig., maps 1-97.